Plant Resources of Tropical Africa 1

Cereals and pulses

Editors: M. Brink
G. Belay

Associate editors: J.M.J. de Wet

O.T. Edje
E. Westphal

General editors: R.H.M.J. Lemmens

L.P.A. Oyen

r\
PROTA Foundation / Backhuys Publishers / CTA
Wageningen, Netherlands, 2006
Correct citation of this publication:
Brink, M. & Belay, G. (Editors), 2006. Plant Resources of Tropical Africa 1. Cereals
and pulses. PROTA Foundation, Wageningen, Netherlands / Backhuys Publishers,
Leiden, Netherlands / CTA, Wageningen, Netherlands. 298 pp.

Correct citation of articles from this publication:

[Author name, initials, 2006. Title of article]. In: Brink, M. & Belay, G. (Editors).
Plant Resources of Tropical Africa 1. Cereals and pulses. PROTA Foundation,
Wageningen, Netherlands / Backhuys Publishers, Leiden, Netherlands / CTA,
Wageningen, Netherlands, pp. ........

ISBN 90-5782-170-2 (book only)

ISBN 90-5782-171-0 (book + CD-Rom)

© PROTA Foundation, Wageningen, Netherlands, 2006.

No part of this publication, apart from bibliographic data and brief quotations em-
bodied in critical reviews, may be reproduced, re-recorded or published in any form
including print, photocopy, microfilm, electric or electromagnetic record without
written permission from the copyright holder: PROTA Foundation, P.O. Box 341,
6700 AH Wageningen, Netherlands.

Printed in the Netherlands by Ponsen & Looijen bv, Wageningen.

Distributed for the PROTA Foundation by Backhuys Publishers, P.O. Box 321, 2300
AH Leiden, Netherlands (worldwide), and CTA, P.O. Box 380, 6700 AJ Wageningen,
Netherlands (ACP countries).
Contents

Contributors 6

PROTA Board of Trustees and Personnel 9

Introduction 11

Alphabetical treatment of cereals and pulses 15

Cereals and pulses with other primary use 239

Literature 242

Index of scientific plant names 289

Index of vernacular plant names 293

PROTA in short 296

CTA in short 297

Map of Tropical Africa for PROTA 298

6 CEREALSANDPULSES

Contributors

E.G. Achigan Dako, IPGRI West and Central Africa, 08 B.P. 0932, Cotonou, Be-
nin (Digitaria exilis, Macrotyloma geocarpum)
S.G. Agong, Department of Horticulture, Jomo Kenyatta University of Agricul-
ture and Technology, P.O. Box 62000-00200, Nairobi, Kenya (Amaranthus cau-
datus)
R. Akromah, Department of Crop and Soil Sciences, College ofAgriculture and
Natural Resources, KNUST, Kumasi, Ghana (Vigna unguiculata)
D.J. Andrews, Department ofAgronomy and Horticulture, University of Ne-
braska, P.O. Box 830817, Lincoln, NE 68583-0817, United States (Pennisetum
glaucum)
I.K. Asante, Department of Botany, P.O. Box LG55, University of Ghana, Legon.
Accra, Ghana (Vigna unguiculata)
G. Assefa, Humboldt University, Berlin, Faculty ofAgriculture and Horticul-
ture, Philipstrasse 13, House no. 9, 10115 Berlin, Germany / Ethiopian Agricul-
tural Research Organization, Holetta Research Centre, P.O. Box, 2003, Addis
Ababa, Ethiopia (Avena sativa)
B. Badu-Apraku, UTA Ibadan, c/o Lambourn (UK) Limited, Carolyn House, 26
Dingwall Road, Croydon, CR9 3EE, United Kingdom (Zea mays)
T.V. Balole, Botswana College of Agriculture, Private Bag 0027, Gaborone, Bot-
swana (Sorghum bicolor)
J.P. Baudoin, Faculté universitaire des Sciences agronomiques de Gembloux
(FUSAGx), Unité de Phytotechnie tropicale et d'Horticulture, Passage des Dé-
portés, 2, 5030 Gembloux, Belgium (Phaseolus lunatus)
G. Bejiga, Green Focus Ethiopia, P.O. Box 802, Addis Ababa, Ethiopia (Cicer
arietinum, Lathyrus sativus, Lens culinaris)
G. Belay, Ethiopian Agricultural Research Organization, Debre Zeit Centre,
P.O. Box 32, Debre Zeit, Ethiopia (Eragrostis tef, Triticum aestivum, Triticum
turgidum, editor)
G. Bezançon, Institut de Recherche pour le Développement (IRD), B.P. 11416,
Niamey, Niger (Oryza glaberrimà)
C.H. Bosch, PROTA Network Office Europe, Wageningen University, P.O. Box
341, 6700 AH Wageningen, Netherlands (Bauhinia petersiana)
M. Brink, PROTA Network Office Europe, Wageningen University, P.O. Box
341, 6700 AH Wageningen, Netherlands (Avena abyssinica, Brachiaria deflexa,
Cenchrus biflorus, Cenchrus prieurii, Cordeauxia edulis, Craibia brownii, Crota-
laria karagwensis, Crotalaria lachnophora, Digitaria iburua, Echinochloa fru-
mentacea, Echinochloa obtusiflora, Echinochloa stagnina, Eragrostis aethiopica,
Eragrostis annulata, Eragrostis nindensis, Eragrostis plana, Limeum obovatum,
Macrotyloma uniflorum, Mucuna gigantea, Oryza barthii, Oryza longistaminata,
Oryza punctata, Panicum kalaharense, Panicum laetum, Panicum turgidum,
Phaseolus coccineus, Secale cereale, Setaria italica, Sporobolus fimbriatus,
Sporobolus panicoides, Tylosema fassoglense, Urochloa mosambicensis, Urochloa
 CONTRIBUTORS 7

trichopus, Vatovaea pseudolablab, Vicia hirsuta, Vigna aconitifolia, Vigna ade-

nantha, Vigna subterranea, editor)
S. Ceccarelli, ICARDA, P.O. Box 5466, Aleppo, Syria (Hordeum vulgare)
K.E. Dashiell, USDA-ARS Northern Grains Insect Research Laboratory, 2923
Medary Avenue, Brookings SD 57006, United States (Glycine max)
J.M.J, de Wet, Department of Crop Sciences, Urbana-Champaign, Turner Hall,
1102 South Goodwin Avenue, Urbana, IL 61801, United States (Eleusine cora-
cana, associate editor)
S. Diallo, ISRA / Zone Fleuve, CRA de Saint-Louis, B.P. 240 Sor Saint-Louis,
Senegal (Oryza glaberrima)
O.T. Edje, Faculty of Agriculture, University of Swaziland, P.O. Luyengo,
Luyengo, Swaziland (associate editor)
M.A.B. Fakorede, Department of Plant Science, Obafemi Awolowo University,
Ile-Ife, Nigeria (Zea mays)
K.E. Giller, Plant Production Systems, Department of Plant Sciences, Wagenin-
gen University, P.O. Box 430, 6700 AK Wageningen, Netherlands (Glycine max)
S. Grando, ICARDA, P.O. Box 5466, Aleppo, Syria (Hordeum vulgare)
G.J.H. Grubben, Boeckweijdt Consult, Prins Hendriklaan 24, 1401AT Bussum,
Netherlands (Vigna unguiculata)
P.C.M. Jansen, PROTA Network Office Europe, Wageningen University, P.O.
Box 341,6700 AH Wageningen, Netherlands (Coix lacryma-jobi, Fagopyrum es-
culentum, Lupinus albus, Vigna aconitifolia, Vigna adenantha, Vigna angularis,
Vigna mungo)
M. Jarso, Ethiopian Agricultural Research Organization, Holetta Research Cen-
ter, P.O. Box 2003, Addis Ababa, Ethiopia (Pisum sativum, Vicia faba)
R.N. Kaume, P.O. Box 583-90200, Kitui, Kenya (Panicum miliaceum)
G. Keneni, Ethiopian Agricultural Research Organization, Holetta Research
Center, P.O. Box 2003, Addis Ababa, Ethiopia (Pisum sativum, Vicia faba)
K.A. Kumar, Agriculture Environmental Renewal Canada Inc., 711 Schäfer
Road, P.O. Box 186, Delhi, ON N4B 2W9, Canada (Pennisetum glaucum)
G.M. Legwaila, Botswana College ofAgriculture, Private Bag 0027, Gaborone,
Botswana (Sorghum bicolor)
R.H.M.J. Lemmens, PROTA Network Office Europe, Wageningen University,
P.O. Box 341,6700 AH Wageningen, Netherlands (general editor)
R. Madamba, Crop Breeding Institute, Department of Research & Specialist
Services, Box CY 550, Causeway, Harare, Zimbabwe (Vigna unguiculata)
H.C.C. Meertens, Pomona 250, 6708 CJ Wageningen, Netherlands (Oryza sa-
tiva)
C.-M. Messiaen, Résidence La Guirlande, Bat. B3, 75, rue de Fontcarrade,
34070 Montpellier, France (Pisum sativum)
K.K. Mogotsi, Botswana College ofAgriculture, Private Bag 0027, Gaborone,
Botswana (Phaseolus acutifolius, Vigna radiata)
B.R. Ntare, ICRISAT, B.P. 320, Bamako, Mali (Arachis hypogaea)
Achmad Satiri Nurhaman, Southeast Asian Regional Centre for Tropical Biol-
ogy (SEAMEO BIOTROP), P.O. Box 17, Bogor, Indonesia (illustrations)
L.P.A. Oyen, PROTA Network Office Europe, Wageningen University, P.O. Box
341, 6700 AH Wageningen, Netherlands (general editor)
8 CKREALSANDPULSES

- R. Rajerison, CNRE, B.P. 1739, Fiadanana, Antananarivo (101), Madagascar

(Vigna umbellata)
- G.M. Ramolemana, Department of Crop Science and Production, Botswana Col-
lege ofAgriculture, Private Bag 0027, Gaborone, Botswana (Vigna subterranea)
- A.A. Seif, ICIPE, P.O. Box 30772, Nyago Stadium, Nairobi, Kenya (Pisum sati-
vum)
- K.P. Sibuga, Department of Crop Science and Production, Sokoine University of
Agriculture, P.O. Box 3005, Morogoro, Tanzania (Vigna subterranea)
- Iskak Syamsudin, Herbarium Bogoriense, Research Centre for Biology - LIPI,
J a l a n Ir. H. J u a n d a 22, Bogor 16122, Indonesia (illustrations)
- H. Tefera, Ethiopian Agricultural Research Organization, Debre Zeit Centre,
P.O. Box 32, Debre Zeit, Ethiopia (Eragrostis tef)
- L.J.G. van der Maesen, Biosystematics Group, Wageningen University, Gen.
Foulkesweg 37, 6703 BL Wageningen, Netherlands (Cajanus cajan, Cicer arieti-
num, Tylosema esculentum)
- S.R. Vodouhè, IPGRI West and Central Africa, 08 B.P. 0932, Cotonou, Benin
(Digitaria exilis, Macrotyloma geocarpum)
- W. Wessel-Brand, Biosystematics Group, Wageningen University, Gen. Foul-
kesweg 37, 6703 BL Wageningen, Netherlands (illustrations)
- E. Westphal, Ritzema Bosweg 13, 6706 BB Wageningen, Netherlands (associate
editor)
- C S . Wortmann, University of Nebraska Lincoln, IANR, Department of Agro-
nomy and Horticulture, 154 Keim Hall, Lincoln, NE 68583-0915, United States
(Phaseolus vulgaris)
- S.S. Yadav, Division of Genetics, Indian Agricultural Research Institute, New
Delhi 110012, India (Lathyrus sativus)

Acknowledgments

- S. van Otterloo-Butler, Bowlespark 21,6701 DR Wageningen, Netherlands

(English language correction)
- N. Wulijarni-Soetjipto, PROSEA Network Office, P.O. Box 332, Bogor 16122,
Indonesia (coordination illustrators)
PROTABoard ofTrustees and Personnel

Board of Trustees

J.R. Cobbinah (FORIG, Ghana), chairman

M.J. Kropff (WU, Netherlands), vice-chairman
H. Andriamialison (PBZT, Madagascar)
P.R. Crane (RBGKEW, United Kingdom)
D. Garrity (ICRAF, Kenya)
L.S. Luboobi (MU, Uganda)
Z.L.K. Magombo (NHBGM, Malawi)
G. Matheron (AGROPOLIS, France)
S. Mbadinga (CENAREST, Gabon)
M. Ouédraogo (CNSF, Burkina Faso)
E. Sukara (PROSEA, Indonesia)

Personnel

Regional Office Central Africa, Gabon

S. Mbadinga, Programme Leader
B. Nziengui, Regional Officer
P.M. Nsole Biteghe, Assistant Regional Officer
D.N. Omokolo, Contact Person Cameroon
M.K.D. Ben-Bala, Contact Person Central African Republic

Regional Office East Africa, Uganda

J.S. Kaboggoza, Programme Leader
R. Bukenya-Ziraba, Regional Officer
M. Atim, Assistant Regional Officer
A. Tsegaye, Contact Person Ethiopia
J. Elia, Contact Person Tanzania

Regional Office Indian Ocean Islands, Madagascar

S. Rapanarivo, Programme Leader

M.E. Rahelivololana, Regional Officer
A. Gurib-Fakim, Contact Person Mauritius
S. Brillant, Contact Person Réunion

Regional Office Southern Africa, Malawi

Z.L.K. Magombo, Programme Leader

N.G. Nyirenda, Regional Officer
V.K. Kawanga, Contact Person Zambia
10 CEREALS AND PULSES

Regional Office (anglophone) West Africa, Ghana

J.R. Cobbinah, Programme Leader

S. Britwum, Regional Officer
A. Armooh, Assistant Regional Officer
O.A. Denton, Contact Person Nigeria

Regional Office (francophone) West Africa, Burkina Faso

M. Ouédraogo, Programme Leader

A. Traoré, Regional Officer
V. Millogo, Assistant Regional Officer
C. Kouamé, Contact Person Côte d'Ivoire
F. Assogba-Komlan, Contact Person Benin

Country Office France

M. Chauvet, Programme Leader

W. Rodrigues, Country Officer

Country Office United Kingdom

S.D. Davis, Programme Leader

O. Grace, Country Officer

Network Office Africa, Kenya

E.A. Omino, Head

J. Chege, Database Officer
D.J. Borus, Dissemination Officer
B.O. Obongoya, Programme Officer
M.W. Kamanda, Secretary
D. Laur, Office Assistant

Network Office Europe, Netherlands

J.S. Siemonsma, Head

A.D. Bosch-Jonkers, Secretary/Management Assistant
R.H.M.J. Lemmens, General Editor
L.P.A. Oyen, General Editor
E.J. Bertrums, Databank Manager
C.H. Bosch, Editor/Dissemination Officer
M. Brink, Editor
A. de Ruijter, Editor
G.H. Schmelzer, Editor/Dissemination Officer
 11

Introduction

Choice of s p e c i e s

PROTA 1:'Cereals and pulses' describes the cultivated and wild species of tropical
Africa used as a cereal or pulse. Cereals can be defined as grasses (family Poaceae)
of which the grain is used for food; they may be cultivated or the grain is collected
from wild plants ('wild cereals'). Three cereals that are not grasses ('pseudo-cereals')
have also been included in this volume: Amaranthus caudatus L. (grain amaranth),
Fagopyrum esculentum Moench (buckwheat) and Limeum obovatum Vicary. Pulses
can be defined as leguminous species (members of the families Papilionaceae, Caes-
alpiniaceae or Mimosaceae, often considered as one family Leguminosae) producing
edible mature seeds. Pulses may also be cultivated or collected from the wild.
Some species are only used as a cereal or pulse, but most have several uses. PROTA
normally assigns a single primary use and, where relevant, one or more secondary
uses to all plant species used in Africa. PROTA 1: 'Cereals and pulses' comprises
only accounts of species whose primary use is as a cereal or pulse. The primary use
of pigeon pea (Cajanus cajan (L.) Millsp.) is as a pulse, and thus it is treated in
PROTA 1, but it has various secondary uses, e.g. the immature seeds and pods are
eaten as a vegetable, the seeds and the by-products of dhal production are used as
animal feed, the vegetative parts are used as fodder, the stems and branches are
used for basketry, thatching, fencing and as fuel, from various plant parts tradi-
tional medicines are prepared, and the plants are grown as a shade crop or cover
crop and in hedges and windbreaks. Winged bean (Psophocarpus tetragonolobus (L.)
D C ) , on the other hand, is eaten as a pulse, but its primary use is its immature
pods being eaten as a vegetable, and consequently winged bean is described in
PROTA 2: 'Vegetables'.
Species that are used as a cereal or pulse in tropical Africa but have another pri-
mary use are listed after the primary use cereals and pulses, and are fully described
in other commodity groups. Some well-known species included in this list are: kodo
millet (Paspalum scrobiculatum L.), lablab (Lablab purpureus (L.) Sweet) and
winged bean (Psophocarpus tetragonolobus (L.) D C ) .
Six species are treated which have two primary uses, including use as cereal or
pulse, and consequently will be described in two commodity groups. These species
are Arachis hypogaea L. (also in PROTA 14: 'Vegetable oils'), Glycine max (L.) Merr.
(also in PROTA 14: 'Vegetable oils'), Phaseolus vulgaris L. (also in PROTA 2 'Vege-
tables'), Pisum sativum L. (also in PROTA 2 'Vegetables'), Vigna unguiculata (L.)
Walp. (also in PROTA 2 'Vegetables'), and Sorghum bicolor (L.) Moench (also in
PROTA 3:'Dyes and tannins').
In PROTA 1: 'Cereals and pulses' comprehensive descriptions are given of 35 impor-
tant species (15 cereals, 19 pulses and 1pseudo-cereal). These major cereals and
pulses comprise most cultivated species, but also several wild or partly domesticated
ones. The accounts are presented in a detailed format and illustrated with a line
drawing and a distribution map. In addition, accounts of 38 species of minor impor-
tance (22 cereals, 14 pulses and 2 pseudo-cereals) are given. Because information on
these species is often scanty, these accounts are in a simplified format. For another
12 CEREALSANDPULSES

9 species (5 cereals and 4 pulses) the information was too scarce to justify an indi-
vidual treatment and they have only been mentioned in the accounts of related spe-
cies.

Plant n a m e s

Family: Apart from the classic family name, the family name in accordance with the
Angiosperm Phylogeny Group (APG) classification is also given where it differs from
the classic name.
Synonyms: Only the most commonly used synonyms and those that may cause con-
fusion are mentioned.
Vernacular names: Only names in official languages of regional importance in Africa
are included: English, French, Portuguese and Swahili. It is beyond the scope of
PROTA to give an extensive account of the names of a species in all languages spo-
ken in its area of distribution. Checking names would require extensive fieldwork by
specialists. Although regional forms ofArabic are spoken in several countries in Af-
rica, the number ofAfrican plant species t h a t have a name in written, classical Ara-
bic is limited. Arabic names are therefore omitted. Names of plant products are
mentioned under the heading 'Uses'.

Origin a n d g e o g r a p h i c distribution

To avoid long lists of countries in the text, a distribution map is added for major
species. The map indicates in which countries a species has been recorded, either
wild or planted. For many species, however, these maps are incomplete because they
are prepared on the basis of published information, the quantity and quality of
which varies greatly from species to species. This is especially the case for wild spe-
cies which are not or incompletely covered by the regional African floras, and for
cultivated species which are only planted on a small scale (e.g. in home gardens).
For some countries (e.g. Central African Republic, Chad, Sudan, Angola) there is
comparatively little information in the literature. Sometimes they are not covered by
recent regional or national floras and although species may be present there, this can-
not be demonstrated or confirmed.

Properties

The food value of the cereals and pulses is mentioned in the species accounts. The
analytical method used to determine the various elements of the nutritional compo-
sition considerably influences the values found. For this reason a few standard
sources were used wherever possible and the sources are mentioned in the text.
These sources are: the USDA Nutrient database for standard reference; McCance &
Widdowson's The composition of foods; FAO Food composition table for use in Af-
rica.
Apart from nutrients, this section includes other properties relevant to the respec-
tive uses.
 INTRODUCTION 13

Description

A morphological characterization of the species is given. The description is in 'tele-

gram' style and uses botanical terms. Providing a description for the general public
is difficult as more generally understood terms often lack the accuracy required in a
botanical description. A line drawing is added for all major and some lesser-known
species to complement and visualize the description.

Management

Descriptions of husbandry methods including fertilizer application, irrigation, and

pest and disease control measures are given under 'Management' and under 'Dis-
eases and pests'. These reflect actual practices or generalized recommendations, opt-
ing for a broad overview but without detailed recommendations adapted to the
widely varying local conditions encountered by farmers. Recommendations on
chemical control of pests and diseases are merely indicative and local regulations
should be given precedence. PROTA will participate in the preparation of derived
materials for extension and education, for which the texts in this volume provide a
basis, but to which specific local information will be added.

Genetic r e s o u r c e s

The genetic diversity of many plant species in Africa is being eroded, sometimes at
an alarming rate, as a consequence of habitat destruction and overexploitation. The
replacement of landraces of cultivated species by modern cultivars marketed by seed
companies is another cause of genetic erosion. Reviews are given of possible threats
for plant species and of the diversity within species. Information on ex-situ germ-
plasm collections is mostly extracted from publications of the International Plant
Genetic Resources Institute (IPGRI).

References

The main objective of the list of references given is to guide readers to additional
information; it is not intended to be complete or exhaustive. Authors and editors
have selected two categories of references; 'major references' are limited to 10 refer-
ences (5for minor species), the number of'other references' is limited to 20 (10 for mi-
nor species). The references listed include those used in writing the account. Where
the internet was used, the website and date are cited.
14 CEREALS AND PULSES
 15

Alphabetical treatment of cereals and pulses

16 CEREALSANDPULSES
 AMARANTHUS 17

AMARANTHUS CAUDATUS L. crop in Asia and Africa. As an ornamental it is

grown throughout much of the tropics and in
Protologue Sp. pi. 2:990 (1753). some temperate regions.
Family Amaranthaceae The exact distribution ofAmaranthus caudatus
Chromosome n u m b e r 2n - 32 in Africa is not known, because it has often
Vernacular names been confounded with other Amaranthus spe-
- Grain amaranth, Inca wheat, jataco (En). cies. It is grown in Ethiopia and Eritrea for its
Amarante-grain, blé des Incas (Fr). Amaran- grain and as an ornamental; it has also been
te de cauda (Po). grown in Uganda and Kenya and has been
- African spinach, Indian spinach (En). Brède recorded from several other countries in Cen-
malabar (Fr). Bredo (Po).Mchicha (Sw). tral, East and southern Africa, and the Mas-
- Love-lies-bleeding, red-hot cattail, foxtail carene Islands, where it may also be found as a
(En). Queue de renard, discipline des reli- weed escaped from cultivation.
gieux (Fr). Cauda de raposa, moncos de peru Uses Amaranthus caudatus seeds are toasted
(Po). and popped, ground into flour or boiled for
Origin and geographic distribution Ama- gruel. For making leavened foods, they must be
ranthus caudatus is not known from the wild. blended with wheat. The seeds are fermented
It originated in the Andes, possibly as a hybrid to make alcoholic beverages, e.g. beer ('tella') in
between the wild Amaranthus hybridus L. Ethiopia. In Ethiopia cooked seeds are made
subsp. quitensis (Kunth) Costea & Carretero into porridge, and ground seeds are mixed with
and the cultivated Amaranthus cruentus L. tef to prepare pancake-like bread ('injera').
(originating from Central America). Amaran- Seeds can be sprouted for use as a nutritious
thus caudatus has long been grown as a food vegetable. The leaves are eaten as a vegetable
crop in the Andes, e.g. by the Incas, and the like those of other amaranth species, e.g. in
greatest genetic variation occurs in this area Peru and Ethiopia. Harvest residues are used
(Ecuador, Peru, Bolivia and Argentina). The for feeding livestock and for thatching. In
earliest archaeological evidence of its cultiva- South America grain amaranths are tradition-
tion dates from 2000-year-old tombs in north- ally used in medicine, folk festivals, and as dye
western Argentina. The chronicler Cobo wrote sources. In Ethiopia the root is used as a laxa-
in 1653 that in the city of Guamanga (now Aya- tive, and the seed for expelling tapeworms and
cucho) delicious sweets were prepared from for treating eye diseases, amoebic dysentery,
amaranth and sugar. Amaranthus caudatus and breast complaints. In India the plant is
was introduced into Europe in the 16th century taken as a diuretic and it is applied to sores.
and it was spread to Africa and Asia later. The Amaranthus caudatus is widely grown as an
cultivated area has notably decreased over the ornamental ('Love-lies-bleeding').
years, but Amaranthus caudatus has remained Production and international trade No sta-
a grain crop in Ecuador, Peru, Bolivia and Ar- tistics are available on production and trade of
gentina. It is occasionally grown as a grain grain amaranths in general and Amaranthus
caudatus in particular. Reports from the 1990s
mention several thousands of ha of grain ama-
ranths in China, similar large production areas
in Argentina, and about 2000 ha in the United
States. Estimates for India and Nepal are up to
4000 ha. In Peru, there are over 1000 ha under
grain amaranths (mainly Amaranthus cauda-
tus) in the high Andean region alone. The
United States imports large quantities of grain
amaranth from Mexico.
P r o p e r t i e s Grain amaranth (species un-
specified) seeds contain per 100 g edible por-
tion: water 9.8 g, energy 1565 kJ (374 kcal),
protein 14.5 g, fat 6.5 g, carbohydrate 66.2 g,
dietary fibre 15.2 g, Ca 153 mg, Mg 266 mg, P
455 mg, Fe 7.6 mg, Zn 3.2 mg, vitamin A 0 IU,
thiamin 0.08 mg, riboflavin 0.21 mg, niacin
Amaranthus caudatus - planted 1.29 mg, vitamin B6 0.22 mg, folate 49 ug and
18 CEREALS AND PULSES

ascorbic acid 4.2 mg. The essential amino-acid

composition per 100 g edible portion is: trypto-
phan 181 mg, lysine 747 mg, methionine 226
mg, phenylalanine 542 mg, threonine 558 mg,
valine 679 mg, leucine 879 mg and isoleucine
582 mg. The main fatty acids are (per 100 g
edible portion): linoleic acid 2834 mg, oleic acid
1433 mg, palmitic acid 1284 mg and stearic
acid 220 mg (USDA, 2004).
Amaranth grain is renowned for the excellent
quality of its protein because of the high lysine
content. Gluten-free types exist and are suit-
able for people with coeliac disease. The starch
mainly consists of amylopectin, with only5—7%
amylose. The rather small starch granules (1-3
|im in diameter) have drawn wide attention for
industrial uses of grain amaranths. The oil
contains 4-11% of the triterpenoid squalene,
which may find a niche market in products
such as lubricants in the computer industry
and in cosmetics. The stems, leaves and seeds
of Amaranthus caudatus contain water-soluble
red-violet betacyanin pigments. In aqueous Amaranthus caudatus - 1,upper part of flower-
plant extracts they consist on average of 81% ingplant; 2, dehisced fruit.
amaranthine and 19% isoamaranthine; in Redrawn and adapted byIskak Syamsudin
dried extracts of 67% amaranthine and 33%
isoamaranthine. Dissolved in water the pig- by 3 stigmas. Fruit an ovoid-globose capsule
ments are unstable except at low temperatures 1.5-2.5 mm long, circumscissile, almost smooth
in the dark and in the absence of air; dried or slightly furrowed, abruptly narrowed to a
pigments are very stable at room temperature. short thick beak, 1-seeded. Seed almost glo-
The seed also contains red-coloured lectins bose, 1—1.5 mm long, smooth and shining, pale
sometimes also referred to as amaranthine. coloured (ivory), reddish or dark brown.
Two peptides isolated from the seed of Ama- Other botanical information Amaranthus
ranthus caudatus (Ac-AMPl and Ac-AMP2) comprises about 70 species, of which about 40
have shown strong antifungal and some anti- are native to the Americas. It includes at least
bacterial activity. 17 species with edible leaves and 3 grain ama-
Description Annual erect herb up to 1.5(— ranths. Amaranthus caudatus is part of the so-
2.5) m tall, commonly reddish or purplish called Amaranthus hybridus aggregate, a
throughout; stem rather stout, not or sparingly group of species in which taxonomie problems
branched, glabrous or thinly furnished with are far from clarified, especially because of
rather long hairs. Leaves arranged spirally, common hybridization and names often being
simple and entire; stipules absent; petiole up to misapplied. Some recognized species of this
8 cm long, but never longer than the blade; aggregate are cultivated taxa. Amaranthus
blade broadly ovate to rhomboid-ovate or ovate- caudatus is one of these, as are the other grain
elliptical, 2.5-15(-20) cm x 1-8 cm, base amaranths, Amaranthus cruentus L. and Ama-
cuneate to attenuate, apex obtuse to acute, ranthus hypochondriacus L., which are in-
glabrous or sparsely hairy on main veins be- cluded in PROTA 2 'Vegetables'. Amaranthus
low, pinnately veined. Inflorescence large (up caudatus can be distinguished by its usually
to 1.5 m) and complex, consisting of numerous long and pendant terminal spike and compara-
agglomerated cymes arranged in axillary and tively broad tepals of female flowers. A classifi-
terminal spikes, the terminal one pendant to cation in cultivar groups might be more appro-
erect; bracts 3—4 mm long, membranous, pale, priate for the cultivated taxa.
with a long awn. Flowers unisexual, sessile; Amaranthus caudatus shows a wide genetic
with 5 mucronate tepals 2—3 mm long; male variation and diversity of plant form, ranging
flowers with 5 stamens c. 1 mm long; female from erect to completely decumbent. Two types
flowers with superior, 1-celled ovary crowned have been distinguished: subsp. caudatus, the
 AMARANTHUS 19

m a i n type, a n d subsp. mantegazzianus (Pass.) Amaranthus caudatus is grown in sole crop-

H a n e l t , grown as a grain crop in t h e valleys of ping as well as in intercropping systems, e.g.
t h e A n d e s in n o r t h - w e s t e r n A r g e n t i n a . The w i t h maize in South America and Ethiopia.
l a t t e r can be distinguished by its d e t e r m i n a t e Sometimes it is p l a n t e d as a guard-row for a
club-shaped inflorescence b r a n c h e s , due to a m a i n crop, e.g. bean, m a i z e or millet. In Ethio-
single recessive gene. According to some, it pia t h e p l a n t s a r e sometimes allowed to sow
should be considered as a s e p a r a t e species themselves.
Amaranthus mantegazzianus Pass., a n opinion M a n a g e m e n t Once t h e y a r e established,
which h a s recently been s u p p o r t e d by t h e re- grain a m a r a n t h s compete well with weeds, b u t
sults of seed protein studies. t h e y m u s t be weeded a t least once d u r i n g t h e
G r o w t h a n d d e v e l o p m e n t G e r m i n a t i o n of first m o n t h . Hilling t h e p l a n t s w h e n t h e y a r e
Amaranthus caudatus seed accelerates w i t h about 30 cm tall helps to control weeds a n d to
increasing t e m p e r a t u r e in t h e r a n g e 5-35°C; reduce lodging; it m a y also control Alternaria
no g e r m i n a t i o n occurs a t 0°C. Seedlings nor- disease. In P e r u Amaranthus caudatus is
mally emerge 3 - 5 days after sowing a n d early sometimes grown with s u p p l e m e n t a l irrigation.
growth is slow. Flowering begins 6 0 - 1 1 0 days R e s u l t s of fertilizer trials a r e inconclusive, a n d
after emergence. Outcrossing r a t e s of 6-29% in general grain a m a r a n t h s grow well u n d e r
h a v e been recorded in Amaranthus caudatus. widely differing n u t r i e n t levels. In P e r u ma-
T h e total crop d u r a t i o n in P e r u r a n g e s from 3— n u r e is usually applied.
4 m o n t h s a t 1800 m a l t i t u d e to 9 m o n t h s a t D i s e a s e s a n d p e s t s In Amaranthus cauda-
3200 m altitude; in K e n y a it is normally 8 0 - 9 0 tus fungal diseases h a v e been observed caused
days. A single p l a n t m a y yield more t h a n by Alternaria, Mycoplasma and Sclerotinia
50,000 seeds. Amaranthus caudatus is a C4- spp. P e s t s c a u s i n g economic d a m a g e to grain
cycle plant, giving h i g h e r yields a t h i g h e r light a m a r a n t h s a r e mainly leaf-eating caterpillars
intensities a n d t e m p e r a t u r e s , a n d being effi- (Heliothis, Hymenia, Spodoptera), stinkbugs
cient in w a t e r use. (e.g. Lygus on t h e inflorescence), stem-boring
E c o l o g y In t h e tropics Amaranthus cauda- l a r v a e of weevils, g r a s s h o p p e r s a n d aphids.
tus performs well u n d e r cool, dry h i g h l a n d con- H a r v e s t i n g H a r v e s t i n g of Amaranthus cau-
ditions. It is more t o l e r a n t to chilling t h a n t h e datus is difficult because of asynchronous rip-
other 2 grain a m a r a n t h s a n d is grown a t ening. T h e crop m a y be h a r v e s t e d once-over by
h i g h e r a l t i t u d e s . I n E a s t Africa it is found a t c u t t i n g t h e inflorescences w h e n t h e p l a n t s a r e
5 0 0 - 2 5 0 0 m a l t i t u d e , in South America a t still green, to avoid seed s h a t t e r i n g . In P e r u
1000-3200 m. In P e r u it is grown in regions t h e p l a n t s a r e cut a t ground level with a sickle,
with a n average a n n u a l rainfall of 550 m m . bundled, a n d left to dry in t h e field for 1—2
T h e photoperiodic response is m a r k e d , with w e e k s . In dry, irrigated a r e a s senescence can
flowering being promoted by s h o r t photo- be induced by stopping irrigation 2 weeks be-
periods. fore t h e h a r v e s t . Leaves m a y be h a r v e s t e d once
Amaranthus caudatus can be grown in s a n d y or several t i m e s to be u s e d as a vegetable, be-
a n d clay soils. I n general grain a m a r a n t h s pre- fore t h e grain or total biomass is h a r v e s t e d a t
fer well-drained n e u t r a l or alkaline soils m a t u r i t y for food or forage.
(pH>6), b u t some types a r e well a d a p t e d to Y i e l d Seed yields of grain a m a r a n t h s v a r y
acid a n d mildly saline soils. widely, from as low a s 500—800 kg/ha to as
Propagation and planting Amaranthus high as 2 5 0 0 - 4 0 0 0 kg/ha. In n o r t h - w e s t e r n
caudatus is p r o p a g a t e d by seed. I t s 1000-seed India, w h e r e all 3 grain a m a r a n t h species a r e
weight is 0.5—1.1 g. Seed scarification with grown, yields of Amaranthus caudatus and
c o n c e n t r a t e d sulphuric acid or s a n d p a p e r en- Amaranthus cruentus a r e lower t h a n those of
h a n c e s germination. Common seed r a t e s in Amaranthus hypochondriacus.
P e r u a r e 8 - 1 8 kg/ha, b u t in Kenya seed r a t e s of H a n d l i n g a f t e r h a r v e s t After drying in t h e
only 1-2 kg/ha a r e common. W i t h improved field, t h e p l a n t s a r e t h r e s h e d by h a n d or ma-
cultivars in P e r u densities of 400,000-500,000 chine, a n d t h e seed is cleaned. Additional dry-
p l a n t s / h a gave t h e h i g h e s t yields. Emergence ing to reduce t h e m o i s t u r e content to 12% m a y
of seedlings is best w h e n t h e seed is sown a t a be necessary for safe storage.
d e p t h of 1—1.5 cm, b u t in dry, hot a r e a s deeper G e n e t i c r e s o u r c e s L a r g e g e r m p l a s m collec-
sowing m a y be necessary. However, emergence tions of Amaranthus caudatus a r e kept in P e r u
is seriously lowered if t h e sowing d e p t h exceeds (Universidad Nacional de S a n Antonio A b a d
5 cm. del Cusco (UNSAAC/CICA), Cusco, 1600 acces-
20 CEREALS AND PULSES

sions; Universidad Nacional Agraria La Moli- Sharma, 1983; Paredes-López (Editor), 1994;
na, Lima, 333 accessions; Estación Experimen- Sauer, 1976; Stallknecht & Schulz-Schaeffer,
tal Agraria Banos del Inca, Cajamarca, 257 1993; Sun, Chen & Leung, 1999; Townsend,
accessions) and the United States (Organic 1985; Townsend, 1994;USDA, 2004.
Gardening and Farming Research Center, Sources of illustration Grubben, 1975.
Kutztown, Pennsylvania, 297 accessions). The Authors S.G. Agong
only germplasm collection of Amaranthus cau-
datus in Africa recorded by IPGRI is in Kenya
(National Genebank of Kenya, Crop Plant Ge- ARACHIS HYPOGAEA L.
netic Resources Centre, KARI, Kikuyu, 4 ac-
cessions). Protologue Sp. pi. 2: 741 (1753).
Breeding Major breeding objectives for Family Papilionaceae (Leguminosae - Papil-
Amaranthus caudatus and other grain ama- ionoideae, Fabaceae)
ranths are improved harvestability (less lodg- Chromosome number 2n = 40
ing, less shattering, better uniformity of ma- Vernacular names Groundnut, peanut, earth-
turity), increased seedling vigour, increased nut, monkey nut (En). Arachide, cacahuète,
resistance to pests and higher yields. In Peru cacahouète, pistache de terre (Fr). Amendoim,
selection in landraces has led to the release of mandobi, caranga (Po). Mjugu nyasa, mnjugu
the Amaranthus caudatus cultivars 'Noel nyasa, karanga (Sw).
Vietmeyer', 'Oscar Blanco' and 'Alan Garcia'. Origin and geographic distribution Ground-
Genetic studies have identified marker loci for nut originated in the area of southern Bolivia
traits such as pigmentation patterns, inflores- and north-western Argentina. It is an ancient
cence morphology and seed characters in Ama- crop of the New World and was widely grown
ranthus caudatus and other grain amaranths. in Mexico, Central America and South America
Research is needed on hybridization barriers in pre-Columbian times. Domesticated ground-
among grain amaranth species as well as on nut had already evolved into several types be-
the biosystematic identity of the species; only fore it was introduced into the Old World by
then can the information be indisputably re- Spanish and Portuguese explorers. Two-seeded
lated to species. types originating from Brazil were brought to
Prospects In tropical Africa Amaranthus West Africa, and 3-seeded types originating
caudatus is presently cultivated on a very lim- from Peru were taken from the west coast of
ited scale and, like the other grain amaranths, South America to the Philippines, from where
it probably does not have much future as a they spread to Japan, China, Indonesia, Ma-
grain crop, because it cannot compete with laysia, India, Madagascar and East Africa. In
cereals that are more productive and easier to the late 1700s 'Spanish' groundnut types were
grow. On a worldwide scale grain amaranths introduced into Europe from Brazil. The first
have some potential, because of their favour- successful introduction in North America con-
able agronomic characteristics, excellent nutri- cerned small-seeded 'runner'-type groundnuts,
tional qualities and diverse food and technical
applications. Amaranths are also considered to
have prospects in food colouring.
Major references Agong & Ayiecho, 1991;
Bale & Kauffman (Editors), 1992; Brenner et
al., 2000; Costea, Sanders & Waines, 2001;
Jain & Sutarno, 1996; National Research
Council, 1984; National Research Council,
1989; Sauer, 1967; Townsend, 2000; Williams
& Brenner, 1995.
Other references Berghofer & Schoen-
lechner, 2002; Broekaert et al., 1992; Cai, Sun
& Corke, 1998; Cai et al., 1998; Coons, 1982;
CSIR, 1950; Drzewiecki, 2001; Getahun, 1976;
Gutterman, Corbineau & Come, 1992; Hanelt
& Institute of Plant Genetics and Crop Plant
Research (Editors), 2001; Hauman, 1951; Jan-
ick & Simon (Editors), 1990; Joshi, Mehra & Arachis hypogaea - planted
 ARACHIS 21

probably originating from northern Brazil or galactagogue, and used as eye-drops to treat
the West Indies. Groundnut is now grown in conjunctivis. Macerations of peeled seeds are
most tropical, subtropical and temperate coun- drunk to treat gonorrhoea, macerations of the
tries between 40°N and 40°S latitude. It is seed coats against syphilis, while macerations
grown throughout tropical Africa and is a ma- of the seed coats and shells are applied against
jor cash crop in Senegal, Gambia, Nigeria and ophthalmia. Sap of ground leaves and seeds is
Sudan. used for ear-drops against ear discharge. Leaf
Uses Groundnut seed is mainly used as food macerations are drunk as a diuretic. Leaf infu-
and for oil extraction. The seeds are eaten raw, sions are drunk against female infertility, and
boiled or roasted, made into peanut butter, used for eye-drops to treat eye injuries and
confectioneries and snack foods, and are used cataract. Plant ash with salt is applied in case
for thickening soups or made into sauces to be of caries. Pod extracts and young plants are
eaten with meat and rice. In northern Nigeria credited with aphrodisiac properties. The plant
groundnut flour is mixed with 'gari' (coarse is also used to relieve cough and is considered
fermented cassava meal) and made into balls emollient and demulcent; emulsions are taken
that are eaten as a snack. In the United States to treat pleurisy, enteritis (including colitis),
and Argentina most of the crop is used as food, and dysuria.
but in most other countries the primary use of Agglutinins (lectins) from groundnut seeds are
groundnut is for the oil market. Worldwide, often used in medical research for histochemi-
more than 50% of groundnut production is cal investigations.
crushed into oil for human consumption or Production and international trade Accord-
industrial use (e.g. in cosmetics). In countries ing to FAO estimates, the average world pro-
such as Senegal, Gambia and Nigeria oil ex- duction of groundnut pods in 1999-2003
traction has been an important cottage indus- amounted to about 34.4 million t/year from
try for years. The use of groundnut in confec- 24.4 million ha. The main producing countries
tionery and for oil and meal production is in- are China (14.0 million t/year in 1999-2003,
creasing, and there is gradual shift taking from 4.9 million ha), India (6.1 million t/year
place from oil and meal to confectionery use, from 6.7 million ha), Nigeria (2.8 million t/year
especially in Latin America and the Caribbean. from 2.7 million ha), the United States (1.7
In South America groundnut seeds are fer- million t/year from 0.5 million ha), Indonesia
mented into alcoholic drinks. (1.3 million t/year from 0.7 million ha) and
The presscake from oil extraction is a feed rich Sudan (1.1 million t/year from 1.7 million ha).
in protein, but it is also made into groundnut The total production in sub-Saharan Africa
flour, which is used in many human foods. was 8.2 million t/year from 9.5 million ha.
Fermented groundnut cake is eaten fried in Average world export of groundnut seeds
Indonesia. The cake finds industrial applica- amounted to 1.1 million t/year in 1998-2002.
tion in the production of glues, sizes for paper The main exporters were China (321,000
and starches for laundering and textile manu- t/year), Argentina (201,000 t/year) and the
facture. Protein from groundnut cake is made United States (171,000 t/year). Export of
into a wool-like fibre, which can be blended groundnut seeds from sub-Saharan Africa was
with wool or rayon. Groundnut shells are used 64,000 t/year, with Gambia as main exporter
as roughage in fodder, as fuel, fertilizer, mulch, (26,000 t/year). Average world export of ground-
in the manufacture of particle board and build- nut pods in 1998-2002 was only 176,000 t/year,
ing blocks, and can be used as a source of acti- with China as main exporter (73,000 t/year).
vated carbon, combustible gases, organic Exports of groundnut pods from sub-Saharan
chemicals, reducing sugars, alcohol and ex- Africa were negligible.
tender resins. The world production of groundnut oil in 1999-
Young groundnut pods and leaves are con- 2003 was 5.1 million t/year. The main produc-
sumed as a vegetable; in West Africa the leaves ers are China (2.0 million t/year), India (1.4
are added to soups. The foliage is an important million t/year), Nigeria (480,000 t/year), Sene-
fodder, especially in the Sahel; it may be eaten gal (178,000 t/year) and Sudan (162,000
fresh or as hay or silage. In southern India the t/year). The production in sub-Saharan Africa
haulms are sometimes applied as a green ma- was 1.2 million t/year. The world groundnut
nure. cake production in 1999—2003 was 6.9 million
Groundnut has a range of uses in traditional t/year, mainly from China (2.6 million t/year),
African medicine. Pod extracts are taken as a India (1.9 million t/year) and Nigeria (750,000
22 CEREALSANDPULSES

t/year). The production in sub-Saharan Africa An important problem in groundnut production

was 1.6 million t/year. is aflatoxin contamination byAspergillus fungi.
Average world export of groundnut oil in 1998- Aflatoxin has immunosuppressive effects and
2002 was 271,000 t/year, with as main export- epidemiological studies, also in Africa, have
ers Senegal (83,000 t/year) and Argentina shown a positive correlation between aflatoxin
(69,000 t/year). The total export of groundnut intake and the incidence of liver cancer. After
oil from sub-Saharan Africa was 114,000 industrial oil extraction, aflatoxin remains in
t/year. The main importers were France the cake, and the refined oil is free of aflatoxin,
(68,000 t/year), Italy (46,000 t/year) and the but in case of small-scale extraction, the non-
United States (25,000 t/year). Average ground- refined oil may be contaminated. Groundnut is
nut cake export amounted to 280,000 t/year. one of the most allergenic foods known and
Major exporters were Senegal (103,000 t/year), may cause anaphylactic reactions. Groundnut
Argentina (51,000 t/year), India (43,000 t/year) seeds contain a haemostatic factor which can
and Sudan (35,000 t/year). Total groundnut be useful in haemophilia. Groundnut oil is
cake export from sub-Saharan Africa was mildly laxative.
143,000 t/year. The main importers were Adulterations and substitutes Groundnut
France (129,000 t/year) and Thailand (53,000 oil can be substituted by other vegetable oils,
t/year). e.g. from maize, soya bean and sunflower.
Properties Mature groundnut seeds contain Description Annual herb, with erect or
per 100 g edible portion (average of several prostrate stem up to 70 cm long; root system
types, which show little difference): water 6.5 consisting of a well-developed taproot with
g, energy 2374 kJ (567 kcal), protein 25.8 g, fat many lateral roots, up to 135 cm deep, but gen-
49.2 g, carbohydrate 16.1 g, dietary fibre 8.5 g, erally restricted to the upper layers of the soil.
Ca 92 mg, Mg 168 mg, P 376 mg, Fe 4.6 mg, Zn Leaves arranged spirally, 4-foliolate with two
3.3 mg, vitamin A 0 IU, thiamin 0.64 mg, ribo- opposite pairs of leaflets; stipules 1.5-4 cm
flavin 0.14 mg, niacin 12.1 mg, vitamin P>60.35 long, with a slender free tip, but fused to the
mg, folate 240 \ig and ascorbic acid 0 mg. The petiole for about half their length; petiole 1.5-7
essential amino-acid composition per 100 g cm long; petiolules 1—2 mm long; leaflets
edible portion is: tryptophan 250 mg, lysine
926 mg, methionine 317 mg, phenylalanine
1337 mg, threonine 883 mg, valine 1082 mg,
leucine 1672 mg and isoleucine 907 mg. The
principal fatty acids are per 100 g edible por-
tion: oleic acid 23.7 g, linoleic acid 15.6 g and
palmitic acid 5.2 g (USDA, 2004).
Groundnut seeds yield 42-56% oil. Groundnut
oil contains 36-72% oleic acid, 13—48% linoleic
acid and 6-20% palmitic acid. The ratio of oleic
to linoleic acid has an important bearing on the
stability of the oil; the higher the ratio, the
more stable the oil and the longer its shelf life.
The ratio in mature seeds can range from less
than 1.0 to greater than 3.0; more than 1.3 is
generally considered satisfactory by processors.
The presscake contains 40-50% easily digesti-
ble protein, 20-25% carbohydrate and 5-15%
residual oil.
Groundnut pods have a thick woody shell con-
taining normally 2-3 seeds ('kernels'). The seed
coat constitutes about 4—5%of the seed weight,
the cotyledons 90-94% and the germ 3-4%.
The major components of the seed coat are
carbohydrate, cellulose and protein. Oil and
protein are the main constituents of the germ Arachis hypogaea - 1, branch with flowers and
and cotyledons. The germ is associated with fruit; 2, inflorescence; 3,fruit; 4, seeds.
bitter components. Source: PROSEA
 AEACHIS 23

obovate or elliptical, 1-7 cm x 0.5-3 cm, high levels of sterility due to ploidy level differ-
cuneate-rounded at base, rounded or emargi- ences and genome incompatibility.
nate and mucronate at apex. Inflorescence an There is considerable variation in Arachis hy-
axillary, 2-5-flowered spike. Flowers bisexual, pogaea and two subspecies have been distin-
papilionaceous, sessile; receptacle long and guished: subsp. hypogaea and subsp. fastigiata
slender, pedicel-like, up to 4 cm long; calyx Waldron. Subsp. hypogaea ('runner type') is
with 4 upper lobes joined, lower lobe free; co- characterized by a more prostrate growth habit
rolla pale yellow to orange-red, rarely white, without flowering branches on the main stem,
standard rounded, c. 1.5 cm x 1.5 cm, wings and with the cotyledonary lateral branches
shorter, keel incurved; stamens (8-)10, alter- carrying alternate pairs of vegetative and re-
nately with small, globular anthers and larger, productive secondary branches; it is usually
oblong anthers, joined at base; ovary superior late-maturing. It includes the 'Virginia' types
but situated at base of receptacle tube, style groundnut. Subsp. fastigiata ('bunch type') is
free within the tube, very long, ending in a characterized by an erect growth habit with
minute club-shaped stigma. Fruit an oblong or flowering branches on the main stem, and
sausage-shaped pod, borne at the tip of an without a regular pattern in the sequence of
elongated fruit stalk ('peg') up to 20 cm long, 1— vegetative and reproductive branches; and it is
8 cm x 0.5—2cm, surface constricted to varying early-maturing. It includes the 'Spanish' and
degrees between the seeds and reticulately 'Valencia' types groundnut.
veined, 1—6-seeded. Seeds cylindrical to ovoid, Most groundnut cultivars grown in West Africa
1-2 cm x 0.5—1 cm, with pointed or flattened belong to subsp. hypogaea; most of those in
ends, enclosed in a thin papery seed coat rang- East Africa to subsp. fastigiata. Subsp. hypo-
ing in colour from white to deep purple. Seed- gaea is mainly used for food, and subsp. fas-
ling with epigeal germination; cotyledons thick tigiata, which has a higher oil content, as a
and fleshy. source ofoil.
Other botanical information Arachis com- Growth and development Seeds of "Vir-
prises about 70 species, all distributed in South ginia' types have a dormancy period of 1-3
America. The centre of origin of Arachis is the months, whereas 'Spanish' and "Valencia' types
Mato Grosso region of Brazil.Arachis hypogaea are without dormancy. The optimum soil tem-
is by far the most economically important spe- perature for seed germination is 25—30°C. Low
cies in this genus, but several other species temperatures retard germination and devel-
have been cultivated for their seeds, including opment and increase the risk of seedling dis-
Arachis villosulicarpa Hoehne and Arachis ste- eases. Upon germination, the primary root
nosperma Krapov. & W.C.Greg. elongates rapidly, reaching 10-12 cm before
High levels of resistance to many diseases and lateral roots appear. As growth proceeds, the
pests of groundnut have been recorded in other outer layer of the primary root of a seedling is
Arachis species. Many of them are closely re- sloughed off so that root hairs do not form.
lated to groundnut and include the other 26 Branching is dimorphic, with vegetative
species in section Arachis. Several diploid spe- branches and reduced reproductive branches.
cies have been suggested as wild progenitors of Secondary and tertiary vegetative branches
groundnut, but molecular and cytogenetic stud- can develop from the primary vegetative
ies indicate that Arachis duranensis Krapov. & branches. Flowering may start as early as 20
W.C.Greg, and Arachis ipaensis Krapov. & days after planting, but 30-40 days after plant-
W.C.Greg, are most closely related to the pro- ing is more usual. The number of flowers pro-
genitors of allotetraploid domesticated ground- duced per day decreases as the seeds mature.
nut. Arachis monticola Krapov. &Rigoni is the Up to 50% of the embryos may abort even un-
only other tetraploid species in the section; it is der ideal environmental conditions, but this
very closely related to Arachis hypogaea and percentage becomes much higher during times
may be the direct descendant of the original of drought or other environmental stress. How-
hybrid between the 2 diploid progenitor spe- ever, plants can produce a 'second crop' of seeds
cies. if adequate moisture becomes available again.
Hybrids between Arachis hypogaea and other Groundnut is self-pollinating, but outcrossing
Arachis species have been produced by direct can occur when bees pollinate the flowers.
hybridization and by first creating auto- Groundnut generally produces more flowers
tetraploids or allotetraploids from the diploid under long day conditions, but reproductive
species before making crosses. Hybrids show efficiency is greater under short days. Only one
24 CEREALSANDPULSES

of the flowers in an inflorescence opens at a weight ranges from 150 g to more than 1300 g.
given time. Flowers wither within 24 hours Sowing high-quality seed in a well-prepared,
after anthesis. Fertilization usually occurs moist seedbed is essential for crop establish-
within 6 hours after pollination, when the ment. Groundnut seeds are often planted at a
basal part of the ovary starts elongating into a depth of 4-7 cm at a rate of 60-80 kg/ha.
structure called 'peg'. The embryo initiates a Groundnut pods intended for sowing are often
growth phase until it reaches an 8-16-cell hand-shelled 1-2 weeks before sowing. Only
stage. It then becomes quiescent during the 5 - fully mature pods are selected. Before sowing,
15 days required for the 'peg' to enter the soil. groundnut seed may be treated with a fungi-
The 'peg' stops elongating within a day or two cide to control seedling diseases. In general,
after soil penetration, the embryo then restart- early sowing improves yields and seed quality.
ing growth. In wild Arachis species the 'peg' Early sown crops also suffer less risk of disease
may continue to grow to a length ofnearly 2 m. such as groundnut rosette virus. However the
Seeds in 'Spanish'-type cultivars usually ma- appropriate sowing date depends on the matur-
ture within 90-120 days after planting, ity period of the cultivar. Small-seeded 'Span-
whereas 'Virginia'-type cultivars take 130 days ish' types are spaced at 60-75 cm between rows
or more. Pods of the same size may differ sig- and 10 cm within the row. This gives an opti-
nificantly in maturity and seed weight. mum plant population of 133,000-167,000
Groundnut is usually effectively nodulated by plants per ha. Large-seeded "Virginia' types are
N2-fixing Bradyrhizobium bacteria. Because spaced at 75 cm between rows and 15 cm
root hairs are absent, the bacteria infect the within the row, giving an optimum plant popu-
root through cracks in the epidermis near mul- lation of 89,000 plants per ha. Groundnut can
ticellular hairs at the basis ofthe root. be grown on the flat, or on ridges as is often the
Ecology The optimum mean daily tempera- case in Malawi. Groundnut grown on ridges
ture for groundnut growth is 27—30°C; growth tends to give higher yields, probably because of
ceases when temperature drops below 15°C. more loose soil favourable for pod development
Groundnut is mainly grown in areas with an and easier uprooting.
average annual rainfall of 500-1000 mm; 500- In tropical Africa groundnut is grown as a sole
600 mm of rain reasonably well distributed crop or intercropped between rows of cereals
over the growing season allows satisfactory such as maize, sorghum or pearl millet.
production. Nevertheless, groundnut is Management Groundnut does not compete
drought-tolerant and can withstand severe lack effectively with weeds, particularly in the early
of water, though yield is generally reduced. A stages of development. The crop should be
dry period is required for ripening and harvest- thoroughly weeded within the first 45 days.
ing. The phenology of groundnut is determined Once the development of the 'peg' begins,
primarily by temperature, with cool tempera- earthing-up is kept to a minimum. Weeds at
tures delaying flowering. In controlled envi- this stage are hand pulled. Pre-and post-
ronments, photoperiod has been shown to in- emergence herbicides may be used to eradicate
fluence the proportion of flowers producing weeds, but they are too expensive for most
pods and distribution of assimilates between small-scale farmers in Africa. In sound rotation
vegetative and reproductive structures (har- systems, groundnut benefits from residual fer-
vest index) in some cultivars. Long photo- tility; in general no additional fertilizer is given
periods (greater than 14 hours) generally in- if the crop is sown on a well-managed soil pre-
crease vegetative growth and short photo- viously treated with a balanced fertilizer. How-
periods (less than 10 hours) increase reproduc- ever, in order to ensure good crop establish-
tive growth. Groundnut can be grown up to ment, high yield and good seed quality, a fertil-
1500 m altitude. izer containing Ca, such as gypsum or single
The best soils for groundnut are deep (at least superphosphate, should be applied. Calcium is
30-40 cm), friable, well-drained sandy loams, absorbed directly by the pods if soil moisture is
well-supplied with calcium and a moderate adequate. A shortage of Ca in the zone where
amount of organic matter. It is important to the pods develop will result in empty pods,
maintain near to neutral soil pH levels and particularly in cultivars of the 'Virginia' type.
Ca:K ratios lower than 3. Groundnut is normally a rainfed crop, but it is
Propagation and planting Groundnut is grown under irrigation in Sudan.
propagated by seed, but vegetative propagation Groundnut should preferably not be grown in
using cuttings is possible. The 1000-seed the same field more than once in 3 years to
 ARACHIS 25

limit damage by soil-borne diseases, nematodes disease of groundnut leading to 30-100% yield
and weeds. It fits into a wide range of rotations loss. Early sowing at high plant populations
and it can follow any clean-weeded crop, e.g. controls the spread of groundnut rosette by
maize, sorghum, pearl millet, cassava, sweet giving complete soil coverage as quickly as
potato or sunflower. To reduce the incidence of possible and restricting the movement of
diseases and pests, groundnut should not be aphids. Cultivars resistant to groundnut ro-
sown after cotton or tobacco. Groundnut does sette are widely grown in Africa. In Malawi it
well on virgin land or immediately following a is common practice for farmers to interplant
grass ley or well-fertilized crop such as maize. groundnut and cowpea to control groundnut
The intensity of management of groundnut rosette. Aspergillus fungi can invade ground-
varies considerably around the world, depend- nut pods and seeds, producing toxic compounds
ing on the economic return for the crop or the known as aflatoxin. Contaminated produce can
role of groundnut in the farming system. In the be poisonous to people and livestock, and can-
United States, Australia and parts of South not be exported. Aflatoxin contamination also
America the crop is grown with intensive man- affects groundnut seed, leading to low germi-
agement, generally with high levels of me- nation percentage and poor seedling estab-
chanical and chemical inputs. In many coun- lishment. It can occur before harvest, during
tries groundnut is grown as a cash crop pri- field drying and curing, and in storage. Pre-
marily for export. harvest contamination is likely to be most seri-
Diseases and pests Groundnut is suscepti- ous under drought. Post-harvest contamination
ble to a number of diseases, such as early leaf occurs if groundnut pods or seeds become moist
spot (Cercospora arachidicola), late leaf spot and/or damaged. Various methods are used to
(Cercosporidium personatum, synonym: Cerco- control aflatoxin. They include avoiding me-
spora personata), rust (Puccinia arachidis), chanical damage to pods or seeds during weed-
groundnut rosette (caused by a complex of 3 ing, harvesting and storage, harvesting as soon
agents: groundnut rosette virus (GRV), ground- as the pods are mature, proper drying and cur-
nut rosette assistor virus (GRAV) and a satel- ing, and storing in the shell at low temperature
lite RNA) and aflatoxin contamination caused under moisture-free conditions.
by Aspergillus fungi. Foliar diseases of ground- Root-knot nematodes (Meloidogyne spp.) may
nut are among the most important yield- cause considerable yield loss in groundnut;
limiting factors in groundnut production. Early they can be controlled by crop rotation.
and late leaf spots and rust together may cause On a global scale the most important insect
up to 70% yield losses; even where fungicides pests include aphids (Aphis craccivora), thrips
are applied significant yield reductions occur. (Frankliniella spp.), jassids (Empoasca
Spraying with fungicide when the disease ap- dolichi), white grubs (larvae ofvarious beetles),
pears controls both leaf spots effectively. Dust- termites (mainly Microtermes sp.) and the red
ing groundnut leaves with sulphur, early in the tea bug Hilda patruelis. False wireworms and
morning when there is still dew on the leaves, millipedes seem to occur less frequently. In
has been reported to control both early and late general, soil pests cause more damage than
leaf spots. The use of sulphur has also been foliage feeders or sucking pests. However,
observed to increase leaf retention, thus in- aphids are particularly harmful because they
creasing the quantity of leafy stems available transmit groundnut rosette virus. In Asia and
for livestock feed. Cultural practices to control Africa white grubs, termites, millipedes and
leaf spots include crop rotation and burning of ants are important pests; in the United States
crop residues. Cultivars with partial resistance the lesser cornstalk borer (Elasmopalpus lig-
to leaf spots have been developed. Rust gener- nosellus) and the southern corn rootworm
ally occurs sporadically and at low severity, (Diabrotica undecimpunctata) are the main
although it can cause crop losses up to 40% insect pests of groundnut. Pests attacking
when an epidemic occurs. The cultural prac- stored groundnut pods and seeds include
tices and fungicidal control measures recom- bruchids (Caryedon serratus, Callosobruchus
mended for leaf spots are also applicable to spp., Acanthoscelides spp.) and flour beetles
rust. Resistant cultivars are available. Ground- (Tribolium spp.).
nut rosette virus, transmitted by the aphid Parasitic plants (Alectra vogelii Benth. and
Aphis craccivora, is endemic to sub-Saharan Striga spp.) are recorded as causing damage to
Africa and widely prevalent in Ghana, Nigeria, groundnut in various African countries.
Malawi and Zambia. It is the most destructive Harvesting The indeterminate flowering
26 CEREALS AND PULSES

pattern of groundnut makes proper timing of 850 kg/ha, which is only slightly higher than
harvest difficult, even though such timing is the average yield in the 1970s (730 kg/ha). Na-
crucial for obtaining maximum yield and qual- tional average yields of groundnut pods in
ity. Harvest at the proper time ensures that tropical Africa range from 300-1000 kg/ha.
the maximum number of pods have attained Average world yields of groundnut pods in-
their greatest weight and that pods are not creased from 0.9 t/ha in the 1970s to 1.4 t/ha in
falling off. Methods to determine the proper the early 2000s. With good management prac-
time for harvesting groundnut are available, tices and proper disease control, yields up to 5
but some are environment-specific or are pro- t/ha can be achieved. On average 100 kg of
hibitively expensive. Presently only the shell- pods yield 70 kg ofseeds, containing 35kg oil.
out method and the hull-scrape method are Handling after harvest Produce quality is
widely used for groundnut maturity determi- closely related to proper harvesting date, har-
nation. The shell-out method is based on colour vesting method and drying; every step is criti-
changes within the pod wall ( 'hull') that occur cal to obtaining or maintaining quality.
as the pod matures. The internal pod wall sur- Groundnut pods are dried to an average mois-
face of most cultivars changes from white to ture content of about 10%. Removing foreign
brown or black blotches covering a large per- materials early helps to maintain quality dur-
centage of the area. The colour of the seed coat ing storage. Cleaning equipment to remove the
changes from white to dark pink or tan at the foreign material has been developed and in-
same time. A sample of plants is taken and cludes sand screens and belt screens.
pods opened. The percentage of pods with dark Groundnut pods are stored in granaries, tanks,
colour inside the pod wall is determined. Har- bins, concrete silos, warehouses or in the open.
vesting should begin when the percentage is In storage, ventilation is crucial to prevent
60-80, but recommendations vary. The shell- moisture build up which can promote mould
out method is widely used because it can di- growth and aflatoxin production. Excessive
rectly be used in the field without further han- heat should be avoided. Storage structures
dling of pods, requires no equipment and pro- should be examined frequently for moisture
vides an immediate answer. The hull-scrape and insect problems as these can greatly re-
method, developed in the early 1990s, is cur- duce quality. Seeds can be protected from me-
rently accepted as the most accurate means of chanical damage by storage and transport in
assessing the maturity of 'runner'-type the pods. In many areas groundnut is only
groundnuts. The method is based on the fact shelled when it is to be used or sold; in local
that the pod mesocarp (the area just beneath markets unshelled pods are often offered for
the pale brown coloured exterior of the sale. Both mechanical and manual shelling are
groundnut pod) changes from white to yellow common.
to orange to brown to black as the crop ma- Groundnut removed from storage is trans-
tures. The method requires colour charts and a ported to shelling centres where the pods are
pocket knife to scrape the pod surface. graded, cleaned and shelled, and the seeds are
Harvesting is carried out manually in most separated into commercial grade sizes. Shelling
parts of Africa, as well as Asia. In the United operations may damage the seeds. Shelling of
States harvesting is normally done using a 100 kg of groundnut pods yields 60-80 kg of
digger shaker inverter. When plants are har- seeds. Generally groundnut seeds can be stored
vested manually, they are loosened with a hoe at 1-5°C and 50-70% relative humidity for 1
and pulled out of the ground, after which they year without loss of quality. Groundnut seeds
are turned to expose the pods to the sun to tend to absorb gases and off-flavours, which
facilitate drying. When dry, the pods are ripped should be avoided.
off the plants. With mechanical harvesting, the Oil is extracted from groundnut seed by expel-
plants are cleanly removed from the soil and ler pressing, hydraulic pressing, solvent extrac-
deposited in inverted windrows. Pods have to tion, or a combination of these methods. Expel-
remain in the windrows until the average ler pressing is most widely used.
moisture content is 18-24%. Pods are then Genetic resources The International Crops
picked using a combine. Rainfall during wind- Research Institute for the Semi-Arid Tropics
rowing may promote mould growth resulting in (ICRISAT), Patancheru, India, holds the larg-
reduced milling quality. est collection of groundnut types, with more
Yield In tropical Africa the average yield of than 15,000 accessions, differing for many
groundnut pods in the early 2000s was about vegetative, reproductive, physiological and
 ARACHIS 27

biochemical traits including their reactions to the close linkage of disease resistance genes
biotic and abiotic stresses. A duplicate sample with loci conferring undesirable pod and seed
is maintained in a regional gene bank at Nia- characteristics; the later maturity, lower parti-
mey, Niger. Other large collections of ground- tioning to seeds, and higher photoperiod-
nut germplasm are held in the United States sensitivity of disease-resistant germplasm com-
(Southern Regional Plant Introduction Station, pared to agronomically elite susceptible mate-
Griffin, Georgia, 9000 accessions), India (Na- rials; the large genotype x environment inter-
tional Research Centre for Groundnut (NRCG), actions for traits of economic importance; and
Junagadh, 8000 accessions), China (Institute of limited gene introgression from wild Arachis
Crop Germplasm Resources (CAAS), Beijing, species to cultivated groundnut.
5400 accessions; Institute of Oil Crops Re- Genetic linkage maps of groundnut have been
search, Wuhan, 5700 accessions). In tropical constructed using various markers, but the
Africa substantial groundnut germplasm col- saturation level is insufficient for routine ap-
lections are held in Senegal (Centre National plication in molecular breeding. An efficient
de Recherche Agronomique, Bambey, 900 ac- tissue culture and transformation system for
cessions), Uganda (Serere Agricultural and groundnut has been developed and transgenic
Animal Production Research Institute, Serere, groundnut plants have been produced using
900 accessions) and Malawi (Plant Genetic biolistic and Agrobacterium-meAiateA methods.
Resources Centre, Chitedze Agricultural Re- Prospects Groundnut remains an extremely
search Station, Lilongwe, 500 accessions). The useful crop, providing food, oil, fodder and fuel
ARC Grain Crops Institute in Potchefstroom, to households and is also an important source
South Africa, has a collection of 850 accessions. of additional income as a cash crop. Important
Core collections that have been developed are problems in groundnut cultivation in tropical
useful for developing models for future germ- Africa are low yields and its susceptibility to
plasm acquisition and evaluation for disease diseases. Many cultivars are still susceptible to
resistance. Additional collections are needed early and late leaf spot and rust, as resistance
for most groundnut-producing regions, as land- tends to be linked with long duration and un-
races in these areas are rapidly being replaced desirable pod and seed characteristics. There-
with modern cultivars. fore, the development of high-yielding cultivars
Breeding Groundnut breeding efforts great- with resistance to disease (especially leaf spots
ly increased when the ICRISAT groundnut and rust) and adaptation to African production
breeding programme was established in 1976. systems remains a major challenge for ground-
Diverse breeding populations are now being nut breeders. The application of DNA markers
tested in regional programmes in sub-Saharan may allow breeders to combine resistance to
Africa and Asia. Most breeding programmes biotic and abiotic stresses with improved pro-
are conducted by public institutions. Ground- ductivity and seed quality. The use of biotech-
nut breeding objectives have concentrated on nology tools will become increasingly important
adaptation to regional markets and production for large-scale germplasm characterization and
systems. All programmes aim at improving the resolving some of the constraints (e.g. disease
productivity of the crop and resistance to dis- problems) in groundnut production.
eases. Large-scale efforts to evaluate wild Ara- Major references Dwivedi et al., 2003;
chis germplasm have resulted in identification Knauft & Ozias-Akins, 1995; Knauft & Wynne,
of useful sources of resistance to many dis- 1995; Kokalis-Burelle et al. (Editors), 1997;
eases. Recently there have been initiatives to Krapovickas & Gregory, 1994; Melouk &
improve flavour and quality. Breeding for re- Shokes (Editors), 1995; Shorter & Patanothai,
sistance to aflatoxin contamination has re- 1989; Smartt (Editor), 1994; Stalker, 1997;
ceived increased attention, and the selection of Wynne, Beute &Nigam, 1991.
short-duration cultivars with drought resis- Other references Burkill, 1995; Clavel,
tance is a high priority in many programmes. 2002; Clavel & Gautreau, 1997; de Waele &
Commonly used breeding methods in ground- Swanevelder, 2001; Gillett et al., 1971; ILDIS,
nut are pedigree selection, bulk-pedigree selec- 2005; Isleib & Wynne, 1992; Kochert et al.,
tion and single-seed descent. Backcross breed- 1996; Lynch & Mack, 1995; McDonald et al.,
ing has not been used extensively as most of 1998; Neuwinger, 2000; Norden, Smith & Cor-
the economically important traits of groundnut bet, 1982; Popelka, Terryn & Higgins, 2004;
are quantitatively inherited. The major con- Purseglove, 1968; Sherwood et al., 1995; Singh,
straints to rapid genetic enhancement include: 1995; Singh & Nigam, 1997; Steinman, 1996;
28 CEREALS AND PULSES

USDA, 2004; Wynne &Gregory, 1981. ana (Malzev) Mordv., resulting in weedy hy-
Sources of illustration Shorter & Patan- brid swarms which shatter easily.
othai, 1989. Ecology Avena abyssinica is cultivated, but
Authors B.R. Ntare is also a weed of arable land, particularly in
barley and wheat fields. In Ethiopia it is found
at 1700-3000 m altitude. Experiments indicate
AVENAABYSSINICA Höchst. that Avena abyssinica is a long-day plant and
that vernalization results in earlier flowering.
Protologue Schimp, iter Abyss, sectio III No Management Avena abyssinica is recorded
1877 (1844). to be grown sometimes in Eritrea and Ethiopia,
Family Poaceae (Gramineae) but it is unclear to what extent this is still the
Chromosome number 2n = 28 case. As a weed in barley and wheat it is often
Vernacular names Ethiopian oat, Abyssin- tolerated and harvested with the main crop.
ian oat (En).Avoine d'Abyssinie (Fr). Avena abyssinica is affected by crown rust or
Origin and geographic distribution Avena leaf rust (Puccinia coronata f.sp. avenae). It is
abyssinica probably originated from Avena also susceptible to infestation with ergot
barbata Pott ex Link. It is native to Eritrea, (Claviceps spp.);consumption of infected grains
Ethiopia and Yemen, and is cultivated for its has led to outbreaks of ergotism in Ethiopia.
grain in northern Ethiopia. It has been tried as Genetic resources and breeding The larg-
a crop in Tanzania and Algeria. est germplasm collections of Avena abyssinica
Uses In Ethiopia the grain of Avena abys- are kept at in the United States (USDA-ARS
sinica is used mixed with barley to make pan- National Small Grains Germplasm Research
cake-like bread ('injera'), local beer ('tella') and Facility, Aberdeen, Idaho, 241 accessions), the
other products. The grain is also eaten roasted United Kingdom (John Innes Centre, Depart-
as a snack ('kollo'). Malt containing an admix- ment of Applied Genetics, Norwich, 65 acces-
ture of Avena abyssinica has been credited sions) and the Russian Federation (N.I. Vavilov
with giving better beer than malt from pure All-Russian Scientific Research Institute of
barley or wheat. Plant Industry, St. Petersburg, 53 accessions).
Botany Erect annual grass up to 1.5 m tall. No germplasm collections are known to exist in
Leaves alternate, simple; leaf sheath long and tropical Africa.
loose; ligule acute, membranous; blade linear, Prospects Avena abyssinica is a semi-
flat, usually glaucous. Inflorescence a terminal domesticated plant in Ethiopia, where it is
panicle 20-35 cm long, loose and open, the used as a component in mixtures for the prepa-
branches slightly rough. Spikelet slender- ration of food and local beer. It has, however,
stalked, pendulous, 2-2.5 cm long, 2-3- not become important and its present status is
flowered, with the uppermost floret reduced or uncertain.
vestigial, non-shattering; glumes almost equal, Major references Baum, 1977; Fröman &
narrowly elliptical, sharply acuminate, several- Persson, 1974; Hanelt & Institute of Plant Ge-
veined; lemma 1.5-2 cm long, smooth and gla- netics and Crop Plant Research (Editors),2001;
brous or with a few bristly hairs near the awn National Research Council, 1996; Phillips,
insertion or margin, narrowly bifid, each lobe 1995.
with 1 vein extended into an apical bristle 1—3 Other references Clayton, 1970; Engels,
mm long, usually also minutely toothed at the Hawkes & Worede (Editors), 1991; Harlan,
base of the bristle, with slender, abruptly bent 1989a; Harlan, 1989b; rung, 1979; Martens &
awn 2.5-3 cm long, arising from the back of the McKenzie, 1973; Sampson & Burrows, 1972;
lemma; palea almost as long as lemma, bifid, 2- Welch (Editor), 1995.
keeled, prickly hairy on the back; stamens 3; Authors M. Brink
ovary superior, villous, with 2 stigmas. Fruit a
caryopsis (grain).
Avena comprises about 30 species, which are AVENASATIVAL.
diploid (2n - 14), tetraploid (2n — 28) or hexap-
loid (2n — 42). The tetraploid Avena abyssinica Protologue Sp. pi. 1:79 (1753).
belongs to section Ethiopica. It can be distin- Family Poaceae (Gramineae)
guished from the common oat (Avena sativa L.) Chromosome number 2n =42
by the presence of two bristles at its lemma tip. Vernacular names Oat, oats, common oat
It crosses easily with the weed Avena vavilovi- (En).Avoine, avoine cultivée (Fr).Aveia, aveia-
 AVENA 29

amarela (Po). for traditional houses. Also in Kenya oat is

Origin and geographic distribution Avena used as food and as fodder. A field sown for
sativa is only known in cultivation and its ex- grain production can be used for grazing if
act origin is unclear. Oat was not cultivated as rains are inadequate; on the other hand fields
early as wheat and barley and probably it per- are sometimes grazed to delay grain develop-
sisted as a weed in fields of these cereals for ment.
centuries before it was taken into cultivation. In Australia oat is planted for sand binding on
Oat seeds have been found in 4000-year-old dunes. An important industrial utilization is
remains in Egypt, but these were probably the use of oat hulls for the production of fur-
from weeds and not from cultivated oat. The fural and other furan compounds, utilized in
oldest known cultivated oat remains were the production of fungicides, disinfectants and
found in caves in Switzerland that date back to preservatives. Oat products also find applica-
around 1000 BC. Avena sativa probably tion in the cosmetic industry as talc replace-
evolved in central or northern Europe from ments and in skin care products. Oat flour has
wild Avena sterilis L. germplasm from south- anti-oxidant properties and has been used for
western Asia. Nowadays oat is extensively cul- food conservation, but it has largely been re-
tivated in northern temperate regions, mainly placed by synthetic chemicals.
in Europe and North America. In tropical Af- Production and international trade Accord-
rica it is mainly grown in Ethiopia and Kenya. ing to FAO statistics the average world oat
It is also cultivated in South Africa, Morocco, grain production in 1999-2003 amounted to
Algeria and Tunisia. about 25.9 million t/year from 12.7 million ha.
Uses Oat has been used as food and fodder The main producing countries are the Russian
since ancient times. Oat grain is an ingredient Federation (5.8 million t/year in 1999-2003,
in a wide range of food products including from 3.8 million ha), Canada (3.3 million t/year
breakfast cereals, porridge, cookies, breads and from 1.4 million ha) and the United States (2.0
muffins, crackers and snacks, beverages, meat million t/year from 0.9 million ha). The average
extenders and baby foods. Oat grain is consid- oat grain production in sub-Sahara Africa in
ered to have potential as a source of good edible 1998-2003 has been estimated at 55,000 t/year
oil. In Ethiopia oat is made into 'injera' (pan- from 53,000 ha, almost entirely from Ethiopia
cake-like bread), 'tella' (local beer) and other (50,000 t/year from 49,000 ha) and Kenya
products. In industrialized countries oat grain (3500 t/year from 3400 ha) and small amounts
mainly finds application as animal feed, espe- from Zimbabwe.
cially for horses, but also for cattle, sheep, tur- Due to the decline of oat as animal feed, partly
keys and other animals. The green plant is as a result of the mechanization of agriculture
good forage; it makes good hay and silage or is and decreased importance of workhorses, world
grazed by animals. The straw too is used as production steadily declined from about 50
forage, e.g. in Ethiopia, where it also serves as million t/year (from about 35 million ha) in the
bedding for livestock, fuel and roofing material early 1960s to about 26 million t/year (from
about 13 million ha) in the early 2000s. In the
same period the production in sub-Sahara Af-
rica increased from about 20,000 t/year to
about 55,000 t/year.
The largest part of the oat production is con-
sumed locally, with about 2.5 million t/year
entering international trade in 1998-2002.
Canada (1.2 million t/year), Sweden (450,000
t/year) and Finland (360,000 t/year) are the
largest exporters; the United States (1.7 mil-
lion t/year) the largest importer. International
trade in oat in tropical Africa is insignificant.
Properties The composition of oat grain per
100 g edible portion is: water 8.2 g, energy
1628 kJ (389 kcal), protein 16.9 g, fat 6.9 g,
carbohydrate 66.3 g, dietary fibre 10.6 g, Ca 54
mg, Mg 177 mg, P 523 mg, Fe 4.7 mg, Zn 4.0
Avena sativa - planted mg, vitamin A 0 IU, thiamin 0.76 mg, ribofla-
30 CEREALS AND PULSES

vin 0.14 mg, niacin 0.96 mg, vitamin B6 0.12

mg, folate 56 |ig and ascorbic acid 0 mg. The
essential amino acid composition per 100 g
edible portion is: tryptophan 234 mg, lysine
701 mg, methionine 312 mg, phenylalanine 895
mg, threonine 575 mg, valine 937 mg, leucine
1284 mg and isoleucine 694 mg. The principal
fatty acids are per 100 g edible portion: linoleic
acid 2424 mg, oleic acid 2165 mg, palmitic acid
1034 mg and linolenic acid 111 mg (USDA,
2004). Compared to other cereals, oat has a
high protein content and a good amino acid
profile, with a high level of lysine. The fat con-
tent is also higher than that of other cereals,
with a high proportion of unsaturated fatty
acids. Starch contents of 43-61% have been
recorded. The amylose content of the starch is
11-34%. Starch granules are irregular to po-
lygonal in shape with an average diameter of
(3.8-)7.0-7.8(-10.5) urn.
The soluble fibre in oat bran is believed to re-
duce blood cholesterol in humans, due to the
presence of ß-glucan. Oat has shown hypogly-
caemic activity and beneficial effects on gastro-
intestinal functions. Oat bran seems to protect
against dental caries. Compounds contributing Avena sativa - 1,part of stem with leaf; 2, in-
to the antioxidant properties of oat flour in- florescence.
clude glyceryl esters of hydroxycinnamic acid, Redrawn and adapted byIskak Syamsudin
ferulic acid and caffeic acids. Oat seems to be
tolerated by most coeliac patients, although slender-stalked, pendulous, 1.5—3.5 cm long,
concerns remain. usually 2-3-flowered, with the uppermost flo-
The unhulled ground grain is highly acceptable rets reduced, non-shattering; glumes almost
for ruminants and horses. Hulled and ground equal, narrowly elliptical-oblong, sharply acute,
oat grain is usually fed to pigs and poultry. Oat several-veined; lemma 1-2.5 cm long, more or
green forage, hay and silage is highly palatable less truncate or minutely 2-4-toothed, awn
to ruminants. In Kenya the crude protein con- present or absent, glabrous or sparsely hairy
tent of oat plants (on dry matter basis) declined around the awn insertion; palea slightly
from 20.2% for 50 cm-tall plants to 8.1%at full shorter than lemma; stamens 3; ovary supe-
flowering, with the in-vitro protein digestibility rior, villous, with 2 laterally exserted stigmas.
declining from 84.9% to 46.7%. The crude fibre Fruit a caryopsis (grain), 0.5-1 cm long, nar-
content increased from 23.3% to 28.1%, the row, with nearly parallel sides, hairy, grooved
carbohydrate content from 42.0% to 56.0%, the lengthwise on the face, tightly enclosed by
ash content decreased from 11.5%to 5.4%, and lemma and palea.
the ether-extract from 3.7% to 2.4%. Straw in Other botanical information Avena com-
Kenya contained on dry matter basis: 5.3% prises about 30 species, which are diploid (2n=
crude protein, 38.0% crude fibre, 10.2% ash, 14), tetraploid (2ra - 28) or hexaploid (2n =42).
1.4% ether extract and 45.1% N-free extract. All hexaploid Avena species belong to section
Description Erect annual grass up to 2 m Avena. The hexaploids Avena sativa, Avena
tall, with a fibrous root system; stems (culms) byzantina C.Koch (red oat), Avena fatua L. and
solitary or tufted, smooth or scabrous beneath Avena sterilis L. are interfertile. Avena byzan-
the inflorescence. Leaves alternate, simple; leaf tina is closely related to Avena sativa and pos-
sheath long and loose, rounded on the back; sibly derived through selection from the latter.
ligule blunt, membranous, 3-5 mm long; blade Some authors include Avena byzantina in
linear, flat, 10-45 cm x 0.3-1.5(-2) cm. Inflo- Avena sativa. Avena byzantina is cultivated
rescence a terminal panicle 15—30(—40) cm mainly in southern Europe. In tropical Africa it
long, loose and open or contracted. Spikelet has been grown experimentally in Kenya and
 AVENA 31

has been recorded as a weed in Tanzania. It than 85% germination. Oat seeds kept under
has naturalized in South Africa, where it is natural cold conditions in the highlands of
found in disturbed locations and on roadsides. Ethiopia still germinated after 15 years of
Avena fatua and Avena sterilis are important storage. Under tropical highland conditions,
weeds of cereals, e.g. in Europe, Ethiopia and the seeds are broadcast or drilled (row spacing
Kenya, and differ from Avena sativa in their 15—20 cm) at a recommended rate of 60-120
shattering spikelets and hairy lemmas. The kg/ha, with drilled crops and crops intended for
tetraploid Avena abyssinica Höchst, can be grain production at the lower end of the range.
distinguished from Avena sativa by the two In Kenya oat is normally sown with a wheat
bristles at its lemma tip. drill in rows 20-25 cm apart, at a seed rate of
Avena sativa is variable, which is reflected in 60-80 kg/ha. In the high-altitude tropics, oat is
elaborate infraspecific classifications, mainly usually sown at the onset of the rainy season.
based on inflorescence and lemma characteris- When grown for forage, oat is sometimes grown
tics. mixed with vetches {Vicia spp.) or pea (Pisum
Growth and development Oat seeds start sativum L.).
to germinate 7 days after sowing. Seedlings Management The vigorous growth of oat
start tillering 35-45 days after sowing. Up to seedlings and the release of allelopathic com-
12leaves are produced per stem. The time from pounds depress weed growth. Hand-weeding
sowing to flowering depends on sowing time, (usually once) and application of broadleaf her-
e.g. in north-western Europe it is 100 days for bicides such as 2,4-D may be used for weed
spring-sown crops to 270 days for autumn- control. In Ethiopia farmers do not weed their
sown crops. Oat is largely self-pollinated with oat fields. Oat fields are seldom fertilized in
up to 1% outcrossing. The time from flowering tropical Africa, although the crop responds well
to harvesting is about 60 days in north-western to application of NPK. In the Ethiopian high-
Europe. The total crop duration is 3—6 months lands the general recommendation is to apply
in Ethiopia and Kenya, and 6-11 months in 18-23 kg N and 20-30 kg P per ha at sowing,
temperate regions. Shattered seeds remain and 35-46 kg N per ha top-dressing at tiller-
viable in the soil for a long time, which may ing. Oat is grown in rotation with barley,
result in weedy growth in subsequent crops. wheat, faba bean, pea and sometimes with
Ecology Oat is mostly grown under cool and fallow or a green manure. Allelopathic com-
moist conditions in cool-temperate regions, pounds can hinder the growth of subsequent
mainly as spring-sown and to some extent au- crops, if they are sown within about 3 weeks
tumn-sown crop. In tropical Africa it is mostly after the harvest of oat. In Kenya an oat crop
grown in mid- to high-altitude areas (1600- may be grazed 1—2times, before it is allowed to
3000 m altitude), with an annual rainfall over mature as a grain crop. Alternatives are 2—4
800 mm and minimum and maximum air tem- grazings of a crop in a season, 2 grazings fol-
peratures of 6°C and 24°C, respectively. In lowed by use as a hay crop, or 1—2 grazings
Ethiopia it is usually grown at 2700-3000 m followed by a hay crop and a grazing. Oat can
altitude. Oat is not as sensitive to frost as be grazed within 6—8 weeks after sowing.
wheat. When moisture is not limiting it also Diseases and pests Leaf (crown) rust (Puc-
performs well in warmer, humid mid-altitude cinia coronata f.sp. avenae) and stem rust
tropical environments. Oat needs more water (Puccinia graminis f.sp. avenae) are the most
than any other cereal except rice. It is gener- important diseases of oat. Systemic fungicides
ally a quantitative long-day plant, but differ- such as triazoles and morpholines are effective
ences in photoperiod-sensitivity exist among in controlling them, but this is seldom eco-
cultivars, with particularly strong responses in nomical. The use of cultivars resistant to rust
northern European cultivars. Vernalization is recommended. Septoria leaf spot (Septoria
responses have also been recorded. avenae), barley yellow dwarf virus (BYDV, also
Oat thrives on a wide range of soil types, as called 'red leaf), halo blight (Pseudomonas
long as drainage is sufficient. It grows on soils coronafaciens), loose smut (Ustilago avenae)
that are sandy, low in fertility, or highly acidic and covered smut (Ustilago hordei) are other
(as low as pH 4.5), but it performs best on well- common diseases of oat.
drained, fertile, loamy soils. Major pests include grasshoppers, army worms
Propagation and planting Oat is propa- and cut worms. Various aphid species are vec-
gated by seed. The 1000-seed weight is 22-37 tors of BYDV. At later stages of maturity birds
g. Seeds 2-3 months old normally have more and rats are important pests. Weevils (Sitophi-
32 CEREALS AND PULSES

lus granarius) and some other beetles attack pia. Oat seeds show orthodox seed storage be-
stored oat grain. haviour.
Harvesting In Africa oat is harvested Breeding The major objectives in oat breed-
manually by sickle or scythe, for forage nor- ing are improved grain and forage yields. The
mally after heading, and for grain when the development of cultivars resistant to fungal
seed is in the hard dough stage, which is nor- and viral diseases, especially crown and stem
mally at the end of the rainy season. The har- rusts, is important too. Sources of resistance to
vest is left in the field for sun-drying and is crown rust are found in wild Avena species,
subsequently threshed (grain crop) or piled especially Avena sterilis. Modern techniques of
(forage crop). A mechanized combine can be breeding have resulted in improved cultivars
utilized for harvesting large-scale grain oat with desirable traits such as resistance to dis-
crops or a mower for forage oat crops. When oat eases, high yield, huskless ('naked') grains,
straw is needed for roofing, the panicle is har- white-coloured large grain, and high contents
vested by sickle for grain, after which the re- of protein and oil in the grain. Molecular
maining stubble is harvested by sickle or marker maps have been constructed and a ge-
scythe at ground level. netic transformation system has been devel-
Yield The world average grain yield of oat is oped that allows the insertion of foreign genes
about 2t/ha with straw yields of about 5.5 t/ha. into oat using particle bombardment. In tropi-
As the result of threshing is not a naked grain, cal Africa there are small-scale breeding activi-
the hull (lemma and palea) generally accounts ties in Ethiopia and Kenya, mainly focusing on
for 25—35%of the total grain weight. The aver- resistance to diseases and increased forage and
age oat grain yield in Ethiopia and Kenya is grain yields. In Ethiopia and Kenya farmers
about 1 t/ha. When oat is harvested for green are mainly interested in dual-purpose culti-
fodder, hay or silage the dry matter yield is 4 - vars.
15 t/ha. Prospects Due to its tolerance to poor soil
Handling after harvest Oat grain should fertility and to frost, its low requirements of
be dried to a moisture content of 12—14% before external inputs such as fertilizers, and its dual-
storage, with a storage temperature below purpose character (food and fodder), oat has
20°C. In industrialized countries oat grain favourable prospects in the highlands of tropi-
processing generally involves cleaning, drying cal Africa, especially for resource-poor farmers.
(to partially inactivate lipolytic enzymes which On a worldwide scale, oat also has potential for
would result in rancidity), hulling, cutting, pharmaceutical and cosmetic uses.
steaming (to complete inactivation of lipolytic Major references Assefa et al., 2003; Baum,
enzymes) and flaking or milling. The cheapest 1977; Boonman, 1993; Coffman (Editor), 1961;
way of conserving oat forage is hay making. In Coffman, 1977;McMullen, 2000; Phillips, 1995;
areas where hay making is difficult, oat can be Suttie, 2004; Thomas, 1995; Welch (Editor),
made into silage, either alone or mixed with 1995.
legumes. Other references Clayton, 1970; Dougall,
Genetic resources Large Avena sativa germ- 1954; Feyissa, 2004; Frey, 1998; Fröman &
plasm collections are maintained in the United Persson, 1974; Gebrehiwot, 1981; Gibbs Rus-
States (National Small Grains Germplasm Re- sell et al., 1990; Gibson & Benson, 2002;
search Facility, USDA-ARS, Aberdeen, Idaho, Hanelt & Institute of Plant Genetics and Crop
10,000 accessions), the Russian Federation Plant Research (Editors), 2001; Hoover et a l ,
(N.I. Vavilov Ail-Russian Scientific Research 2003; Jellen & Beard, 2000; Jutzi & Grysels,
Institute of Plant Industry, St. Petersburg, 1984; Kassam et a l , 1991;Mailu, 1997; Mulat
8800 accessions), Canada (Soil and Crops Re- & Damesa, 1996; Peltonen-Sainio, 1998;
search and Development Centre, Sainte-Foy, Rogerson, 1956; USDA, 2004; Wight et a l ,
Quebec, 7500 accessions) and Kenya (National 2003; Zhou, Jellen &Murphy, 1999.
Genebank of Kenya, Crop Plant Genetic Re- Sources of illustration Hanelt & Institute
sources Centre, KARI, Kikuyu, 3700 acces- of Plant Genetics and Crop Plant Research
sions). A total of 656 accessions are kept at (Editors), 2001;Hegi, 1906.
ICARDA (Syria) and ILRI (Ethiopia). About Authors G. Assefa
835 oat accessions (mostly from Europe, United
States and Ethiopia) are available at the
EARO (Ethiopian Agricultural Research Or-
ganization) Holetta Research Centre in Ethio-
 BAUHINIA 33

B A U H I N I A PETERSIANA Bolle 2-lobed to one-third to two-thirds down, lobes

elliptical to ovate or rounded. Inflorescence an
Protologue Peters, Naturw. Reise Mossam- axillary, leaf-opposed or terminal raceme, 1—
bique 6(1): 24 (1861). 10-flowered. Flowers bisexual, almost regular,
Family Caesalpiniaceae (Leguminosae - Cae- 5-merous; hypanthium (1.5-)2-5.5(-6.5) cm
salpinioideae) long; sepals linear to linear-lanceolate, 1.5-5
Chromosome number 2n =28 cm long; petals narrowly elliptical to ovate, 2 -
Synonyms Bauhinia macrantha Oliv. (1871). 8.5 cm x 0.5-4 cm, white throughout or some-
Vernacular names Kalahari white bauhi- times base of midrib pink; fertile stamens (4-)
nia, wild coffee bean, coffee neat's foot, camel's 5(-6), slightly unequal in length, staminodes
foot (En). Bauhinia blanc du Kalahari (Fr). 4-5; ovary superior, slender, hairy, style 2-4
Chingando (Po). cm long. Fruit a linear-oblong to oblanceolate-
Origin and geographic distribution Bau- oblong pod 10-24 cm x 1.5-5 cm, woody, dehis-
hinia petersiana occurs in south-eastern DR cent, 5-6-seeded. Seeds 1-3 cm x 0.5-2 cm,
Congo and Tanzania, and throughout southern deep chestnut-brown to blackish.
Africa. Bauhinia is a widespread tropical genus with
Uses The meal of pounded seeds of Bauhinia about 250 species. In Bauhinia petersiana 2
petersiana is eaten. The seeds are also eaten as subspecies are distinguished. Subsp. petersiana
nuts after roasting and are considered a deli- has 2—10-flowered inflorescences, appressed
cacy in parts of Botswana. Roasted and ground hairs at the lower side of the leaves, and it is
seeds are used as a substitute for coffee. Un- distributed in the more eastern and northern
ripe seeds can also be eaten. The pods are parts ofthe area of the species. Subsp. macran-
eaten either roasted (Namibia) or boiled (Zam- tha (Oliv.) Brummitt & J.H.Ross has l-3(-4)-
bia). Seed oil is extracted in Botswana for local flowered inflorescences and curved or spread-
use. In DR Congo the bark fibres are used to ing hairs at the lower side of the leaves, and it
make rope and the roots to produce a dye. is found from southern Zambia and western
Bauhinia petersiana is widely browsed by live- Zimbabwe towards the south and west.
stock. In Zimbabwe, South Africa and the Bauhinia petersiana does not possess root nod-
United States it is grown as an ornamental ules and relies on soil nitrogen.
shrub. In much of its area of distribution the Ecology Bauhinia petersiana is found in
leaves of Bauhinia petersiana are boiled, the open grassland, wooded grassland and wood-
steam inhaled and the cooled-down liquid land. In East Africa it is found at altitudes of
drunk to cure common cough. The Shona peo- 150—1850 m. In southern Africa it is found in
ple of Zimbabwe take an infusion of the roots to dry localities, e.g. in the Kalahari with an an-
treat dysmenorrhoea and female infertility. In nual rainfall of about 350 mm only, and it tol-
South Africa the pounded leaves mixed with erates frost.
salt are boiled and the warm liquid is sprinkled Management In the Kalahari the seeds of
on wounds to promote healing. A decoction of Bauhinia petersiana are harvested from April
the macerated roots is drunk as a remedy for to July. For use as an ornamental Bauhinia
diarrhoea. petersiana is propagated by seed, cuttings or
Properties Dry seeds of Bauhinia petersi- layering. The 100-seed weight is about 670 g.
ana contain per 100 g: water 6.8 g, energy 1554 Genetic resources and breeding There
kJ (371 kcal), protein 22.9 g, fat 13.1 g, carbo- are indications that Bauhinia petersiana has
hydrate 40.2 g, fibre 13.0 g, Ca 237 mg, P 317 disappeared completely from communal graz-
mg, Fe 3.9 mg, thiamin 0.58 mg, riboflavin 0.2 ing land in southern Botswana, possibly as a
mg and niacin 1.6 mg (Arnold, Wells & Weh- result of increased grazing pressure. Two ac-
meyer, 1985). The principal fatty acids in the cessions from Botswana are stored in the Mil-
seed oil are linoleic acid (45%), oleic acid (26%), lenium Seedbank (Ardingly, West Sussex,
palmitic acid (16%) and stearic acid (7%). The United Kingdom), a single accession from Zim-
roots and leaves contain tannins. babwe is kept by the Desert Legume Pro-
Botany Shrub or small tree up to 10 m tall; gramme in the United States.
young branchlets pubescent and with many Prospects Although Bauhinia petersiana
small orange glands or scales, some branchlets has been considered a candidate for cultivation
coiled apically, tendril-like. Leaves alternate, as a food crop for a long time, no attempts have
simple; stipules 3-5 mm x 1-2 mm, deciduous; been made to domesticate the species, nor to
petiole 0.5-3 cm long; blade 2-8 cm x 2-10 cm, exploit or even explore its genetic variation.
34 CEREALS AND PULSES

Efforts to conserve the southern populations, in to 15 mm long stalked, broadly elliptical, 2-3.5
situ or ex situ, are urgently needed to avoid mm long, glabrous to pubescent, acute, 2-
loss of variation. flowered with lower floret male or sterile and
Major references Arnold, Wells & Weh- upper bisexual; lower glume up to half as long
meyer, 1985; Coates Palgrave, 1983;Gelfand et as spikelet, upper glume as long as spikelet,
al., 1985;Leger, 1997;von Koenen, 2001. membranous, 7-veined; lemma of lower floret
Other references Brenan, 1967; Brummitt membranous, lemma of upper floret wrinkled
& Ross, 1982; Dakora, Lawlor & Sibuga, 1999; and acute; palea ofupper floret obtuse to acute;
Ketshajwang, Holmback & Yeboah, 1998; Na- stamens 3; ovary superior, with 2 stigmas.
tional Academy of Sciences, 1979; Neuwinger, Fruit a caryopsis (grain), ellipsoid, compressed.
2000; Ross, 1977; Story, 1958; van Wyk & Brachiaria comprises about 100 species dis-
Gericke, 2000;Watt &Breyer-Brandwijk, 1962. tributed in the tropics and subtropics, mainly
Authors C.H. Bosch in the Old World. It has been proposed that
Brachiaria be nearly completely reduced to
Urochloa. Brachiaria deflexa is usually easily
BRACHIARIA DEFLEXA (Schumach.) distinguishable from other Brachiaria species
C.E.Hubb. ex Robyns by its panicle-like inflorescence, which resem-
bles that of Panicum spp. It intergrades with
Protologue Bull. Jard. Bot. Etat. 9(3): 181 Brachiaria ramosa (L.) Stapf and is sometimes
(1932). included in the latter. Guinea millet is often
Family Poaceae (Gramineae) confused with fonio (Digitaria exilis (Kippist)
Chromosome number 2n = 18, 36 Stapf). Compared to fonio, it has larger grains
Synonyms Pseudobrachiaria deflexa (Schu- and it grows faster, but it requires higher soil
mach.) Launert (1970). fertility and better drainage. The cultivated
Vernacular names Guinea millet, animal type sown in the Fouta Djallon Highlands
fonio, false signal grass (En). Fonio à grosses (called var. sativa Portères) differs from the
graines, gros fonio, millet de Guinée, kolo rassé wild types harvested elsewhere particularly by
(Fr). Jégé (Po). being totally glabrous and by having a
Origin and geographic distribution Guinea branched stem and much larger grains; fur-
millet is a semi-domesticated weed of the Afri- thermore, it is non-shattering.
can savanna. It is found from Cape Verde and Some Guinea millet types mature in as little as
Senegal eastward to Ethiopia, Eritrea and So- 70-75 days, but most types take 90-130 days
malia and southward to South Africa; it also to reach maturity. Guinea millet follows the C4-
occurs in western Asia to Pakistan and India. cycle photosynthetic pathway.
Uses Guinea millet is considered to belong to Ecology Guinea millet is found from sea-
the 'kreb' grasses, a group of grasses occurring level up to 1500 m altitude in open woodland,
in the Sahel region and collected for human forest margins and as a weed of cultivated land
consumption, especially in time of food short- and disturbed soils, often preferring slightly
age. In the Fouta Djallon Highlands on the shady locations. It is considered drought-
Guinea-Mali border the grain of a cultivated resistant. Brachiaria deflexa needs fertile and
type is ground into flour used to make cakes well-drained soils for optimum growth.
and fritters. Guinea grass provides excellent Management Guinea millet is mostly col-
forage. lected from the wild, but farmers sometimes
Properties Guinea millet has soft grains encourage its invasion into cereal fields and it
that are easily ground into flour. is sown as a cereal in the Fouta Djallon High-
Botany Annual grass up to 70(-100) cm tall; lands. Farmers also sometimes sow fast-
stems (culms) solitary or tufted, slender, often maturing Guinea millet types to fill in gaps in
weak and ascending. Leaves alternate, simple a field sown with fonio, sorghum, maize or
and entire; leaf sheath pale, striate, finely pu- other cereals.
bescent; ligule ciliate; blade broadly linear to Genetic resources and breeding Guinea
narrowly lanceolate, 4-25 cm x 0.5-2.5 cm, millet collections are held at CIAT (Centro
velvety pubescent. Inflorescence panicle-like, Internacional de Agricultura Tropical, Cali,
composed of 5-15 racemes borne on an axis 6 - Colombia, 16 accessions) and in Kenya (Na-
15 cm long; racemes distant, widely spreading, tional Genebank of Kenya, Crop Plant Genetic
2-10 cm long, often with side-branches, bear- Resources Centre, Kikuyu, Muguga, 5 acces-
ing mostly paired distant spikelets. Spikelet up sions). In view of its wide distribution, Guinea
 CAJANUS 35

millet seems not threatened by genetic erosion.

The Guinea millet cultivar sown in the Fouta
Djallon Highlands may have potential for fur-
ther selection.
Prospects Too little is known about Guinea
millet to make an accurate assessment of its
potential as a food plant. More information is
needed on its nutritional properties, agronomy,
ecological requirements and genetic diversity.
The wild type will remain a valuable fodder
plant for dry regions, due to its drought resis-
tance and excellent fodder characteristics.
Major references Burkill, 1994; Clayton &
Renvoize, 1982; Gibbs Russell et al., 1990; Na-
tional Research Council, 1996;Portères, 1976.
Other references Basappa, Muniyamma &
Chinnappa, 1987; Baudet, 1981;Clayton, 1972;
Clayton, 1989; Cope, 1995; de Wet, 1995c; Cajanus cajan - planted
Froman &Persson, 1974; Hanelt &Institute of
Plant Genetics and Crop Plant Research (Edi- dried, hulled and split seeds), and this use is
tors), 2001; Phillips, 1995; van der Hoek & carried on by Indian communities in Africa.
Jansen, 1996a. The use of immature pigeon pea seeds and
Authors M. Brink pods as a vegetable in soups and sauces is
common in many African countries. Canning
and freezing of the green seed is mainly done
CAJANUS CAJAN (L.) Millsp. in Central America. In Asia pigeon pea may be
used instead of soya bean to make tempeh or
Protologue Publ. Field Columbian Mus., tofu.
Bot. Ser. 2(1): 53 (1900). Vegetative parts are excellent fodder and seeds
Family Papilionaceae (Leguminosae - Papil- are also used as animal feed. The by-products
ionoideae, Fabaceace) of dhal production (seed coat and broken coty-
Chromosome number In — 22 ledons) are used as cattle and poultry feed in
Synonyms Cajanus indicus Spreng. (1826). India, and also in Kenya and Malawi.
Vernacular names Pigeon pea, Congo pea, Pigeon pea is useful in hedges and windbreaks
red gram (En). Pois cajan, pois d'Angole, am- on dry soils and in agroforestry (e.g. in alley
brevade (Fr). Ervilha do Congo, feijào guandu, cropping systems, where it is pruned to supply
ervilha de Angola (Po).Mbaazi (Sw). green manure). It is also grown as a shade
Origin and geographic distribution Pigeon crop, cover crop, or as support for vanilla. Pi-
pea originated in India, where it has been geon pea improves the soil through its exten-
grown for thousands of years. It reached Africa sive root system, nitrogen fixation and the
about 2000 BC or earlier, and a secondary cen- mulch provided by the fallen leaves. It serves
tre of diversity developed in East Africa. With as a host for silkworm (Madagascar) and the
the conquests and slave trade it reached the lac insect. Stems and branches, especially
Americas probably via both the Atlantic and those of medium- and long-duration cultivars,
the Pacific. Nowadays it is grown all over the are used for basketry, thatching, fencing and
tropics, but is most important in the Indian as fuel. In Nigeria the stems serve as stakes for
subcontinent and East Africa. It is not known yam. Pigeon pea finds wide application in tra-
in the wild, but often occurs naturalized as an ditional medicine. Diarrhoea, gonorrhoea, mea-
escape from cultivation. sles, burns, eye infections, earache, sore throat,
Uses In Africa dry pigeon pea seeds are of- sore gums, toothache, anaemia, intestinal
ten used for sauces accompanying staple food worms, dizziness and epilepsy are treated with
preparations such as cassava, yam and rice. leaf preparations. Root preparations are taken
Ripe seeds are eaten fried or boiled, often after to treat cough, stomach problems and syphilis.
being soaked first, or boiled into porridge. In Stem ash is applied on wounds, and stalks and
the Indian subcontinent pigeon pea is mainly roots are chewed against toothache. Powdered
used as a pulse, in the form of 'dhal' (soaked, seeds serve as a poultice on swellings. In
36 CEREALS AND PULSES

Madagascar the leaves are used to clean teeth. in Nigerian pigeon pea seed include trypsin
Production and international trade Accord- inhibitor activity, tannins and phytate.
ing to FAO statistics the world pigeon pea pro- The composition of raw immature seeds per
duction in 1999-2003 amounted to 3.1 million 100 g edible portion is: water 65.9 g, energy
t/year from 4.3 million ha; the main producing 569 kJ (136 kcal), protein 7.2 g, fat 1.6 g, car-
country is India (2.5 million t from 3.4 million bohydrate 23.9 g, dietary fibre 5.1 g, Ca 42 mg,
ha). The main producers in tropical Africa in Mg 68 mg, P 127 mg, Fe 1.6 mg, Zn 1.0 mg,
1999-2003 were Malawi (79,000 t from 123,000 vitamin A 67 IU, thiamin 0.40 mg, riboflavin
ha), Uganda (78,000 t from 78,000 ha), Kenya 0.17 mg, niacin 2.2 mg, folate 173 |Jg and
(59,000 t from 152,000 ha) and Tanzania ascorbic acid 39 mg (USDA, 2004). The leaves
(47,000 t from 66,000 ha). The annual produc- contain 15-24% crude protein. Extracts of pi-
tion in Mozambique is estimated at 40,000 t. geon pea seeds have shown antisickling action
Worldwide the area under pigeon pea has in- on red blood cells. This activity has been re-
creased steadily from about 2.8 million ha in lated to the presence of phenylalanine and hy-
the early 1960s to about 4.3 million ha at pre- droxybenzoic acid; related compounds have an
sent, whereas the production increased from even more pronounced effect.
1.5-2 million t to around 3 million t in the Description Erect shrub or subshrub, but
same period. regularly grown as an annual, up to 4 m tall,
Pigeon pea is mostly consumed locally, with with roots up to 2 m deep; stem erect, ribbed,
limited amounts entering international trade, up to 15 cm in diameter; branches many, slen-
and trade statistics are hardly available. Occa- der. Leaves alternate, 3-foliolate; stipules lin-
sional export demand may boost cultivation. In ear, 2-4 mm long; petiole (l-)1.5-6(-8) cm
Malawi and Kenya an estimated 65% of the long, grooved above; rachis 0.5-3 cm long,
pigeon pea production in 1996-1998 was con- slightly winged; stipels filiform, 1-4 mm long;
sumed on-farm, 10% traded on the domestic petiolules 1-4 mm long; leaflets elliptical to
market, and 25%exported. For Tanzania these lanceolate, 2.5-13.5 cm x 1-5.5 cm, acute, cov-
amounts were 35%, 10%and 55%, respectively. ered with small yellow glands, green above,
By far the most important export market is
India, followed by the Middle East. In the In-
dian market African pigeon pea has to compete
mainly with pigeon pea from Myanmar and pea
from Canada and France. Pigeon pea is widely,
but often informally, traded within Africa, for
instance between Mozambique and Malawi and
between Tanzania and Kenya. In northern
Tanzania most of the pigeon pea produced is
sold in Kenya, where it is very popular among
the Indian community.
Properties The composition of raw mature
pigeon pea seeds per 100 g edible portion is:
water 10.6 g, energy 1435 kJ (343kcal), protein
21.7 g, fat 1.5 g, carbohydrate 62.8 g, dietary
fibre 15.0 g, Ca 130 mg, Mg 183 mg, P 367 mg,
Fe 5.2 mg, Zn 2.8 mg, vitamin A 28 IU, thiamin
0.64 mg, riboflavin 0.19 mg, niacin 3.0 mg,
vitamin Ik 0.28 mg, folate 456 |Jg and ascorbic
acid 0 mg. The essential amino acid composi-
tion per 100 g edible portion is: tryptophan 212
mg, lysine 1521 mg, methionine 243 mg,
phenylalanine 1858 mg, threonine 767 mg,
valine 937 mg, leucine 1549 mg and isoleucine
785 mg. The principal fatty acids per 100 g
edible portion are: linoleic acid 778 mg and Cajanus cajan - 1,part of flowering and fruit-
palmitic acid 307 mg (USDA, 2004). Methion- ing branch; 2,seed.
ine is the limiting amino acid, followed by tryp- Redrawn and adapted by Achmad Satiri Nur-
tophan and threonine. Antinutritional factors haman
 CAJANUS 37

silvery grey-green beneath. Inflorescence an nating insects. Pigeon pea roots are nodulated
axillary false raceme 3.5-12 cm long; peduncle and fix nitrogen in association with Brady-
up to 8 cm long; bracts deciduous, ovate, c. 8 rhizobium and Rhizobium strains.
mm x 5 mm, acute. Flowers bisexual, papil- Ecology Pigeon pea is grown in the tropics
ionaceous; pedicel 0.5-1.5 cm long; calyx cam- and subtropics between 30°N and 30°S lati-
panulate, yellowish velvety and glandular, tudes. Optimum average temperatures range
tube (3-)4-5(-6) mm long, lobes 3-5(-7) mm from 18-29°C; frost is not tolerated. Above
long; corolla yellow or cream, standard almost 29°C, soil moisture and fertility may be limit-
round, 12-22 mm in diameter, dorsally yellow- ing. The optimum annual rainfall is 600-1000
red, orange or purple, wings obovate, 15—20 mm, but pigeon pea is tolerant to drought and
mm x 6—7 mm, yellow, clawed, keel petals 14- can be grown in areas with less than 600 mm
17 mm x 5-7 mm, yellow-green, clawed; sta- rainfall. It also grows in regions with an an-
mens 10, 9 fused and 1 free; ovary superior, 1- nual rainfall of over 2500 mm. Flowering is
celled, sessile, style curved. Fruit a straight or accelerated by short days; there are very few
sickle-shaped pod 2-10(-13) cm x (0.5-)l-1.5 truly day-neutral types. In Africa pigeon pea is
cm, hairy, glandular-punctate, splitting into 2 grown at altitudes up to 2000(-2400) m. It can
spiralling valves, septate between the seeds, be grown on a wide range of soil types, but wa-
(2-)4—9-seeded. Seeds globose to ellipsoid or terlogging is harmful. Drained soils of inter-
squarish, 4—9mm x 3-8 mm x 3-6 mm, white, mediate water-holding capacity and with pH
cream, brown, purplish to almost black, plain 5-7 are favourable. A soil salinity of 6-12 dS/m
or mottled. Seedling with hypogeal germina- is tolerated by many cultivars.
tion; first leaves simple. Propagation and planting Pigeon pea is
Other botanical information Cajanus com- propagated by seed. Stem cuttings rarely suc-
prises 34 species. Two wild Cajanus species are ceed. Longevity of seeds depends on storage
known in Africa: Cajanus kerstingii Harms conditions; in gene banks at low temperature
from West Africa, and Cajanus scarabaeoides and moisture the seed survives for decades.
(L.) Thouars occurring along the coasts of Af- The 1000-seed weight is 30-280 g; in Africa
rica and Madagascar, and at some locations large-seeded cultivars are the commonest.
more inland. The former does not cross with Planting arrangements vary widely, and seeds
Cajanus cajan, but the latter can produce hy- may be broadcast or sown in rows with plant
brids with it; spontaneous hybrids are known spacings of 40-200 cm x 20-180 cm. Seedlings
but rare. Although the use of Cajanus kerst- are difficult to transplant. In Africa and India
ingii as human food and animal feed seems not pigeon pea is often grown in intercropping sys-
recorded, it could be of value in ways similar to tems, usually with cereals, but also with cas-
pigeon pea. In Senegal the branches of Cajanus sava and cotton. It fits well in intercropping
kerstingii are used for making temporary hut systems because its slow initial growth reduces
walls. Other relatives of pigeon pea are found competition for the associated crop and its late
in Asia and Australia. maturity spreads labour requirements at har-
In India 10 maturity groups are distinguished vest time. After harvest of the intercrop, long-
in pigeon pea, usually combined into four cate- duration pigeon pea continues to grow and to
gories: extra early, early, medium and late- produce seed and to protect the soil. Pigeon pea
maturing cultivars (120, 145, 185, more than performs well when grown in single rows al-
200 days after sowing, respectively). ternating with 2 rows of cereals (e.g. sorghum,
Growth and development Pigeon pea seeds millets), cotton or groundnut. In Uganda and
germinate at temperatures of 19-43°C, but Mauritius, it is generally planted as a restora-
most rapidly at 20-30°C. Emergence is com- tive crop towards the end of a rotation cycle.
plete 2-3 weeks after sowing. Vegetative de- In vitro cultures have been initiated success-
velopment starts slowly, but after 2-3 months fully from different tissue sources, including
growth accelerates. Flowering (of 50% of the leaves, shoots and roots, and organogenesis as
plants) starts 56-210 days after sowing; seed well as somatic embryogenesis is possible. Re-
maturity normally ranges from 95-260 days. In generated plants have also been obtained via
humid areas, flowering and fruiting may con- callus and by direct differentiation from leaves.
tinue throughout the year. The flower struc- Management As a field crop, pigeon pea
ture of pigeon pea favours self-pollination, but may be typified as rather primitive; the tall
up to 82% out-crossing has been recorded, de- genotypes in particular are quite cumbersome
pending on the presence and activity of polli- in cultivation. Weed control is necessary be-
38 CEREALS AND PULSES

cause of the slow initial growth. Response to pod-sucking bugs (mainly Clavigralla spp.),
fertilizers is rarely economic; a phosphate pod borers (including Helicoverpa armigera
dressing is generally recommended at 20-100 and Maruca vitrata, synonym: Maruca testula-
kg/ha. In tropical Africa fertilizer application to lis) and the pigeon pea pod fly (Melanagromyza
pigeon pea is not common. Residual nitrogen chalcosoma). The use of insecticides is recom-
after a crop of pigeon pea can be about 40 mended, but chemical control is cumbersome
kg/ha. In India the nutrient uptake of a pigeon and expensive in tall indeterminate forms.
pea crop yielding 1.2 t seeds and 6.3 t straw Pigeon pea lines have been recorded with resis-
per ha has been calculated as 85 kg N, 8 kg P, tance to either or both Helicoverpa armigera
16 kg K, 23 kg Ca, 15 kg Mg, 9 kg S, 38 g Zn, and Melanagromyza chalcosoma, and to Ma-
31 g Cu, 128 g Mn and 1440 g Fe per ha. Wind ruca vitrata, but resistance has not been incor-
may bend the plants but staking is not prac- porated into cultivars that are acceptable to
tised. Irrigation as a lifesaver can be economic; farmers with respect to taste, seed colour and
in intensive cropping of short-duration culti- size. Integrated pest management (IPM), com-
vars, irrigation may be required. Pigeon pea is prising the judicious application of insecticides,
also grown as a ratoon crop, e.g. in Central and the use of tolerant or resistant cultivars, agro-
East Africa. For fast regrowth, the pruning nomic practices (planting date) and biological
height should not be lower than 50 cm. Part of control with natural enemies, is receiving at-
the shoots may be removed to reduce competi- tention. Because of its long flowering period,
tion. damage by pests such as Helicoverpa armigera
Diseases and pests The most important and other borers, and Agromyza fruit flies, may
fungal diseases of pigeon pea in tropical Africa be compensated for by new flushes. Bruchids
are leaf spot (Mycovellosiella cajani, synonym: (Callosobruchus spp.) are important pests of
Cercospora cajani), Fusarium wilt (Fusarium pigeon pea, infecting pods in the field as well
udum) and powdery mildew (Leveillula tau- as stored seeds. Seed storage in clean bins and
rica). Leaf spot is not important in drier areas, in the form of split seeds reduces bruchid at-
but can cause serious losses in humid regions. tack. Other control measures include sun-
It can be controlled by periodic sprays of fungi- drying, treatment with insecticides and storage
cides such as benomyl and mancozeb and by in admixture with ash and various plant prod-
the use of disease-free seed and the selection of ucts, such as tobacco and neem (Azadirachta
fields away from perennial pigeon pea, which indica A.Juss.) extracts or leaves.
may act as a source of inoculum. Pigeon pea Harvesting Immature pigeon pea pods are
lines with resistance to leaf spot have been picked over a long period of time in home gar-
identified. Recommended control measures a- dens or hedge crops, as is usual in Africa. For
gainst seed-borne and soil-borne Fusarium wilt ripe seeds the crop is usually cut near the
are intercropping and crop rotation with cere- ground when most pods are mature; many
als, fallow, removal of diseased plants, seed leaves are still green at that stage. Alterna-
treatment with fungicides, and the use of dis- tively, the ripe pods are picked from the stand-
ease-free seed and fields, but the best strategy ing crop, sometimes in several rounds as the
is the use of resistant cultivars. Moderately crop often matures unevenly. Mechanical har-
resistant lines in all maturity groups are avail- vesting of ripe pods is possible with combine-
able. Fusarium wilt is generally more severe in harvesters, but only for short-statured culti-
ratoon crops, from the second year onwards. vars maturing uniformly with pods at a uni-
Suggested control measures against powdery form level above the soil.
mildew include the use of fungicides and the Yield In Africa pigeon pea seed yield aver-
selection of fields not near perennial pigeon ages around 600 kg/ha with traditional land-
pea; resistant lines have been identified. Pi- races in mixed intercropping. For Uganda 1000
geon pea sterility mosaic virus (PPSMV) is the kg/ha has been recorded. Under optimum con-
most important disease of pigeon pea in India, ditions in sole cropping, yields of more than
but it appears to be restricted to Asia. Pigeon 5000 kg/ha are possible. Low yields may be
pea is susceptible to root-knot nematodes (Me- partly due to the fact that a considerable part
loidogyne spp.) and reniform nematodes (Roty- of the seeds is harvested and eaten before ma-
lenchus spp.). Resistant lines have been identi- turity. Forage yields range from 3—8 t/ha, but
fied in India. in experiments 50 t/ha have been obtained.
Insect pests are important in all pigeon pea- Fuel yields are usually 7-10 t/ha, but yields up
growing areas. The most important ones are to 30 t/ha have been recorded.
 CAJANUS 39

Handling after harvest Entire air-dried environment interaction. Genetic male sterility
pigeon pea plants are threshed, usually by is available in Cajanus cajanifolius (Haines)
hand or with cattle, and seed is cleaned. Maesen and is now used in hybrid breeding
Shelled fresh peas are sold on markets as a programmes. However, the production costs of
vegetable. Processing includes dhal-making, hybrid seeds are high, and a search is going on
either wet, after sprinkling heaps of seed, or for cytoplasmic male sterility. Short-duration
dry, by milling. Indian cultivars include 'Prabhat', 'T21', 'UPAS-
Genetic resources The world pigeon pea 120'; medium-duration cultivars are 'C 11', 'BDN-
germplasm collection covers India and several 1', 'Pusa Ageti', 'Sharda' and several 'ICP' lines
African countries, and some Caribbean islands. developed by ICRISAT. Hybrid cultivars are
More than 13,000 samples of Cajanus cajan also available. Improved cultivars in Kenya
are available in the collection of the Interna- include 'NPP 670', 'KAT 60/8' (both developed
tional Crops Research Institute for the Semi- in Kenya) and 'ICPL 87091'. However, they are
Arid Tropics (ICRISAT, Patancheru, India), more susceptible to insect pests than the local
and various breeders and institutes have parts landraces, due to their determinate growth
of this collection. Some 18wild Cajanus species habit and the fact that they start flowering
and at least 39 other species of the subtribe when pest populations are high. Furthermore,
Cajaninae are represented. Related perennial the seeds of 'KAT 60/8' and 'ICPL 87091' are
species are regularly rejuvenated in ICRISAT's relatively small, making them less wanted on
Botanical Garden. Attempts are continuing to the local markets than the larger seeds from
cover all taxa and areas of occurrence. In tropi- landraces. The Fusarium wilt-resistant culti-
cal Africa the National Genebank of Kenya in var TCP 9145', derived from a landrace col-
Kikuyu has a collection of more than 1200 ac- lected in Kenya by ICRISAT, has been success-
cessions of pigeon pea. Pigeon pea has orthodox fully released in Malawi. 'ICPL 87' and 'ICPL
seed storage behaviour. 146' lines are recommended as sole-cropping,
Breeding Pigeon pea breeding work started multiple-harvest cultivars for Tanzania, Ma-
in India in the early 20 th century, and mainly lawi and Zimbabwe. The transfer of insect re-
involved the selection of more productive and sistance (from Cajanus scarabaeoides (L.)
early-maturing landraces. Concerted interna- Thouars), high protein content (several spe-
tional pigeon pea improvement work began in cies), improved drought resistance (Cajanus
the 1970s at the International Institute of acutifolius (Benth.) Maesen), soil salinity tol-
Tropical Agriculture (UTA), where short- erance (Cajanus albicans (Wight &Arn.) Mae-
statured cultivars of determinate growth habit sen) or annual behaviour (Cajanus platycarpus
were developed. Within the Consultative Group (Benth.) Maesen) has not yet materialized.
on International Agricultural Research (CGIAR) Insect-tolerant lines have been identified, how-
system, ICRISAT now has the mandate for pi- ever.
geon pea improvement. So far, national breed- Transgenic pigeon pea plants, expressing a
ing programmes in Africa have mainly relied cowpea protease inhibitor gene or a protective
on ICRISAT lines and selections from local antigen of the Rinderpest virus, have been ob-
landraces, but some countries (e.g. Kenya, tained using Agrobacterium-medi&ted gene
Uganda and Rwanda) have developed their transfer or bombardment with micro-particles.
own crossing programmes. Prospects As a multi-purpose crop pigeon
Breeding of pigeon pea for high yield, and for pea is well known but ought to be promoted
consumer and miller preference are prime cri- especially in more semi-arid regions, for which
teria. Stability of yield may be obtained by the crop is well suited due to its tolerance to
selecting for photoperiod insensitivity, disease drought and low soil fertility and its ability to
and pest resistance, and suitability for inter- recover after environmental or biotic stress. Its
cropping and multiple harvests. Improved large seed-yield potential offers promise in
genotypes are now available for most of these more favourable environments. Pigeon pea fits
characteristics. Resistance is available in wild well in agro-forestry, in smallholder garden
relatives and there are promising pest- cropping and in hedge cultivation, and is suit-
resistant and disease-resistant types. Because able for improved short-duration fallows. It
out-crossing is frequent, traditional methods also fits in more intensive systems. Short-
for self-pollinated crops, such as pedigree duration, photoperiod-insensitive, short-stat-
breeding, have not been very effective. Selec- ured cultivars of determinate growth habit
tion is further complicated by large genotype x with fast growth and elevated harvest index
40 CEREALS AND PULSES

may be the ideotype to aim for. In the export Cenchrus biflorus is considered a valuable for-
market (mainly India) African pigeon pea faces age grass in the Sahel; it is mainly browsed in
strong competition and higher productivity; the juvenile stage and when the grains have
more efficient marketing arrangements are nec- fallen off. It can be cut several times during the
essary to remain competitive in this market. rainy season and made into hay or silage. The
Major references Hillocks et al., 2000; lo spiny involucres are sufficiently softened by
Monaco, 2003; Nene, Hall & Sheila, 1990; ensiling to make consumption of the whole
Reddy, Raju & Lenné, 1998; Silim, Mergeai & plant possible. Cenchrus biflorus persists until
Kimani (Editors), 2001;Silim, Tuwafe & Singh the end of the dry season and thus is important
(Editors), 1994; Singh et al., 2001; van der as a reliable source of fodder. Also in India the
Maesen, 1985; van der Maesen, 1989a; White- plant is used as a fodder and it is sown against
man, Byth &Wallis, 1985. desertification; in northern Australia it is sown
Other references Akojie & Fung, 1992; as a forage. The leaves are eaten during famine
Aulakh et al., 1985; Berhaut, 1976; du Puy et in the Thar desert in India.
al., 2002; Ene-Obong, 1995; Gillett et a l , 1971; The root of Cenchrus biflorus is an ingredient
Hanelt & Institute of Plant Genetics and Crop oftraditional aphrodisiac prescriptions.
Plant Research (Editors), 2001; Joshi et al., Properties The composition of hulled grains
2001; Kay, 1979; Mackinder et al., 2001; Mer- of Cenchrus biflorus per 100 g is: water 9.8 g,
geai et al., 2001;Neuwinger, 2000; Polhill, 1990; energy 1549 kJ (370 kcal), protein 17.8 g, fat
Popelka, Terryn &Higgins, 2004; Remanandan 8.5 g, carbohydrate 62.3 g, Ca 144 mg, P 270
& Singh, 1997; Tabo et al., 1995;Thulin, 1989a; mg and Fe 22 mg (Leung, Busson & Jardin,
USDA, 2004; van der Maesen, 2003; Westphal, 1968). The essential amino-acid composition
1974. per 100 g hulled grain is: lysine 214 mg, me-
Sources of illustration Busson, 1965. thionine 393 mg, phenylalanine 926 mg,
Authors L.J.G. van der Maesen threonine 658 mg, valine 1052 mg, leucine
2745 mg and isoleucine 892 mg (FAO, 1970).
The protein and fat contents are high com-
CENCHRUS BIFLORUS Roxb. pared to other cereals.
Cenchrus biflorus plants in the Sahel contain
Protologue FL ind. 1:238 (1820). crude protein 10.0%, crude fibre 34.6%, crude
Family Poaceae (Gramineae) fat 1.5%, nitrogen-free extracts 42.8%, P 0.35%,
Chromosome number n = 15, 16, 17, 18, 24 K 4.18%, Ca 0.28%, Mg 0.21%and Na 0.01%.
Vernacular names Cram-cram, Indian sand- In spite of its usefulness, Cenchrus biflorus is
bur (En). Cram-cram (Fr). often considered a noxious weed; the spiny
Origin and geographic distribution Cen- inflorescences may injure humans and live-
chrus biflorus is found throughout tropical stock and cause infection.
Africa, extending eastwards through Arabia Botany Loosely tufted, annual grass, with
and Iran to Pakistan and India. It has been ascending stems (culms) up to 1 m tall. Leaves
introduced elsewhere, e.g. in North America alternate, simple and entire; ligule a line of
and Australia. hairs; blade linear, flat, 2-25(-35) cm x 2-7(-
Uses The grain of Cenchrus biflorus is edible 10) mm, apex filiform. Inflorescence a spike-
and highly nutritious. People in areas of mar- like panicle 2-15 cm x 9-12 mm, with 1-3
ginal subsistence regularly collect the seed; spikelets enclosed by an involucre of prickly
elsewhere it is considered a famine food. In the bristles; rachis angular, sinuous; involucre
Sahel it is collected as a wild cereal, e.g. by the ovoid, 4-11 mm long with numerous spines,
Tuareg people. The grains are pounded and inner spines erect, fused at base, retrorsely
eaten raw, made into porridge, or mixed and hairy on the pungent, recurving apex, outer
cooked with other foods. The grain is also made spines shorter, spreading. Spikelet lanceolate
into a drink. In Sudan a thin bread ('kisra') is 3.5-6 mm long, acute, consisting of 2 glumes
made from the grain and in Mauritania the and usually 2 florets; glumes shorter than
ground grains are made into cakes. The grain spikelet; lower floret male or sterile, its lemma
of Cenchrus biflorus is also a famine food in as long as spikelet, membranous, upper floret
India, where it is eaten raw or used, mixed bisexual, its lemma as long as spikelet, thinly
with pearl millet, to make bread. In normal leathery; stamens 3, ovary superior, glabrous,
years it is mixed with sugar and 'ghee', and with 2 hairy stigmas. Fruit a dorsally com-
eaten as a children's food. pressed caryopsis (grain), 2-2.5 mm x 1.5-2
 CENCHRUS 41

Major references Burkill, 1994; Clayton &

Cenchrus comprises about 20 species in tropi- Renvoize, 1982; Naegele, 1977; National Re-
cal and warm temperate regions, mainly in search Council, 1996;Phillips, 1995.
Africa andtheAmericas. It isclosely related to Other references Bartha, 1970; Batello,
Pennisetum, which differs in non-spiny inner Marzot &Touré, 2004; Bhandari, 1974; Bouw-
involucral bristles free tothebase. man, 1979; Clayton, 1989; Cope, 1995; FAO,
The spiny spikelets ofCenchrus biflorus adhere 1970; Harlan, 1989b; Leung, Busson & Jardin,
to hairs of animals and clothes, making possi- 1968; Peyre deFabrègues,1992.
ble wide dispersal. Cenchrus biflorus follows Authors M. Brink
the C4-cycle photosynthetic pathway.
Ecology Cenchrus biflorus is mostly found
in semi-arid and arid regions with an annual CENCHRUS P R I E U E I I (Kunth) Maire
rainfall of250-650 mm,upto 1300 m altitude,
usually on dry sandy soils and in cultivated, Protologue Bull. Mus. natn. Hist, nat.,
overgrazed or otherwise disturbed areas. It is Paris, sér. 2, 3:523 (1931).
extremely abundant in the Sahel and southern Family Poaceae (Gramineae)
Sahara, where it mayform massive stands. A Chromosome number 2n- 34
study in western Niger showed that it had be- Origin and geographic distribution Cen-
come much more abundant and dominant in chrus prieurii is distributed from Mauritania
the late 1980s than itwas intheearly 1960s. and Senegal through the Sahel zone to Ethio-
Management Cenchrus biflorus can be pia; it also occurs in Arabia, Pakistan and
propagated byseed. Theoptimum temperature northern India.
for seed germination is 35°C. In tropical Africa Uses The grain of Cenchrus prieurii is an
the grains are collected from the wild. The important food for some desert nomads; it
spiny spikelets shatter easily at maturity and serves as a famine food in Africa and India.
are often allowed to fall, after which they are The crushed or ground grain is made into por-
swept into piles with a bunch ofstraw, or they ridge. In India the grains are eaten raw and
are raked with a big'comb' with a handle.The are used, mixed with pearl millet, for making
plants may be beaten with a stick if not all bread.
spikelets have fallen. The spikelets are Cenchrus prieurii is valued for grazing; it also
pounded in a mortar andthe grains are sepa- makes suitable hayandsilage. It persists until
rated bywinnowing. IntheLake Chad area the the endofthe dryseason andthus is important
inflorescences are cut off with a knife, after as a reliable source of fodder. In northern Ni-
which the grains are dried, threshed and win- geria Cenchrus prieurii isplanted asa forage.
nowed. In Kordofan (Sudan) the grains are Properties The fodder value of Cenchrus
hulled by rubbing them between two pieces of prieurii plants in the Sahel is: crude protein
leather. 9.2%, crude fibre 37.1%, crude fat 1.8%, nitro-
Genetic resources and breeding TheIn- gen-free extractives 42.8%, P 0.15%, K 3.36%,
ternational Livestock Research Institute Ca 0.23%, Mg 0.19%and Na 0.02%. Informa-
(ILRI), Addis Ababa, Ethiopia, holds 10 acces- tion on the nutrititional characteristics ofthe
sions of Cenchrus biflorus. In view of its wide grain isnotavailable.
distribution and abundance, Cenchrus biflorus Botany Loosely tufted, annual grass, with
is certainly notthreatened bygenetic erosion. stems (culms) up to 80 cm tall. Leaves alter-
Prospects Cenchrus biflorus yields a highly nate, simple and entire; ligule a line of hairs;
nutritious grain, with unusually high protein blade linear, flat, 10-30 cm x 3.5-10 mm,
and fat contents. Formerly it wasimportant as finely acute. Inflorescence a cylindrical spike-
a wild cereal, but nowadays it seems to play a like panicle 5-12 cm x 2-4 cm, with 1-2
role in human nutrition in times of shortage spikelets enclosed byan involucre oflong bris-
only. As a forage it has remained important, tles; rachis angular, scabrid, sinuous; involucre
especially because ofits persistence throughout with many slender scabrid bristles 15-27 mm
the dryseason. Cenchrus biflorus is unlikely to long and furnished with spines directed up-
become more important in the future, mainly wards, far exceeding the spikelet, fused at
due to its spiny spikelets which adhere to base. Spikelet lanceolate, 4-5 mm long, acute,
clothes and cause injuries to humans and ani- consisting of 2 glumes and usually 2 florets;
mals, and result in the plant often being con- glumes shorter than spikelet; lower floret male
sidered a noxious weed. or sterile, its lemma as long as spikelet,mem-
42 CEREALS AND PULSES

branous; upper floret bisexual, its lemma as

long as spikelet, thinly leathery. Fruit a dor-
sally compressed caryopsis (grain).
Cenchrus comprises about 20 species in tropi-
cal and warm temperate regions, mainly in
Africa and the Americas. It is closely related to
Pennisetum, which differs in non-spiny inner
involucral bristles free to the base.
Cenchrus prieurii follows the C4-cycle photo-
synthetic pathway.
Ecology Cenchrus prieurii is found in semi-
arid and arid regions with an average annual
rainfall of 200-500 mm, in open sandy loca-
tions up to 1000 m altitude. A study in western
Niger showed that Cenchrus prieurii had be-
come much more abundant and dominant in
the late 1980s than it was in the early 1960s.
Management Cenchrus prieurii is collected Cicerarietinum -planted
from the wild. The 1000-seed weight is 0.2 g.
Genetic resources and breeding A few pea is not known in a wild state. Its origin is
accessions of Cenchrus prieurii are held in Aus- believed to be in south-eastern Turkey and
tralia (Australian Tropical Crops & Forages adjoining Syria and Iran. The earliest remains
Genetic Resources Centre, Biloela, Queen- of chickpea seeds date back to around 7000 BC
sland, 3 accessions) and the United Kingdom (Syria and Turkey). Chickpea was gradually
(Welsh Plant Breeding Station, Institute of introduced to the western Mediterranean re-
Grassland and Environmental Research, Aber- gion, to eastern and southern Asia and East
ystwyth, Wales, 2 accessions). In view of its Africa. It reached the Indian subcontinent be-
wide distribution, Cenchrus prieurii is not fore 2000 BC. Chickpea cultivation is expand-
threatened by genetic erosion. ing where it has been recently introduced, e.g.
Prospects Cenchrus prieurii has some value in Australia, New Zealand, the United States
as a source of food in times of scarcity and as a and Canada. In tropical Africa it is mainly cul-
fodder grass, but it is unlikely to increase in tivated in East Africa (Sudan, Eritrea, Ethio-
importance in the future. Investigations are pia, Kenya, Tanzania) and in Malawi; it is
necessary to find out if the nutritional quality grown particularly in areas with a marked cool
of the grain of Cenchrus prieurii is as high as season. Lesotho and South Africa have recently
that of the grain of Cenchrus biflorus Roxb. introduced chickpea at experimental level.
Major references Bogdan, 1977; Burkill, Chickpea is found semi-naturalized as an es-
1994; Hanelt & Institute of Plant Genetics and cape, e.g. in Tanzania.
Crop Plant Research (Editors), 2001; Peyre de Uses Chickpea is primarily grown for its
Fabrègues, 1992;Phillips, 1995. mature seeds, which are used as human food.
Other references Bartha, 1970; Breman & These are consumed alone or together with
de Ridder, 1991; Clayton, 1972; Freedman, cereals as a side dish in the form of a sauce or
undated. soup. In Ethiopia sauces ('wot') made of ground
Authors M. Brink seeds ('shiro') and split seeds ('kik') are com-
monly eaten with 'injera' (unleavened, pan-
cake-like bread). Chickpea is also an ingredient
ClCERARIETINUML. of weaning foods. The immature seeds are con-
sumed fresh, or roasted and salted as snacks.
Protologue Sp. pi. 2: 738 (1753). In India the whole dried seeds are eaten boiled
Family Papilionaceae (Leguminosae - Papil- or made into dhal, prepared by splitting the
ionoideae, Fabaceace) seed and separating the husk. In Mediterra-
Chromosome number 2n = 16 nean countries, chickpea is eaten whole in sal-
Vernacular names Chickpea, Bengal gram, ads, or in stews, and flour mixed with sesame
gram, garbanzo (En). Pois chiche (Fr). Grâo de paste yields the well-known appetizer 'hum-
bico, gravanço, ervanço (Po).Mdengu (Sw). mus'. Canned chickpea seeds are popular in the
Origin and geographic distribution Chick- United States and in Europe.
 ClCER 43

In India young chickpea sprouts are eaten as a Fe 6.2 mg, Zn 3.4 mg, vitamin A 67 IU, thiamin
vegetable. Broken seeds and residues from 0.48 mg, riboflavin 0.21 mg, niacin 1.5 mg,
dhal production are used as feed, the straw vitamin Be 0.54 mg, folate 557 (ig and ascorbic
serves as fodder and dried stems and roots are acid 4.0 mg. The essential amino-acid composi-
used as fuel for cooking. Chickpea starch is tion per 100 g edible portion is: tryptophan 185
suitable for textile sizing, giving a light finish mg, lysine 1291 mg, methionine 253 mg,
to silk, wool and cotton clothes, and can also be phenylalanine 1034 mg, threonine 716 mg,
used in the manufacturing of plywood. An in- valine 809 mg, leucine 1374 mg and isoleucine
digo-like dye is obtained from chickpea leaves. 828 mg. The principal fatty acids are per 100 g
Production and international trade Accord- edible portion: linoleic acid 2593 mg, oleic acid
ing to FAO statistics, the annual world produc- 1346 mg, palmitic acid 501 mg, linolenic acid
tion and harvested area of chickpea from 1961 101 mg and stearic acid 85 mg (USDA, 2004).
to 2003 has remained relatively stable at The protein content of chickpeas is lower than
around 7 million t and 10 million ha, respec- that of most other pulses, but this is compen-
tively. The production in 1999-2003 amounted sated for by the higher protein digestibility.
to 7.9 million t per year from 10.3 million ha. Antinutritional factors include trypsin inhibi-
The main producing countries were India (4.1 tors, haemagglutinins, tannins and oligosac-
million t per year from 6.3 million ha), Turkey charides.
(600,000 t from 600,000 ha), Pakistan (500,000 Adulterations and substitutes In India
t from 1.1 million ha), Canada (250,000 t from chickpea is sometimes adulterated with the
200,000 ha), Mexico (250,000 t from 150,000 cheaper, but potentially toxic, grass pea
ha) and Australia (200,000 t from 200,000 ha). (Lathyrus sativus L.).
The annual production in sub-Saharan Africa Description Spreading to erect, annual
was about 280,000 t from 430,000 ha, the main herb up to 100 cm tall; stem simple or
producers being Ethiopia (168,000 t from
191,000 ha), Malawi (35,000 t from 88,000 ha),
Sudan (25,000 t from 13,000 ha), Tanzania
(25,000 t from 63,000 ha) and Kenya (20,000 t
from 55,000 ha). In tropical Africa the area and
production of chickpea have been increasing
recently, whereas they are declining in north-
ern Africa. The decline in Ethiopia was ar-
rested and has been reversed due to the release
of improved cultivars, liberalized markets and
intensive extension activities. In Zambia
chickpea is grown by commercial farmers
around urban areas.
The world trade in chickpea steadily increased
from 100,000-150,000 t per year in the 1970s
to about 700,000 t per year in 1998-2002. The
main exporters in 1998-2002 were Australia
(192,000 t per year), Mexico (155,000 t), Turkey
(114,000 t), Canada (85,000 t) and Iran (75,000
t). Ethiopia exported about 50,000 t in 2002,
but insignificant amounts in 1998-2001. Tan-
zania exported about 20,000 t in 2002 and less
than 10,000 t per year in 1998-2001.The main
importers in this period were India (183,000 t
per year), Pakistan (98,000 t), Spain (57,000 t),
Algeria (43,000 t) and Bangladesh (40,000 t).
Imports into sub-Saharan Africa were very low.
Properties The composition of mature raw
chickpea seeds per 100 g edible portion is: wa-
ter 11.5 g, energy 1525 k j (364 kcal), protein Cicer arietinum • 1, flowering and fruiting
19.3 g, fat 6.0 g, carbohydrate 60.7 g, dietary branch; 2, seed.
fibre 17.4 g, Ca 105 mg, Mg 115 mg, P 366 mg, Source: PROSEA
44 CEREALS AND PULSES

branched from the base; taproot reaching 1-2 types. Some intermediate cultivars also exist.
m depth, secondary roots mostly spreading in Desi-type chickpeas are bushy plants with
the top 15-30 cm soil layer. Leaves alternate, relatively small leaflets and flowers, purplish
imparipinnate, with (7—)11—15(—17) leaflets; anthocyanin pigments in their stems and blue-
stipules 2—5-fid, ovate to triangular, 3-5 mm x violet flowers, and are primarily grown in
2-4 mm; rachis 2.5-8 cm long, grooved above; southern Asia and Ethiopia. The Kabuli types
leaflets sessile, ovate to elliptical, 5-20 mm x have erect growth and white flowers and are
2-15 mm, the upper two-thirds of the margins grown in the Mediterranean region. Cultivars
sharply toothed, glandular pubescent on both of the Kabuli type cook faster and have less
sides. Inflorescence reduced to a single axillary dietary fibre than those of the Desi type. Seed
flower; peduncle 3-20(-37) mm long; bracts 1- colour is an important characteristic of chick-
3, linear to triangular, up to 3 mm long. Flow- pea, determining its quality and acceptance in
ers bisexual, papilionaceous; pedicel 3-12 mm many countries. In East Africa brown Desi
long, recurved at fruiting; calyx campanulate, chickpeas are most popular.
tube 3—4 mm long, teeth lanceolate, 4—5 mm Growth and development Seedlings nor-
long, with prominent midribs; corolla white, mally emerge 7—15 days after sowing. Flower-
pink, purplish or blue, standard obovate, 8-10 ing starts after 30—60 days. Chickpea is self-
mm x 7—10(—17)mm, with a broad claw, wings pollinated, with less than 2% out-crossing. The
obovate, 6-9 mm x c. 4 mm, auriculate, keel 6— crop duration is normally 3-6 months, but
8 mm x c. 3 mm with a 2—3 mm long claw; chickpea is indeterminate by nature and may
stamens 10, 9 united for 4-5 mm and 1 free, continue to grow as long as moisture is not
anthers basi-dorsifixed; ovary superior, sessile, limiting. The deep, strong taproot serves as a
ovate, 2—3 mm x 1-1.5 mm, 1-celled, style in- water-storage organ for the growing plants,
curved, 3-4 mm long, stigma small. Fruit an extending growth into the dry season. Chick-
inflated rhomboid-ellipsoid pod 12-35 mm x 8- pea is effectively nodulated by Mesorhizobium
20 mm, densely glandular pubescent, 1—2(-4)- ciceri and Mesorhizobium mediterraneum.
seeded. Seeds globular to angular obovoid, 5— Ecology Chickpea is cultivated in tropical,
14 mm x 4-10 mm, with a median groove and a subtropical and warm temperate zones. Its
conspicuous beak overhanging the hilum, production is concentrated in the cool, dry sea-
creamy to brown, green or black, surface son of the semi-arid tropics. It grows from sea-
smooth or wrinkled. Seedling with hypogeal level to over 2500 m altitude, but it is not suit-
germination; the first two leaves scale-like. able for the humid and hot lowland tropics,
Other botanical information Cicer com- where it often fails to flower. In East Africa it
prises 43 species, 9 annual and 34 perennial. is grown at 1200-2700 m altitude, in areas
The wild Cicer species most closely related to with an annual rainfall of 500-1800 mm and
Cicer arietinum are the annuals Cicer reticula- an average temperature of 10-25°C during the
tum Ladiz. and Cicer echinospermum P.H. growing period. Under these conditions the
Davis. Cicer reticulatum, a rare species from crop takes 120-150 days to mature. It performs
Turkey, is sometimes regarded as a subspecies well when planted during the rainy season
of Cicer arietinum; it is morphologically, bio- provided that the field is well drained, but hu-
chemically and karyologically very similar and midity favours the development of aseochyta
completely cross-compatible. Fertile hybrids blight. Rain during flowering hampers seed set.
have also been produced in crosses of chickpea The drought tolerance varies from moderate to
and Cicer echinospermum, though fertility bar- considerable. Chickpea is generally a quantita-
riers do exist. Other related species are Cicer tive long-day plant. Soils need to be well
bijugum Rech.f., Cicer chorassanicum (Bunge) drained, with pH 5-7 or more. Salinity is
Popov, Cicer cuneatum Höchst, ex A.Rich., Ci- hardly tolerated, if at all. Soils vary from sandy
cerJudaicum Boiss., Cicer pinnatifidum Jaub. to sandy loam and black cotton soils. In Ethio-
& Spach, Cicer yamashitae Kitam. (all annu- pia and Kenya chickpea is mainly grown to-
als) and Cicer anatolicum Alef. (a perennial), wards the end of the rainy season on black
and some of these have been used for crossing cotton soils with declining soil moisture.
with cultivated chickpea. Propagation and planting Chickpea is prop-
Within cultivated chickpea 2 main groups are agated by seed. Seed can be stored for 4—5
commonly discerned: the large-seeded, cream- years at a temperature of 4°C. The 1000-seed
coloured Kabuli types and the small-seeded, weight varies widely, from 20 g for some Desi
darker-coloured, smooth or wrinkled Desi types, to more than 600 g for the larger Kabuli
 ClCER 45

types. Seed does not show dormancy. Because (Helicouerpa armigera) and cutworm (Agrotis
of the large variation in seed size, seed rates ipsilon) are common insect pests on chickpea in
may range from 30-40 kg/ha for small-seeded East Africa and India. Insecticides such as
cultivars to 90-100 kg/ha for large-seeded ones. endosulfan are recommended for the control of
In East Africa the seeds are broadcast and these insects. Integrated Pest Management
then ploughed under by animal-drawn ploughs. (IPM) practices, including tolerant cultivars,
In southern Asia spacings of 25-30 cm between pest population monitoring, bio-pesticides and
rows and 10-30 cm between plants in a row are natural enemies, have been developed to re-
common. Chickpea is grown as a sole crop or in duce the reliance on insecticides. Calloso-
intercropping with linseed, sorghum or other bruchus spp. are important storage insects in
crops. It is often sown as a relay crop, e.g. in chickpea.
rice paddies. Harvesting Mature seeds are harvested
Management Chickpea is very sensitive to when the pod tips of the uppermost branch of
weed competition, especially in the first 4-6 the plant turn yellow. Harvesting is done
weeks after sowing. Weed control is usually by manually by pulling the plants, after which
mechanical methods. The weed population can they are sun-dried in the field. Where chickpea
be high if the crop receives late rains after sow- is machine-harvested, a tall and erect plant
ing and in that case immediate weeding is re- type is preferred.
quired. The use of P-fertilizer high in S is rec- Yield The average yield of chickpea is less
ommended, but normally no inorganic fertiliz- than 1 t/ha in sub-Saharan Africa, except un-
ers are applied. Some farmers in Sudan apply der irrigation in Sudan, where average yields
100 kg diammonium phosphate (DAP) per ha. up to 1.9 t/ha are obtained. In the Central
In Sudan and in India chickpea is sometimes Highlands of Ethiopia, farmers harvest over 2
grown under irrigation. In Ethiopia chickpea is t/ha due to a favourable growing environment,
grown in rotation with cereals, mainly tef (Era- while research shows a potential yield up to 5.5
grostis te/(Zuccagni) Trotter); in India it is also t/ha in this area. The average yield of chickpea
grown in rotation with cereals, including pearl in India is also less than 1 t/ha.
millet, sorghum, wheat, barley and rice. Handling after harvest When the har-
Diseases and pests The most important vested pods have been sun-dried to a moisture
chickpea disease worldwide is ascochyta blight content of 12—13%, they are taken to the
caused by the seed-borne Ascochyta rabiei, but threshing ground. In Ethiopia, Eritrea and
this disease is of lesser importance in Ethiopia, Sudan, threshing is done by trampling using
except when chickpea is sown early in the draught animals. Small harvests are threshed
rainy season. Important diseases in Ethiopia, by beating with sticks. Seeds are separated
Sudan and Eritrea are fusarium wilt caused by from the chaff by winnowing. The cleaned
Fusarium oxysporum f.sp. ciceris, dry root rot seeds are kept in stores or taken to the market.
caused by Macrophomina phaseolina (Rhizoc- Bruchid attack makes storage for more than 6
tonia bataticola), and collar rot caused by Scle- months difficult. Ethiopian farmers mix chick-
rotium rolfsii. Control measures against the pea with tef to keep the seeds for a longer pe-
seed- and soil-borne fusarium wilt include the riod. Seed for planting may be protected with
use of seed from disease-free plants, seed Pyrethrine insecticides.
treatment with fungicides, and the use of resis- Genetic resources ICRISAT (Patancheru,
tant cultivars. Crop rotation is not effective, India) has a collection of about 17,000 chickpea
because the fungus survives in the soil for long accessions, from which a representative core
periods. Control measures for dry root rot and collection of 1956 accessions has been formed.
collar rot include the use of disease-free seed, Other large collections are kept by ICARDA
the removal of crop residues and the elimina- (Aleppo, Syria, about 10,000 accessions) and in
tion of weed hosts. Effective control with crop Australia (Australian Temperate Field Crops
rotation is difficult, because both pathogens Collection, Horsham, 7700 accessions) and the
have a wide host range. Resistance to Macro- United States (USDA-ARS Western Regional
phomina phaseolina has been observed, but the Plant Introduction Station, Pullman, Washing-
disease may affect even resistant cultivars if ton, 4400 accessions). ICRISAT and ICARDA
these are grown in infected soil for a long pe- have 58 and 268 accessions of wild Cicer spe-
riod. cies, respectively. Many species from Central
Root-knot nematode (Meloidogyne javanica) is Asia are not yet represented in the collections.
an important parasite of chickpea. Pod borer In Ethiopia, which is considered a secondary
46 CEREALS AND PULSES

centre of diversity for chickpea, the Institute of moderate to high drought tolerance. Chickpea
Biodiversity Conservation has the largest col- production in the North African countries is
lection (of about 1000 accessions) in Africa. declining due to its low yield compared to cere-
Chickpea shows orthodox seed storage behav- als. Therefore, countries elsewhere in Africa,
iour and can be stored for long periods without such as Ethiopia, Sudan and Tanzania, may
loss of viability. For long-term storage a tem- increase their production to take advantage of
perature of-20°C is used. the markets in North Africa and the Indian
Breeding Genetic improvement of chickpea subcontinent. Chickpea is one of the main
aims at higher yield and resistance to diseases, pulses providing affordable protein for the
insects and other stresses such as drought, Ethiopian, Eritrean, and Sudanese popula-
waterlogging and cold. Sources of resis- tions, whereas more promotion is required in
tance/tolerance have been identified for dis- Kenya, Tanzania and Malawi. In Zimbabwe,
eases (including ascochyta blight, fusarium Uganda, South Africa and Lesotho chickpea
wilt and dry root rot), insect pests (including trials, received from ICARDA and ICRISAT,
pod borer) and abiotic stress factors (including have recently been started. The development of
cold and drought). Cultivars have been re- new food products and recipes will help to in-
leased with resistance to ascochyta blight, crease chickpea consumption in sub-Saharan
fusarium wilt and cold, but limited success has Africa.
been attained in the development of cultivars Major references Bejiga, Eshete & Anbes-
tolerant to insect pests. Conventional breeding sa, 1996; Haware, 1998; Pope, Polhill & Mar-
techniques for self-pollinated crops are used in tins (Editors), 2003; Saxena & Singh (Editors),
chickpea breeding. Mutation breeding has been 1987; Saxena et al. (Editors), 1996; Singh &
carried out to create new variability, e.g. for Saxena, 1999; Singh et al., 1997b; Smithson,
ascochyta blight resistance. Wild Cicer species Thompson & Summerfield, 1985; Telaye et al.
have been used in interspecific hybridization (Editors), 1994;van der Maesen, 1989b.
programmes, but few perennials have been Other references Ahmad, 1999; Anbessa &
tried for use in chickpea improvement. Chick- Bejiga, 2002; Bejiga, 1990; Bejiga & Degago,
pea crossing techniques are tedious. Within the 2000; Bejiga et al., 1998; Choumane et a l ,
CGIAR system ICARDA and ICRISAT are in- 2000; Flandez-Galvez et al., 2003; Gillett et al.,
volved in chickpea breeding. The only substan- 1971; Hanelt & Institute of Plant Genetics and
tial national chickpea programmes in the sub- Crop Plant Research (Editors), 2001; Joshi et
Saharan region are those of Ethiopia and Su- al., 2001;McPhee &Muehlbauer, 2002; Polhill,
dan, which have made much progress in devel- 1990; Popelka, Terryn & Higgins, 2004; Singh,
oping high-yielding and disease-resistant culti- 1993; Thulin, 1989a; Upadhyaya, Bramel &
vars for commercial production. In Ethiopia 10 Singh, 2002; USDA, 2004; van der Maesen,
cultivars (5 each from Kabuli and Desi types) 1972; Westphal, 1974;Williams et al., 1994.
were released and more than 7 cultivars were Sources of illustration van der Maesen,
promoted by extension for multiplication. 1989b.
Kenya so far released one cultivar from germ- Authors G. Bejiga &L.J.G. van der Maesen
plasm received from ICRISAT.
Linkage maps of chickpea have been devel-
oped, and molecular markers associated with CODCLACRYMA-JOBIL.
quantitative trait loci for resistance to as-
cochyta blight, fusarium wilt and various mor- Protologue Sp. pi. 2: 972 (1753).
phological traits have been located on these Family Poaceae (Gramineae)
maps. Transgenic chickpea plants, e.g. showing Chromosome number 2n = 20
inhibitory effects on development of Calloso- Vernacular names Job's tears, adlay (En).
bruchus spp. due to an a-amylase inhibitory Larmes de Job, larmilles, herbe à chapelets
gene from common bean, or inhibitory effects (Fr). Lâgrimas de Job, lâgrimas de Nossa Sen-
on the growth on larvae of Helicoverpa ar- hora, erva dos rosârios (Po). Mtasubihu, mtas-
migera due to gene transfer from Bacillus thur- bihi (Sw).
ingiensis, have been obtained using Agrobacte- Origin and geographic distribution Job's
rium-mediated gene transfer or biolistic trans- tears is indigenous to southern and eastern
formation. Asia. It has been cultivated since ancient
Prospects Chickpea is a very suitable crop times, 3000-4000 years ago in India, 2000
for the semi-arid regions of Africa, due to its years ago in China, and was very important
 COIX 47

before maize and rice became widespread sta- niacin 4.7 mg (Leung, Busson & Jardin, 1968).
ple foods. At present Job's tears is cultivated as The content of essential amino acids per 100 g
a minor cereal crop throughout the tropics and protein (16 g N) is: tryptophan 0.5 g, lysine 1.9
subtropics, especially in Asia. Plants escaped g, methionine 2.6 g, phenylalanine 4.9 g,
from cultivation occur as weeds. In Africa Job's threonine 3.0 g, valine 5.7 g, leucine 13.6 g and
tears is naturalized in most countries but only isoleucine 3.9 g (Busson, 1965).
very occasionally cultivated (e.g.in Liberia). The root contains coixol, which is analgesic and
Uses Types of Job's tears with soft-shelled sedative. Methanolic extracts of the grains
false fruits can be easily husked and have large showed an antiproliferative effect on human
grains which are eaten in the same way as rice, lung cancer cells in vitro and in vivo and might
alone or mixed with it. They can be substituted reduce the risk of tobacco-induced lung tumori-
for rice in all foodstuffs. The grain can also be genesis. The grains might also be beneficial for
roasted before husking and then used in por- the treatment of allergic disorders.
ridge, cakes, soups and other foods or in the Botany Erect, perennial, strongly tillering
preparation of sweets. Dough made from the grass up to 3 m tall, often cultivated as an an-
flour will not rise because of the absence of nual; stem (culm) filled with pith, glabrous,
gluten. A good mixture for bakery purposes is branched in the upper part. Leaves alternate,
70% wheat flour and 30%Job's tears flour. The simple and entire; leaf sheath short, glabrous
raw grain tastes sweet and is often eaten as a or with long hairs at apex; ligule short and
snack. In Africa Job's tears is considered a fam- membranous; blade linear to ovate-lanceolate,
ine food. Both alcoholic and non-alcoholic 8-100 cm x 1.5-7 cm, base rounded to almost
drinks are prepared from it. A beer made from cordate, apex acute, margins rough, upper sur-
the pounded grain is popular among Indian hill face smooth or scabrid, midrib prominent. In-
tribes and in the Philippines. florescences in axils of upper leaves, solitary or
The whole grain and the bran are fed to poul-
try and the flour can replace maize flour in
poultry feed. Job's tears is often given as a fod-
der, especially for cattle and horses. It is suit-
able for silage, and straw and leaves are used
for thatching.
The grain and flour of Job's tears are easily
digestible and given to people in weak condi-
tion. They are believed to have medicinal value
with diuretic, depurative, anti-inflammatory
and antitumour activity. A decoction of the
leaves is drunk against headache, rheumatism
and diabetes. Sap of the stem is applied
against insect bites. A decoction of the roots is
used as a vermifuge and to treat dysentery,
gonorrhoea and menstrual disorders.
Almost everywhere where Job's tears grows,
the decorative, hard-shelled false fruits of the
wild types are used as beads for necklaces,
rosaries, rattles, curtains etc., and in Africa
they are often worn at ritual and religious oc-
casions. The whole inflorescence is sometimes
used in dried flower arrangements.
Properties Whole grain of Job's tears con-
tains per 100 g edible portion: water 8.9 g, en-
ergy 1394 kJ (333kcal) protein 10.4 g, fat 5.3 g,
carbohydrate 66.5 g and fibre 10.5 g. The
hulled grain contains per 100 g edible portion:
water 11.6 g, energy 1511kJ (361kcal), protein Coix lacryma-jobi - 1, flowering stem; 2, male
14.8 g, fat 4.9 g, carbohydrate 66.9 g, fibre 0.5 inflorescence; 3, male spikelet; 4, female inflo-
g, Ca 47 mg, P 254 mg, Fe 6.0 mg, ß-carotene 0 rescence with cupule partly removed.
mg, thiamin 0.26 mg, riboflavin 0.19 mg and Source: PROSEA
48 CEREALS AND PULSES

2-7-fascicled and arranged panicle-like, on recommended for fodder production. Propaga-

peduncle 3-6 cm long, consisting of 2 unisexual tion by seed gives deeper rooting, and, conse-
racemes; female raceme enclosed by a hollow, quently, better drought tolerance and higher
bony, globular to ovoid-ellipsoid cupule 5—15 grain yield.
mm long, shiny, white, pale brown, grey, bluish Weeding is necessary up to 60 days after sow-
or black, with a sessile spikelet accompanied by ing or until Job's tears has reached a plant
2 barren pedicels; male raceme 3-5 cm long, height of 40 cm. In general, plants are not
exserted from the mouth of the cupule, with given much care, but when young they need
about 10 spikelets borne in pairs or threes, one abundant water. They respond well to applica-
pedicelled, the other(s) sessile. Female spikelet tion of manure; chemical fertilizers or insecti-
2-flowered, with orbicular glumes, lower floret cides are not used.
reduced to an orbicular lemma, upper floret The most serious disease of Job's tears is smut
with membranous lemma and palea and supe- (Ustilago coicis) which destroys the ovaries.
rior ovary with 2 stigmas exserted from the Smut can severely damage crops and therefore
mouth of the cupule; male spikelet lanceolate seed treatment with fungicide or with hot wa-
to ellipsoid, 7-8 mm long, 1-2-flowered, lower ter (60-70°C) for at least 10 minutes before
glume winged, upper glume boat-shaped, each sowing is recommended. Another important
floret with membranous lemma and palea and disease of Job's tears is leaf blight (Bipolaris
3 stamens. Fruit a caryopsis (grain) enclosed coicis); control measures include burning of
by the cupule (shell of false fruit), globose, dark crop residues, spraying of fungicides and the
red in hard-shelled types, pale brown in soft- use of more resistant cultivars. Tar leaf spot
shelled types. (Phyllachora coicis), rust (Puccinia operata)
Coix comprises about 5 closely related species, and Ustilago lachrymae-jobi (synonym: Spori-
all native in Asia, but some have been intro- sorium lachrymae-job) are some of the other
duced elsewhere. Mainly based on characteris- diseases attacking Job's tears. Rats, birds and
tics of the false fruit, 4 varieties have been sometimes grasshoppers and termites may
distinguished in Coix lacryma-jobi, but only cause considerable losses.
var. lacryma-jobi occurs in Africa; it is charac- Job's tears is normally harvested 4-6 months
terized by ovoid, hard, smooth false fruits. after sowing, depending on the cultivar and the
Most agricultural information (from outside season. Usually, whole plants are cut at the
Africa) concerns var. ma-yuen (Rom.Caill.) base when the grain is ripe. The stubble can be
Stapf, with ovoid to pear-shaped, quite soft, left in the field and will then tiller again; the
striate false fruits; it is cultivated as a cereal. new fresh leaves are an excellent fodder. Nor-
Job's tears takes about 1-2 weeks to germi- mal yield of husked grain varies from 2-4 t/ha.
nate, depending on the moisture content of the The hulling percentage is 30-50%. If cultivated
soil. Both self-pollination and cross-pollination for fodder, several cuts per year are possible.
are possible, with the latter usually being pre- After threshing and husking, which is done
dominant. Total crop duration is 4—6(—8) manually or with the same tools as for rice, the
months. When most of the seeds are ripe, the grain is sun-dried on mats. Under humid con-
plant starts to dry. Job's tears follow the C4- ditions, the storability of the grain is limited,
cycle photosynthetic pathway. but is better for whole than for husked grain.
Ecology Job's tears occurs wild in swampy Genetic resources and breeding The larg-
locations and along watercourses. It is a quan- est germplasm collections of Job's tears are
titative short-day plant and requires high tem- held in China (Institute of Crop Germplasm
peratures, abundant rainfall and reasonably Resources (CAAS), Beijing, 87 accessions) and
fertile soils. In the tropics it occurs from sea- the Philippines (Institute of Plant Breeding,
level up to 2000 m altitude, in Africa often University of the Philippines Los Banos
around villages and on abandoned fields. (UPLB), College, Laguna, 31 accessions). The
Management Job's tears is usually propa- greatest variation in wild forms occurs in India
gated by seed. The 1000-seed weight is 80-90 and Myanmar, in cultivated Job's tears in
g. Seed is dibbled, 5 cm deep, at the start of the South-East Asia. In the course of time, Job's
rains, after ploughing or hoeing the field. Row tears has been selected by farmers for easy
spacing is 40-80 cm, and seed rate 7-15 kg/ha. husking, resulting in var. ma-yuen. However,
When cultivated as an intercrop, it is sown at the crop has a relatively long growing season,
random or plants are grown along field bor- shows uneven ripening and variable yields.
ders. Propagation by cuttings is possible and Nevertheless, the large variability in Job's
 CORDEAUXIA 49

tears offers opportunities for breeding pro-

grammes, e.g. to obtain resistance against
smut disease. In Japan selection work focuses
on the use as a fodder. In Brazil a high-yielding
'dwarf cultivar, probably introduced from Ja-
pan, has been selected and distributed.
Prospects Although enjoyed locally by many
people, Job's tears is still decreasing in popu-
larity in favour of higher-yielding cereals,
mainly maize and rice. However, because it is
less susceptible to diseases and pests, it can be
grown where other crops are difficult to culti-
vate, it does not need much care, it is highly
nutritious and has promising medicinal proper-
ties, Job's tears deserves more research atten-
tion.
Major references Burkill, 1994; Chang,
Huang & Hung, 2003; Clayton & Renvoize, Cordeauxia edulis - wild
1982; Mello et a l , 1995;van den Bergh & Iam-
supasit, 1996. sweetish, agreeable, chestnut-like taste after
Other references Busson, 1965; Chang & roasting. The water in which the seeds have
Hwang, 2002; Gurib-Fakim, Guého & Bissoon- been boiled is sweet and is sometimes con-
doyal, 1997; Hsu et al., 2003; Leung, Busson & sumed. The seed oil is useful for soap making.
Jardin, 1968; Naku Mbumba, Walangululu & The seeds have been mentioned as a coffee
Basiloko, 1984; Neuwinger, 2000;Numata et al., substitute. Cordeauxia edulis is said to regu-
1989; Purseglove, 1972; Watt & Breyer-Brand- late gastric secretion and to permit treatment
wijk, 1962. of ulcers due to hot food. It is also believed to
Sources of illustration van den Bergh & alleviate anaemia by augmenting the number
Iamsupasit, 1996. of red blood cells. The leaves are made into a
Authors P.C.M. Jansen tea. Cordeauxia edulis is an important dry-
Based on PROSEA 10: Cereals. season fodder for camels, goats, sheep and cat-
tle, but in the rainy season other plants are
preferred. The red pigment in the glands on
CORDEAUXIA EDULIS H e m s l . several plant parts forms vividly coloured, fast
and insoluble combinations with many metal
Protologue Bull. Misc. Inform. Kew: 361 mordants, and is locally used for dyeing tex-
(1907). tiles. The wood is used as firewood.
Family Caesalpiniaceae (Leguminosae - Cae- Production and international trade The
salpinioideae) seeds of Cordeauxia edulis are mostly con-
Chromosome number 2M = 24 sumed locally, but are also sold in towns. De-
Vernacular names Yeheb nut, yeheb bush mand exceeds supply because of rapidly dimin-
(En).Yeheb (Fr). ishing plant populations. From Ethiopia the
Origin and geographic distribution Cor- seeds are exported to Somalia and Arab coun-
deauxia edulis is endemic to south-eastern tries, but no quantitative information is avail-
Ethiopia (eastern Ogaden) and central Soma- able. Cordeauxia edulis seeds have export po-
lia. It is cultivated on a small scale in Somalia tential for European markets as 'dessert nuts'.
and near Voi in Kenya, where it was intro- Properties Shelled Cordeauxia edulis seeds
duced in the 1950s. It has been introduced on contain per 100 g edible portion: water 11.1 g,
an experimental scale into Sudan, Tanzania, energy 1666 k j (398 kcal), protein 10.8 g, fat
Yemen, Israel and the United States. 12.0 g, carbohydrate 63.9 g, fibre 1.4 g, Ca 32
Uses In its native region the seeds ('yeheb mg, P 185 mg and Fe 6.4 mg (Leung, Busson &
nuts') of Cordeauxia edulis are an important Jardin, 1968). The protein content is consid-
food for pastoralists, especially as a famine erably less than that of most pulses, but the fat
food during drought periods. They are eaten content is higher. In various studies the pro-
fresh, dried, roasted or boiled. The seeds taste tein was found to resemble that of other pulses
sour when eaten fresh or dried, but have a in containing considerable and well-balanced
50 CEREALSAND PULSES

amounts of essential amino acids, especially

lysine (3.9-6.9%), and being deficient in me-
thionine. The seed lipids contain palmitic acid
(26-31%), stearic acid (12-13%), oleic acid (31-
32%), linoleic acid (25-30%) and linolenic acid
(traces). The seed oil is yellow. The seeds con-
tain trypsin inhibitors, which can be inacti-
vated by boiling. Cordeauxia edulis leaves from
Somalia have a low crude protein content (7.5—
11.8%), energy content (559-586 k j per 100 g
dry matter) and in vitro dry matter digestibil-
ity (27.2-39.8%). Furthermore they have a high
tannin content (2.5—2.7%), which reduces their
feed quality. The leaves contain N 1.2-1.5%, P
0.1%, Ca 0.7-1.8%, Mg 0.1-0.2% and S 0 . 1 -
0.2%. According to herdsmen the meat of ani-
mals fed with Cordeauxia edulis is particularly
tasty. Cordeauxia edulis is reputed to cause
intestinal disorders in goats when eaten as the
sole diet.
The red pigment of Cordeauxia edulis is cor-
deauxione (cordeauxiaquinone), a naphtha-
quinone which is unknown in other plants. The
leaves contain 0.7-0.8% ofthis dye. When fresh
leaves are handled, they stain the hands red.
When animals eat the leaves, their teeth are
stained orange-red and the dye is also depos- Cordeauxia edulis - 1, flowering twig; 2, fruit;
ited as a calcium complex in their bones, which 3,seed;4,seed kernel.
become pink. This colouration is considered a Redrawn and adapted by Achmad Satiri Nur-
sign of good meat quality, e.g. in Somalia and haman
Saudi Arabia. The wood of Cordeauxia edulis
has been described as good firewood, inflam- comprises a single species. It is closely related
mable even when wet. to Caesalpinia and Stuhlmannia. Cordeauxia
Description Densely branched, evergreen edulis is rather variable, and sometimes two
shrub or small tree up to 2.5(-4) m tall, with a types are distinguished: 'suley' ('sulei') and
long taproot up to 3 m deep and lateral roots at 'moqley' ('mogollo'). 'Suley' is pale green, with a
10-40 cm below the soil surface extending up large stem diameter and large leaflets. 'Moqley'
to 2.5 m; stem with conspicuous red glands. is dark green, with small stem diameter and
Leaves alternate, paripinnate, without stip- small leaflets. Pods of the 'moqley' type contain
ules; leaflets (2-)4-8(-12), elliptical-oblong, up a large single seed. Pods of the 'suley' type con-
to 3(-5) cm x 1.5(-2.5) cm, leathery, olive-green tain several seeds, which are compressed later-
above, paler with many red glands beneath. ally and smaller in size. The seeds of the 'mo-
Inflorescence a terminal few-flowered raceme. qley' type are claimed to be sweeter. Mixed
Flowers bisexual, almost regular, 5-merous, c. stands ofthe two types exist but are rare.
2.5 cm in diameter; sepals oblong, c. 1 cm long, The leaves of Cordeauxia edulis have an ex-
obtuse, green with red glands; petals almost tremely thick cuticle and mesophyll consisting
equal, c. 1.5 cm long, yellow, clawed; stamens of palisade cells with lateral walls capable of
10, free, straight, filaments hairy below the folding in a concertina-like way. This may en-
middle; ovary superior, 1-celled, shortly stalk- able the leaves to survive extended periods of
ed, densely glandular; stigma obtuse. Fruit an drought and store water quickly when avail-
ovoid pod, 4-6 cm x 2 cm, shortly stalked, with able, thus allowing them to remain evergreen.
a curved beak, opening by 2 hard valves, l(-6)- Growth and development Germination of
seeded. Seed ovoid, 2.0—4.5 cm long, with thin, Cordeauxia edulis is rapid. Subsequent growth
easily cracked testa. Seedling with epigeal of the aerial parts is very slow, especially in
germination; cotyledons thick. the seedling stage, whereas the root system
Other botanical information Cordeauxia grows rapidly. Plants 60 cm tall may already
 CORDEAUXIA 51

have roots 2 m long. Under natural conditions Diseases and pests Cordeauxia edulis
flowering starts just before the onset of the shrubs are essentially free of insect pests, but
rains, when the relative humidity rises, or im- storage pests, such as weevils and larvae of
mediately after the first rains, whereas some moths, heavily attack the seeds.
sources indicate flowering is year-round but Harvesting Cordeauxia edulis fruits are
more profuse during the rainy season. The flo- picked from the plant, the fruit wall is peeled
ral parts fall soon after pollination, leaving off and the seeds are placed in sacks. The seeds
only the fertilized ovary. The fruit ripens 10-15 are often harvested before maturity, which
days after flowering. Unlike in many other may be a factor in the low seed viability often
plants, seeds of Cordeauxia edulis mature encountered. Usually all seeds are removed
when the plant moisture content is at its peak. from the plant at the same time, hampering
Fruit development stops when the rain ceases regeneration of natural stands. Fruits can be
and the ovaries remain dormant for 4-5 harvested twice a year, provided rainfall is
months, resuming ripening after the rains have adequate during both rainy seasons.
returned. Cordeauxia edulis requires 3—4 years Yield The seed yield of Cordeauxia edulis is
to bear fruit. The plants are long-lived, some 5—8 kg per plant per year, but may be zero in
reaching more than 200 years according to drought years. The estimated average forage
Somali sources, and they coppice well. It is production is 325—450kg/ha (1.4-2 kg/plant).
unclear whether Cordeauxia edulis is able to Handling after harvest To prevent Cor-
fix atmospheric nitrogen. deauxia edulis seeds from being attacked by
Ecology In Ethiopia and Somalia Cor- insects, freshly picked seeds are roasted or
deauxia edulis is found in semi-desert regions boiled to kill insects and harden the seed coat.
in Acacia-Commiphora deciduous bushland In this form they fetch a higher price on the
and shrub vegetation at (100-)200-500(-1000) market, but the practice contributes to the
m altitude, with a minimum distance of 100 difficulty of obtaining viable seed for planting.
km to the Indian Ocean. These regions have an Pastoralists keep the seeds for many years in
average annual temperature of 26-30°C, an containers made of tanned, dried camel
average annual rainfall of 100-300(-400) mm leather. For dyeing, Somalis pulverize about
and two rainy seasons. 200 g of dried leaves in water to dye about 10
Cordeauxia edulis is resistant to normal m2 cotton cloth. Alkaline extracts develop a
drought periods of 4-5 months and occasional more intense violet colour than neutral or
drought of up to 15 months. It does not tolerate slightly acid extracts.
frost. It grows on deep, permeable, reddish, Genetic resources The populations of Cor-
sandy, slightly alkaline (pH up to 8-8.5), non- deauxia edulis declined in the 20 th century due
calcareous soils with a very low nutrient to deterioration of the vegetation caused by
status. Cordeauxia edulis does not tolerate overgrazing, and overharvesting of the seed. In
waterlogging. the IUCN Red List of Threatened Plants (1997)
Propagation and planting Cordeauxia edu- Cordeauxia edulis is classified in the category
lis is normally propagated by seed, but vegeta- 'rare', which includes taxa with small world
tive propagation through stem cuttings is also populations that are not at present considered
possible. The 100-seed weight is 100-300 g. The endangered or vulnerable, but are at risk. Re-
seed is often said to be viable for a few months generation and protection of natural stands
only, but seed coated in wood ash and stored in and cultivation in afforestation projects, in and
a sack is reputed to remain viable for at least a outside its native region, are recommended.
year. Direct sowing in the field seems prefer- Germplasm collections are seriously lacking,
able, as the fast-growing taproot is easily dam- with one accession kept in Ethiopia at the In-
aged in transplanting, with mortality rates of ternational Livestock Research Institute
up to 100%. No information is available on op- (ILRI), Addis Ababa, one in Kenya at the Na-
timum plant densities and spacings. Under tional Genebank of Kenya, Kikuyu and one in
natural conditions in Somalia there are up to the United States at the Southern Regional
320 plants/ha, depending on growing conditions Plant Introduction Station, Griffin, Georgia.
and distance from villages and water points. Prospects Cordeauxia edulis is a useful
Management Cordeauxia edulis is usually multipurpose plant in dry areas of Ethiopia
collected from the wild. Ample water is needed and Somalia. Natural stands are threatened by
for seedling establishment, but once the plants over-exploitation, and protection of natural
are established, little care is needed. stands as well as expansion of its cultivation
52 CEREALS AND PULSES

are called for. Cordeauxia edulis could bea leaflets alternate, elliptical, lanceolate or ob-
promising potential source of food and dry- long, 4-15cm x 1.5-7cm, base cuneate to
season fodder forother hot, arid regions. How- rounded, apex gradually acuminate, glabrous,
ever, the limited availability of viable seed and shiny. Inflorescence a many-flowered terminal
the shortage of knowledge onthecrop, espe- or axillary raceme 4-15 cm long, densely
cially its propagation, agronomic practices and brown-pubescent; bracts oblong, up to 5 mm
potential forselection and breeding, are impor- long, caducous. Flowers bisexual, papiliona-
tant constraints. ceous; pedicel up to 12(-19) mm long; calyx
Major references Assefa, Bollini & Kleiner, brown-pubescent, tube 4-5 mm long, lobes very
1997; Bally, 1966; Booth & Wickens,1988; broad, c.2 mm long; corolla bluish, pinkishor
Drechsel &Zech, 1988; Miège &Miège, 1978; white, 18-22 mm long, standard with a short
National Academy ofSciences, 1979; Seegeler, claw, wings andkeel with well-marked auri-
1983; Thulin, 1989a; Thulin, 1993; Yahya & cles; stamens 10, 9 fused and 1free, sheath 10-
Durand, 1991. 13 mmlong, free parts 5—6 mmlong; ovary
Other references Bekele-Tesemma, Birnie superior, shortly stalked, 10-11mm long,
& Tengnäs, 1993; Brenan, 1967; Curtis,Ler- densely hairy, style cylindrical, 5-6mmlong.
sten &Lewis, 1996; Drechsel, 1988; Drechsel & Fruit a shortly stalked, flat pod, 4.5-10 cm x
Assefa, 1991; El-Zeany & Gutale, 1982; Eug- 1.5—3.5 cm,asymmetric, shortly beaked, gla-
ster, 1967; Greenway, 1947; Hanelt & Institute brescent, dehiscing with twisted valves, 2-3-
of Plant Genetics and Crop Plant Research seeded. Seeds ellipsoid, c. 17mmx 15 mm,
(Editors), 2001; Hedberg, 1979; International black. Seedling with hypogeal germination.
Centre for Research in Agroforestry (ICRAF), Craibia comprises 10 species and is confinedto
undated; Kebebew, 1988; Lepidi, Nuti & tropical Africa.
Capretti, 1979; Leung, Busson & Jardin, 1968; Ecology Craibia brownii isfound indry and
Lewis, 1996; Miège, Crapon deCaprona & La- moist forest and along rivers, at 1100-2200m
cotte, 1978; Walter & Gillett (Editors), 1998; altitude, in areas with an annual rainfall of
Wickens, 1998; Zemede Asfaw & Mesfin Ta- 850-1400 mm.
desse, 2001;Zimsky, 1990. Genetic resources and breeding Craibia
Sources of illustration Booth &Wickens, brownii isnot threatened by genetic erosionas
1988; Thulin, 1983. it is widespread and locally common.
Authors M. Brink Prospects Since very little is known about
Craibia brownii, itsprospects are unclear. Re-
search isneeded on the nutritional and chemi-
CRAIBIA BROWNII Dunn cal properties ofthe seeds.
Major references Beentje, 1994; Gillett et
Protologue Journ. Bot. 49: 108 (1911). al., 1971; Hauman et al., 1954b; Lovett, Ruffo
Family Papilionaceae (Leguminosae -Papil- & Gereau, 2003; Troupin, 1982.
ionoideae, Fabaceae) Other references ILDIS, 2002; USDA, ARS
Synonyms Craibia elliottii Dunn (1911). & National Genetic Resources Program, 2001.
Origin and geographic distribution Craibia Authors M. Brink
brownii is found in north-eastern DRCongo,
Kenya, Uganda, Tanzania, Zambia andpossi-
bly in Rwanda. CROTALARIA KARAGWENSIS Taub.
Uses The seeds ofCraibia brownii are edible
after long cooking. The wood isused forbuild- Protologue Engl., Pflanzenw. Ost-Afrikas
ing poles, tool handles and wooden spoons, and C: 204 (1895).
as firewood and to make charcoal. Craibia Family Papilionaceae (Leguminosae -Papil-
brownii is also planted as ashade tree. ionoideae, Fabaceae)
Properties The wood of Craibia brownii is Synonyms Crotalaria lugardiorum Bullock
whitish and hard, and has afine texture. (1932).
Botany Small to medium-sized tree up to24 Origin and geographic distribution Crota-
m tall, with apale grey bark. Leaves alternate, laria karagwensis isdistributed inCentral and
imparipinnate with 3—8 leaflets; stipulesob- East Africa, from Cameroon to Ethiopia and
long, often semi-persistent; petiole 1.5-2.5 cm southward to DR Congo and Tanzania.
long, rachis up to 12 cm long; stipels sometimes Uses The seeds of Crotalaria karagwensis
present; petiolules l-2(-5) mm long, wrinkled; are considered edible inKenya.
 CROTALAEIA 53

Properties Various toxic compounds (alka- CROTALARIA LACHNOPHORA Höchst, ex

loids and non-protein amino acids) are present A.Rich
in Crotalaria spp., but toxin levels in Crotala-
ria karagwensis are not known. Protologue Tent. fl. abyss. 1: 151 (1847).
Botany Erect, annual herb up to 1 m tall, Family Papilionaceae (Leguminosae - Papil-
often with spreading, weakly ascending ionoideae, Fabaceae)
branches from the base; stem appressed hairy. Chromosome number n = 8
Leaves alternate, simple; stipules linear- Vernacular names Crotalaire à toison (Fr).
subulate, up to 3.5 mm long; petiole 1-2 mm Origin and geographic distribution Crota-
long; blade linear-lanceolate to elliptical, 1.5- laria lachnophora is widespread in tropical
11.5 cm x 2-12 mm, acute to rounded at apex, Africa, from Senegal east to Ethiopia and south
appressed hairy beneath. Inflorescence a ter- to Angola and Zimbabwe. It has recently been
minal or axillary lax raceme 9-24 cm long, (6-) introduced into Madagascar from Rwanda.
10-24-flowered. Flowers bisexual, papiliona- Uses The seeds of Crotalaria lachnophora
ceous; pedicel c. 5 mm long; calyx (4.5-)6-8 mm are considered edible in DR Congo. Crotalaria
long, upper lobes narrowly attenuate-tri- lachnophora is promoted in Rwanda as a green
angular, longer than the tube; corolla yellow, manure crop in rotation systems, together with
standard elliptical, c. 9 mm x 7 mm, with red- pigeon pea (Cajanus cajan (L.) Millsp.) and
dish-purple veins, wings c. 7 mm x 2—3 mm, Tephrosia vogelii Hook.f. In Madagascar it is
keel angular, 7-11 mm x 4 mm, with a long being used experimentally as a cover crop for
straight twisted beak; stamens 10, all joined; fallow land and in contour hedges. In Guate-
ovary superior, oblong, c. 3.5 mm long, 1-celled, mala it has been recommended as a shade
style c. 7 mm long. Fruit an oblong, club- plant for coffee plantations and for soil conser-
shaped pod, narrowed basally into a 2—3 mm vation. Leaf sap is dropped into the ear or
long stipe, c. 2.5 cm x 3.5 cm x 0.5 cm, 15-34- drunk to treat otitis.
seeded. Seeds obliquely heart-shaped, 1.5-4 Properties Various alkaloids and non-
mm in diameter, smooth. protein amino acids (y-glutamyltyrosine, isow-
Crotalaria comprises about 600 species dis- illardiine, 2-piperudinecarboxylic acid) have
tributed throughout the tropics and subtropics, been detected in Crotalaria lachnophora seeds
with about 500 species in tropical Africa. Cro- and may cause toxicity. However, amino acids
talaria karagwensis belongs to section Crotala- known to be toxic to mammals and birds and
ria, subsection Longirostres. In this subsection present in many Crotalaria species were not
levels of toxic compounds are in general rela- detected in Crotalaria lachnophora.
tively low, although most species contain the Botany Perennial herb or shrub up to 3 m
free amino acid y-glutamyltyrosine. tall, much-branched above; branches densely
Ecology Crotalaria karagwensis occurs at hairy. Leaves alternate, 3-foliolate; stipules
1100-2300 m altitude in grassland and wood- oblong-falcate, 1-2.5 cm x 2-8 mm, caudate;
land; it also persists on roadsides and in culti- petiole 0.5-3.5(-5) cm long; petiolules 1—2.5
vated land. mm long; leaflets oblanceolate to obovate, 3 -
Genetic resources and breeding No germ- 7.5(—10) cm x 0.5-3.5 cm, base cuneate, apex
plasm collections of Crotalaria karagwensis are acute to rounded, densely appressed pubescent
known to exist. In view of its wide distribution beneath. Inflorescence a terminal, lax raceme
Crotalaria karagwensis is not threatened by 10—30 cm long, few-many-flowered. Flowers
genetic erosion. bisexual, papilionaceous; pedicel 5-11 mm
Prospects It is unclear to what extent Cro- long; calyx 11-15(-18) mm long, spreading
talaria karagwensis seeds are eaten in tropical hairy, lobes twice as long as the tube; corolla
Africa. More information is needed on the lev- yellow, fading to orange-red, standard circular,
els of toxic compounds in the seeds and appro- c. 20 mm x 20-30 mm, wings broadly oblong,
priate processing methods to eliminate these 20-30 mm x 10—12mm, keel abruptly rounded
compounds. in lower half, (13-)20-24(-26) mm x 11 mm,
Major references Burkill, 1995; Gillett et with a rather short, blunt, slightly incurved
a l , 1971; Pilbeam & Bell, 1979; Polhill, 1982; beak; stamens 10, all joined; ovary superior, 1-
Thulin, 1989a. celled, style 20-23 mm long. Fruit a broadly
Other references Hepper, 1958; ILDIS, cylindrical pod, 2—4.5 cm x 1—2 cm, hairy, 16—
2005; Toussaint et al., 1953. 18-seeded. Seeds oblong to kidney-shaped, 4.5—
Authors M. Brink 5 mm long, granulate, orange-yellow.
54 CEREALS AND PULSES

Crotalaria comprises about 600 species dis-

tributed throughout the tropics and subtropics,
with about 500 species in tropical Africa. Cro-
talaria lachnophora belongs to section Chryso-
calycinae, subsection Stipulosae.
Ecology Crotalaria lachnophora occurs in
grassland and woodland, sometimes in thorn
scrub; it is also found on roadsides and in dis-
turbed or cultivated locations, at 900-2200 m
altitude. In Nigeria it occurs in regions with an
average annual rainfall of 500-1300 mm, on
acidic, ferruginous soils.
Genetic r e s o u r c e s a n d b r e e d i n g One ac-
cession of Crotalaria lachnophora from Kenya
is kept in the National Genebank of Kenya,
Crop Plant Genetic Resources Centre, KARI,
Kikuyu. Crotalaria lachnophora is widely dis-
tributed and not threatened by genetic erosion. Digitaria exilis -planted
P r o s p e c t s Although the seeds of Crotalaria
lachnophora are considered edible, possibly north-western Benin. It is also grown in the
toxic compounds have been detected in the Dominican Republic.
seeds. More information is needed on the toxic- Uses Fonio is a staple food in various parts
ity of the seeds and appropriate processing of West Africa, where it is also known as 'acha'
methods to eliminate the toxic compounds. or 'fundi', but it is also a prestige food ('chiefs
Crotalaria lachnophora has some potential as a food') and a gourmet item. In the Hausa re-
green manure. gions of Nigeria, Benin, Togo and Ghana, spe-
Major references Beentje, 1994; Burkill, cial couscous types ('wusu-wusu') are prepared
1995; Gillett et a l , 1971;Pilbeam &Bell, 1979; with fonio. In southern Togo, the Akposso and
Polhill, 1982. Akebou people prepare fonio with beans in a
O t h e r references Berhaut, 1976; du Puy et dish for special occasions. In Nigeria fonio flour
al., 2002; Hepper, 1958; Husaini & Gill, 1985; is made into thick, unfermented porridges
ILDIS, 2005; Moller, 1990; Neuwinger, 2000; ('tuwo acha'), and fermented grains are used
Thulin, 1989a; Toussaint et al., 1953; Wieden- for thin porridges ('kunu acha'). Boiled whole
roth, 1991. grains are eaten with vegetables, fish or meat.
A u t h o r s M. Brink In northern Togo, the Lamba people brew beer
('tchapalo') from fonio. It is also popped and can
be mixed with other flours to make bread. In
DlGITARIAEXILIS (Kippist) Stapf the Dominican Republic fonio flour is made
into porridges and creams, mixed with other
Protologue Bull. Misc. Inform. Kew 1915: cereal flours to make cookies, and it is used in
385 (1915). the preparation of candy and fermented bever-
Family Poaceae (Gramineae) ages; aside from everyday meals, fonio is also
Chromosome n u m b e r In = 54 associated with various religious festivities
V e r n a c u l a r n a m e s Fonio, hungry rice, white inherited from African ancestors.
fonio (En). Fonio, fonio blanc, petit mil (Fr). Fonio grain is a valuable, easily digested feed
Origin and geographic distribution Fonio for farm animals. The straw and chaff are ex-
is only known from cultivation and its exact cellent fodder and are often sold in markets for
origin is unknown, but it is of ancient cultiva- this purpose. Chopped fonio straw is mixed
tion in West Africa. It may have derived from with clay to build walls of houses. The straw is
Digitaria longiflora (Retz.) Pers. in the inland also used as fuel for cooking or to produce ash
delta region of the Niger. Historical records of for potash. Fonio grain is considered to have
the use of fonio as a cereal go back to the 14th medicinal properties; it is recommended for
century. Nowadays fonio is grown scattered lactating women and diabetic people.
from Cape Verde and Senegal to Lake Chad, Production and international trade Accord-
especially on the Fouta Djallon Plateau in ing to FAO statistics the average world produc-
Guinea, the Bauchi Plateau in Nigeria and in tion of fonio (the major share) and black fonio
 DlGITAEIA 55

(Digitaria iburua Stapf) together in 1999-2003

amounted to 257,000 t per year from 360,000
ha, all in West Africa. The main producing
countries are Guinea (128,000 t per year in
1999-2003, from 137,000 ha), Nigeria (78,000 t
from 142,000 ha), Mali (21,000 t from 33,000
ha), Burkina Faso (13,000 t from 16,000 ha)
and Côte d'Ivoire (11,000 t from 22,000 ha).
The production in the Dominican Republic is
not known. FAO statistics show an increase in
world production from around 180,000 t per
year in the early 1960s to around 260,000 t in
the early 2000s, with an increase in acreage
from around 280,000 ha to about 360,000 ha.
Fonio is hardly traded outside West Africa,
except for small quantities sold as luxury
product in Europe.
Properties The composition of whole fonio
grain per 100 g edible portion is: water 11.2 g,
energy 1390 k j (332 kcal), protein 7.1 g, fat 3.0
g, carbohydrate 74.4 g, fibre 7.4 g, Ca 41 mg, P
191 mg, Fe 8.5 mg, thiamin 0.24 mg, riboflavin
0.10 mg and niacin 1.9 mg (Leung, Busson &
Jardin, 1968). The essential amino-acid content
per 100 g grain is: tryptophan 111 mg, lysine
205 mg, methionine 441 mg, phenylalanine 402 Digitaria exilis - 1, plant habit; 2, spikelet; 3,
mg, threonine 315 mg, valine 457 mg, leucine grain-
772 mg and isoleucine 315 mg (FAO, 1970). Redrawn and adapted by Achmad Satiri Nur-
The amino acid composition of fonio is compa- haman
rable with that of other cereals, but it has a
relatively high methionine content. The palat- a taxonomically difficult genus comprising
ability offonio grain is considered high. about 230 species in tropical, subtropical and
Adulterations and substitutes Black fonio warm-temperate regions, particularly in the
and Guinea millet (Brachiaria deflexa (Schu- Old World. Digitaria barbinodis Henr., occur-
mach.) Robyns) are used as substitutes of fonio. ring in Mali and Nigeria, is harvested as a wild
Description Ascending, free-tillering an- cereal during times of scarcity and is occasion-
nual grass up to 80 cm tall, with delicate kneed ally grown in Nigeria. Digitaria ciliaris (Retz.)
stems. Leaves alternate, simple; leaf sheath Koeler is sometimes eaten as a supplementary
glabrous, smooth, striate; ligule membranous, food (Chad) or as a famine food. Digitaria debi-
broad, c. 2 mm long; blade linear to lanceolate, lis (Desf.) Willd., Digitaria fuscescens (Presl)
gradually tapering to an acute apex, 5-15 cm x Henrard, Digitaria leptorhachis (Pilg.) Stapf,
0.3-0.9 cm, glabrous. Inflorescence a terminal Digitaria longiflora (Retz.) Pers., Digitaria
digitate panicle of 2-5 slender, spike-like pri- nuda Schumach. and Digitaria ternata (A.Rich)
mary branches up to 15 cm long. Spikelet up to Stapf are also known to be eaten as famine
1 mm stalked, narrowly ellipsoid, 1.5-2 mm foods in tropical Africa, but are considered
long, acute, glabrous, pale green, 2-flowered; more important as forage or auxiliary plant. In
lower glume hyaline, minute, upper glume India (Assam) and Vietnam Digitaria cruciata
broadly oblong, slightly shorter than spikelet, (Nees ex Steud.) A.Camus ('raishan') is grown
hyaline between the 3-5 green veins; lower for food and fodder, whereas Digitaria sangui-
floret sterile, upper floret bisexual; lemma nalis (L.) Scop, ('crabgrass') is or was grown as
about as long as spikelet, 7-9-veined; palea a cereal in Europe, Asia and America.
slightly shorter than lemma; stamens 3; ovary Diversity within Digitaria exilis is broad, with
superior, with 2 stigmas. Fruit a caryopsis a large number of locally cultivated landraces,
(grain), oblong to globose-ellipsoid, c. 0.5 mm differing in plant habit, plant colour, glume
long, white to pale brown or purplish. colour, grain size and length of the crop cycle.
Other botanical information Digitaria is Based on morphology, 5 varieties have been
56 CEREALS AND PULSES

distinguished: can be grown on poor, shallow, sandy or rocky

- var. gracilis Portères: leaf margin curled, soils unsuitable for other cereals, but does not
inflorescence with 2 primary branches, each prosper in saline or heavy soils. On the Fouta
with 50-100 spikelets per 10 cm, spikelets in Djallon Plateau of Guinea, it grows on acidic
groups of (2-)3(-4) and in 1-2 rows, stalks soils with very high aluminium contents.
rough; early-maturing; Kankan region Propagation and planting Fonio is propa-
(Guinea). gated by seed. The 1000-seed weight is 400-
- var. striata Portères: leaf margin slightly 600 mg. Fonio is usually sown at the beginning
curled, inflorescence with 2 primary of the rainy season. Soil preparation is mini-
branches, each with 50—100 spikelets per 10 mal: the fallow vegetation is burnt and the
cm, spikelets in groups of (2-)3(-4) and ashes spread, and the soil may be loosened by
mostly in 1 row, stalks smooth; early- superficial cultivation. Seed, mixed with an
maturing; Casamance (Senegal), Guinea, equal quantity of sand or ashes, is usually
Mali and Burkina Faso. broadcast, and covered with soil by a light hoe-
- var. rustica Portères: robust plants, inflores- ing or brushing with tree branches. The seed
cence with (2—)3-4(—5) primary branches, rate is 10-30 kg per ha. Fonio is sometimes
each with 90—120 spikelets per 10 cm, raised in a nursery and planted out in the field.
spikelets in groups of (3—)4 and in 2—3rows, Fonio is normally grown as a sole crop, but
stalks smooth; late-maturing; Casamance sometimes intercropped with sorghum or pearl
(Senegal), Guinea, Mali and Burkina Faso. millet. Farmers in Guinea commonly sow vari-
- var. mixta Portères: robust plants, vegetative ous fonio types together and later fill in any
parts reddish pigmented, inflorescence with gaps with Guinea millet.
(2—)3-4(-5) primary branches, each with 90- Management Although it has been stated
120 spikelets per 10 cm, spikelets in groups that fonio seldom needs weeding due to its
of (3—)4and in 2—3rows, stalks smooth; late- quick establishment and the high seed rates
maturing; Guinea. applied, other sources indicate that a weeding
- var. densa Portères: tall, strong plants, with at 4-5 weeks after sowing is necessary for good
a long vegetative cycle, inflorescence with 3— yields. Fonio is usually not fertilized and little
4 primary branches, each with 120-140 is known of its nutrient requirements. In crop
spikelets per 10 cm, spikelets in groups of rotations fonio often follows rainfed rice, as a
2(-3) and in 2—3 rows; late-maturing; Togo. short-cycle crop before another crop is sown in
Growth and development Fonio normally the same season. It is also grown at the end of
germinates 2-4 days after sowing and grows a rotation.
rapidly. Flowering usually occurs 6—8 weeks Diseases and pests Fonio is susceptible to
after emergence. The time from sowing to ma- rust (Puccinia oahuensis). Resistance to the
turity is normally 2—5(-6) months. Certain nematodes Meloidogyne incognita and Meloi-
landraces mature so quickly that they produce dogyne javanica has been recorded in soils
grain already 6-8 weeks after planting, long where other plant species were infected. Birds
before all other cereals, and provide food early can cause serious losses, so bird scaring is usu-
in the growing season. At maturity the stems ally necessary. Fonio is attacked by parasitic
bend down due to the weight of the grains. plants of the genus Striga. Fonio seed is not
Fonio follows the C4photosynthetic pathway. liable to damage by storage pests and stores
Ecology Fonio is grown at sea level in Gam- well.
bia, Guinea-Bissau and Sierra Leone, but more Harvesting Fonio is usually cut with a knife
often it is cultivated at 600-1500 m altitude. or sickle, tied into sheaves, dried and stored
The average temperature in the growing sea- under cover. Mechanization is difficult because
son ranges from 20°C at higher altitudes to 2 5 - of lodging. When plants are dry, the grain
30°C near sea level. Fonio is grown in areas shatters easily, and therefore it is better to
with an average annual rainfall of 150-3000 harvest before the dry season has fully estab-
mm, but its cultivation is concentrated in re- lished and the relative air humidity has con-
gions with an average annual rainfall of 900- siderably declined. Harvesting is often stag-
1000 mm. It is not as drought resistant as gered, to suit the immediate needs of the
pearl millet, but fast-maturing landraces are farmer.
suited to areas with short and unreliable rains. Yield Grain yields of fonio are normally
In areas with very low rainfall it is grown in 600-900 kg/ha, but yields of over 1000 kg/ha
valleys benefiting from run-off water. Fonio have been recorded. In marginal areas yields
 DlGITAEIA 57

may be as low as 150-200 kg/ha. prevent lodging, photoperiod-sensitivity, culti-

H a n d l i n g after h a r v e s t Fonio is normally vation techniques, grain size, the development
threshed at about 8 days after harvesting, tra- of less laborious processing methods and im-
ditionally by beating or trampling. The husks provement of farmers' seed systems. Study of
remain on the grains, which therefore retain the genetic diversity of fonio and multiloca-
moisture and must be dried further. The grains tional screening of germplasm are also highly
are sufficiently dry when they run easily recommended.
through the fingers. The product after thresh- Major references Burkill, 1994; Froment &
ing ('fonio paddy' or 'raw fonio') is further proc- Renard, 2001; Haq & Dania Ogbe, 1995; Hilu
essed in 2 stages: husking (removal of the et al., 1997; National Research Council, 1996;
husks from the grains) and whitening (removal Ndoye & Nwasike, 1993; Portères, 1976; van
of the fruitwall and the germ). Husking and der Hoek &Jansen, 1996a; Vodouhè, Zannou &
whitening are done manually and require 4-5 Achigan Dako (Editors), 2003;Vodouhè &Achi-
beatings with pestle and mortar, alternated gan Dako (Editors), 2003.
with winnowing. To obtain a product of good O t h e r references Busson, 1965; Chevalier,
quality, all dirt and sand must be removed by 1950; Cissé, 1974-1975; Clayton, 1972; Cruz,
repeated washings. The processing cycle is 2004; de Wet, 1995c; FAO, 1970; Hanelt & In-
difficult and time-consuming and efforts are stitute of Plant Genetics and Crop Plant Re-
being made to develop equipment that will search (Editors), 2001; Harlan, 1993; Jideani,
make processing easier. Small-scale fonio proc- 1990; Jideani, 1999; Konkobo-Yaméogo et al.,
essing enterprises can be found in towns, e.g. 2004; Kuta et al., 2003; Kwon-Ndung, Misari &
in Mali and Burkina Faso, aiming at urban Dachi, 1998; Leung, Busson & Jardin, 1968;
and export markets. Lewicki, 1974; Morales-Payân et al., 2002;
Genetic r e s o u r c e s Fonio seems not threat- Purseglove, 1972; Sarr & Prot, 1985; van der
ened by genetic erosion. RAPD-analysis has Zon, 1992.
shown a relatively high level of genetic diver- Sources of illustration Henrard, 1950; Stapf,
sity in fonio compared to other millets, possibly 1916.
due to its outbreeding nature. Most germplasm A u t h o r s S.R. Vodouhè &E.G. Achigan Dako
collections made before 1990 and kept in na-
tional genebanks have been lost, but duplicates
exist at IRD (Institut de Recherche pour le DlGITARIAIBURUA Stapf
Développement), Montpellier, France, which
keeps more than 400 fonio accessions. Fonio Protologue Bull. Misc. Inform. Kew 1915:
accessions are also conserved in Senegal, 382 (1915).
Guinea, Mali, Burkina Faso, Togo, Benin and Family Poaceae (Gramineae)
Nigeria. In Nigeria, Benin and Togo germ- Chromosome n u m b e r In =54
plasm characterization work for a better un- V e r n a c u l a r names Black fonio, iburu, black
derstanding and utilization of the fonio gene acha (En).Fonio noir, manne noire, ibourou (Fr).
pool has started. Origin and geographic distribution Black
B r e e d i n g So far fonio has been largely ne- fonio is cultivated as a cereal in scattered lo-
glected in breeding programmes. Breeding ef- calities from Côte d'Ivoire to northern Nigeria
forts are being undertaken in Guinea, but no and southern Niger, and in Cameroon. It has
results are available so far. Improvement of also been reported to be grown in Guinea and
fonio through traditional hybridization does DR Congo. Black fonio is only known from cul-
not seem attractive because of insufficient tivation. Its origin is uncertain, but it may
knowledge on its floral biology and the ex- have been derived from Digitaria ternata
traordinarily miniature nature of its floral or- (A.Rich.) Stapf.
gans. Uses Black fonio is a staple food of the Bi-
P r o s p e c t s Fonio is wrongly named 'hungry rom people of the Jos Plateau in northern Ni-
rice', because it is not grown to relieve hunger geria and an important supplementary food to
but because of its quality and contribution to people in the Atakora mountains of Togo and
food security. It is a crop with a short cycle, Benin. It is eaten as porridge or mixed with
able to produce on very poor soils. It is appreci- meal of other cereals. The grain is also eaten
ated as a food in West Africa, and its nutri- cooked like rice or in stews. In Benin and Nige-
tional quality is excellent. Interesting research ria black fonio is made into couscous. In Togo it
topics include improved plant architecture to is used for brewing beer ('tchapalo').
58 CEREALSANDPULSES

P r o p e r t i e s The composition of whole black lected in Togo for the Institut Togolais de Re-
fonio grain per 100 g edible portion is: water cherche Agronomique (ITRA). Information on
10.3 g, energy 1436 kJ (343 kcal), protein 8.9 g, the genetic variation within the species and its
fat 3.0 g, carbohydrate 75.6 g, fibre 6.2 g, P 234 liability to genetic erosion is not available.
mg and Fe 10.0 mg (Leung, Busson & Jardin, P r o s p e c t s Black fonio is clearly less impor-
1968). The essential amino-acid content per tant than fonio, but is valued as a traditional
100 g grain is: tryptophan 215 mg, lysine 225 cereal in some parts of West Africa. Its impor-
mg, methionine 355 mg, phenylalanine 803 mg, tance is unlikely to increase, also in view of the
threonine 389 mg, valine 614 mg, leucine 1395 difficult husking. Little is known about its eco-
mg and isoleucine 508 mg (FAO, 1970). logical requirements, agronomy and potential
Botany Loosely tufted, erect, annual grass for genetic improvement, and research in these
up to 1.4 m tall, with glabrous stems. Leaves fields is recommended.
alternate, simple; leaf sheath glabrous, smooth, Major references Burkill, 1994; Haq &
striate; ligule membranous, rounded, broad, 2 - Dania Ogbe, 1995; Hilu et a l , 1997; Portères,
3 mm long; blade linear, tapering upwards, up 1976;van der Zon, 1992.
to 30 cm x 1 cm, glabrous except for some long Other references Adoukonou-Sagbadja et
hairs near the base. Inflorescence a terminal al., 2004; Busson, 1965; FAO, 1970; Hanelt &
digitate panicle of (2-)4—10(-11) sessile ra- Institute of Plant Genetics and Crop Plant Re-
ceme-like primary branches 12-14 cm long. search (Editors), 2001; Jideani, 1999; Leung,
Spikelet up to 2.5 mm stalked, elliptical- Busson & Jardin, 1968; National Research
lanceolate to oblong, up to 2 mm x 1 mm, Council, 1996; Prasada Rao & de Wet, 1997;
acute, glabrous, green to dark brown, 2- Stapf, 1917-1934; van der Hoek & Jansen,
flowered; lower glume hyaline, minute; upper 1996a.
glume ovate-oblong, 1-1.5 mm long, hyaline, 3- Authors M. Brink
veined; lower floret sterile, upper floret bisex-
ual; lemma of lower floret 7-veined, lemma of
upper floret brownish to black; palea slightly ECHINOCHLOAOBTUSIFLORA Stapf
shorter than lemma; stamens 3; ovary supe-
rior, with 2 stigmas. Fruit a caryopsis (grain), Protologue Prain, FI. trop. Afr. 9(4): 606
ellipsoid, 1.5-2 mm x 1mm. (1920).
Digitaria iburua mainly differs from its possi- Family Poaceae (Gramineae)
ble ancestor Digitaria ternata by its glabrous Chromosome number 2n = 18
spikelets. Digitaria iburua greatly resembles Origin and geographic distribution Echi-
Digitaria exilis (Kippist) Stapf (fonio). It is nochloa obtusiflora is only known from Niger,
called black fonio because of its dark spikelets, northern Nigeria, northern Cameroon and Su-
but its grain is white. dan.
Ecology Black fonio is grown at 400-1300 m Uses The grains of Echinochloa obtusiflora
altitude in areas with an annual rainfall of are eaten in Sudan (Kordofan) in times of food
900-1000 mm. It is credited with yielding a scarcity; they are collected from the wild. Echi-
crop where fonio fails due to drought. Though nochloa obtusiflora is a good forage plant.
it reputedly grows well on poor soils, it is Botany Erect, tufted, annual (sometimes
planted on more fertile soils in northern Nige- perennial) grass up to 1 m tall; stem (culm)
ria. erect or ascending. Leaves alternate, simple
Management The 1000-seed weight of black and entire; leaf sheath glabrous, smooth; ligule
fonio is about 500 g. In northern Nigeria it is ciliate; blade linear, 20-40 cm x 2-9 mm, hairy
usually planted towards the end of June and at base and margins or glabrous. Inflorescence
harvested in November—December. It is fre- composed of 2—12 racemes along a central axis
quently grown intercropped with fonio, pearl 5—17cm long; racemes erect, 1-4 cm long, with
millet or sorghum. Black fonio is difficult to spikelets in 4 rows along the rachis. Spikelet
husk and it is mostly eaten imperfectly elliptical, 2-3.5 mm long, obtuse, somewhat
cleaned. rough towards the tip, 2-flowered with lower
Genetic resources and breeding No germ- floret male and upper bisexual; lower glume
plasm collections or breeding programmes of obtuse, about one-third as long as the spikelet,
black fonio seem to exist, but germplasm of 2 upper glume as long as the spikelet; lemma
landraces ('Tchibam' and 'Tripka'), tentatively and palea of upper floret with incurved tip;
identified as Digitaria iburua, has been col- stamens 3; ovary superior, stigmas 2. Fruit a
 ECHINOCHLOA 59

caryopsis (grain). herds of pastoralists in West Africa. In Chad

Echinochloa comprises 30—40 species. It is a Echinochloa stagnina is sown to improve pas-
taxonomically difficult genus, because clear-cut tures; it is also sown as a fodder grass in
boundaries between the species seldom exist Egypt. It can bemade into hay. The stemsare
and the species are very variable. Introgression used for thatching and mat-making, and the
between species is common. Examined strains leaves for caulking boats. Theash of burnt
of Echinochloa obtusiflora were partially self- leaves has been used in the manufacture of
incompatible. soap and as amordant with indigo dye.
Ecology Echinochloa obtusiflora is found in Properties Echinochloa stagnina plants in
shallow pools, inundation plains and other wet mid-bloom in Niger contain crude protein
locations.Itis aweed of rice. 11.3%, crude fibre 32.5%, crude fat2.2%, nitro-
Genetic resources and breeding Although gen-free extracts 44.2%, Ca 0.31%, Mg 0.31%
Echinochloa obtusiflora has a limited distribu- and P 0.25%. Thepith ofthe culms contains
tion, itdoes not seem liable to genetic erosion. 10% saccharose and 7—8% reducing sugars.
Prospects Therole of Echinochloa obtusi- Because ofits high sugar content Echinochloa
flora as a cereal is limited to providing some stagnina is considered an excellent fodder
food in times of scarcity, andit is unlikely to grass.
become more important inthe future. Informa- Botany Perennial aquatic grass upto2.5m
tion islacking on the nutritional quality ofthe tall, ortaller (up to10m) when floating, with
grain. stout, often floating rhizomes; stem (culm) de-
Major references Burkill, 1994; Clayton, cumbent, with a diameter upto2.5cm, often
1972; van derZon, 1992; Yabuno, 1983; Yabu- spongy, rooting and branching at the lower
no, 1988. nodes. Leaves alternate, simple and entire; leaf
Other references Phillips, 1995; Stapf, sheath 15—25cm long, glabrous or rarely hairy,
1917-1934. loose at base of plant; ligule a line of hairs,
Authors M. Brink often absent inupper leaves; blade linear,10-
60 cmx 0.5-3 cm, firm, with scabrid margin
and filiform tip. Inflorescence composed of ra-
E C H I N O C H L O A STAGNINA (Retz.) P.Beauv. cemes along a central axis 6-35 cm long, erect
or nodding; racemes upto 15cmlong, closely
Protologue Ess.Agrostogr.: 53, 161, 171 overlapping ordistant, with spikelets in pairs.
(1812). Spikelets narrowly ovate, 3.5-6 mm x1-2mm,
Family Poaceae (Gramineae) slightly hairy but with prickly hairs onthe
Chromosome number 2n= 18, 36,54, 63, veins, 2-flowered with lower floret male or ster-
72, 108, 126 ile and upper bisexual; lower glume c. %of
Synonyms Echinochloa scabra (Lam.) Roem. spikelet length, sharply acuminate to mucro-
& Schult. (1817). nate, upper glume aslong asspikelet, awnless
Vernacular names Hippo grass, long-awn- or with an awnupto 4 mmlong; lemma of
ed water grass, burgu grass (En). Bourgou, lower floret with astout awn up to25(-50) mm
roseau sucré, roseau àmiel du Niger (Fr). long, lemma ofupper floret 3-5mm long; sta-
Origin and geographic distribution Echino- mens 3, anthers violet; ovary superior, stigmas
chloa stagnina occurs throughout tropical Af- 2. Fruit acaryopsis (grain).
rica and isalso found intropical Asia, whereit Echinochloa comprises 30-40 species. It is a
has possibly been introduced. Occasionally, it taxonomically difficult genus, because clear-cut
is naturalized in other tropical regions. boundaries between the species seldom exist
Uses Intropical Africa the grains of Echino- and the species are very variable. Introgression
chloa stagnina aretraditionally collected asa between species is common. Echinochloa stag-
cereal, especially in times of food shortage. nina is extremely variable.
Echinochloa stagnina is sown as a cereal in Stem elongation enables Echinochloa stagnina
India. The sweet stems and rhizomes have to support a water level increase of4cm per
been used toproduce alcoholic or non-alcoholic day, and it can befound inwater depths ofup
beverages and are still used forthe extraction to 4m.Inthe central Niger delta the biomass
of sugar for making confectionery and liqueurs. accumulated in theflooding season canbe as
Children suck the stems forthesugar. Grass- high as 15-30(-40) t dry matter per ha. Stems
lands ofEchinochloa stagnina ('bourgoutières') trampled byanimals and covered bysoil form
are important dry-season grazing areas forthe roots atthe nodes, which isanimportant mode
60 CEREALS AND PULSES

of natural regeneration of Echinochloa stag- but since then much has been replaced by rice
nina. Echinochloa stagnina is self-pollinating. fields. Around 1970 the area was estimated at
It follows the Ci-cycle photosynthetic pathway. 8000-10,000 ha. Since 1970 further reduction
Ecology In tropical Africa Echinochloa has taken place due to rice cultivation, reduced
stagnina occurs from sea-level up to 2300 m rainfall, reduced water levels in the river,
altitude, in shallow water, swamps and on pe- overharvesting and overgrazing, resulting in a
riodically inundated clay soils. It often forms disturbance of traditional pastoral systems.
large floating mats, rooting in the mud. Echi- The International Livestock Research Institute
nochloa stagnina is frequently the dominant (ILRI), Addis Ababa, Ethiopia, holds 9 acces-
species of the natural flood-plain grasslands in sions ofEchinochloa stagnina.
the central Niger delta and the shores of Lake Prospects Echinochloa stagnina is a useful
Chad. It may occur in massive, nearly pure multipurpose plant in semi-arid regions of
stands or together with Echinochloa colona (L.) West Africa, especially in the central delta of
Link, Echinochloa pyramidalis (Lam.) Hitchc. the Niger river. Its area and importance have
& Chase and Oryza longistaminata A.Chev. & declined due to various factors, and this trend
Roehr. will be difficult to reverse. In many other re-
Echinochloa stagnina is an important weed of gions Echinochloa stagnina is considered a
rice in tropical Africa, the Indian subcontinent weed, and therefore it does not seem advisable
and Thailand, sometimes obstructing water- to promote it elsewhere. Information is lacking
ways. on the nutritional quality ofthe grain.
Management Echinochloa stagnina is Major references Bonis Charancle, 1994;
propagated by seed, stem cuttings or plant Burkill, 1994; François, Rivas & Compère,
division. The 1000-seed weight is about 2.4 g. 1989; Harlan, 1989b; Phillips, 1995.
Under natural conditions the seeds are shed in Other references Bartha, 1970; Busson,
water. In experiments, seeds stored under wa- 1965; Clayton, 1989; François et a l , 1991;
ter in the dark at a temperature of 20CC Gibbs Russell et al., 1990; Hanelt & Institute
showed no dormancy and had a germination of Plant Genetics and Crop Plant Research
percentage of almost 100%, whereas seeds kept (Editors), 2001;McKenzie et al., 1993; van der
under dry conditions had a dormancy period of Zon, 1992;Yabuno, 1968;Yabuno, 1983.
6-7 months. The dormancy is broken by remov- Authors M. Brink
ing the glumes, but this results in rapidly re-
duced viability. Seeds germinate within a week
after sowing. In regeneration programmes in ELEUSINE CORACANA(L.) Gaertn.
Mali seedlings or rooted cuttings are planted
out into the field at densities of 10,000-16,000 Protologue Fruct. sem. pi. 1: 8(1788).
plants/ha. Family Poaceae (Gramineae)
In the central Niger delta in Mali the grains of Chromosome number 2n =36
Echinochloa stagnina are traditionally har- Synonyms Eleusine indica (L.) Gaertn.
vested using boats and by beating the inflores- subsp. coracana (L.) Lye (1999).
cences over a net. As the grains shatter easily, Vernacular names Finger millet, African
they are harvested at an early stage. To obtain millet, koracan (En). Eleusine, coracan, mil
sugar, the harvested plants are traditionally rouge (Fr). Luco, capim colonial, nachenim
dried in the sun, after which the leaves are (Po). Mwimbi, ulezi (Sw).
burnt off. The stems are washed and ground, Origin and geographic distribution Finger
and sugar is extracted from them by filtrating millet was domesticated in the East African
with warm water. Vegetative material for for- highlands. The oldest known archaeological
age is cut using boats, and is eaten green or as remains were excavated at Axum, Ethiopia and
hay. The forage is not only used locally, but date back an estimated 5000 years. These re-
also traded commercially, with an important semble types of highly evolved finger millet
market in Tombouctou. After the water has that are still grown in Ethiopia. Cultivation of
receded, animals are allowed to graze on the finger millet spread across the eastern and
remaining plant material until the end of the southern African savanna during the expan-
dry season. sion of iron working technology, to eventually
Genetic resources and breeding Around reach South Africa some 800 years ago. In
100 years ago, the 'bourgou' area in the central tropical Africa it is now grown from Ethiopia
Niger delta was estimated at about 250,000 ha, and Eritrea south to Mozambique, Zimbabwe
 ELEUSINE 61

plaiting, and in China for papermaking. In

Sudan the leaves are made into string.
Production and international trade In
trade and production statistics, finger millet is
usually combined with other millets, such as
pearl millet (Pennisetum glaucum (L.) R.Br.),
foxtail millet (Setaria italica (L.) P.Beauv.) and
proso millet (Panicum miliaceum L.). The es-
timated world area of finger millet is about 3
million ha yielding about 2.5 million t of grain
annually. India is the largest producer. The
total area planted annually in Africa is fairly
constant, and slightly less than 1 million ha.
Major producers in Africa are Ethiopia,
Uganda, Malawi and Zimbabwe. In Africa fin-
T" o?
ger millet grain is produced for local consump-
tion. Surpluses are sold in local markets. In-
Eleusine coracana -planted ternational trade, even among neighbouring
countries in Africa, is negligible.
and Namibia. It is also recorded from Mada- Properties Whole grain of finger millet con-
gascar. Finger millet is of little importance in tains per 100 g edible portion: water 10.9 g,
West Africa, but is recorded from a low-rainfall energy 1377 kJ (329 kcal), protein 7.4 g, fat 1.3
zone from Senegal eastwards, especially in g, carbohydrate 77.7 g, fibre 4.3 g, Ca 397 mg,
Niger and northern Nigeria. Finger millet P 190 mg, Fe 17.1 mg, ß-carotene traces, thia-
reached India 2000-3000 years ago. From In- min 0.18 mg, riboflavin 0.11 mg and niacin 0.8
dia it spread across South-East Asia to China mg (Leung, Busson &Jardin, 1968). The essen-
and Japan. In the United States it is grown on tial amino-acid composition per 100 g food is:
a small scale for bird-seed. tryptophan 107 mg, lysine 213 mg, methionine
Uses The principal use of finger millet in 229 mg, phenylalanine 383 mg, threonine 310
Africa is to provide malt for making local beer mg, valine 487 mg, leucine 701 mg and isoleu-
and other alcoholic or non-alcoholic beverages. cine 324 mg (FAO, 1970). Whole grain is used
In Ethiopia a distilled liquor known as 'areki' is in grinding, resulting in a high fibre content of
produced from finger millet. Finger millet is the flour, making it hard to digest. For food,
also widely used as a food cereal, especially white-coloured grain is preferred. The more
during times of scarcity. The most common use bitter dark-coloured grain is preferred for beer-
of finger millet flour is to prepare porridge, making. In malting, finger millet grain has a
usually served with a side dish of vegetables, higher enzyme activity than all other major
meat or fish. Freshly ground, slightly wet flour cereals except barley, making it very suitable
is made into 'cakes', which are wrapped in for brewing. Finger millet straw has an in-vitro
maize husks or banana leaves and roasted. digestibility of 40-60%.
Raw 'cakes' can be stored for several days; Description Robust, free-tillering, tufted
when needed, water is added to form a refresh- annual grass up to 170 cm tall; stem slender,
ing thin gruel. Flour is also pounded with ba- erect or geniculately ascending, glabrous and
nanas and the mixture is made into flat cakes smooth, sometimes branching, rooting at lower
that are fried in oil or baked in a dry pan. nodes; root system shallow, branched, fibrous.
Finger millet straw is used as forage for cattle, Leaves alternate, distichous, simple and entire;
sheep and goats. It produces excellent hay, and leaf sheath flattened, overlapping, split along
in India it is cultivated as a fodder grass. In the entire length; ligule 1-2 mm long, fimbri-
Uganda the by-products of finger millet beer ate; blade linear to linear-lanceolate, up to 75
production are fed to chickens, pigs and other cm x 2 cm, usually folded, scabrous above. In-
animals. Finger millet is used medicinally, e.g. florescence a terminal digitate panicle, often
the seed as a prophylaxis for dysentery. In with one or a few branches ('thumbs') below the
southern Africa the juice of a mixture of finger main cluster of 4-19 branches ('fingers');
millet leaves and leaves of Plumbago zeylanica branches slender to robust, linear to oblong, up
L. are taken as an internal remedy for leprosy. to 24 cm long, reflexed when slender or
Finger millet straw is used for thatching and straight to incurved at the tip when robust,
62 CEREALS AND PULSES

(subsp. africana (Kenn.-O'Byrne) Hilu & de

Wet) because it is also tetraploid (2n =36) and
crosses with finger millet produce fertile hy-
brids. It is an aggressive colonizer and forms
large continuous populations in disturbed habi-
tats, from where it is still harvested as a wild
cereal in times of scarcity. It is a noxious weed
of agriculture in Africa and invades fields of
finger millet where, although predominantly
self-fertilized, it occasionally crosses with the
cereal to form extensively variable, weedy hy-
brid swarms.
Primitive finger millet cultivars resemble wild
finger millet in inflorescence morphology but
lack the ability of natural seed dispersal. They
are characterized by inflorescences with
spreading branches that are straight or slightly
incurved at the tip when mature. These culti-
vars are widespread in Africa and are also
grown in southern and eastern India.
A second group of highland African and Indian
cultivars is also characterized by spreading
inflorescence branches. Cultivars from East
Africa typically have inflorescence branches
that are up to 24 cm long, while others from
Eleusine coracana - 1,stem part with leaves; 2, East Africa and from southern India have 10—
inflorescence; 3,part of inflorescence branch; 4, 18 cm long inflorescence branches. Cultivars
spikelet; 5, floret without lemma and palea; 6, from Africa commonly have more slender inflo-
grain within lemma and palea; 7,grain. rescences than those from India, allowing the
Source: PROSEA branches to become reflexed at maturity. Indi-
viduals in fields of these cultivars sometimes
sometimes with secondary branches, each have their spikelets arranged in clusters along
branch with 60—80 spikelets. Spikelets ovoid- the rachis.
ellipsoid, up to 10 mm x 4 mm, mostly ar- A morphologically distinct group of cultivars is
ranged in two rows along one side of the rachis, widely grown from Ethiopia to Zambia. These
(3-)6-9(-12)-flowered; lower glume 1-4 mm cultivars are characterized by spreading inflo-
long, with a 3-veined keel, upper glume 2-5 rescence branches with large, narrowly lanceo-
mm long, with a (3-)5-7-veined keel; florets late spikelets that are arranged in two even
bisexual, but terminal ones sometimes sterile rows along one side of the rachis. Morphologi-
or male, arranged in 2 opposite rows; lemma cally related finger millets are grown in the
narrowly ovate, 2-5 mm long, palea slightly mountains of eastern India, but in these culti-
shorter than lemma; stamens 3; ovary supe- vars the large spikelets are irregularly ar-
rior, with 2 free styles ending in plumose stig- ranged and essentially surround the rachis.
mas. Fruit a grain with free, soft fruit wall The most advanced cultivars have highly pro-
(utricle), 4-7 per spikelet, more or less globose, liferated inflorescence branches that are
up to 2 mm in diameter, white, red, brown or clumped together to form a fist-like structure.
black; pericarp remaining distinct during de- These cultivars are grown across the range of
velopment and at maturity appearing as a pa- finger millet cultivation in Africa and the In-
pery structure surrounding the seed. dian sub-continent. The most commonly grown
Other botanical information Eleusine com- finger millet cultivars in Africa and India have
prises about 10 species, distributed in the trop- much smaller inflorescences with more or less
ical and subtropical parts of Africa, Asia and spreading branches that may become some-
South America. The probable wild ancestor of what incurved or reflexed at maturity.
finger millet is Eleusine africana Kenn.- Growth and development Finger millet
O'Byrne (wild finger millet), commonly consid- seeds lack dormancy. However, they will not
ered as a subspecies of Eleusine coracana germinate in soil that lacks sufficient moisture
 ELEUSINE 63

to support seedling growth. Seedlings are sen- sown or planted 1-2 weeks before the expected
sitive to drought, but mature plants go dor- onset of rain.
mant during short periods of drought and pro- Finger millet is often intercropped with other
duce new tillers when conditions become fa- cereals, pulses or vegetables. In Ethiopia sole
vourable again. Plants tiller strongly and root cropping of finger millet is common. In Africa
from lower nodes, and provide excellent protec- finger millet is grown most commonly by small-
tion against soil erosion. Time from planting to holders.
flowering is 50-120 days; the complete crop Management Weeds are a major problem in
cycle is 3-6 months. Flowering on individual finger millet, the first two weeks after germi-
inflorescences lasts for 8-10 days and proceeds nation being critical. Several rounds of manual
from top to bottom on branches. Finger millet weeding are common, requiring much labour.
is predominantly self-pollinated, with about 1% When finger millet is planted in rows, animal-
out-crossing. Heavy rain at flowering reduces drawn weeders are often used. Finger millet
seed set. Finger millet follows the C4 photosyn- responds well to fertilizer. Recommended rates
thetic pathway. of application are 40-60 kg N, 26-40 kg P and
Ecology Finger millet grows best at an av- 30—50kg K per ha. Smallholders, however, can
erage temperature around 23°C. In eastern rarely afford chemical fertilizers. Finger millet
and southern Africa finger millet is grown from also responds well to the addition of organic
sea-level up to about 2500 m altitude, most manure or ash. In parts of Africa finger millet
commonly at 1000-2000 m. It is mostly grown is grown in a shifting-agriculture system, e.g.
in areas with 750-1200 mm of rainfall during in the 'chitemene' system in Zambia. In Kenya
the growing season. The minimum rainfall for and Tanzania it is often grown as the first crop
finger millet is 300-500 mm, but below 750 after clearing the land, when weed pressure is
mm sorghum and pearl millet are more com- low and soil fertility relatively high. Finger
monly grown because of their superior drought millet is commonly grown in rotation with
tolerance. Finger millet is a short-day plant other annual crops, preferably pulses. In
with a critical daylength mostly close to 12 Uganda it is grown after tobacco or cotton.
hours. Diseases and pests Finger millet is rela-
Finger millet grows on a range of soils, but tively free of diseases and pests. The most seri-
prefers fertile, well-drained, sandy to sandy- ous disease is head blast caused by the fungus
loam soils with reasonable water-holding ca- Magnaporthe grisea (synonym: Pyricularia
pacity. It prefers a pH of 5—7, but tolerates grisea). It attacks finger millet across its range
very alkaline (pH 11) soils. It does not tolerate of cultivation. All aerial parts are affected from
waterlogging. seedling to maturity. Serious reduction in yield
Propagation and planting Finger millet is occurs when inflorescences are infected during
propagated from seed. The weight of 1000 grain development. Control methods include
seeds is 2-3 g. Fields are prepared by hoe or crop rotation and the use of tolerant or resis-
animal-drawn plough. To control weeds, fields tant cultivars. Bipolaris nodulosa (synonym:
may be ploughed at the onset of rains, weeds Helminthosporium nodulosum) causes a dark
are allowed to germinate and the fields are brown leaf blight and foot and root rot,
ploughed a second time or even as many as six whereas Helminthosporium leucostylum causes
times, before the cereal crop is planted. Har- leaf shredding, seedling blight and head blight.
rowing before planting also helps to reduce Insect pests include shoot fly (Atherigona soc-
weeds. Seeds are broadcast or planted in rows catd), stem borers, caterpillars, grasshoppers
behind the plough. Seed rates up to 35 kg/ha and locusts; the phytophagous ladybird (Epila-
may be used when the crop is broadcast; in row chna similis) sporadically causes serious dam-
planting seed rates are only 3—10kg/ha. In row age. Quelea birds are pests in some areas. Ma-
planting, seeds are sown 2-3 cm deep in rows jor weeds of finger millet in tropical Africa in-
20-35 cm apart. As soon as convenient, seed- clude wild finger millet, Eleusine indica (L.)
lings are thinned to 5-12 cm apart within the Gaertn. and Brachiaria deflexa (Schumach.)
row. In India seeds are sometimes germinated Robyns. These species are difficult to distin-
in nurseries, and the seedlings planted in the guish from finger millet in early stages of de-
field when they are 3-4 weeks old. Although velopment and almost impossible to weed out
labour intensive, this practice provides fresh successfully. The broad-leafed weed Guizotia
cereal grain well before direct-sown finger mil- scabra (Vis.) Chiov. is a problem in Ethiopia,
let matures. Alternatively, finger millet may be but is commonly weeded out by hand. The root
64 CEREALS AND PULSES

parasite Striga hermonthica (Delile) Benth. diversity Conservation (IBC), formerly known
occurs across the range of finger millet cultiva- as Plant Genetic Resources Center of Ethiopia
tion in Africa, but rarely seems to cause serious (PGRC/E), in Addis Ababa, Ethiopia. Germ-
problems. Stored finger millet is insect- plasm from the rest of tropical Africa and trop-
resistant due to the grains being too small for ical Asia needs to be collected.
weevils to squeeze inside, and can be stored for Breeding Finger millet breeders need to
several years without serious damage. identify resistance to head blast and to incor-
Harvesting In Africa finger millet fields are porate such resistance into cultivars with ac-
often harvested in several rounds to prevent ceptable yield. Screening for resistance is mak-
loss of grain through shattering because of ing progress. Progress is also being made to
uneven ripening. Harvesting usually starts reduce susceptibility to lodging and shorten the
when grain of the earliest genotypes contains growing cycle, and to increase tolerance to
about 10% moisture. Inflorescences are indi- moisture stress and yield under traditional
vidually cut and allowed to dry. farming systems. Hand emasculation and pol-
Yield The average finger millet grain yield lination of finger millet florets are tedious and
under local practices of agriculture in tropical hamper rapid progress through trait recombi-
Africa is 0.25—1.5 t/ha. With improved culti- nation. A male sterile line has been developed
vars, optimal weed control and use of fertilizers in Zimbabwe. Improved cultivars released in
yields of up to 5 t/ha are obtained under ex- Africa include 'Tadesse', 'Padet' and 'Boneya' in
perimental conditions. Straw yields range from Ethiopia, 'P-283', 'P-224', 'P-221' and 'Serere-1'
1-2.5 t/ha for rainfed crops to 9 t/ha for irri- in Kenya, 'Engeny', 'Serere-1', 'Gulu-E' and 'P-
gated crops. 224' in Uganda, 'Steadfast' and 'M-144' in
Handling after harvest Finger millet grain Zambia and 'FMV-1' and 'FMV-2'in Zimbabwe.
is stored after threshing or in inflorescences Finger millet has limited levels of polymor-
that are threshed as needed. Threshing is usu- phism for DNA-based markers within the cul-
ally by beating the inflorescences with a stick. tivated and wild species. A genetic map based
Grain is ground on a grinding stone or in a on restriction fragment length polymorphism
mill. A little water may be added during the (RFLP) and amplified fragment length poly-
grinding process to keep the grains together morphism (AFLP) markers is being con-
and to avois fragmentation of the bran. The structed using crosses between finger millet
coarse bran is winnowed off and may be used and wild finger millet.
in making beer. Straw is commonly grazed by Prospects The area under finger millet cul-
cattle. In East Africa grain to be used in brew- tivation varies from year to year in both Africa
ing is typically soaked in water and left for 2-3 and Asia. The trend, however, shows stability
days to germinate, after which the germinated or increases in most countries where finger
seeds are ground, mixed with fried fermented millet is a staple cereal. A major constraint in
maize, sorghum or finger millet flour, and finger millet production is the high labour re-
placed in water to further ferment for 2-5 quirement, especially for weeding, harvesting
days. and milling. However, its excellent storage
Genetic resources Major finger millet germ- quality and the fact that in Africa it is pre-
plasm collections are being maintained and ferred over other cereals in the production of
evaluated by ICRISAT Asia Center (Patan- local beer assure finger millet a place in agri-
cheru, India), with selected duplicate speci- culture.
mens at ICRISAT Southern and Eastern Africa Major references Anand Kumar & Renard,
(Bulawayo, Zimbabwe and Nairobi, Kenya) and 2001; Bisht & Mukai, 2002; Burkill, 1994; de
SADC (Bulawayo, Zimbabwe). Germplasm Wet, 1995a; de Wet, 2000; de Wet et al., 1984;
collections include about 2800 accessions from Hilu & Johnson, 1997; Hilu, de Wet & Harlan,
Africa and about 2100 from Asia. Major African 1979; Jansen & Ong, 1996; Prasada Rao & de
collections are from Uganda, Zimbabwe, Kenya, Wet, 1997.
Malawi and Zambia, and most Asian collec- Other references Acland, 1971; Clayton,
tions are from India and Nepal. A large collec- Phillips &Renvoize, 1974; Cope, 1999; Davie &
tion (about 2000 accessions, mainly from Gordon-Gray, 1977; Dida, Gale & Devos, 2001;
Kenya) is maintained at the National Gene- Doggett, 1998; FAO, 1970; Harlan, de Wet &
bank of Kenya, Muguga. Another extensive Stemler, 1976; Hilu & de Wet, 1976a; Hilu &
collection, numbering 1300 accessions from de Wet, 1976b; Hussaini, Goodman & Timothy,
Ethiopia, is maintained by the Institute of Bio- 1977; Kennedy-O'Byrne, 1957; Leung, Busson
 ERAGROSTIS 65

& Jardin, 1968; McDonough, Rooney & Serna- silky hairs in the lower panicle axis. Further-
Saldivar, 2000; Ministry of Agriculture and more, Eragrostis pilosa is never glandular. In
Rural Development, 2002; National Research southern Africa Eragrostis aethiopica flowers
Council, 1996; Phillips, 1972; Phillips, 1995; from January toMay.
Riley et al. (Editors), 1993;Weher, 1991. Ecology Eragrostis aethiopica isfound upto
Sources of illustration Jansen & Ong, 1600 m altitude in semi-desert and savanna
1996. areas on sand, silt or clay, e.g. in floodplain
Authors J.M.J, de Wet grassland, small vleis, pan edges and banks
and beds of rivers, but also in disturbed habi-
tats, such as roadsides and cultivated land. It
ERAGROSTIS AETHIOPICA Chiov. is sometimes considered a weed, e.g. in Mo-
zambique.
Protologue Rob.-Brich., Somalia &Benadir; Management Eragrostis aethiopica is col-
726 (1899). lected from thewild.
Family Poaceae (Gramineae) Genetic resources and breeding The Na-
Chromosome number 2n= 20 tional Genebank ofKenya, Kikuyu, Kenya,and
Origin and geographic distribution Era- the USDA-ARS Western Regional Plant Intro-
grostis aethiopica is distributed from Eritrea, duction Station, Pullman, Washington, United
Ethiopia and Djibouti southwards to South States, each have 1 accession of Eragrostis
Africa. It is also found in the southern part of aethiopica. This species is widespread and in
the Arabian peninsula. many areas common and thus not liable to
Uses The grain of Eragrostis aethiopica is genetic erosion.
consumed in Ethiopia. In the Turkana area of Prospects The role of Eragrostis aethiopica
Kenya theplant iseaten bycattle, goats, sheep will not extend beyond being a local source of
and donkeys, butin Ethiopia it is consideredof food andforage.
little importance for grazing. Major references Clayton, Phillips & Ren-
Botany Annual grass upto 75cmtall, erect voize, 1974;Cope, 1999;Gibbs Russell et al.,
or ascending; stem (culm) slender, solitary or 1990; Phillips, 1995; Zemede Asfaw & Mesfin
tufted, often with pitted glands below the Tadesse, 2001.
nodes. Leaves alternate, simple; leaf sheath Other references Cope, 1995; Fröman &
glabrous; ligule a line ofhairs; blade linear, 3 - Persson, 1974; Holm, Pancho & Herberger,
20 cm x 1-3 mm, flat or involute, glabrous. 1979; IPGRI, undated; Morgan, 1981;SEPA-
Inflorescence a loose, open, ellipsoid panicle up SAL, 2003; USDA, ARS & National Genetic
to 26 cm long, branches and pedicels slender Resources Program, 2001.
and flexible, lowermost primary branches usu- Authors M. Brink
ally in whorls butsometimes solitary or paired.
Spikelet linear to oblong, 1.5—7 mm x 0.5-1
mm, 4-9(-28)-flowered, with bisexual florets; ERAGROSTIS ANNULATA Rendle ex Scott-
glumes unequal, hyaline, the lower veinless Elliot
and up to 0.5 mm long, the upper lanceolate,
up to 1mm long; lemma c. 1mm long, thinly Protologue Journ. Bot. 29:72(1891).
membranaceous, obtuse; palea with smooth Family Poaceae (Gramineae)
keel; stamens 3,anthers c.0.2mmlong; ovary Vernacular names Ringed lovegrass, ring
superior, with 2 stigmas. Fruit a caryopsis windgrass (En). Eragrostis annelé (Fr).
(grain), ellipsoid, c.0.5mmlong. Origin and geographic distribution Era-
Eragrostis is a large and taxonomically com- grostis annulata is found in Angola, Namibia,
plex genus comprising more than 350 species Botswana andSouth Africa.
mainly in tropical and subtropical regions, of Uses The grain of Eragrostis annulata is
which 14 are said to be endemic to Ethiopia. eaten in Namibia.
The diploid Eragrostis aethiopica is very simi- Properties Eragrostis annulata has anun-
lar to the tetraploid Eragrostis pilosa (L.) pleasant smell.
P.Beauv., a forage species ofwhich thegrain is Botany Annual, loosely tufted grass upto 40
sometimes eaten by humans. The former can cm tall; stem (culm) ascending, with a glandu-
be distinguished from the latter by its more lar ring below the nodes. Leaves alternate,
delicate habit, smaller spikelets with blunter simple; leaf sheath thinly hairy, the slender
lemmas, smaller grain, and absence of long hairs mixed with shorter gland-tipped hairs;
66 CEREALSAND PULSES

ligule aline ofhairs; blade linear, 2-12 cm x1 - E R A G R O S T I S NINDENSIS Ficalho & Hiern
5 mm, flat, thinly hairy, with scattered crater-
shaped glands along themargins and a lineof Protologue Trans. Linn. Soc. London,Bot.
glandular pits along themidvein below. Inflo- 2: 32 (1881).
rescence an ovoid panicle 4-20 cmlong, fairly Family Poaceae (Gramineae)
dense to open, stiffly branched, primary Synonyms Eragrostis denudata Hack.
branches not inwhorls, terminating ina fertile (1895).
spikelet. Spikelet on a pedicel 1-3 mm long Vernacular names Perennial lovegrass,
with a distinct annular gland, narrowly oblong wether lovegrass (En). Eragrostis vivace (Fr).
or sometimes linear, laterally compressed, 3— Origin and geographic distribution Era-
9(-15) mmx 1.5-2.5 mm, 6-16(-40)-flowered, grostis nindensis isdistributed from DR Congo
with bisexual florets; glumes almost equal, and Tanzania southwards to South Africa.
narrowly ovate, upto1mm long, keeled, apex Uses In Namibia the grain of Eragrostis
acute; lemma ovate to broadly ovate, 1.5-2 mm nindensis is eaten. Eragrostis nindensis is a
long, keeled, papery, glabrous, apex obtuse; palatable pasture grass and iswell utilizedby
palea glabrous on the sides, persistent; sta- sheep in particular. The young leaves are
mens 3, anthers 0.5-1 mmlong; ovary supe- sucked to treat colds.
rior, with 2 stigmas. Fruit a more or less Botany Perennial, tufted grass upto90cm
square caryopsis (grain) c.0.5 mm long, with a tall, with a short oblique rhizome; stem (culm)
shallow to deep depression along the back. erect, unbranched, glabrous at the nodes.
Eragrostis is a large andtaxonomically com- Leaves alternate, simple, mainly forming a
plex genus comprising more than 350species basal tuft; leaf sheath glabrous or with straight
mainly in tropical and subtropical regions. Era- silky hairs, terete; ligule a line ofhairs; blade
grostis annulata resembles theforage species linear, 5-30 cm x2-3 mm, involute, rarely flat.
Eragrostis cilianensis (All.) F.T.Hubb., but the Inflorescence a panicle 5-20cm long, ovoid
latter lacks gland-tipped hairs andhasa glo- with stiffly spreading primary branches, or
bose grain. narrowly lanceolate and densely contracted,or
In its native region Eragrostis annulata flow- linear and interrupted with the spikelets in
ers from February to May. clusters onstubby side branches, the primary
Ecology Eragrostis annulata is found ona branches not inwhorls, terminating ina fertile
range ofsoil types, especially onsandy, stony spikelet. Spikelet almost sessile, ovate to nar-
or calcareous soils where thegroundwater ta- rowly oblong, strongly laterally compressed,4—
bleis high, and indisturbed locations. 20 mmx 1.5-4mm, 7-30-flowered, dark yel-
Management Eragrostis annulata is col- lowish green todull grey, with bisexual florets;
lected from the wild. glumes almost equal, ovate, 1-2 mm long,
Genetic resources andbreeding TheDi- keeled, glabrous, apex acute; lemma ovate,2—
vision ofPlant andSeed Control, Department 3.5 mmlong, keeled, leathery, apex acute to
of Agriculture Technical Service, Pretoria, acuminate; palea oblong-elliptical, glabrouson
South Africa, holds 2 accessions of Eragrostis the sides; stamens 3, anthers 1—1.5 mmlong;
annulata. This species occurs ina diversityof ovary superior, with 2 stigmas. Fruit anellip-
habitats inafairly large region, and thus isnot soid caryopsis (grain) 1-1.5 mm long.
easily liable to genetic erosion. Eragrostis is a large and taxonomically com-
Prospects Therole of Eragrostis annulata plex genus comprising more than 350species
as asource of food is very limited and will most mainly in tropical and subtropical regions. Era-
probably remain so. grostis nindensis is a polymorphic species,
Major references Cope, 1999; de Villiers& varying widely in the shape ofthe inflorescence
Kok, 1988; Gibbs Russell etal., 1990; Klaassen and spikelet.
6 Craven, 2003; Launert, 1970. In southern Africa Eragrostis nindensis flowers
Other references IPGRI, undated; Mis- from October to June. It is a so-called 'resur-
souri Botanical Garden, undated; USDA, ARS rection plant', able to survive near-complete
& National Genetic Resources Program, 2001. desiccation ofits tissues. It retains mobile wa-
Authors M. Brink ter inits leaves even when dried naturally to
less than 20% water content. It also disassem-
bles chloroplasts when too dry to maintain
photosynthesis toavoid light-induced oxidative
stress. Young seedlings, however, aresensitive
 ERAGROSTIS 67

to drought. along the midvein. Inflorescence a narrowly

Ecology Eragrostis nindensis is found in oblong to narrowly ovoid panicle 10-35 cm
bare, exposed or disturbed locations at 600- long, branches ascending or spreading; primary
2400 m altitude, often on moist sandy and branches not in whorls, but sometimes loosely
stony soils and on granite outcrops. clustered, terminating in a fertile spikelet.
Management Eragrostis nindensis is col- Spikelet on a pedicel 1.5-2 mm long, linear to
lected from the wild. narrowly oblong, 6-13.5 mm x 0.5-2 mm, 9-13-
Genetic resources and breeding The In- flowered, with bisexual florets; glumes un-
ternational Livestock Research Institute equal, the lower 0.5-1 mm long, the upper 1-
(ILRI), Addis Ababa, Ethiopia, holds 3 acces- 1.5 mm long, keeled; lemma 2-2.5 mm long,
sions ofEragrostis nindensis. keeled, membranous with prominent lateral
Prospects The role of Eragrostis nindensis veins, olive green; palea with slender keel, per-
as a food or fodder plant will remain limited, sistent; stamens 3, anthers (1-)1.5-2 mm long;
although its ability to survive in dry conditions ovary superior, with 2 stigmas. Fruit an oblong
offers some prospect in semi-arid and arid re- to ellipsoid caryopsis (grain) c. 1mm long.
gions. Eragrostis is a large and taxonomically com-
Major references Clayton, Phillips & Ren- plex genus comprising more than 350 species
voize, 1974; Cope, 1999; Gibbs Russell et al., mainly in tropical and subtropical regions.
1990; Klaassen & Craven, 2003; van Oudts- Eragrostis plana flowers from September to
hoorn, 1999. May. It has the C4-cycle photosynthetic path-
Other references Balsamo et al., 2005;IPGRI, way.
undated; Mundree et al., 2002; USDA, ARS & Ecology Eragrostis plana occurs from 400-
National Genetic Resources Program, 2001; 2000 m altitude in grassland on sandy soils
vander Willigen et al., 2003. and shallow latérite pans, in dry areas on wet
Authors M. Brink soils around vleis and rivers. Its common oc-
currence in pastures is considered an indicator
ofovergrazing or too much burning.
ERAGROSTIS PLANA Nées Management Eragrostis plana is collected
from the wild.
Protologue FLAfr. austr. ill: 390 (1841). Genetic resources and breeding The USDA-
Family Poaceae (Gramineae) ARS Western Regional Plant Introduction Sta-
Chromosome number n = 10 tion, Pullman, Washington, United States, holds
Vernacular names Tough lovegrass, South- 3 accessions of Eragrostis plana, the Royal
African lovegrass (En). Eragrostis d'Afrique du Botanic Gardens, Kew, United Kingdom, 2
Sud (Fr). Capim choräo, capim teff (Po). accessions. This species is common in disturbed
Origin and geographic distribution In trop- areas and thus not liable to genetic erosion.
ical Africa Eragrostis plana occurs in Malawi, Prospects Eragrostis plana is eaten only in
Zambia, Zimbabwe and Mozambique. It is also times of famine and is a poor grazing grass.
found in South Africa, Lesotho and Swaziland. Therefore it is unlikely that it will become of
It has been introduced and has naturalized more than minor importance in the future.
elsewhere, e.g. in India and Brazil. Major references Cope, 1999; Gibbs Rus-
Uses The grain of Eragrostis plana is eaten sell et al., 1990; Jacot Guillarmod, 1971; van
as a famine food. Eragrostis plana is consid- Oudtshoorn, 1999;van Wyk &Gericke, 2000.
ered a poor grazing grass, but is utilized late in Other references Botha, 1992; IPGRI, un-
the rainy season in more arid regions. In Leso- dated; Neuwinger, 2000; O'Reagain & Grau,
tho it is woven into hats, baskets, necklaces 1995; Spies & Jonker, 1987; Steenkamp, 2003;
and bangles, and made into ropes and plaited USDA, ARS & National Genetic Resources
items used in funerals. In South Africa the root Program, 2001.
is used to treat menorrhagia and impotence. Authors M. Brink
Botany Densely tufted perennial grass up to
1 m tall, without rhizomes or stolons; stem
(culm) erect, unbranched. Leaves alternate,
simple; leaf sheath glabrous, strongly com-
pressed, keeled; ligule a line of hairs; blade
linear, 10—80 cm x 1.5—5 mm, flat or folded,
glabrous, sometimes with punctate glands
68 CEREALS AND PULSES

ERAGROSTIS TEF (Zuccagni) Trotter range of sizes and is consumed with various
sauces ('wot'), based on meat or pulses. Tef
P r o t o l o g u e Boll. Soc. Bot. Ital. 1918: 62 flour produces the best quality 'injera': pliable,
(1918). soft with glossy appearance, which does not fall
Family Poaceae (Gramineae) apart under handling or stick to the fingers,
Chromosome n u m b e r In =40 and has a slightly sour taste. Fenugreek
Synonyms Eragrostis abyssinica (Jacq.) (Trigonella foenum-graecum L.) can be added
Link (1827), Eragrostis pilosa (L.) P.Beauv. to tef flour in a small proportion to improve the
subsp. abyssinica (Jacq.) Asch. & Graebn. 'injera' flavour. It also increases the lysine con-
(1900). tent. Tef flour is also mixed with barley or sor-
V e r n a c u l a r n a m e s Tef, teff, teff grass (En). ghum flour to make 'injera'. Other traditional
Tef, teff (Fr). Tef(Po). preparations from tef flour include 'kitta'
Origin and geographic distribution Tef ori- (unleavened bread), 'atmit' or 'muk' (gruel),
ginated in northern Ethiopia, where it is porridge and local alcoholic beverages. Several
widely cultivated. Details of its domestication recipes that fit Western tastes have been de-
are unknown, but it may predate the introduc- veloped from tef flour particularly in the
tion of wheat and barley to the region. Tef is United States, where it has found niches in the
perhaps descended from the closely related health food market and as a gourmet food. Tef
wild Eragrostis pilosa (L.) P.Beauv., which is a flour is used as a thickening agent in a range of
tetraploid (2n = 40) annual like tef, and which products, including soups, stews, gravies and
has a cosmopolitan distribution. Grain cultiva- puddings.
tion of tef has been confined mainly to Ethiopia In Ethiopia tef straw is used as forage, espe-
and to some extent the highlands of Eritrea. It cially during the dry season. Mixed with clay it
is also grown in northern Kenya. Small-scale is used as plastering material for local houses
commercial tef production takes place in South and to make bricks, stoves, granaries, beds and
Africa, the United States, Canada, Australia, pottery.
Europe (the Netherlands) and Yemen. Tef is Outside Ethiopia tef is mainly grown for hay
grown as a forage grass, for instance in South (e.g. in South Africa) and as green fodder (e.g.
Africa, Morocco, Australia, India and Pakistan. in Morocco and India). In South Africa it is
It has been introduced experimentally into planted for erosion control, often in mixtures
other tropical countries, either for its grain or with Eragrostis curvula (Schrad.) Nees or other
for hay, e.g. in other parts of East Africa and in grasses.
southern Africa. It is commonly found as an Production and international trade In
escape from cultivation. 1992-1998 tef was annually cultivated on 1.9
Uses In Ethiopia and Eritrea tef flour is million ha in Ethiopia, which is about 30% of
mainly used to prepare 'injera', a thin, flat, the total acreage of cereals in the country. With
pancake-like bread, made from dough fer- an average annual production of 1.6 million t of
mented for 2-3 days. 'Injera' is prepared in a grain, tef constitutes 22% of the annual cereal
grain production in Ethiopia. Annually, an
average of 4 million t of forage (27%of national
production) is produced from tef. In Ethiopia
tef is grown by smallholders, mainly for the
local market and home consumption. Statistics
for 1997/98 and 1998/99 indicate that 1800 t of
tef grain was exported each year. Though re-
cent statistics are not available, there is an
export market for this crop in the Middle East,
North America and Europe, mainly for Ethio-
pian expatriates.
P r o p e r t i e s The composition of whole tef
grain per 100 g edible portion is: water 11 g,
energy 1407 kJ (336 kcal), protein 9.6 g, fat 2.0
g, carbohydrate 73 g, fibre 3.0 g, Ca 159 mg,
Mg 170 mg, P 378 mg, Fe 5.8 mg, Zn 2 mg,
thiamin 0.3 mg, riboflavin 0.2 mg, niacin 2.5
Eragrostis tef - planted mg and ascorbic acid 88 mg (National Research
 ERAGROSTIS 69

Council, 1996). The essentia] amino-acid com-

position per 100 g edible portion is: tryptophan
146 mg, lysine 273 mg, methionine 246 mg,
phenylalanine 474 mg, threonine 334 mg,
valine 491 mg, leucine 724 mg and isoleucine
378 mg (FAO, 1970). Tef starch grains are con-
glomerates of many polygonal simple granules
2-6 |lm in diameter. Their amylose content is
25-30%. The Kofler hot stage gelatinization
temperature range is 68°C (onset) - 74°C
(peak) - 80°C (conclusion), which is similar to
that of other tropical cereal starches, but nar-
rower than that of maize. The viscosity of the
starch is considerably lower than that of maize
starch, its water absorption index is higher,
and its water solubility index lower.
Due to the small size of its grains, tef is almost
always made into a whole-grain flour (bran and
germ included), resulting in a high nutrient
content. The amino acid composition of tef flour
is favourable and its protein is easily digesti-
ble. It is a good source of minerals, particularly
Ca and Fe, and tef has been implicated in the
low incidence of anaemia in Ethiopia. Tef does
not contain gluten, making it a suitable substi-
tute for wheat in foods for people with coeliac
disease. Several species of yeasts and bacteria Eragrostis tef- 1, upper part of flowering culm;
are involved in the preparation of 'injera', but 2,part of inflorescence with spikelets.
little is known about their identity and relative Redrawn and adapted byIskak Syamsudin
importance. In Ethiopia, white-grained types
are preferred for food, but consumption of 'in- pale green to dark purple; palea similar to
jera' from red- or brown-grained types is on the lemma, but with 2 veins; stamens 3, anthers
rise, especially for health-conscious urban peo- up to 0.5 mm long, 2-celled; ovary superior
ple. with 2 stigmas. Fruit a caryopsis (grain), ovoid
Tef straw is preferred by cattle over straw of to ellipsoid, 1-1.5 mm x 0.5-1 mm, yellowish-
other cereals, and its quality is comparable to white to deep brown.
good natural pasture. Analyses have shown a Other botanical information Little is known
relatively high digestibility (65%), but a rela- on the biosystematics ofEragrostis, a large and
tively low protein content (1.9-5.2%). taxonomically complex genus comprising more
Description Annual, tufted grass, up to than 350 species mainly in tropical and sub-
150(-200) cm tall, with a shallow, fibrous root tropical regions, of which 14 are said to be en-
system; stem (culm) usually erect, simple or demic to Ethiopia. Tef is the only Eragrostis
sparsely branched. Leaves 2-6 per culm, alter- species cultivated for its grain. The grains of
nate, simple; leaf sheath glabrous; ligule 0.5-1 several forage species are sometimes eaten by
mm long, ciliate; blade linear, 25-45 cm x 0 . 1 - humans, mainly as famine food, particularly
0.5 cm, glabrous. Inflorescence a panicle 10-65 Eragrostis cilianensis (All.) F.T.Hubb., Era-
cm long, with 10-40 slender primary branches, grostis ciliaris (L.) R.Br., Eragrostis curvula
very loose with central rachis fully exposed to (Schrad.) Nees, Eragrostis cylindriflora
very compact with central rachis completely Höchst., Eragrostis gangetica (Roxb.) Steud.,
hidden, with 30-1100 spikelets per panicle. Eragrostis pilosa (L.) P.Beauv., Eragrostis
Spikelet long-stalked, narrowly oblong, 4—9 tremula Steud. and Eragrostis turgida
mm x 1-3 mm, 2-12(-20)-flowered; florets bi- (Schumach.) De Wild.
sexual; glumes unequal, lanceolate, acuminate, Crossability relationships among the species
the lower 1-2.5 mm long, the upper 1.5-3 mm are largely unknown. Hybridization of tef is a
long; lemma 2-3 mm long, 3-veined, scaberu- tedious process which is a disincentive to mak-
lous on the keel and towards the acuminate tip, ing large numbers of crossing attempts. Era-
70 CEREALSANDPULSES

grostis tef is an allotetraploid of which the dip- weathered from volcanic ash under humid con-
loid progenitors are unknown. ditions). Vertisol-grown tef gives higher yields
Tef cultivars have been recognized and de- provided that prolonged waterlogging does not
scribed based on the colour of the grains and prevail and sufficient nutrients, particularly N,
inflorescences, ramification of the inflorescen- are available. Farmers usually alleviate the
ces and the size of plants. For marketing pur- effects of waterlogging by adjusting their plant-
poses, tef is classified on the basis of seed col- ing date or using surface drainage systems
our: 'netch' (white), 'tikur/ka'y' (red-brown) and (furrows). Micronutrient deficiencies can also
'sergegna' (mixed). The molecular variation as be limiting factors on Vertisols. Tef is normally
determined using DNA markers (RFLP, RAPD grown on soils of neutral pH, but it has been
and AFLP) is not commensurate with the mor- observed that it tolerates soil acidities below
phological variation. pH 5. Differences exist between cultivars for
Growth and development Germination of response to saline conditions. Tef can be found
tef normally takes place in 3—4(-12) days after as an escape from cultivation along roadsides
sowing. In experiments germination was above and railway lines, and in dry grassland on
90% at temperatures of 15-35°C; no germina- sandy loam soils.
tion occurred at 10°C. A booting stage is not Propagation and planting Tef is propa-
noticeable in tef: the inflorescences emerge gated by seed. There is no seed dormancy and
suddenly from the upper leaf sheath without germination is rapid. The 1000-seed weight is
boot formation. The flowers open in the morn- 200-500 mg.A single inflorescence can produce
ing (7—9am) in response to light and tempera- over 1000 seeds and a single plant over 10,000
ture. Tef is predominantly self-pollinating, seeds. Tef seeds remain viable for several years
with a very low degree of outcrossing (up to provided direct contact with moisture and sun-
1%), and pollen is set free in the early morning. shine is avoided. In Ethiopia centuries-old tra-
In the inflorescence floral maturity starts from ditional practices are used in tef production.
the top and progresses downward, whereas in An oxen-drawn plough ('maresha') is used to
the spikelet it progresses from the base up- till the land, with 2-5 passes made before sow-
ward. Seeds mature within a month after pol- ing. Studies indicate that tef can be success-
lination. The total growth cycle from sowing to fully grown under reduced tillage conditions
maturity is 2—5(—6)months. Tef follows the C4- (one ploughing to bring the seeds in contact
cycle photosynthetic pathway. with the soil) provided non-selective herbicides
Ecology Tef is a very versatile cereal and are used. To enhance germination and seedling
grows in a wide range of environments, from establishment on Vertisols, a firm seedbed is
sea-level up to 2800 m altitude. The highest made by trampling with farm animals. Nor-
yields are obtained at altitudes of 1800-2100 mally farmers sow tef by broadcasting on a
m, an annual rainfall of 750-850 mm, a sea- moist, fine seedbed. A seed rate of 15-30 kg/ha
sonal (July-December) rainfall of 450-550 mm is sufficient, but farmers often use 40-50
and an average daily temperature range of 15— kg/ha, because it is difficult to distribute the
27°C. Yields decline when the seasonal rainfall seed evenly, the viability of farmers' own seed
drops below 250 mm, the mean temperature is reduced, and it helps to suppress weeds at
during pollination exceeds 22°C, and when the early stages. The seeds are left uncovered or
growing period drops below 90 days, in which covered lightly by pulling branches over the
case early-maturing cultivars become neces- field using oxen. Tef can also be drilled in rows
sary. Despite its shallow root system, tef is using adjusted machinery. Row planting mini-
drought resistant, due to its ability to regener- mizes lodging under good growth conditions.
ate quickly after moderate water stress and to Sole cropping is the usual practice, but occa-
produce grain in a relatively short period. Its sionally early-maturing tef cultivars are used
rapid vegetative growth and short life cycle in intercropping systems, including relay- and
make tef particularly suitable for areas subject alley-cropping.
to drought after short rains. Flowering in tef is Successful in-vitro somatic embryogenesis and
delayed at long daylengths. In Ethiopia the plant regeneration procedures have been de-
bulk of tef production takes place during the veloped, using leaf, root or seed material to
main rainy season ('meher') between July and initiate callus cultures on Murashige and
November. Tef is mostly grown on Vertisols Skoog medium.
(dark, heavy clay soils with well-developed Management After crop establishment,
horizons) and Andosols (young, shallow soils, most farmers control weeds through hand
 ERAGROSTIS 71

weeding once or twice. Some farmers use her- a good indicator of maturity. If harvesting is
bicides such as 2,4-D to control broadleaf done after physiological maturity, shattering of
weeds, supplemented with hand weeding to seed is inevitable, particularly in windy and
remove grass weeds. On light soils the follow- rainy conditions. In Ethiopia harvesting starts
ing applications are recommended: 25-40 kg N in November and continues until early Janu-
and 10-18 kg P per ha; on heavy clay soils 50- ary. Harvesting is done by hand using sickles.
60 kg N and 10-15 kg P per ha. Tef responds to Farmers cut the plants at ground level, heap
N more than to P, by producing tall plants and them in the field and transport them to a
large amounts of biomass; as a result, high threshing ground. When grown for hay, tef can
N-rates promote lodging. To reduce the risk of normally be harvested 9—12 weeks after sow-
lodging, farmers reduce the N-application or ing.
plant tef after pulses with no additional fertili- Yield The average tef grain yield is less than
zation, and they delay the planting time so 1 t/ha, but farmers using improved cultivars
that rains have stopped by heading stage. Ro- and recommended management practices eas-
tation of tef with other cereals, pulses and Ni- ily get 1.7-2.5 t/ha. Yields over 2.5 t/ha have
ger seed (Guizotia abyssinica (L.f.) Cass.) is been recorded from several regions in recent
common practice in Ethiopia. extension programmes in Ethiopia. In experi-
Diseases and pests A number of diseases ments, grain yields up to 4.6 t/ha have been
(mainly caused by fungi) and pests are known obtained. Normal straw yields are about 3 t/ha,
to attack tef, but only a few are of economic but straw yields up to 20 t/ha have been re-
importance, mostly in specific localities and corded.
production years. Among the diseases, leaf rust Handling after harvest Threshing is car-
(Uromyces eragrostidis), head smudge (Helmin- ried out by trampling by farm animals. Some
thosporium miyakei) and damping off (Drech- farmers rent combines used for other cereals
slereispp. and Epicoccum nigrum) are the most for threshing. Tef is stored in any locally avail-
important. Low plant densities and early sow- able storage facility. Because tef is not at-
ing reduce the damage caused by leaf rust and tacked by storage insects, no chemical protec-
damping off, respectively. Fungicides that con- tion is required. Sometimes farmers even mix
trol these two diseases have been identified at tef seeds with pulses to protect the latter from
experimental level, although there are no weevils. Tef in Ethiopia is traditionally sold in
known cases of field applications. Breeding for the form of grains, not as flour. The straw is
resistance has not been carried out because of piled up near farmers' houses to be fed to ani-
limited genetic variation in resistance and the mals during the dry season; a small proportion
sporadic nature and environmental specificity may be sold.
of the diseases. No viral or bacterial diseases Genetic resources The Institute of Biodi-
are known. versity Conservation (IBC), formerly known as
Pests known to attack germinating tef seeds Plant Genetic Resources Center of Ethiopia
and seedlings include the Wollo bush-cricket (PGRC/E), holds 2541 tef accessions collected
(Decticoides brevipennis), the red tef worm from different agro-ecological regions and 1497
(Mentaxya ignicollis), grasshoppers, ants and accessions acquired through donations and
termites. The black tef beetle (Erlangerius ni- repatriations. IBC currently has no collection
ger) attacks the inflorescence. Among the of the wild Eragrostis spp. The majority of the
weeds, annual grasses cause the biggest dam- tef germplasm collections are conserved ex
age. The parasitic weed Striga hermonthica situ; seeds are dried to a moisture content of 3 -
(Delile) Benth., the recently introduced inva- 7% and stored in laminated aluminium foil
sive weed Parthenium hysterophorus L. and bags at —10°C for long-term and at 4°C for
the cosmopolitan weed Convolvulus arvensis L. short-term storage. In-situ conservation and
have also become problematic. Hand weeding enhancement programmes are used in some
and crop rotation, particularly with pulses, are regions of Ethiopia, primarily to help farmers
the most common methods in dealing with maintain crop diversity and to protect major
these weeds in tef; the use of herbicides is very cultigens from extinction while improving the
limited. Stored tef grains are not attacked by yield potential. Debre Zeit Agricultural Re-
storage insects, but rodents can be a problem. search Center, part of the Ethiopian Agricul-
Harvesting Tef is harvested 2-5(-6) months tural Research Organization (EARO), has se-
after sowing, when the vegetative parts turn lected 320 core-accessions that represent the
yellow. Yellowing of the stalk of the spikelet is phenotypic diversity of tef for genetic studies
72 CEREALSAND PULSES

and breeding purposes. Outside Ethiopia, has started to a limited extent in the United
smaller tef collections are held in Brazil States and Europe, targeting immigrant
(Centro de Pesquisa Agropecuaria dos Cerra- Ethiopian communities and its use as a glu-
dos (CPAC), Planaltina; 400 accessions), the ten-free substitute for wheat. There is reason-
United States (Western Regional Plant Intro- able optimism that, if investments are made in
duction Station, USDA-ARS, Washington State scientific and developmental research, tef can
University, Pullman), Germany (Federal Cen- rise to the status of a specialty crop in devel-
tre for Breeding Research on Cultivated Plants oped nations. Proneness to lodging is the big-
(BAZ), Braunschweig; 30 accessions) and Ja- gest drawback of tef; the use of appropriate
pan (National Institute of Crop Science, Tsu- machinery and agronomic practices may be
kuba; 30accessions). temporary solutions. In the long run, biotech-
Breeding Major breeding work on tef has nological approaches - using cloned dwarfing
been going on at Debre Zeit Agricultural Re- genes from other cereals - seem necessary to
search Center in Ethiopia since the 1960s.The arrive at non-lodging tef genotypes in the field.
main objectives have been the development of Increasing the inflorescence:culm ratio could
high-yielding cultivars for the major also be a suitable approach, although tef straw
tef-growing agro-ecological zones and the im- is important too.Knowledge ofthe influence of
provement of resistance to lodging. Conven- environmental factors on the nutritional qual-
tional breeding hasnot solved thelodging prob- ity oftef andofvariation in feed quality is still
lem. So far 15 cultivars have been developed very limited.
through direct selection from thelandraces and Major references Berhe, 1975;Deckers et
trait recombination. A crossing technique for al., 2001; Ebba, 1975; Ketema, 1997; Lovis,
the crop was developed in 1974and since then 2003; National Research Council, 1996; Phil-
hybridization of selected parents has resulted lips, 1995;Tefera, Belay & Sorrels (Editors),
in the release of 5 cultivars. The majority of 2001; Tefera, Ayele & Assefa, 1995;van der
farmers still grow landraces. Among the im- Hoek &Jansen, 1996a.
proved cultivars, 'Magna' (DZ-01-196), Other references Assefa, 2003; Assefa, Gaj
'Enatite' (DZ-01-354), 'Dukem' (DZ-01-974), & Maluszynski, 1998;Ayele et al., 1999; Baiet
'Tseday' (DZ-Cr-37) and'Ziquala' (DZ-Cr-358) a l , 2000; Bai et al., 1999; Bekele, Klöck &
are the most widely grown. Genotype-envi- Zimmermann, 1995; Bultosa, Hall & Taylor,
ronment interaction is high in tef production, 2002; Clayton, Phillips &Renvoize, 1974; Cope,
especially due to environmental effects on the 1999; FAO, 1970; Gibbs Russell et al., 1990;
time toflowering and maturity. Ingram & Doyle, 2003; Kebebew, Gaj &
Interspecific hybridization with wild Eragrostis Maluszynski, 1998;Kedir, Jones & Mengiste,
spp. has been tried, but success was obtained 1993; Lazarides, 1997;Lemordant, 1971a; Le-
only with Eragrostis pilosa; short-stature and mordant, 1971b; Mamo &Parsons, 1987; Mek-
early maturity were the favourable traits bib, Mantell & BuchananWollaston, 1997;
transferred to tef. Eragrostis curvula maypo- Tefera, Assefa & Belay, 2003; Vecchio, Simoni
tentially provide stalk strength and large seed & Casini, 1996;Yizengaw &Verheye,1994.
size, but hybrids with tef do not set seed. At- Sources of illustration Hanelt & Institute
tempts are being made to construct a genetic of Plant Genetics and Crop Plant Research
linkage map for tef. Anther/microspore culture (Editors),2001.
and subsequent breeding of double-haploid Authors H.Tefera &G. Belay
cultivars is also being attempted. Inter simple
sequence repeats are more promising than
other DNA-based markers for the quantifica- FAGOPYRUM ESCULENTUM Moench
tion ofgenetic diversity andidentification oftef
genotypes. Protologue Methodus: 290(1794).
Prospects In Ethiopia the expansion of tef Family Polygonaceae
to new production areas has continued un- Chromosome number 2n=16, 32
abated, despite farmers being encouraged to Vernacular names Buckwheat, beech wheat
cultivate other well-known cereals instead of (En). Sarrasin, blé noir (Fr). Trigo sarraceno,
tef. Tef cultivation has expanded to the low- fagópiro, trigo-mourisco(Po).
land areas, where sorghum and maize cultiva- Origin and geographic distribution Buck-
tion has frequently failed due to severe mois- wheat is native to central and northern Asia
ture stress. Outside Ethiopia, tef cultivation and was domesticated in south-western China
 FAGOPYRUM 73

(Yunnan, Sichuan provinces) from wild types. manure and cover crop, e.g. in Uganda.
For over a thousand years buckwheat has been Fresh leaves and inflorescences are used for
an important subsistence and cash crop from industrial extraction of rutin, which is applied
northern India and southern China to Korea to strengthen the inner lining of blood vessels
and Japan. In the early Middle Ages it was (however, it is rather the related species
introduced into Europe and became a leading Fagopyrum tataricum (L.) Gaertn., which is
crop on poor soils and an important staple food. commonly grown for rutin production). Rutin is
European emigrants introduced buckwheat in also industrially used as a natural pigment,
the United States and Canada. The increased antioxidant, stabilizer, food préservant and
use of chemical fertilizer in the beginning of absorber of UV light. In East Africa leaves are
the 20 th century led to an enormous decrease of chewed or its juice is drunk against fever.
the buckwheat area in Europe and North Production and international trade Accord-
America and replacement by higher yielding ing to FAO estimates, the average world pro-
crops such as rye, oats, maize, wheat, and Irish duction of buckwheat seed in 1999-2003
potato. Buckwheat is still important in India, amounted to 2.7 million t/year from 2.7 million
China, Korea, Japan and eastern Europe. In ha. The main producing countries are China
tropical Africa (e.g. DR Congo, Ethiopia, (1.4 million t/year from 980,000 ha), the Rus-
Uganda, Zimbabwe, Réunion) and South Africa sian Federation (600,000 t/year from 930,000
it is cultivated sporadically and it also occurs ha) and Ukraine (320,000 t/year from 440,000
as an introduced weed. ha). Buckwheat is mainly commercialized lo-
Uses The seed of buckwheat is cooked like cally. Average world export of buckwheat was
rice or made into flour for the preparation of only 160,000 t/year in 1998-2002, with as main
noodles, pancakes, porridge, cakes and biscuits. exporter China (104,000 t/year). No production
It is an ingredient in breakfast cereals. Groats or trade statistics are available for tropical
is the part of the seed left after hulling. Buck- Africa. At present, buckwheat is regaining
wheat seed is often ground or milled coarsely, some importance in Western countries because
to produce broken groats. Many consumers like of its excellent nutritional qualities. In Brazil,
the coarsely milled flour which is brownish Canada, the United States and South Africa
because of the high content of hull particles. At buckwheat is grown as an export crop on
present, high fibre content is considered a fa- highly mechanized farms.
vourable character, and buckwheat is gaining Properties The composition of buckwheat
importance as a health food. Buckwheat has a seed per 100 g edible portion is: water 9.8 g,
particular taste, liked by some, rejected by energy 1435 kJ (343 kcal), protein 13.3 g, fat
others. When sieved for almost white buck- 3.4 g, carbohydrate 71.5 g, dietary fibre 10.0 g,
wheat flour, the extraction rate is quite low Ca 18 mg, Mg 231 mg, P 347 mg, Fe 2.2 mg, Zn
(60-70%), the waste being used for fodder. Al- 2.4 mg, thiamin 0.10 mg, riboflavin 0.43 mg,
though pure buckwheat flour is sometimes niacin 7.0 mg, vitamin B60.21 mg, folate 30 |ig
used for baking bread, the absence of gluten and ascorbic acid 0 mg. The essential amino-
prevents the dough from rising. It is popular acid composition per 100 g edible portion is:
for use in mixtures with wheat, barley or rye tryptophan 192 mg, lysine 672 mg, methionine
flour to improve the taste and nutritional value 172 mg, phenylalanine 520 mg, threonine 506
of bread and other foodstuffs. Up to 30% of mg, valine 678 mg, leucine 832 mg and isoleu-
buckwheat flour may be mixed in wheat dough cine 498 mg. The principal fatty acids are per
for baking bread. In the Himalaya buckwheat 100 g edible portion: oleic acid 988 mg, linoleic
is processed into alcoholic drinks. acid 961 mg and palmitic acid 450 mg (USDA,
The seed is also fed to animals, especially pigs 2005). Whole buckwheat fruits are rich in fibre,
and chickens, and buckwheat is sometimes the hull providing most of it. Stored flour may
considered a fodder crop rather than a food become rancid because of the high fat content.
crop, e.g. in southern Africa. The plants are Buckwheat differs from true cereals in the high
occasionally used for silage, but must be mixed biological value of the protein, caused by the
with other fodders. The tender shoots make a high content of essential amino acids, in par-
palatable green leaf vegetable. Honeybees in ticular lysine. Due to the absence of gluten,
buckwheat fields produce a dark-coloured fra- buckwheat is suitable for the diet of people
grant honey. The fruit hulls are used as litter with coeliac disease. On the other hand, buck-
in poultry houses, for stuffing pillows, as fuel wheat seed is considered to be one of the most
or for compost. Buckwheat is grown as green important food allergens. It also contains com-
74 CEREALSANDPULSES

pounds which can cause irritating skin disor- celled, trigonous, with 3 styles ending in head-
ders ('fagopyrism') mainly in sheep and pigs shaped stigmas. Fruit a 3-sided nutlet, 5—7.5
and occasionally in humans in case of heavy mm x 3 mm, sometimes winged, grey-brown,
consumption and exposure to sunlight. Fago- dark brown to almost black, 1-seeded. Seed
pyrism has also been observed in humans after pale green turning reddish brown, slightly
consumption of buckwheat honey. It may also smaller than fruit.
affect cattle when fed pure buckwheat silage. Fagopyrum comprises about 15 species most of
The flavonoid rutin is present in all above- them from eastern Asia. Harpagocarpus, com-
ground plant parts (leaves, stems, inflorescence, prising a single species, Harpagocarpus snow-
fruit). It has antioxidative, anti-inflammatory denii Hutch. & Dandy from Central and East
and antihypertensive activity; it strengthens Africa, is closely related and should possibly be
the inner lining of blood vessels, reduces cho- included in Fagopyrum. Numerous landraces
lesterol levels, protects the blood vessels from and cultivars of Fagopyrum esculentum are
rupturing, and blood from forming clots. known, differing in fruit shape, adapted to
Botany Erect annual herb up to 120 cm tall summer or winter cultivation and comprising
with angular, hollow stem. Leaves alternate, special-purpose types for grain, fodder, vegeta-
simple and entire; stipules fused into a tubu- ble or medicine.
lar, short, truncate ocrea; petiole up to 10 cm At soil temperatures above 10°C the seed ger-
long in lower leaves, upper leaves almost ses- minates fast, and seedlings emerge within 7
sile; blade triangular, hastate or cordate, 2-10 days. The crop grows fast, reaching the full
cm x 2-10 cm, acute, 5-7-veined from the base. height of 60-100 cm in 4-6 weeks. Flower for-
Inflorescence an axillary or terminal cluster of mation starts 20 days after emergence, anthe-
flowers combined into false racemes. Flowers sis starts a week later and continues until
bisexual, regular, small, rose-red to white, het- complete senescence and death of the whole
erostylous; pedicel slender; tepals 5, 3-4 mm plant. Buckwheat is self-incompatible. Cross-
long, persistent; stamens 8, alternating at the pollination occurs by insects, mostly bees and
base with 8 honey glands; ovary superior, 1- flies. After the onset of flowering, the vegeta-
tive organs (leaves and stems) continue to grow
while fruits develop, hence seed ripening is
very uneven. From the middle of the flowering
period onwards, when the leaf area has
reached its maximum, further growth of vege-
tative organs is slow, and the seed becomes the
main sink for assimilates. The seed is ready for
harvesting 70-130 days after emergence, de-
pending on cultivar and ecological conditions.
Ecology Buckwheat is a crop of temperate
and subtropical areas, but may be grown suc-
cessfully at higher elevations in the tropics. In
Ethiopia it is grown at about 1500 m altitude.
Exact data on optimal temperatures for buck-
wheat cultivation are scarce, but climate de-
scriptions indicate a range of 18-30°C for day
temperatures, and night temperatures 5-10°C
lower. Because the leaf mass dries slowly, a dry
period is required at maturity and harvest.
Buckwheat is very sensitive to frost. Strong
winds cause lodging during crop growth and
seed shattering at maturity. Buckwheat is
rather sensitive to drought because of its
poorly developed root system. During flower-
ing, drought combined with high temperatures
Fagopyrum esculentum - 1, flowering branch; will cause poor seed set. Much rain during the
2, flower; 3, unwinged fruit; 4, winged fruit; 5, crop cycle stimulates vegetative growth, but
top view of winged fruit. inhibits seed setting, also because it hampers
Source: PROSEA pollination by insects. Buckwheat cultivars are
 FAGOPYRUM 75

either day-neutral or short-day plants. spora sp.), powdery mildew (Erysiphe polygoni)
Buckwheat performs best on nitrogen-poor and Rhizoctonia root rot {Rhizoctonia sp.) are
light sandy soils, from neutral to rather acid the major ones. Cultivars differ markedly in
(pH 4.5-7). It is suitable for newly cleared in- susceptibility. Several viral diseases have been
fertile land, drained marshland, rough land or recorded, but they do not cause much damage.
acid soils with a high content of decomposing Insect damage is rare, but grasshoppers, bean
organic matter. Buckwheat has the reputation weevils, cutworms, aphids, grain moths and
of producing an acceptable yield on marginal, storage beetles may feed on the crop. The worst
infertile land. On wet soils or soils rich in ni- problem for buckwheat production is damage
trogen, luxuriant growth leads to lodging, poor by birds at maturity and after harvest, when
fruit set, considerable losses during harvest, the crop is left to dry in the field. Rats are also
and thus reduced yields. When used for silage sometimes destructive.
or as green manure, a low seed yield is unim- When most (at least 75%) seed is mature and
portant, and more biomass will be produced on most leaves have yellowed and dropped, the
wetter, heavier soils. crop is harvested by mowing, after which the
Management Buckwheat is propagated by stems are bundled and put in heaps to dry.
seed. The 1000-seed weight is 12-35 g, averag- Farmers prefer to harvest early in the morning
ing about 22 g. Before sowing, the seedbed or late in the afternoon, or even at night, when
should be finely crumbed. Afirm soil at about 5 the plants are slightly damp from dew, to re-
cm depth reduces drought injury and lodging. duce grain shattering. The bundles are stacked
Very crusted land and heavy clay soil will re- alternately head-to-tail in the heaps, to reduce
sult in poor field emergence. Most growers use bird damage. If the leaves are not dry enough,
farm-saved seed. In mechanized cultivation, they may stick together, causing problems for
seed is drilled in rows about 30 cm apart, at a threshing. Combine harvesting is practised in
depth of 2-4 cm, requiring 40-60 kg of seed per more industrialized countries.
ha. The crop compensates for a thin stand by Seed yields normally vary from 0.6-2.5 t/ha,
branching more. Thin stands produce more but 3 t/ha is occasionally obtained. Research
inflorescences and seeds per plant. In manual has not succeeded in raising yields of buck-
cultivation, seed is broadcast, followed by har- wheat; they remain about the same as a cen-
rowing to cover the seed with topsoil. Broad- tury ago.
casting requires 10-20 kg seed more per ha Thorough drying to a moisture content below
than row drilling. Buckwheat is a crop with a 16% facilitates the removal of straw fragments
short growing season, easily fitting in cropping and immature seed. Small farmers usually
patterns with cereals, root crops, pulses, and thresh manually. Mechanical threshing re-
forages. It is sometimes intercropped with quires careful regulation of the threshing cyl-
vegetables. inder to avoid damaging the seed. Processing
Buckwheat competes well with most weeds, starts with hulling and separation of the hulls
but some fast-growing weeds can be a problem. from the groats, followed by milling. Formerly,
Some growers sow more densely on purpose, processing was done by individual households
and then weed mechanically by harrowing or in small village workshops. At present, most
about 4 weeks after emergence, killing most buckwheat is processed in factories that apply
weed plants together with a number of buck- advanced food technology to make specific
wheat seedlings. foodstuffs.
The uptake of minerals per ha for a seed yield Genetic resources and breeding The larg-
of 2t/ha is about 45 kg N, 10 kg P and 50 kg K. est collections ofbuckwheat germplasm are held
Growers usually apply no organic manure and in the Russian Federation (N.I. Vavilov All-
no or little chemical fertilizer, e.g. 10-30 kg N, Russian Scientific Research Institute of Plant
0-15 kg P and 15-30 kg K. Only P and K fertil- Industry, St. Petersburg, 2010 accessions),
izers should be applied if there is a risk of lodg- China (Institute of Crop Germplasm Resources
ing. In crop rotations with buckwheat, any crop (CAAS), Beijing, 1495 accessions) and Canada
is suitable as preceding crop provided that it (Agriculture Canada Research Station, Mor-
does not leave much nitrogen or weed infesta- den, Manitoba, 570 accessions). Germplasm is
tion. also available in national collections in the
Many fungal diseases have been recorded to United States, South Africa, Japan, Korea,
affect buckwheat, but they only occasionally India, Pakistan, Nepal, Slovenia, Poland and
cause serious damage. Downy mildew (Perono- Germany. All these countries are part of a net-
76 CEREALS AND PULSES

work under the International Plant Genetic Sources of illustration Grubben & Sie-
Resources Institute (IPGRI), responsible for monsma, 1996.
characterization and documentation. There are Authors P.CM. Jansen
numerous landraces and many have already Based on PROSEA 10: Cereals.
been collected for selection, testing and storage
in genebanks. Buckwheat is not threatened by
genetic erosion. GLYCINE MAX (L.) Merr.
Breeding of buckwheat has been carried out in,
for example, the United States, Russia, Japan, Protologue Interpr. Herb, amboin. 274
India and former Yugoslavia. Uniform, highly (1917).
self-compatible diploid lines have been isolated. Family Papilionaceae (Leguminosae - Papil-
They revealed a severe inbreeding depression, ionoideae, Fabaceae)
and heterosis in Fi generations. Breeders have Chromosome number 2n - 40
selected improved cultivars with higher yields, Synonyms Glycine hispida (Moench) Maxim.
e.g. by improving the plant habit (shorter (1873).
stems with reduced liability to lodging). Auto- Vernacular names Soya bean, soybean
tetraploid buckwheat selections show superior (En). Soja, soya (Fr).Soja (Po). Soya (Sw).
characters in many aspects (self-fertile, higher Origin and geographic distribution Soya
rutin content, increased dry matter production, bean was domesticated in the north-east of
improved nitrogen uptake, no seed shedding). China around the 11 thcentury BC. From there,
Improvement is also expected from breeding it spread to Manchuria, Korea, Japan and
programmes with close relatives of buckwheat other parts of Asia. Soya bean was introduced
such as Fagopyrum tataricum (L.) Gaertn. into Korea between 30 BC and 70 AD, and it
(Tatary buckwheat) and Fagopyrum homo- was mentioned in Japanese literature around
tropicum Ohnishi, e.g. to increase the rutin 712 AD. It reached Europe before 1737. Soya
content and to increase self-compatibility. So- bean was introduced into the United States in
matic embryogenesis of buckwheat is possible 1765 and into Brazil in 1882. It is unclear
using immature embryos, protoplasts, cotyle- when soya bean first reached tropical Africa.
dons, hypocotyls, leaf segments or stem seg- There are reports of its cultivation in Tanzania
ments as expiants. in 1907 and Malawi in 1909, but it is likely
Prospects Internationally, the interest in that soya bean was introduced during the 19th
buckwheat as a health food is increasing. With century by Chinese traders who were active
a higher price compensating for the lower yield along the east coast of Africa. Nowadays, soya
level compared to cereals, the acreage under bean is widely cultivated in tropical, subtropi-
buckwheat may increase. It is potentially an cal and temperate regions throughout the
interesting crop for marginal land in highland world. The slow distribution outside Asia is
areas in Africa, especially as a low-input sub- explained by the absence of soya bean specific
sistence or cash crop in rotation with other rhizobia in the soils of other regions; the crop
crops. An interesting feature is that at the
moment buckwheat is hardly affected by dis-
eases and pests. The main disadvantages of the
crop are lodging, seed shattering and low
yields. Given the existing genetic variability, it
is likely that breeding will result in cultivars
better adapted to tropical conditions, with less
lodging and seed shattering, and improved
seed set, hence with higher yield levels.
Major references Biacs et al., 2002; Camp-
bell, 1997a; Grubben & Siemonsma, 1996; Oh-
nishi, 1998;Zeiler &Hsam, 2004.
Other references Edwardson, 1996; Gu-
merova et al., 2003; Hedberg, 2000; Joshi &
Rana, 1995; Kim et al., 2005; Kokwaro, 1993;
Ohnishi & Asano, 1999; Sohn, Lee & Kim,
2003; USDA, 2005; Watt & Breyer-Brandwijk,
1962. Glycine max-planted
 GLYCINE 77

only developed in the United States at the be- Production and international trade Accord-
ginning of the 20 th century, following the dis- ing to FAO estimates, the average world pro-
covery ofthe nodulation process by scientists. duction of soya bean seeds is 173 million t/year
Uses In tropical Africa dry soya bean seeds from 77 million ha (mean of 1999-2003). The
are boiled for use in relishes, or used in the main producing countries are the United
preparation of milk substitutes and flour. A States (73.5 million t/year in 1999-2003, from
popular use of soya bean milk in Nigeria is to 29.4 million ha), Brazil (39.0 million t/year
make a tofu-like product that is deep fried and from 15.1 million ha), Argentina (26.4 million
sold as a snack or breakfast food. The flour is t/year from 10.2 million ha), China (15.4 mil-
used as a component of bread or mixed with lion t/year from 9.0 million ha), India (5.9 mil-
maize flour to make a fortified porridge ('ugali', lion t/year from 6.3 million ha), Paraguay (3.4
'sadza'). In West Africa soya bean flour is used million t/year from 1.3 million ha) and Canada
to thicken soup and to replace a traditional (2.3 million t/year from 1.0 million ha). South
flour that is made from the seed of egusi melon Africa produced 188,000 t/year from 121,000
(Citrullus lanatus (Thunb.) Matsum. & Nakai). ha. The soya bean production in tropical Africa
'Okara' is the pulp and bran left over from in 1999-2003 was 790,000 t/year from 895,000
making soya milk; this cake is used in almost ha, the main producers being Nigeria (439,000
all the same ways as soya bean flour. Soya t/year from 601,000 ha), Uganda (139,000
bean seeds are dry roasted and used directly as t/year from 124,000 ha) and Zimbabwe
a snack or as a coffee substitute. The seed is (119,000 t/year from 62,000 ha).
also milled into flour and mixed with maize Average world export of soya bean seeds
meal to serve as a relief food during famine. In amounted to 47.4 million t/year in 1998-2002,
Asia soya bean is used in the preparation of a the main exporters being the United States
variety of fresh, fermented and dried food prod- (25.4 million t/year), Brazil (12.3 million t/year)
ucts like milk, tofu, tempeh, miso, yuba, soya and Argentina (4.7 million t/year). Export of
sauce and bean sprouts (soya bean sprouts are soya beans from tropical Africa was only
meant here, and not mung bean sprouts, which 27,000 t/year, with Zimbabwe as main exporter
are more common in Western countries, and (11,000 t/year). The main importer was China
which are often called 'germes de soja' in (11.0 million t/year). Soya bean import in tropi-
French). Immature soya bean seeds are eaten cal Africa was 37,000 t/year. Average world
as a vegetable. export of soya bean oil in 1998-2002 was 8.2
Soya bean seed is processed to extract oil for million t/year, with as main exporters Argen-
food and for numerous industrial purposes; the tina (3.0 million t/year), Brazil (1.5 million
crop is currently the world's most important t/year) and the United States (0.9 million
source of vegetable oil.The edible oil enters the t/year). The export of soya bean oil from tropi-
market as cooking oil, salad oil, margarine and cal Africa was negligible. The main importers
shortening. Soya bean lecithins are used as in 1998-2002 were China (975,000 t/year), In-
emulsifier in the food industry, in pharmacy, dia (837,000 t/year), Iran (701,000 t/year) and
and in the industrial production of decorating Bangladesh (522,000 t/year). Soya bean oil
materials, printing inks and pesticides. Soya import in tropical Africa in 1998—2002
bean oil is the main commercial source of oe- amounted to 338,000 t/year, the main import-
tocopherol (natural vitamin E) and contains ing countries being Senegal (83,000 t/year),
stigmasterol, which is used for the commercial Angola (39,000 t/year), Mauritius (25,000
synthesis of steroid hormones and other phar- t/year), Madagascar (22,000 t/year) and Zim-
maceutical products. The cake remaining after babwe (22,000 t/year). Average soya bean cake
oil extraction is rich in protein and is an impor- export amounted to 40.8 million t/year, with as
tant animal feed. Uses of soya bean proteins in major exporters Argentina (13.6 million t/year),
food include defatted flours and grits, concen- Brazil (10.8 million t/year) and the United
trates, isolates, textured flours and textured States (6.4 million t/year). Soya bean cake ex-
concentrates (commonly used as meat ex- port from tropical Africa was 30,000 t/year,
tender). The protein is also used in the produc- mainly from Zimbabwe (14,000 t year) and
tion of synthetic fibres, glues and foams. Zambia (12,000 t/year). The main importers
Soya bean is also grown as fodder and as green were countries of the European Union. Tropical
manure; it is suitable for haymaking and silag- Africa imported 72,000 t/year.
ing. The leafy stems remaining after pod re- Soya bean is grown by smallholders in many
moval can alsobe used as fodder. countries of West, East and southern Africa,
78 CEREALS AND PULSES

though normally as a minor food crop. Com- nisms are not clear. There are also indications
mercial soya bean production on large farms that soya bean has a positive effect on bone
and estates is common in Zambia and Zim- health. The relation between soya bean con-
babwe, and also in South Africa. sumption and reduced risk of cancer is more
Properties The composition of mature raw uncertain.
soya bean seeds per 100 g edible portion is: Description Usually erect, bushy annual
water 8.5 g, energy 1742 kJ (416 kcal), protein herb up to 2 m tall, sometimes viny; taproot
36.5 g, fat 19.9 g, carbohydrate 30.2 g, dietary branched, up to 2 m long, lateral roots spread-
fibre 9.3 g, Ca 277 mg, Mg 280 mg, P 704 mg, ing horizontally to a distance of up to 2.5 m in
Fe 15.7 mg, Zn 4.9 mg,vitamin A 0 IU, thiamin the upper 20 cm of the soil; stem brownish or
0.87 mg, riboflavin 0.87 mg, niacin 1.6 mg, greyish pubescent. Leaves alternate, 3(—7)-
vitamin BÖ 0.38 mg, folate 375 (ig and ascorbic foliolate; stipules broadly ovate, 3-7 mm long;
acid 6.0 mg. The essential amino-acid composi- petiole 2-20 cm long, especially in lower leaves;
tion per 100 g edible portion is:tryptophan 530 leaflets ovate to lanceolate, 3-15 cm x 2-6(-10)
mg, lysine 2429 mg, methionine 492 mg, cm, base cuneate or rounded, apex acute to
phenylalanine 1905 mg, threonine 1585 mg, obtuse, entire, glabrous to pubescent. Inflores-
valine 1821 mg, leucine 2972 mg and isoleucine cence an axillary false raceme up to 3.5 cm
1770 mg. The principal fatty acids are per 100 long, often compact, densely hairy, (2-)5-8(-
g edible portion: linoleic acid 9925 mg, oleic 35)-flowered. Flowers bisexual, papilionaceous;
acid 4348 mg, palmitic acid 2116 mg, linolenic pedicel up to 3 mm long; calyx tubular, with 2
acid 1330 mg and stearic acid 712 mg (USDA, upper and 3 lower lobes, hairy; corolla 5-7 mm
2004). Soya bean seeds have a protein content long, white, pink, purple or bluish, standard
higher than any other pulse. The seeds have a obovate to rounded, c. 5 mm long, glabrous,
high lysine content; the limiting amino-acid is wings obovate, keel shorter than the wings;
methionine. Mature soya bean seeds are not stamens 10, 9 fused and 1free; ovary superior,
easily digested, contain toxic compounds and style curved with head-shaped stigma. Fruit a
have an unpleasant taste. Therefore they must slightly curved and usually compressed pod
be soaked and cooked for a long time before 2.5-8(-15) cm x 1-1.5 cm, hairy, dehiscent, (1-)
being edible, or be processed by techniques 2-3(-5)-seeded. Seeds globose to ovoid or rhom-
such as roasting, fermentation or sprouting. boid, 6—11mm x 5—8mm, yellow, green, brown
Heat-labile antinutritional factors of soya bean
are trypsin inhibitors, haemagglutinins, goi-
trogens, antivitamins and phytates, and heat-
stable ones are saponins, oestrogens, flatulence
factors and lysinoalanine.
Yield of meal from soya bean seeds is 80% and
of oil 18%. The meal contains about 50% pro-
tein. The average fatty acid composition of
commercial soya bean oil is: linoleic acid 54%,
oleic acid 22%,palmitic acid 10%,linolenic acid
10% and stearic acid 4%. Soya bean oil is rich
in vitamin E and contains 1.1-3.2% lecithins.
Soya bean seeds are always heat-treated before
oil extraction, because of the presence of
antinutritional compounds. Soya bean oil tends
to become rancid when exposed to air or light,
due to the instability of the linolenic acid. The
protein and oil concentrations of soya bean are
negatively correlated, and efforts to raise both
simultaneously have been unsuccessful. The oil
content tends to increase with increasing tem-
perature during growth, whereas the protein
content tends to decrease.
Consumption of soya bean is associated with Glycine max - 1, flowering branch; 2, fruiting
decreased risk of atherosclerosis and cardio- branch; 3, seeds.
vascular disease, although the exact mecha- Source: PROSEA
 GLYCINE 79

or black, or blotched and mottled in combina- Although older literature indicates that soya
tions of those colours; hilum small, black, bean is nodulated exclusively by slow-growing
brown or yellow. Seedling with epigeal germi- rhizobia (Bradyrhizobium spp.; initially called
nation; cotyledons thick and fleshy, yellow or 'cowpea-type rhizobia') it is now well estab-
green; first leaves simple and opposite. lished that the fast-growing Sinorhizobium
Other botanical information Glycine com- fredii can also form effective nodules with the
prises about 20 species distributed in the trop- crop. Soya bean genotypes differ enormously in
ics and subtropics of Asia and Australia. It is their ability to nodulate with indigenous rhizo-
divided into 2 subgenera; Glycine (perennials) bia in soils. The ability to nodulate spontane-
and Soja (annuals), with the latter including 2 ously and prolifically with indigenous rhizobia
species: Glycine soja Sieb. & Zucc. (wild types, is known as the 'promiscuous' character, com-
occurring in eastern Asia) and Glycine max pared with the 'specific' character of soya bean
(cultivated types). Glycine soja is considered types that normally require inoculation with a
the wild ancestor of Glycine max. The 2 taxa specific type or with a few specific types of
hybridize easily and may also be considered a rhizobia in order to grow well. However, it has
single species with 2 subspecies, Glycine max now been established that all soya bean geno-
(L.) Merr. subsp. max and subsp. soja (Sieb. & types nodulate to some extent with indigenous
Zucc.) Ohashi. rhizobia, but the diversity of strains with
Numerous cultivars are recognized in tropical which they can nodulate determines the extent
Asia that vary in time to maturity, size, plant of their promiscuity. Rates of N2-fixation in
habit, colour, content of oil and protein in the soya bean are greatest in the more luxuriant
seed, and uses to which they are put. For oil and late maturing genotypes. Studies con-
production, yellow seeds are preferred. For ducted in Nigeria have measured a N2-fixation
immature seeds to be used as a vegetable, rate of 126 kg of N per ha for an uninoculated
types with large yellow or green seeds are pre- late-maturing soya bean line.
ferred. Hay and fodder cultivars usually have Ecology Soya bean is grown from the equa-
brown or black seeds and the plants often tor to latitudes 55°N or 55°S, at altitudes from
twine. In tropical Africa the older cultivars close to sea level up to 2000 m. Although the
that originated from Asia tend to be tall and crop grows well under a wide range of tempera-
indeterminate in growth habit, take compara- tures, the optimum temperature for growth
tively long to mature (about 120 days) and are and development is in general around 30°C.
'promiscuous' in their ability to nodulate with Both excessively high (>32°C) and low (<20°C)
rhizobia indigenous to African soils. These cul- temperatures can reduce floral initiation and
tivars can be contrasted with soya bean culti- pod set. Soya bean requires at least 500 mm
vars that have emerged from breeding pro- water during the growing season for a good
grammes and tend to be short-statured, deter- crop; water consumption under optimal condi-
minate, and relatively fast-maturing (70-90 tions is 850 mm. Drought stress during flower-
days). ing reduces pod-set but drought during pod-
Growth and development Soya bean seed- filling reduces yield even more. Soya bean can
lings emerge within 5—15 days after sowing; a tolerate brief waterlogging but weathering of
seedbed temperature of 25-33°C is optimal. seed is a serious problem under humid condi-
Flowering starts from 25 days to more than tions. Soya bean is considered a quantitative
150 days after sowing, depending on daylength, short-day plant, but some cultivars are insensi-
temperature and cultivar. Flowering can take tive to photoperiod. The response to photope-
1—15 days. Soya bean is normally self- riod interacts strongly with temperature, and
pollinated and completely self-fertile with less given the relatively small variation in day-
than 1% cross-pollination. Pollen is normally length in the tropics, temperature is the major
shed in the morning, before the flowers have factor influencing the rate of phenological de-
completely expanded. At higher altitudes with velopment. The photoperiod sensitivity means
lower temperatures, flowers are usually cleis- that types brought directly into tropical Africa
togamous. The time from flowering to pod ma- from North America will often flower and set
turity is 30-50 days. The total crop cycle from seed before they have fully developed, restrict-
sowing to maturity is 65-200 days. Develop- ing their yields.
ment to maturity is usually shorter with short Soya bean grows well on most soils, except very
days than with long days. The number of pods coarse sands. The optimum pH is 5.5-7.5, and
per plant varies from a few to more than 1000. soya bean is sensitive to soil acidity, in particu-
80 C E R E A L S AND PULSES

lar to aluminium toxicity. Where soya bean has treatment (e.g. with thiram), deep-ploughing of
not grown previously, or where P is limiting, crop residues, crop rotation and application of
symbiotic N2-fixation may be inadequate to fungicides. Resistant cultivars are available.
meet the N requirement ofthe plants. Purple seed stain and leaf blight are caused by
Propagation and planting Soya bean is Cercospora kikuchii, also occurring worldwide.
propagated by seed. The 1000-seed weight is Recommended control measures are crop rota-
100—250 g. The seed can be sown before the tion, the use of clean seed, ploughing back of
start of the rainy season, or when the soil is crop residues, spraying with fungicides and the
moist. Seed rates are 40-120 kg/ha. Soya bean use of tolerant cultivars. Among the bacterial
is sown in rows (20-)40(-75) cm apart. Within diseases of soya bean, bacterial blight (Pseu-
the rows, 2-3 seeds are sown in hills spaced at domonas syringae pv. glycinea, synonym:
7.5-10 cm intervals, at a depth of 2-5 cm. With Pseudomonas savastanoi pv. glycinea) is com-
intercropping, sowing rates are less than for mon wherever soya bean is grown. Control
sole cropping. In traditional agriculture the practices of this foliar disease include the use
land is prepared by hand or animal traction of resistant cultivars, the use of clean seed,
before sowing. Soya bean is grown mainly on crop rotation and burying of crop residues.
the flat, but sowing on hills or ridges may be Bacterial pustule (Xanthomonas campestris pv.
practised where the soil is heavy, the water glycines, synonym: Xanthomonas axonopodis
table high, or rainfall heavy. Small-scale farm- pv. glycines) is also widespread. It is seed-
ers in tropical Africa grow soya bean as a sole transmitted and survives on crop debris. Con-
crop or in mixed cropping with maize, sorghum trol measures are similar to those of bacterial
or cassava. blight. Virus diseases of soya bean include soya
Management Soya bean is usually weeded bean mosaic virus (SMV), cowpea mild mottle
1-3 times during the first 6-8 weeks after virus (CPMMV) and bean yellow mosaic virus
planting, after which its canopy should be suf- (BYMV), but these are of little importance in
ficiently developed to suppress weeds. Irriga- tropical Africa.
tion is uncommon except for dry season produc- Soya bean cyst nematode (Heterodera glycines)
tion. Basal fertilization with 20-25 kg P per ha and root-knot nematodes (Meloidogyne spp.)
is often required for adequate symbiotic N2- can cause severe damage, especially on sandy
fixation and general growth. Soya bean is soils. Therefore, soya bean should not be grown
commonly grown in rotation with cereals, such continuously or in rotation with other suscepti-
as maize, rice, sorghum, wheat and finger mil- ble crops, such as tobacco. Soya bean cultivars
let, whereby all fertilizer may be applied to the resistant to nematodes are available.
cereal. The most widespread and probably most seri-
Diseases and pests Various fungal diseases ous pest of soya bean in tropical Africa is the
affect soya bean. Soya bean rust (Phakopsora southern green stink bug or soya bean green
pachyrhizi and Phakopsora meibomiae) is a stink bug (Nezara viridula), of which the
devastating disease that can reduce yields by nymphs and adults feed on soya been seeds.
as much as 90%. It is widespread; in tropical Control is by using insecticides. The most im-
Africa it is recorded from Sierra Leone, Ghana, portant leaf-eating pest is probably the soya
Nigeria, DR Congo, Uganda, Tanzania and bean looper (Xanthodes graellsii). Bean flies
Zambia. Partial resistance has been found in (mainly Melanagromyza sojae and Ophiomyia
various cultivars; fungicides may reduce dam- centrosematis) can cause complete yield loss.
age. Red leaf blotch (Dactuliochaeta glycines, Soya bean seedlings are occasionally damaged
synonym: Pyrenochaeta glycines) is confined to by cutworms (Agrotis spp.). No major storage
Africa; it is economically important in Zambia pests are recorded from Africa, except rodents.
and Zimbabwe, where yield losses of up to 50% Harvesting Mature seeds of early-maturing
have been recorded. Seeds are not infected, but soya bean cultivars can be harvested 65 days
the fungus can survive in the soil for many after planting; late maturing cultivars may
years. Tolerant cultivars have been developed need more than 150 days. In tropical Africa the
in Zimbabwe. Frogeye leaf spot (Cercospora plants are generally allowed to dry in the field
sojina, synonym: Passalora sojina) occurs and the whole plants (above ground) are col-
worldwide. It is primarily a leaf disease, but it lected by hand when most leaves have turned
may also affect stems, pods and seeds. It sur- yellow and fallen, and when the pods have
vives on stored seeds and crop residues and is turned brown. The moisture content of the
spread by wind. Control measures include seed seeds at harvesting should be 14-15%. Pods of
 GLYCINE 81

older cultivars have a tendency to shatter in used for the improvement of resistance to dis-
the field when drying and plants need to be eases and pests, plant morphology and seed
harvested on time to prevent major loss of composition. The genetic variation of soya bean
yield. Combine-harvesting is used on large cultivars is rather narrow. For instance, about
farms and estates. Soya bean seed for vegeta- 80% of the genepool of the soya bean cultivars
ble use is harvested when the pods are still grown in the United States can be traced to
green but the seeds fill the pod. only 7-10 introductions from the same geo-
Yield Average world soya bean yields are graphical area. It is therefore considered nec-
2.25 t/ha; those in the United States 2.5 t/ha. essary to broaden the genetic base of cultivated
Under smallholder farming conditions in tropi- soya bean by using wild relatives.
cal Africa yields are often as low as 0.5 t/ha due Breeding Breeding work on soya bean in
to a combination of poor soil conditions and tropical Africa aims at the development of im-
poor management. However, yields of more proved cultivars with high and stable seed
than 2 t/ha have been recorded on smallholder yield, resistance to major diseases and pests,
farms in Zimbabwe and Nigeria, particularly tolerance to aluminium toxicity, resistance to
when farmers are growing soya bean as a cash lodging and pod shattering, promiscuous nodu-
crop to sell in urban food markets or for proc- lation, improved seed longevity and acceptable
essing for oil and feed. The average yield of seed colour, oil and protein content. A breeding
commercial, large-scale farmers hovers around programme at UTA has focused since the early
2 t/ha. Under optimal growing conditions yields 1980s on combining the yield potential of culti-
of more than 4.5 t/ha have been recorded in vars bred in North America with the 'promis-
Zimbabwe. In Nigeria and most of West Africa cuous' or 'naturally-nodulating' ability of land-
the yield potential of soya bean is about 3 t/ha. races from Asia to form nodules and fix nitro-
Handling after harvest The whole plants gen without inoculation in African soils. This
are dried in the sun. They are then threshed by breeding programme has produced a series of
beating with sticks. The seeds are winnowed, excellent multi-purpose cultivars that combine
cleaned and prepared for store or market. For a leafy growth habit with appropriate seed type
on-farm storage a seed moisture content of10— and high yield potential. These cultivars are
12% must be maintained. Deterioration of seed liked by smallholder farmers because they pro-
in storage is a major problem in the humid vide biomass for forage or to improve soil fertil-
tropics and is attributable to poor storage con- ity in addition to having high seed yields. They
ditions and pests. In the savanna region of are being actively promoted in many countries
West Africa producers have developed appro- in East and West Africa at present. In south-
priate seed handling methods that ensure good ern Africa similar benefits of a largely unim-
seed germination when they save their own proved cultivar, 'Magoye', were recognized.
seeds. 'Magoye' is a leafy, indeterminate cultivar,
Genetic resources The largest germplasm relatively resistant to stresses and mid-season
collections of soya bean are held in China (In- drought, that grows better on poor soils than
stitute of Crop Germplasm Resources (CAAS), some of the improved cultivars, nodulating well
Beijing, 23,600 accessions; Nanjing Agricul- with indigenous rhizobia. Despite its smaller,
tural University, Nanjing, 13,000 accessions), yellow seed, and susceptibility to some diseases
the United States (USDA-ARS Soybean Germ- such as bacterial pustule, this makes it an at-
plasm Collection, Urbana, Illinois, 18,400 ac- tractive cultivar for use by smallholder farmers
cessions) and Taiwan (Asian Vegetable Re- in southern Africa.
search and Development Centre (AVRDC), Research at UTA has identified soya bean
Shanhua, 12,500 accessions). In tropical Africa breeding lines that favour the germination of
substantial germplasm collections are held in Striga hermonthica (Delile) Benth., a parasitic
Zimbabwe (Crop Breeding Institute, Harare, weed that infects maize, sorghum and pearl
2250 accessions), Nigeria (International Insti- millet, and one of the major constraints to pro-
tute of Tropical Agriculture (UTA), Ibadan, duction of these crops in Africa. The probable
1800 accessions), Rwanda (Institut des Sci- cause of this effect of soya bean is the presence
ences Agronomiques du Rwanda (ISAR), Bu- of root exudates. The inclusion of these soya
tare, 550 accessions) and Kenya (National bean cultivars in crop rotations stimulates
Genebank of Kenya, Crop Plant Genetic Re- Striga germination and reduces infestation
sources Centre, KARI, Kikuyu, 130 accessions). levels in following sorghum, maize or pearl
Genebank accessions have been successfully millet crops as a result of the decline of Striga
82 CEREALS AND PULSES

seed numbers in the soil.After germination the tofu which rapidly became one of the most
Striga plants are unable to infest the soya bean popular snacks in markets in the region and is
crop, and die without producing seed. A 3-year widely used by the food processing industry. In
trial conducted in Benin showed that 2 seasons some areas, the low world prices may depress
of soya bean followed by maize reduced Striga opportunities for local producers to respond to
hermonthica emergence by about 80-90% and increased local demand for soya bean. Soya
increased maize yield from 1.5 t/ha to 3 t/ha. bean can play an increasingly important role in
Similar results have been obtained in farmers' diversifying cereal-based farming systems in
fields in Nigeria. As soya bean becomes more tropical Africa. Apart from being a source of
popular in areas where maize, sorghum and residual nitrogen for subsequent cereal crops in
pearl millet are grown, the amount of damage crop rotations, the new multi-purpose cultivars
caused by Striga hermonthica should become bred by UTA provide the additional benefit
significantly less. that they help to reduce Striga hermonthica
A number of private seed companies are in- damage on maize, sorghum and millet, thus
volved in breeding soya bean in southern Af- representing a major opportunity to provide
rica, with particular emphasis on cultivars sustainable crop rotations for smallholder
suitable for mechanized production. The com- farmers. It is therefore very likely that soya
panies are targeting a number of traits includ- bean production will expand in many tropical
ing high seed yield, resistance to lodging, resis- African countries in the future.
tance to pod shattering, rapid stem dehydra- Major references Boerma & Specht, 2004;
tion, seed quality and resistance to diseases Carsky et al., 2000; Dashiell & Fatokun, 1997;
(particularly red leaf blotch and frogeye leaf Hymowitz, 1995; Javaheri & Baudoin, 2001;
spot). New cultivars are 'Solitaire', 'Soma', 'So- Mpepereki et al., 2000; Sanginga et al., 2003;
prano' and 'Viking', all of which have some Shanmugasundaram & Sumarno, 1989; Sin-
resistance to frogeye leaf spot. These cultivars clair, 1998; Singh, Rachie & Dashiell (Editors),
are all specific in their nodulation ability and 1987.
require inoculation with the appropriate rhizo- O t h e r references Akem & Dashiell, 1996;
bia. Inoculants for soya bean are produced, sold Aljanabi, 2001; Dashiell & Akem, 1991; FAO,
and used on a large scale in both Zimbabwe 1998; Giller, 2001; Hanelt & Institute of Plant
and South Africa. Genetics and Crop Plant Research (Editors),
Soya bean is a leading crop in the field of ge- 2001; Hume, Shanmugasundaram & Bevers-
netic transformation. In 2001 the world area dorf, 1985; ILDIS, 2005; James, 2002; Mackin-
under transgenic herbicide-tolerant soya bean der et al., 2001;Musiyiwa, Mpepereki & Giller,
was estimated at 33 million ha; it was grown in 2005; Rehm & Espig, 1991;Sanginga, Thottap-
the United States, Argentina, Canada, Mexico, pilly & Dashiell, 2000; Sanginga et al., 1997;
Uruguay, Romania and South Africa. Genetic Sanginga et al., 1999; Shannon &Kalala, 1994;
linkage maps have been constructed for soya Thulin, 1989a; Tindall, 1983; USDA, 2004;
bean on the basis of various markers (RFLP, Weiss, 2000.
SSR, RAPD, AFLP), and several moderate- to S o u r c e s of i l l u s t r a t i o n Shanmugasunda-
high-density genetic maps are now available. ram &Sumarno, 1989.
In-vitro regeneration of soya bean is possible A u t h o r s K.E. Giller &K.E. Dashiell
through organogenesis and somatic embryo- Based on PROSEA 1:Pulses.
genesis.
P r o s p e c t s Soya bean is a relatively new
crop in tropical Africa. It has long been thought HORDEUMVULGARE L.
that soya bean was not a suitable food crop for
the region, because of the long cooking time P r o t o l o g u e Sp. pi. 1:84 (1753).
needed and the unacceptable taste. However, Family Poaceae (Gramineae)
the importance of the crop in tropical Africa Chromosome n u m b e r In = 14
has grown rapidly during the past decades. Synonyms Hordeum sativum Jess. (1863).
Especially Nigeria witnessed a rapid expansion V e r n a c u l a r n a m e s Barley (En). Orge (Fr).
in soya bean production in the smallholder Cevada (Po). Shayiri (Sw).
farming sector in the savanna zone during the Origin and geographic distribution Barley
1990s. The driving force for this expansion was was domesticated in western Asia before 7000
the use of soya bean in the preparation of many BC. Cultivation spread to northern Africa and,
traditional foods and the introduction of soya moving upwards along the Nile, into Ethiopia,
 HORDEUM 83

is also fed to animals. By-products from the

brewing process are also used in livestock feed.
Production and international trade Annual
world production of barley is 136 million t
grains (mean of 1999-2003) from 54 million ha.
Major producing countries are the Russian
Federation, Germany and Canada with 16.2,
12.1, and 11.4 million t per year, respectively.
In tropical Africa the main barley-producing
country is Ethiopia, with 950,000 t of grain
from 870,000 ha in 1999-2003, followed by
Kenya (45,000 t from 20,000 ha) and Eritrea
(24,000 t from 44,000 ha). In Ethiopia and Eri-
trea barley production is mainly for subsis-
tence, and in Ethiopia the share of malting
barley is about 2% of the total production.
Small areas of barley (less than 4000 ha) are
Hordeum vulgare - planted planted in Mauritania, DR Congo, Tanzania,
Zambia and Zimbabwe. Most barley is con-
where it became one of the major cereals. It is sumed locally, with about 20 million t per year
not known exactly when barley reached Ethio- entering international trade in 1998-2002. The
pia, but it has been grown there for at least European Union, Australia and Canada are the
5000 years. Barley reached southern Spain largest exporters; Saudi Arabia, China and
4000-5000 BC and central and northern Japan the largest importers. In 1998-2002 the
Europe, as well as India, 2000-3000 BC. It largest importers in tropical Africa were Zim-
reached China 1000-2000 BC. Barley was cul- babwe and Ethiopia, with average annual im-
tivated in oases in the Sahara 100-300 BC, but ports of 8000 t and 3000 t, respectively.
seems not to have migrated southwards into Properties The composition of barley per
West Africa before the 16th Century AD. Co- 100 g edible portion is: water 9.4 g, energy
lumbus took it to the New World. Nowadays it 1482 kJ (354 kcal), protein 12.5 g, fat 2.3 g,
is grown over a broader environmental range carbohydrate 73.5 g, dietary fibre 17.3 g, Ca 33
than any other cereal, from 70°N in Norway to mg, Mg 133 mg, P 264 mg, Fe 3.6 mg, Zn 2.8
44°S in New Zealand. In Ethiopia, Tibet and mg, vitamin A 22 IU, thiamin 0.65 mg, ribofla-
the Andes it is cultivated higher on the moun- vin 0.29 mg, niacin 4.6 mg, vitamin B60.32 mg,
tain slopes than other cereals. In tropical Af- folate 19 |Xgand ascorbic acid 0 mg. The essen-
rica it is mainly found in East Africa. In West tial amino acid composition per 100 g edible
Africa barley is grown as a cold season crop in portion is: tryptophan 208 mg, lysine 465 mg,
the Sahel region and northern Nigeria. In methionine 240 mg, phenylalanine 700 mg,
Madagascar it is grown in the dry season. threonine 424 mg, valine 612 mg, leucine 848
Uses On a worldwide scale barley is used, in mg and isoleucine 456 mg. The principal fatty
order of importance, as animal feed, for acids are per 100 g edible portion: linoleic acid
malting (especially for beer brewing) and as 999 mg, palmitic acid 411 mg, oleic acid 241 mg
human food. In the tropics and subtropics it is and linolenic acid 110 mg (USDA, 2004). Bar-
mainly grown for human food. In Ethiopia and ley cannot be used to make leavened bread
Eritrea most of the grain is used for making because of its low gluten content. The rela-
the local pancake-like bread ('injera'), but it is tively high dietary fibre content makes barley
also made into porridge, soup and home-made interesting from a nutritional point of view, as
alcoholic and non-alcoholic drinks. In Kenya increased amounts of dietary fibre have been
and Tanzania it is more important for brewing. shown to help in controlling diabetes, hyper-
Grains are roasted or fried, and consumed as lipidaemia, obesity, hypertension, coronary
snack, particularly during social gatherings. heart disease and various gastro-intestinal
Barley straw is used as animal feed, for animal disorders. Barley is preferred to wheat and rye
bedding, and as cover material for huts. Barley for malting because of the cemented lemma
can be grazed during tillering or cut before and palea, which protect the young plumule
maturity and directly fed to the animals or during germination, produce a firmer grain at
used for silage. In temperate areas barley grain the high moisture content needed for steeping
84 CEREALS AND PULSES

and malting, and facilitate filtering. barley has been developed from original wild
Description Annual grass up to 120(-150) populations. Fertile hybrids between wild and
cm tall, tillering freely; root system consisting cultivated types are easily obtained and occur
of 3—9 primary roots and adventitious roots; naturally where the two grow together. Wild
stem (culm) usually erect. Leaves 5-10 per barley has been classified as subsp. sponta-
culm, alternate, simple; leaf sheath glabrous, neum (C.Koch) Thell. (synonym: Hordeum
with large, overlapping auricles at apex; ligule spontaneum C.Koch), distributed in northern
1-3 mm long, hyaline, ciliate; blade lin- Africa, the eastern Mediterranean area and
ear-lanceolate, 5-40 cm x 0.5-1.5 cm. Inflores- western Asia, and cultivated types as subsp.
cence a terminal cylindrical spike 5-10(-30) cm vulgare.
long, with groups of 3 spikelets attached alter- The variation in cultivated barley is over-
nately. Spikelets 1-flowered, with bisexual flo- whelming, with thousands of landraces and
ret; glumes narrow, about half the length of the hundreds of cultivars. Cultivars can be divided
lemma, with fine bristles at the tip; lemma according to the number of rows of grains (2 or
ovate, 9-11 mm x 3 mm, 5-veined, usually end- 6), lax and compact spikes, or awned and
ing in an awn up to 15 cm long; palea as long awnless lemmas. All wild types have 2-rowed
as lemma, awnless; stamens 3; ovary superior, spikes, i.e. of the 3 spikelets at each node the
with 2 stigmas. Fruit a caryopsis (grain), ellip- two lateral ones are sterile, and only the cen-
soid, flattened and grooved on one side, varying tral one develops a grain. Under domestication
in size according to cultivar, hairy at the tip. 6-rowed types appeared in which all 3 spikelets
Other botanical information Hordeum com- produce grains. Here, 2 genes are involved,
prises 32 species. Hybrids from crosses of Hor- both with multiple allelic series, but a single
deum vulgare with other Hordeum species are recessive mutation is enough for a 2-rowed
sterile or anomalous. The large variability in barley to become 6-rowed. Cultivars grown in
barley led to many barley species being distin- West Africa have 6 rows of grains in the spike,
guished in the past. At present, the accepted but in Ethiopia, especially in the upper high-
view is that in barley evolution only a single lands, a type ofbarley is found with 2 full rows
species is involved, Hordeum vulgare, forming and parts of other rows ('irregular barley'). In
a crop-weed complex, in which the cultivated 'husked barley' lemma and palea adhere to the
grain at threshing, whereas in 'naked barley'
the grain threshes free. A single recessive gene
controls the latter character. Based on vernali-
zation requirements barley is classified in win-
ter and spring types.
Growth and development The seedling
emerges from the soil 5-6 days after germina-
tion. Tillers are produced on the main shoot
until flower initiation. The number of tillers
per plant is influenced by plant density, culti-
var and environmental factors: a single plant
usually develops 1—6 stems, but at low densi-
ties it may be several times as high. Time to
flower initiation varies among cultivars, but in
general barley flowers earlier than wheat. Bar-
ley is a quantitative long-day species, flowering
earlier under longer photoperiods, but the pho-
toperiod sensitivity differs between cultivars,
ranging from very sensitive to almost insensi-
tive. Flowers are predominantly self-pollinated,
but cross-pollination can be as high as 10%.
The grain ripens in 20-40 days. Barley can
mature in a short season of 3-4 months, which
is shorter than the season needed for other
Hordeum vulgare 1, lower part of plant; 2, major cereals.
spike; 3, spikelets. Ecology Barley grows under a wide range of
Source: PROSEA photoperiod, temperature and rainfall condi-
 HORDEUM 85

tions, but is best adapted to temperate cli- can be applied before seeding or as topdressing
mates. It withstands high temperatures in dry after planting. Under dry conditions high rates
climates and humidity in cool climates, but it is of N application can cause yield reductions,
ill adapted to hot, humid climates, primarily whereas under favourable conditions high
because of its susceptibility to diseases. In rates of N increase the risk of lodging and dis-
Ethiopia, barley is cultivated at 1500—3500 m eases. Excessive rates of N fertilization of malt-
altitude, but mainly between 2000 m and 3000 ing barley may increase the protein level in the
m. In Kenya it is grown at 1500-3000 m alti- grain above the acceptable level.
tude. In Ethiopia farmers usually grow barley in low-
Winter barley types need vernalization by a input systems, with minimal seedbed prepara-
period of low temperature (3-12°C). Barley is tion and weeding, and without applying herbi-
adapted to an annual rainfall ranging from 200 cides, fertilizers or insecticides. Low soil fertil-
mm to more than 1000 mm. It is more drought- ity and insufficient weed management are ma-
escaping, due to its early maturity, than jor constraints. In experiments at Holetta, ap-
drought-tolerant. plication of 57 kg N and 25 kg P per ha led to a
Well-drained fertile loams or light clay soils are grain yield increase of up to 200%, whereas a
best for barley production. Barley is more tol- single hand weeding at 35 days after emer-
erant of alkaline soils than other cereals, but gence gave a grain yield increase of up to 20%
does not tolerate acid soils; a pH of 6.0-8.5 is compared to an unweeded crop. Lodging is also
generally acceptable. It is very sensitive to a common problem in Ethiopian barley cultiva-
waterlogging. Some cultivars withstand up to tion. In West Africa barley is grown as a dry-
1%salt in the soil. season crop, often under irrigation.
Propagation and planting Barley is plant- Diseases and pests Barley is affected by
ed through direct seeding. The 1000-seed weight several viral and fungal diseases. The most
is 25-50 g. Before sowing, the land is ploughed important viral diseases are barley yellow
to a depth of 10-15 cm. On smallholdings the dwarf virus (BYDV), transmitted by various
land is prepared by animal traction. The seed aphid species, and barley stripe mosaic virus
should preferably be treated with fungicide to (BSMV), transmitted through seed or plant
protect the crop from seed- and soil-borne dis- contact. Control measures for BYDV include
eases. Barley can be planted with drills, but on the use of tolerant or resistant cultivars and
smallholdings seeding is usually by hand. aphid control, whereas BSMV can be controlled
When sowing with drills, the distance between by using virus-free seed and resistant culti-
rows is 15—35cm, and the seed rate is 50-150 vars. Epidemics of African cereal streak,
kg/ha. The sowing depth is 2—6cm. caused by maize streak virus (MSV) and
In the highlands of Ethiopia, barley is cropped transmitted by leafhoppers (Cicadulina spp.),
twice a year. The main cropping season ('me- have occurred in Kenya. Important fungal dis-
ner') is from June—December (with most of the eases include powdery mildew (Blumeria
rain in June-September), while the minor graminis f.sp. hordei, synonym: Erysiphe
cropping season ('belg') is from February—June graminis f.sp. hordei), spot blotch (Bipolaris
(most rain in March—April). Barley is the most sorokiniana, synonym: Helminthosporium sati-
suitable crop for 'belg'-season production. Bar- vum), scald (Rhynchosporium secalis f.sp. hor-
ley is usually sole-cropped, but in Eritrea and dei), scab (Fusarium spp.), rusts (Puccinia
northern Ethiopia barley is often intercropped spp.), net blotch (Pyrenophora teres), barley
with wheat (the 'hanfetz' cropping system). In stripe (Pyrenophora graminea) and smuts
Eritrea farmers traditionally broadcast mix- (Ustilago spp.). In Ethiopia and Eritrea, scald,
tures of 67%barley and 33%wheat; sometimes blotches and rusts are the most important fun-
50:50 mixtures are sown. In Ethiopia and Eri- gal diseases; the use of resistant cultivars is
trea barley is a smallholder crop, but in Kenya often the most effective control measure. Other
it is grown on large-scale mechanized farms. control measures include crop rotation, the use
Management Weeds cause economic losses of clean seed, seed treatment, fungicides, the
in barley due to a reduced number of tillers destruction of infected plant material and deep
and grains per spike. Resource-poor farmers ploughing. Several nematodes can parasitize
weed by hand. The most widely used herbicide barley: cereal cyst nematodes (Heterodera
to control broad-leaved weeds in post-emer- spp.), root-knot nematodes (Meloidogyne spp.),
gence spraying is 2,4-D. Barley requires about root-gall nematode (Subanguina radicicola)
25-35 kg N to produce 1 t of grain. Nitrogen and root-lesion nematodes (Pratylenchus spp.).
86 CEREALS AND PULSES

Control measures include crop rotation and haviour.

fallow. Resistance breeding and the use of fun- Breeding In barley breeding methods are
gal pathogens have been successful against used that are typical for self-pollinated crops.
cereal cyst nematodes. Until 1950 the main breeding method was ei-
Barley is susceptible to attack by many kinds ther mass selection or pure line selection
of pests, including aphids, shoot flies, grass- within landraces, which are still cultivated
hoppers, crickets, thrips, army worms, cut- today in many countries, e.g. Ethiopia. New
worms, and beetles and their larvae. Control variability has been created by crossing, back-
may involve adjusting the planting date, and crossing and mutation. Recurrent selection is
applying insecticides. In general, losses due to applied to increase recombination by crossing
pests are relatively limited, with many of the among Frs and by repeating the crosses among
pests causing greater damage as vectors of a number of F2plants or by using male-sterility
viruses, notably BYDV. Major storage pests of genes. Mutation breeding with radiation or
barley are insects and rodents. chemicals and double haploids have been
Harvesting Barley is ready for harvesting widely used in barley breeding. Among the
after reaching 35-40% kernel moisture. The International Centres, ICARDA has the global
crop is harvested by hand using a sickle or by mandate for assisting barley improvement pro-
combine. Threshing of malting and 'naked' grammes of National Agricultural Research
barley requires special care to avoid too much Systems (NARS) in developing countries. Mod-
broken seed. ern breeding has resulted in barley types with
Yield Barley yields vary from 0.3 t/ha in dry improved amino-acid composition (4.1%of the
years and in marginal environments to 10 t/ha protein as lysine). The major emphasis is on
in high input agriculture. In Africa average producing cultivars resistant to diseases and
yields are 0.3-2.5 t/ha. In Asia and South pests and with adaptation to specific agro-eco-
America average yields are 1.5—1.7 t/ha, in logical environments. Resistance to aphids has
North America 2.9 t/ha and in Europe 4 t/ha. been incorporated into some cultivars. Ethio-
Straw yields are equally important in many pian barley has been particularly useful in the
developing countries, but yield statistics are improvement of nutritional quality and in sup-
not available. plying resistance to fungal and viral diseases
Handling after harvest A high moisture (including powdery mildew, smuts, rusts, net
content of barley grains at harvest favours the blotch, scald, BYDV and BSMV). In Ethiopia, 9
development of mycotoxins dangerous to hu- improved barley cultivars were released be-
mans and livestock. Before storing, grain has tween 1980 and 2000, but the adoption level is
to be dried to 14% moisture or less. When bar- low (11% ofthe area).
ley is cultivated on small areas, it is common The first genome-wide maps of barley were
practice to keep selected spikes to provide the published in 1991, and more than 40 genome-
seed for the next crop. wide maps have now been published.
Genetic resources Current barley germ- Prospects In general large improvements in
plasm collections at ICARDA (International barley production are possible through breed-
Center for Agricultural Research in the Dry ing of cultivars resistant to lodging and dis-
Areas, Aleppo, Syria) contain more than 25,000 eases. As most barley breeding has been for
accessions. Since 1989 the 'International Bar- favourable environments, the potential of the
ley Core Collection' has been developed by an crop in low-input agriculture is largely under-
international consortium including ICARDA. exploited. In the Ethiopian highlands, where
Many national programmes maintain their barley is the preferred food crop, soil fertility
own working collections. The Institute of Bio- has declined as a result of erosion and continu-
diversity Conservation (IBC, Addis Ababa, ous cultivation. Under these circumstances it
Ethiopia) has a collection of more than 12,500 has been difficult to improve the locally
accessions. Smaller collections in sub-Saharan adapted farmers' cultivars. The staple Ethio-
Africa are held in South Africa (National De- pian cereal tef (Eragrostis tef (Zuccagni) Trot-
partment of Agriculture, Pretoria; Small Grain ter), is taking over from barley as a result of
Institute, Bethlehem), Madagascar (Départe- favourable market prices, and high-yielding 6-
ment de Recherches Agronomiques de la Re- rowed barley cultivars are becoming less im-
publique Malgache, Antananarivo) and Kenya portant. In these areas breeding for early-
(National Genebank of Kenya, KARI, Kikuyu, maturing cultivars with higher resistance/tol-
Muguga). Barley shows orthodox storage be- erance to moisture stress than the existing
 LATHYRUS 87

ones might reverse the trend. Ethiopia has the

potential to meet the local demand for malt
barley and to produce for the African market.
This can be achieved with the breeding and
production of high-quality malt barley that
meets the breweries' standards, a concomitant
increase in the number and capacity of malt-
processing factories, and an efficient market
structure.
Major references Ceccarelli & Grando,
1996; Gebre & van Leur (Editors), 1996;
Hanelt & Institute of Plant Genetics and Crop
Plant Research (Editors), 2001; Harlan, 1995;
Hockett, 2000; Mathre, 1997; Nevo, 1992; Ras-
musson (Editor), 1985; Slafer et a l , 2002; von
Bothmer, Jacobsen &Baden, 1995.
Other references Asfaw, 1990; Aw-Hassan
& Shideed, 2003; Berhane, Yitbarek & Fekadu : Lathyrus sativus -planted
1995; Briggs, 1978; Burkill, 1994; Clayton.
1972; Engels, Hawkes & Worede (Editors) dia. Nowadays grass pea is widely cultivated in
1991; Launert, 1971; National Research Coun large parts of Asia (especially Bangladesh, In-
cil, 1996; Phillips, 1995; Purseglove, 1972: dia, Nepal, Pakistan and the Middle East),
Rehm & Espig, 1991; Sharpley, 1988; Tareke- southern Europe and northern Africa, and to a
gne, Gebre & Francis, 1997; Thomas, 2003 lesser extent in America, Australia and South
USDA, 2004; von Bothmer et a l , 2003; Wil- Africa. In tropical Africa it is mainly grown in
liams, 2003; Woldeamlak, 2001; Yirga, Ale- Ethiopia, but also in Sudan, Eritrea, Kenya,
mayehu &Sinebo (Editors), 1998. Tanzania, Angola and Mauritius.
Sources of illustration Ceccarelli & Gran- Uses In Ethiopia and Eritrea grass pea
do, 1996. seeds are mainly consumed in the form of
Authors S. Ceccarelli &S. Grando sauces ('wot'); 'shiro wot' (sauce made of flour)
Based on PROSEA 10: Cereals. and 'kik wot' (sauce made of hulled split seeds)
are eaten together with 'injera' (a pancake-like
unleavened bread). Boiled grass pea seeds ('ni-
LATHYRUS SATIVUS L. fro') are also consumed in most areas, whereas
'kitta' (an unleavened bread) made from grass
Protologue Sp. pi. 2: 730 (1753). pea seeds is consumed mainly during times of
Family Papilionaceae (Leguminosae - Papil- acute food shortage. In India the seeds are
ionoideae, Fabaceae) sometimes boiled whole, but are most often
Chromosome number 2n = 14 processed into dhal. The flour, made by grind-
Vernacular names Grass pea, chickling pea, ing either the whole or split seeds, is sold as
chickling vetch, white pea (En). Gesse, gesse 'besan'. In Bangladesh 'roti' made out of grass
blanche, gesse commune, pois carré, lentille pea flour is a staple for landless labourers. In
d'Espagne (Fr). Chicharo, chicharo comun, India grass pea is sometimes used to adulter-
sincho (Po). ate more expensive pulses, such as chickpea or
Origin and geographic distribution The or- pigeon pea. Care should be taken in the con-
igin of Lathyrus sativus is unknown. Records sumption of grass pea seeds, as excessive con-
exist of wild Lathyrus sativus plants in Iraq, sumption leads to a neurological disorder in
but it is not clear if these are truly wild or es- people and animals, called lathyrism and char-
capes from cultivation. Lathyrus sativus is acterized by paralysis of the lower limbs. In
perhaps a derivative from Lathyrus cicera L., many countries grass pea seeds are used as
which occurs wild in southern Europe, north- animal feed, e.g. as an ingredient in pig starter
ern Africa and western Asia and is sometimes and grower diets.
grown there. Domestication of grass pea In Asia immature pods are cooked and eaten as
probably took place in the Balkan around 6000 a vegetable, or are boiled, salted and consumed
BC. Remains of Lathyrus sativus dating back as a snack. Young vegetative parts are cooked
to 2000-1500 BC have been recorded from In- as a green vegetable; they are also dried for off-
88 CEREALS AND PULSES

season use as a vegetable. Young grass pea if water is changed after soaking and during
plants are used as fodder for cattle or for graz- cooking. When the seeds are ground into flour,
ing in many countries. The stems and chaff which is then used in baking or cooking, ODAP
remaining after harvest are often the most may not be removed. Unfortunately, effective
important reason for growing the crop in Asia. detoxification treatments often also result in
As fodder, the plants can be eaten green or as decrease of nutritional quality. Other antinu-
hay; they are not suitable for silage. Grass pea tritional factors in grass pea include trypsin
is grown as a green manure, e.g. in Australia inhibitors, tannins, lectins, phytate and oligo-
and Canada. Oil from the seeds is used medici- saccharides.
nally as a powerful cathartic. Grass pea hay contains: water 14.6%, protein
Production and international trade Accord- 9.9%, fat 1.9%, fibre 36.5%, nitrogen-free ex-
ing to estimates India produced about 0.8 mil- tract 31.0% and ash 6.1%. The seeds of culti-
lion t grass pea seeds per year from 1.5 million vars with up to 0.22 g ODAP per 100 g seed
ha in the mid 1990s, whereas production was could be included in the diets of growing chicks
lower in Bangladesh (175,000 t from 240,000 at a rate of 400 g grass pea seeds per kg feed
ha) and Pakistan (45,000 t from 130,000 ha). In without negative effects on weight gain or fat
the late 1990s production in Ethiopia was es- or protein digestibility.
timated at 105,000 t from 142,000 ha. As a food Description Much-branched, erect, strag-
grain, grass pea is traditionally traded within gling or climbing, glabrous annual herb; stem
the region of production, and it does not enter slender, quadrangular, winged, up to 90(—170)
international trade. cm long; taproot well-developed. Leaves alter-
P r o p e r t i e s The composition of whole grass nate, 2- or 4-foliolate, ending in a simple or
pea seed per 100 g edible portion is: water 8.4 branched tendril; stipules prominent, leaf-like,
g, energy 1457 k j (348 kcal), protein 27.4 g, fat narrowly triangular, with a smaller but simi-
1.1 g, carbohydrate 59.8 g, fibre 7.3 g, Ca 127 larly shaped basal appendage and often with a
mg, P 410 mg and Fe 10.0 mg (Leung, Busson small tooth between the lobes; petiole mostly
& Jardin, 1968). Grass pea is highly deficient
in methionine and tryptophan. Raw whole
seeds contain 41% starch on a dry matter ba-
sis; the starch granules are oval and on aver-
age 25 (im long and 17(xmwide.
The neurological disorder lathyrism is caused
by the water-soluble non-protein amino acid
ODAP (ß-N-oxalyl-L-a,ß-diaminopropionic acid),
also known as BOAA (ß-N-oxalylamino-L-
alanine) and OAP (L-3-oxalylamino-2-amino-
propionic acid). ODAP is present in all parts of
the plant and affects various parts of the cen-
tral nervous system, disrupting neurotrans-
mission and thus impairing muscular activity.
The onset of lathyrism can be slow or sudden,
and is often indicated by a feeling of heaviness
and pain in the lower limbs. Lathyrism is often
irreversible, but not fatal. Lathyrism seems to
occur when food ratios containing at least 25%
grass pea are consumed continuously over 1.5—
6 months and may then affect up to 5% of the
population. Outbreaks of lathyrism often occur
during near-famine conditions that force people
to rely too heavily on grass pea. The ODAP
content of grass pea seeds typically ranges
from 0.1-1.4(-2.5) g per 100 g seed. ODAP lev-
els are not only genetically determined, but
also highly influenced by growing conditions. Lathyrus sativus - 1, flowering and fruiting
In general, soaking and boiling reduce ODAP branch; 2, seeds.
levels in the seeds, and this effect is enhanced Source: PROSEA
 LATHYRUS 89

winged, (l-)1.5-2.5(-3.5) cm long; leaflets ses- Total crop duration is 3-6 months. Grass pea
sile, narrowly elliptical-oblong, (3-)4-5(-7.5) effectively nodulates with Rhizobium legumi-
cm x 3-5(-13) mm, cuneate at base, acute or nosarum.
acuminate at apex. Flowers solitary in leaf Ecology Grass pea is grown successfully in
axils, bisexual, papilionaceous, pedicel with regions with an average annual rainfall of 400-
joint, lower part (l-)3-3.5(-5) cm long, upper 650 mm/year and an average temperature of
part (2—)5—7(—8) mm long; calyx campanulate, 10-25°C. It withstands heavy rains in early
tube c. 3 mm long, lobes 5, almost equal, nar- growth stages and prolonged drought during
rowly triangular, 3-6 mm long; corolla blue, grain-filling. It grows well in the subtropics as
reddish-purple, red, pink or white, standard a winter crop. Grass pea can be grown on a
erect and spreading, very broadly obovate, c. 15 wide range of soil types, including poor soils
mm x 18 mm, clawed, retuse at top, wings and heavy clays. It tolerates waterlogging and
broadly obovate, c. 14 mm x 8 mm, clawed, moderate salinity. In Ethiopia grass pea is
with auricle, keel slightly twisted, boat-shaped, often grown in the dry season on residual soil
c. 10 mm x 7 mm, clawed, with 2 auricles; sta- moisture in heavy black clay soils at 1700-
mens 10, 9 united and 1 free; ovary superior, 2700 m altitude. In India grass pea is grown as
sessile, c. 6 mm long, style abruptly upturned, a cold-season crop up to 1200 m altitude.
c. 7 mm long, stigma spoon-shaped. Fruit an Propagation and planting Grass pea is
oblong, flattened pod (1.5-)2.5-4.5(-5.5) cm x propagated by seed. The 1000-seed weight
0.5-2 cm, upper margin 2-winged, shortly ranges from 30-300 g. In Ethiopia it does not
beaked, glabrous, (l-)2-5(-7)-seeded. Seeds require a fine seedbed; 1-2 ploughings are
wedge-shaped, 4—7 mm in diameter, white, enough. The average seed rate is normally 4 5 -
pale green, grey or brown, marbled; hilum el- 60 kg/ha for a sole crop, and about 35 kg/ha
liptical. Seedling with hypogeal germination. when intercropped. Seeds that may have been
Other botanical information Lathyrus com- soaked in water overnight are broadcast or
prises about 150 species, mainly in the tem- drilled in furrows. Plant densities of 200,000-
perate regions of the northern hemisphere and 250,000 plants/ha are common for grass pea. In
South America, with a few species in Africa. Ethiopia grass pea is usually sown in Septem-
Lathyrus sativus is placed in section Lathyrus ber-November and harvested in January-
along with about 30 other annual or perennial April.
species. Based on crossability and cytological Grass pea is grown as a sole crop or inter-
evidence, Lathyrus amphicarpos Gouan and cropped, e.g. with barley, linseed or chickpea.
Lathyrus cicera L. have been placed in the sec- In many countries grass pea is produced in
ondary gene pool of grass pea. More recently, rice-based cropping systems before the rice
successful crosses between Lathyrus sativus crop or alternately with a rice crop. In India
and Lathyrus pseudocicera Pamp. have been grass pea is often grown as a relay crop: it is
made. Apart from Lathyrus sativus, other La- broadcast into a standing rice crop about 2
thyrus species cultivated in Ethiopia are the weeks before the rice harvest and left to grow
ornamental Lathyrus odoratus L. and the for- on the residual moisture.
age Lathyrus aphaca L. Management Grass pea often receives
Infraspecific classification is mainly based on hardly any attention after sowing, although for
colour of flowers, markings on pods and size optimum yields it should be kept reasonably
and colour of seeds. In general, white seed is free from weeds. In a well-prepared field, the
most popular for human consumption. The crop comes up as a thick mass over the entire
level of infraspecific variation for RAPD mark- surface, smothering out weeds. Grass pea is
ers is low compared to other grain legumes not normally fertilized, but atmospheric nitro-
such as lentil and pea. Based on isozyme analy- gen fixation rates of 25—50 kg/ha have been
sis variation was found to be highest in west- recorded. In Ethiopia grass pea is grown in
ern Asia and northern Africa. rotation after barley or sometimes after a pulse
Growth and development Germination of crop, such as pea or chickpea, which has been
grass pea seeds is most rapid around 20°C. sown in April and harvested in July.
Flowering time is 1.5-4 months after sowing. Diseases and pests The main diseases of
The floral biology of grass pea favours self- grass pea are powdery mildew (Erysiphe pisi)
pollination (anthers usually dehisce before full and downy mildew (Peronospora spp.), but the
opening of the flower), but there are many re- latter not in Ethiopia. Rust (Uromyces fabae)
cords of substantial outcrossing (up to 28%). and Fusarium wilt (Fusarium oxysporum) have
90 CEREALSANDPULSES

been recorded from Ethiopia. Faba bean ne- potential is an important goal. Other breeding
crotic yellows virus (FBNYV) has been ob- objectives are the incorporation of disease re-
served on grass pea in Ethiopia; it is transmit- sistance and increase of seed size, earlier ma-
ted by the aphids Acyrtosiphon pisum and turity and a higher harvest index. Lines with
Aphis cracciuora. In host-range studies grass moderate resistance to powdery mildew have
pea was found to be susceptible to pea seed- been identified. In Ethiopia a large number of
borne mosaic virus (PSbMV). Insect pests of accessions and breeding lines introduced from
grass pea include aphids and thrips. The pea ICARDA are resistant to powdery mildew.
aphid (Acyrthosiphon pisum) is the main pest Improvement has been slow in grass pea. High-
of grass pea in Ethiopia. yielding improved cultivars low in ODAP and
Harvesting Harvesting of grass pea should with resistance against biotic and abiotic
be done when leaves turn yellow and pods turn stresses have not been released in Africa. Some
grey, to avoid shattering. The plants are pulled attempts to provide improved cultivars with
out by hand or cut with a sickle near the base. low ODAP content have been made in India. In
They are then stacked and allowed to dry for Chile and Bangladesh some promising lines
7-8 days in the field or on the threshing floor. have also been identified with low ODAP and
Yield The average seed yield of grass pea is high yield. Recently, of the 13 lines with low
350-700 kg/ha; in Ethiopia it is about 700 ODAP content identified in Ethiopia, three
kg/ha. Yield trials conducted recently in vari- lines introduced from ICARDA have consis-
ous countries recorded yield levels of 1500— tently shown low ODAP and reasonable yield
3000 kg/ha. over three years. However, the substantial
Handling after harvest Grass pea pods are outcrossing rate in grass pea has limited the
threshed by animal trampling or by beating progress in identifying stable lines with low
with sticks, after which the seed is winnowed ODAP content; seeds of selected lines must be
and cleaned. The seeds may be dried for a few multiplied in isolation and be provided to
days before storage. farmers every year.
Genetic resources ICARDA (Aleppo, Syria) Indirect somatic embryogenesis (from callus) is
holds a Lathyrus collection of about 1880 ac- possible in grass pea using shoot tips, axillary
cessions, of which 1560 belong to Lathyrus sa- buds, and stem, leaf and root expiants. Direct
tivus. Large grass pea germplasm collections somatic embryogenesis has been achieved from
are also kept in France (IBEAS, Laboratoire immature leaflets and nodal segments. Soma-
d'Ecologie Moleculaire, Université de Pau; clones with low ODAP combined with high
1810 accessions), Australia (Australian Tem- yield have been developed. Other biotechno-
perate Field Crops Collection, Horsham, Victo- logical approaches applied in breeding for low
ria; 844 accessions), Russia (N.I. Vavilov All- ODAP grass pea types include incorporation
Russian Scientific Research Institute of Plant into grass pea of ODAP-degrading genes from
Industry, St. Petersburg; 688 accessions), microbes, and application of antisense technol-
Bangladesh (Plant Genetic Resources Centre, ogy to silence the genes involved in the bio-
Bangladesh Agricultural Research Institute synthesis of ODAP. Transgenic grass pea
(BARI), Joydebpur, Gazipur; 584 accessions) plants have been produced using bombardment
and the United States (USDA/ARS Western of expiants with DNA-coated particles. Genetic
Regional Plant Introduction Station, Pullman, linkage maps of the Lathyrus sativus genome
Washington; 248 accessions). In tropical Africa have been developed using various molecular
germplasm collections are kept in Ethiopia markers (RAPD, STMS and STS/CAPS), and
(197 accessions at the Institute of Biodiversity quantitative trait loci associated with resis-
Conservation, Addis Ababa; 13 accessions at tance to ascochyta blight (Mycosphaerella pi-
the International Livestock Research Institute nodes) have been located for possible future
(ILRI), Addis Ababa) and Kenya (National transfer of this trait into the closely related
Genebank of Kenya, Crop Plant Genetic Re- Pisum sativum L.
sources Centre, KARI, Kikuyu; 4 accessions). Prospects Grass pea is the least preferred
Grass pea seeds show orthodox seed storage among the common food legumes, but it has a
behaviour. number of features that make it attractive par-
Breeding The major objective in grass pea ticularly to resource-poor farmers, because of
breeding is reduction of ODAP levels, which is its adaptation to harsh conditions such as
the most feasible method of producing a safe drought and waterlogging. Therefore, grass pea
crop. Secondly, increasing the genetic yield is a useful crop for dry and poor soils and a
 LENS 91

rescue crop when other crops have failed. How-

ever, the presence in the seeds of the toxin
ODAP is a serious disadvantage, as it poses a
real danger to the health of consumers. Culti-
vation of grass pea is often discouraged or has
sometimes even been forbidden, e.g. in certain
states of India, but this has not been successful
due to the absence of cheap alternatives. The
first priority in grass pea breeding therefore is
the development of high-yielding cultivars with
low ODAP content, which can safely be con-
sumed. Also, more research is needed on effec-
tive detoxification methods without reducing
the nutritional value ofthe seeds.
Major references Campbell, 1997b; Camp-
bell et a l , 1994; Jansen, 1989a; Kay, 1979;
Kearney & Smartt, 1995; Kislev, 1989; Knight
(Editor), 2000; Muehlbauer & Kaiser (Editors), Lens culinaris -planted
1994; Smartt, 1984;Westphal, 1974.
Other references Akalu et al., 1998; Barna associated with many legends, tales and cus-
& Mehta, 1995; Chowdhury & Slinkard, 2000; toms, and it is the first pulse crop mentioned in
Croft, Pang & Taylor, 1999; Dadi et a l , 2003; the Bible. The oldest archaeological remains of
Getahun, Lambein & Vanhoorne, 2002; Geta- lentil are from Greece, dated 11,000 BC, and
hun et al., 2002; Hanbury et al., 2000; ILDIS, Syria, dated 8500-7500 BC. However, it is un-
2002; Leung, Busson & Jardin, 1968; Mehta, certain whether they were from cultivated
Ali & Barna, 1994; Mehta & Santha, 1996; plants or from wild ones. It is from the 5 th mil-
Mondai et al., 1998; Rotter, Marquardt & lennium BC that unequivocally domesticated
Campbell, 1991; Skiba, Ford & Pang, 2004 lentil seeds have been found. Lentil has been
Spencer, 1994; Tekle Haimanot et a l , 1993 introduced into the Americas, New Zealand
Thulin, 1989a; Wuletaw & Endashaw, 2003 and Australia. It is now widely cultivated in
Yunus &Jackson, 1991. temperate and subtropical regions, and in the
Sources of illustration Jansen, 1989a. tropics at higher elevations and in cool seasons.
Authors S.S. Yadav &G. Bejiga In tropical Africa it is grown in Sudan, Eritrea,
Ethiopia (mainly in the northern, central and
eastern Highlands), Kenya, Tanzania, Malawi,
L E N S CULINARIS Medik. Zimbabwe, Madagascar, Réunion and Mauri-
tius. It is also cultivated in Morocco, Tunisia,
Protologue Vorles. Churpfälz. Phys.-Okon. Algeria, Libya, Egypt and South Africa.
Ges. 2: 361 (1787). Uses Lentil is primarily grown for its ma-
Family Papilionaceae (Leguminosae - Papil- ture seeds, which are consumed mainly in
ionoideae, Fabaceace) sauces and soups. In Ethiopia they are used in
Chromosome number In — 14 'kik wot' (sauce of split seeds), soup (from
Synonyms Lens esculenta Moench (1794), whole seeds or flour), 'nufro' (boiled and
Vicia lens (L.) Coss. &Germ. (1845). salted), 'azifa' (cooked and mashed) and 'elbet'
Vernacular names Lentil, common lentil (paste from flour). Many other dishes are pre-
(En). Lentille, lentillon (Fr). Lentilha (Po). pared from lentil in different countries. Some
Mdengu (Sw). of these are: spicy lentil salad, lentil burgers
Origin and geographic distribution Lentil with coriander-yoghurt sauce, lentil and mush-
is one of the oldest pulse crops and of ancient room cottage pie and lentil potatoes. In India
cultivation in western Asia, Egypt and south- split seeds (dhal) are used in soups and the
ern Europe. It probably originated in western whole seed is eaten salted and fried. The seeds
Asia, from where it spread into the Mediterra- are ground into flour used for cakes and bread
nean region, Asia, Africa and Europe. Lentil and for the preparation of special foods, e.g. for
was a common part of the diet of the ancient infants and invalids. Young pods, sprouted
Greeks, Jews and Romans and was the main- seeds and leaves are eaten as vegetable.
stay of the poor, especially in Egypt. It was Lentil seeds are occasionally fed to animals as
92 CEREALSANDPULSES

a source of protein, particularly to poultry. hay contains moisture 10.2%, protein 4.4%, fat
They are sometimes used as a source of starch 1.8%, carbohydrate 50.0%, fibre 21.4% and ash
for the textile and printing industries. The 12.2%.
husks, bran and fresh or dried leafy stems pro- Description Erect, pale green annual herb
vide fodder for livestock. Lentil is sometimes up to 60(-75) cm tall; stem square, much-
grown for forage or as green manure, though branched; taproot slender. Leaves alternate,
the dry matter production is low. Lentil straw pinnately compound, with 5-16 leaflets; rachis
is used as fuel. The seeds are believed to rem- (1-)2.5-3.5(-5) cm long, usually ending in a
edy constipation and other intestinal problems. tendril or bristle; stipules entire, 2.5-6 mm
In India they are applied as a poultice to long; leaflets opposite or alternate, sessile, ob-
slow-healing sores. In Ethiopia the seeds are long or elliptical, (3-)10-15(-20) mm x (1.5-)2-
credited with aphrodisiac properties. 5(-8) mm, entire. Inflorescence an axillary ra-
Production and international trade Accord- ceme, l-4(-7)-flowered; peduncle slender, (2-)
ing to FAO statistics, the annual world lentil 3—4(-5.5) cm long. Flowers bisexual, papiliona-
production in 1999-2003 amounted to 3.1 mil- ceous; pedicel short; calyx campanulate, nar-
lion t/year from 3.8 million ha. The main pro- rowly 5-lobed, tube c. 1.5 mm long, lobes c. 3
ducers were India (948,000 t/year from 1.43 mm long; corolla pale blue, white or pink,
million ha), Canada (616,000 t/year from standard 5-7 mm x 4-5 mm, wings c. 4.5 mm x
554,000 ha) and Turkey (473,000 t/year from 1.5 mm, keel c. 4.5 mm x 2 mm; stamens 10, 9
490,000 ha). In tropical Africa the main pro- united and 1 free, anthers uniform; ovary su-
ducer is Ethiopia (47,000 t/year from 78,000 perior, 1-celled, style inflexed, inner surface
ha). About 60% of the lentil production in Af- bearded. Fruit a rhomboid, laterally com-
rica (including North Africa) comes from pressed pod, 6-20 mm x 3.5-12 mm, short-
Ethiopia, where the area under lentil has de- beaked, l-2(-3)-seeded. Seeds lens-shaped, 2-9
creased since the mid-1980s, but in the late mm x 2-3 mm, grey, green, brownish green,
1990s this trend reversed due to the release of pale red speckled with black, or black; hilum
cultivars with resistance to rust and fusarium
wilt. In Malawi lentil is grown in the northern
part (near Mzimba) to supply the Indian com-
munity. World lentil exports in 1998-2002
amounted to about 1 million t/year. The main
exporters were Canada (430,000 t/year), Tur-
key (127,000 t/year), Australia (124,000 t/year)
and India (120,000 t/year). The main importers
were Egypt (90,000 t/year), Sri Lanka (86,000
t/year) and Turkey (81,000 t/year).
Properties The composition of mature raw
lentil seeds per 100 g edible portion is: water
11.2 g, energy 1413 kJ (338 kcal), protein 28.1
g, fat 1.0 g, carbohydrate 57.1 g, dietary fibre
30.5 g, Ca 51 mg, Mg 107 mg, P 454 mg, Fe 9.0
mg, Zn 3.6 mg, vitamin A 39 IU, thiamin 0.48
mg, riboflavin 0.25 mg, niacin 2.6 mg, vitamin
B60.54 mg, folate 433 (ig and ascorbic acid 6.2
mg. The essential amino-acid composition per
100 g edible portion is: tryptophan 251 mg,
lysine 1957 mg, methionine 238 mg, phenyla-
lanine 1383 mg, threonine 1006 mg, valine
1392 mg, leucine 2034 mg and isoleucine 1212
mg (USDA, 2004). The main limiting amino
acids are methionine and cystine. Antinutri-
tional factors include trypsin inhibitors, hae-
magglutinins, tannins, phytate and oligosac-
charides, but the levels are considerably lower Lens culinaris - 1, flowering and fruiting
than those in e.g. pea and faba bean, and len- branch; 2, seeds.
tils are considered more easily digested. Lentil Source: PROSEA
 LENS 93

minute. Seedling with hypogeal germination. cross-pollination by insects may occur. The
Other botanical information In a recent growth cycle ranges from 80—110 days for
revision of Lens 4 species are recognized on the early-maturing cultivars to 125-130 days for
basis of morphological characters, crossability late-maturing ones. Lentil is effectively nodu-
relationships and cytogenetic, biochemical and lated by Rhizobium leguminosarum.
molecular evidence; these are Lens culinaris Ecology Lentil is grown as a summer an-
(containing wild and cultivated types) and 3 nual in temperate regions and as a winter an-
wild species: Lens ervoides (Brign.) Grande, nual in subtropical regions. In the tropics it is
Lens nigricans (M.Bieb.) Godr. and Lens lamot- cultivated at higher elevations (1800—2500(-
tei Czefr. Lens ervoides is found in East Africa 2700) m in Ethiopia) or as a cool season crop. It
(Ethiopia and Uganda). grows at mean temperatures of 6-27°C, but
Lens culinaris has been divided into 4 subspe- lentil is not suited to the hot and humid trop-
cies (1cultivated and 3wild): ics. Intense or prolonged frost and tempera-
- subsp. culinaris: stipules entire, lanceolate, tures much higher than 27°C seriously affect
pod indéhiscent, glabrous, seed coat spotted; growth. Lentil requires an annual rainfall of
the cultivated lentil; about 750 mm, with dry conditions around
- subsp. odemensis (Ladiz.) M.E.Ferguson et harvest time, but an annual rainfall of 300-
al. (synonym: Lens odemensis Ladiz.): stip- 2400 mm is tolerated. It is moderately tolerant
ules slightly hastate, at least the lower ones to drought, but differences exist between culti-
slightly toothed, pod dehiscent, glabrous, vars. Lentil normally requires long daylengths
seed coat with W-shaped pattern; native to for flowering, but the response varies among
Libya, Israel, Turkey and Greece; genotypes, and some cultivars are daylength
- subsp. orientalis (Boiss.) Ponert (synonym: insensitive. In Ethiopia lentil is grown in the
Lens orientalis (Boiss.) Hand.-Mazz.): stip- short rainy season ('belg', February-May) and
ules entire, obliquely lanceolate, pod dehis- during the main rainy season ('kiremt', J u n e -
cent, glabrous, seed coat usually spotted; the December), the latter being predominant. To
wild progenitor of the cultivated lentil, dis- avoid waterlogging the 'kiremt' crop is sown on
tributed from Greece to Uzbekistan and from Vertisols at the end of the rainy season (Sep-
the Crimean Peninsula to Jordan; tember) and grown on residual soil moisture.
- subsp. tomentosus (Ladiz.) M.E.Ferguson et In India lentil is grown during winter on resid-
al. (synonym: Lens tomentosus Ladiz.): stip- ual soil moisture. Lentil can be grown on a
ules entire, obliquely lanceolate, pod dehis- wide range of soil types, from sandy to fairly
cent, tomentose, seed coat spotted; native to heavy clay soils, but does not tolerate flooded
Syria and Turkey. or waterlogged soils. A pH near 7.0 is best for
Lentil cultivars have been divided into 2 culti- lentil production, but it tolerates a pH of4.5—
var groups, based mainly on seed size: 9.0. Lentil is generally very sensitive to salin-
- Microsperma Group: flowers small (5-7 mm ity.
long), violet-blue to white or pink, pods Propagation and planting Lentil is prop-
small, convex, seeds small (diameter less agated by seed. The 1000-seed weight ranges
than 6 mm, 1000-seed weight less than 45g), from 10—90 g. Seeds remain viable for more
convex, cotyledons red, orange or yellow; than 5 years under cool and dry storage condi-
- Macrosperma Group: flowers large (7-8 mm tions. A dormancy period of 4—6weeks is com-
long), white, rarely blue, pods large, gener- mon, and some cultivars have been found to be
ally flat, seeds large (diameter more than 6 responsive to vernalization. The minimum
mm, 1000-seed weight more than 45 g), flat- temperature for germination is 15°C and the
tened, cotyledons generally yellow, some- optimum temperature 18—21°C; temperatures
times orange. above 27°C are harmful. A firm, smooth seed-
Macrosperma Group predominates in North bed is best for lentil. The seed is broadcast, or
Africa, Europe and America, Microsperma planted in rows 20-90 cm apart with 5-25 cm
Group in Asia, Egypt and Ethiopia. In western between plants within the row. Seed rates
Asia and south-eastern Europe both cultivar range from only 10 kg/ha in intercropping sys-
groups are grown. tems to 150 kg/ha for sole-cropped large-seeded
Growth and development At optimum cultivars. The sowing depth is 1-6 cm depend-
temperatures lentil seeds germinate in 5-6 ing on seed size and moisture availability. Len-
days. Flowering starts 6—7weeks after sowing. til is mainly grown as a sole crop, but some-
Lentil is usually self-fertilized, but up to 1% times mixed with other crops, e.g. in India with
94 CEREALS AND PULSES

barley, mustard or castor. virus (AMV) and tomato spotted wilt virus
Management Lentil is a poor competitor (TSWV). Pea seed-borne mosaic virus (PSbMV)
with weeds, especially when young. It should is common in Ethiopia.
be sown in a clean field and weeding should Among the common insect pests of lentil,
generally be done within 3 weeks after sowing. aphids are important. The pea aphid (Acrytho-
Lentil normally responds well to P fertilizer. siphon pisum) is the most important aphid in
Effectively nodulated lentil seldom responds to Ethiopia, causing up to 25% yield loss. Stored
application of N fertilizer. Alentil crop yielding seeds are attractive to bruchids (Calloso-
about 2 t seed per ha takes up about 100 kg N, bruchus spp.). Broomrape (Orobanche spp.) is
12 kg P and 65 kg K per ha. In Sudan lentil is an important parasitic weed on lentil in the
grown under irrigation, but elsewhere in tropi- Mediterranean region and western Asia; it is
cal Africa it is a rainfed crop. In Ethiopia it is difficult to control by management practices or
often grown in rotation with the major small- genetic means.
grain cereals. In crop rotations planting lentil Harvesting Lentil is harvested when the
after other legumes, Brassica crops, sunflower pods turn yellow-brown and the lower ones are
or potato should be avoided because these are still firm. Further delay may lead to shatter-
susceptible to the same diseases. ing. In many areas the plant is cut down
Diseases and pests The economically most manually to ground level and left to dry for
important diseases of lentil are rust (Uromyces about 10 days, before being threshed and win-
viciae-fabae), Ascochyta blight (Ascochyta fabae nowed. Alternatively, for instance in Ethiopia,
f.sp. lentis), grey mould (Botrytis cinerea), lentils are harvested by hand-pulling the
Stemphylium blight (Stemphylium botryosum), plants, after which they are left in the field to
collar rot (Sclerotium rolfsii) and fusarium wilt dry to a seed moisture content of 12-13%. In
(Fusarium oxysporum f.sp. lentis). Other fungal the United States lentil is harvested mechani-
diseases include Rhizoctonia root rot (Rhizoc- cally, preferably at a moisture level of 18-20%
tonia solani), powdery mildew (Erysiphe poly- to prevent excessive shattering and seed dam-
goni, Leveillula taurica), anthracnose (Colleto- age.
trichum spp.), leaf spot (Alternaria alternata) Yield The average lentil seed yield in Ethio-
and Sclerotinia stem and root rot (Sclerotinia pia is about 600 kg/ha, which is below the av-
sclerotiorum). Rust, fusarium wilt and root rot erage world yield of about 800 kg/ha. In the
are the most important diseases in Sudan, Eri- Ethiopian highlands, where the growing period
trea and Ethiopia. Yield losses of 10% due to is long, yields of about 4 t/ha have been ob-
rust and 50% due to fusarium wilt and root rot tained in experiments, and more than 2 t/ha in
have been recorded on Vertisol-grown lentil in farmer's fields when the recommended agro-
Ethiopia. Symptoms of rust are leaves and nomic package was applied. In Asia average
stems losing their green colour and turning seed yields are 300—600 kg/ha in mixed crops
purple, in case of serious infection leading to and 900—1100 kg/ha for sole crops. Leafy stem
death of the plant. The spread of rust is fa- yields of up to 7 t/ha are possible for late-type
voured by high humidity and moderate tem- lentils in Ethiopia.
peratures (17-25°C). Control measures include Handling after harvest Harvested lentil
destruction of diseased plants, treatment of should be dried to a moisture content of11—
seed with fungicide, and the use of resistant 14%; at a lower moisture content seeds tend to
cultivars. Fusarium wilt causes leaf curling, break. In Ethiopia the dried plants are spread
followed by wilting of individual branches or on a cemented area, where they are threshed
the whole plant. It is favoured by light, dry by animals, after which the seeds are sepa-
soils. Suggested control measures are crop ro- rated from the residues by winnowing. The
tation, treatment of seed with fungicide, and clean seeds are stored as whole seeds or in de-
the use of resistant cultivars. Integrated dis- hulled form. Because of storage insects, mainly
ease management packages have been devel- Callosobruchus spp., lentil seeds are not stored
oped to control wilt and root rot in Ethiopia for more than half a year, except where pit
and Sudan. Seed treatment compounds should (underground) storage is used. Mechanically
be selected and used with care as they can in- harvested lentil seeds can be dried in heated
terfere with the nodulation process. Several air dryers, but the temperature should not
virus diseases affect lentil, the most important exceed 43°C to reduce cracking ofthe seed coat.
being cucumber mosaic virus (CMV), faba bean Genetic resources The largest lentil germ-
necrotic yellows virus (FBNYV), alfalfa mosaic plasm collection is that of ICARDA (Interna-
 LENS 95

tional Centre for Agricultural Research in Dry Wild relatives are considered potentially valu-
Areas, Aleppo, Syria), with about 10,000 acces- able to improve the tolerance to environmental
sions, including wild Lens. Large collections stresses. Different institutions are studying
are also kept at the Australian Temperate crossability of these wild relatives among
Field Crops Collection (Horsham, Victoria, themselves and with cultivated lentil. Crosses
Australia, about 4800 accessions), the USDA- between Lens culinaris and Lens ervoides or
ARS Western Regional Plant Introduction Sta- Lens nigricans usually abort, but Fi hybrids
tion (Pullman, Washington, United States, can be rescued and produce viable and largely
about 2800 accessions), and the N.I. Vavilov fertile Fa segregates.
Ail-Russian Scientific Research Institute of Successful tissue culture of lentil has been
Plant Industry (St. Petersburg, Russia, about achieved with shoot apical meristem tips, nodal
2400 accessions). The largest lentil germplasm segments and intact seedlings. Genetic trans-
collection in tropical Africa (about 370 acces- formation of lentil plants has been achieved by
sions) is held by the Institute of Biodiversity electroporation, particle bombardment and
Conservation (IBC), Addis Ababa, Ethiopia, a Agrobacterium-mediated methods. Fertile trans-
country which is considered as a secondary genic plants have been obtained using particle
centre of diversity for lentil. Some accessions of bombardment. Genetic linkage maps for lentil
Lens ervoides have been collected by IBC. Cul- have been developed.
tivated lentil shows a wide range of morpho- P r o s p e c t s Lentil seeds are tasty, relatively
logical variation, in vegetative as well as in easily cooked and have excellent nutritional
generative parts. Analyses using biochemical qualities because of the high protein content
and molecular markers such as RFLPs and and good digestibility. The susceptibility of
RAPDs generally show little genetic variation, lentil to diseases, especially rust and wilt, has
but more variation is revealed when ISSR limited its development, but considerable pro-
markers are used. gress has been achieved in breeding for resis-
Breeding Like in many other self-polli- tance to major diseases. In North and East
nating crops, the genetic variation in lentil has Africa the demand for lentil remains high
been structured into true-breeding landraces while the area and production remained con-
endemic to restricted areas. Since the 1920s stant or declined until the late 1990s, but re-
breeding work has focused on the collection covered thereafter. Currently, there is also an
and evaluation of landraces, on the basis of increasing export demand, which can be met
yield, seed size and disease resistance. Selec- with research and development efforts towards
tion is now complemented by crossing pro- increasing yield, setting up seed supply
grammes, the main breeding objective being schemes and improving quality through proc-
yield, but also with attention to broad adapta- essing industries. The crop can be grown in
tion, tolerance to environmental stresses, resis- various agro-ecological zones and is useful in
tance to diseases and pests, and nutritional rotations with cereals. Therefore, its role in
quality. Considerable progress has been crop production systems remains important,
achieved in breeding for resistance to rust, especially in Ethiopia.
wilt, Ascochyta blight and Stemphylium blight. Major references Bayaa & Erskine, 1998;
ICARDA has the global mandate for research Ferguson et al., 2000; Jansen, 1989b; Kay,
on lentil improvement. National lentil im- 1979; Knight (Editor), 2000; Muehlbauer,
provement programmes in lentil-producing Cubero & Summerfield, 1985; Muehlbauer &
countries use their own germplasm collections Kaiser (Editors), 1994; Telaye et al. (Editors),
and introductions from other institutes for 1994; Webb &Hawtin, 1981;Zohary, 1995.
their breeding programmes. The national pro- Other references Abraham & Makkouk,
grammes of lentil-producing countries have 2002; Bejiga, Tsegaye & Tullu, 1995; Bejiga et
released many cultivars. Ethiopia, for instance, al., 1996; Durân et a l , 2004; Erskine, 1997;
has released 10 cultivars ('EL-142', 'R-186', Gulati, Schryer & McHughen, 2002; Hawtin &
'Chalew', 'Chekol', 'Adaa', 'Gudo', 'Alemaya', Chancellor (Editors), 1979; ICARDA, 2002;
'Assano', 'Alem Tena' and 'Teshale') and others Lock, 1989; Polhill, 1990; Popelka, Terryn &
are being developed for different agro- Higgins, 2004; Rubeena, Ford & Taylor, 2003;
ecological zones. Cultivars derived from hy- Smartt, 1976; Sonnante & Pignone, 2001;
bridization schemes are also being developed. Summerfield (Editor), 1988; Tadesse et al.,
Sudan has released some cultivars for its irri- 1999; Thulin, 1983; USDA, 2004; Westphal,
gated agriculture. 1974;Williams et a l , 1994.
96 CEREALSANDPULSES

Sources of illustration Jansen, 1989b. 2000; Gast, 2000; Gilbert, 2000; Keay, 1954.
Authors G. Bejiga Other references Ozenda, 1977.
Authors M. Brink

LlMEUM OBOVATUM V i c a r y
LUPINUSALBUSL.
Protologue Journ. As. Soc. Beng. 16: 1163
(1847). Protologue Sp. pi. 2: 721 (1753).
Family Molluginaceae Family Papilionaceae (Leguminosae - Papil-
Chromosome number n = 9 ionoideae, Fabaceae)
Synonyms Limeum indicum Stocks ex Chromosome number 2n = 50
T.Anderson (1861). Vernacular names White lupin, Egyptian
Origin and geographic distribution Lime- lupin (En). Lupin blanc, lupin (Fr). Tremoceiro,
um obovatum is distributed in the desert re- tremoceiro branco, tremoceiro da Beira, tre-
gions of Mauritania, Mali, Niger, Chad, Sudan moço(Po).
and Eritrea and through northern Africa and Origin and geographic distribution White
Arabia to Pakistan. lupin originates from south-eastern Europe
Uses In Tibesti (northern Chad) the seeds of and western Asia where wild types still occur.
Limeum obovatum are collected as food. They It is known to have been cultivated since an-
are a famine food for the Touareg in the Hog- cient times in Greece, Italy, Egypt and Cyprus.
gar Mountains in southern Algeria. In Kordo- The importance of white lupin has fluctuated
fan (Sudan) the plant in all growth stages is often during the history of its cultivation; at
reportedly grazed by livestock, especially present it has almost disappeared in central
sheep. In Chad the plant is used for the treat- Europe, while it is becoming more widely
ment of burns. grown in the Americas. Today it is a traditional
Botany Annual or short-lived perennial, minor pulse crop, grown around the Mediter-
glandular-pubescent herb; stems prostrate, up ranean and the Black Sea, and in the Nile val-
to 40 cm long, pale brown, strongly branched. ley, extending to Sudan and Ethiopia. It is also
Leaves opposite, simple and entire; stipules occasionally grown elsewhere, e.g. in Kenya,
absent; petiole up to 5 mm long;blade orbicular Tanzania, Zimbabwe, South Africa, Mauritius,
to obovate or elliptical, up to 12 mm x 10 mm, United States and South America (mainly Bra-
cuneate at base, rounded at apex. Inflorescence zil and Chile).
an apparently axillary cyme up to 5 mm across. Uses White lupin is traditionally cultivated
Flowers bisexual, regular, small, green; sepals for human consumption, green manuring and
5, ovate, c. 2.5 mm long, acuminate; petals ab- as forage. Before consumption, seeds are first
sent; stamens 7, inserted on a disk; ovary su- soaked for 1-3 days in running water to re-
perior, 2-celled, styles 2. Fruit splitting into 2 move the bitter, toxic alkaloids, then cooked
mericarps; mericarp indéhiscent, hemispheri- and eaten as a pulse or pickled in brine and
cal, smooth, pale brown, 1-seeded. served as a snack. In Ethiopia a high-quality
Limeum comprises about 20 species and is dis- spirit ('araki') is distilled from fermented seeds.
tributed in the Old World tropics, with the cen- In general, consumption of white lupin seeds is
tre of distribution in south-western Africa. restricted to low-income classes and to times of
Limeum is sometimes included in the family drought, because of their bitter taste. Modern
Aizoaceae and occasionally in Phytolaccaceae. sweet cultivars have very low alkaloid con-
Ecology Limeum obovatum occurs on dry tents, and their seeds do not require laborious
sandy soils, often in dry riverbeds. In Eritrea it detoxification; they are a promising nutritive
is found in sandy locations on coastal plains. pulse and can be used as a rich additive for
Genetic resources and breeding It is un- human food and livestock feed products.
likely that Limeum obovatum is threatened by White lupin plants are fed to livestock as fresh
genetic erosion in the light of its wide occur- or dry fodder. In southern Europe it is a tradi-
rence and habitat conditions. tional green manure crop in vineyards and
Prospects Limeum obovatum seems to be a olive plantations. White lupin is a good honey
useful wild source of food and fodder in desert plant and an attractive annual ornamental. In
regions. However, research on the nutritional traditional medicine it is used for various ail-
and chemical properties of the seeds is needed. ments, e.g. as an anthelmintic, carminative,
Major references Burkill, 1985; Burkill, deobstruent, diuretic and pectoral. Lupin meal
 LUPINUS 97

mixed with honey or vinegar is used as a cure ingestion of toxins, known as phomopsins, pro-
for worms, while infusions or poultices are ap- duced by the fungus Diaporthe toxica that colo-
plied to treat boils and skin complaints. Burn- nizes lupin plants. It is primarily a disease of
ing seeds are used as an insect repellant. sheep but can also occur in other livestock and
Production and international trade No spe- is characterized by severe liver damage, which
cific statistics are available for Lupinus albus. results in loss of appetite and condition, leth-
About 2 million ha are cultivated with lupin argy, jaundice and often death. Most of the
(all species) worldwide, of which 60% is mainly problem can be solved by using Diaporthe-
for seed production and 40% for forage and resistant cultivars such as 'Kiev' and 'Ultra';
green manure. The major producer of lupin where other cultivars are used, overfeeding
seed is Australia, with about 1.4 million t/year must be avoided.
from 1.2 million ha in the early 1990s, mainly Botany Annual, erect, branched, bushy,
Lupinus angustifolius L. for livestock feed. short-hairy herb up to 120 cm tall, with a
Properties Mature, raw Lupinus albus strong taproot. Leaves alternate, digitately
seeds contain per 100 g edible portion: water compound with 5-9 leaflets; stipules linear to
10.4 g, energy 1552 k j (371 kcal), protein 36.2 narrowly triangular, up to 1 cm adnate to the
g, fat 9.7 g, carbohydrate 40.4 g, Ca 176 mg, base of the petiole; petiole 3.5—7(—12) cm long;
Mg 198 mg, P 440 mg, Fe 4.4 mg, Zn 4.8 mg, leaflets obovate, 2-6 cm x 0.5-2 cm, cuneate at
vitamin A 23 IU, thiamin 0.64 mg, riboflavin base, rounded and mucronate at apex, nearly
0.22 mg, niacin 2.2 mg, vitamin He 0.36 mg, glabrous above, hairy beneath. Inflorescence a
folate 355 |ig and ascorbic acid 4.8 mg. The terminal false raceme 3—30 cm long, many-
essential amino-acid composition per 100 g flowered, lower flowers alternate, upper ones in
edible portion is: tryptophan 289 mg, lysine whorls; peduncle short or absent. Flowers bi-
1933 mg, methionine 255 mg, phenylalanine sexual, papilionaceous; pedicel 1—2 mm long;
1435 mg, threonine 1331 mg, valine 1510 mg, calyx 8-14 mm long, densely hairy outside,
leucine 2743 mg and isoleucine 1615 mg. The tube c. 4 mm long, 2-lipped, upper lip entire,
principal fatty acids are per 100 g edible por- lower lip entire or slightly 3-toothed; corolla
tion: oleic acid 3558 mg, linoleic acid 1995 mg, white to violet-blue, standard obovate, 15—18
palmitic acid 742 mg, linolenic acid 446 mg, mm x 8-12 mm, margins partly reflexed, wings
stearic acid 316 mg and eicosenoic acid 255 mg obovate, 13-17 mm x 6-10 mm, keel ladle-
(USDA, 2005). The seed coat makes up about shaped, 12-15 mm x 4 mm, beaked; stamens
15% of the seed weight. The net protein utiliza- 10, all joined into a tube; ovary superior, 1-
tion for humans is 77%and the protein fraction celled, style c. 7.5 mm long with a ring of small
is low in lysine and methionine. The levels of hairs below the stigma. Fruit a narrowly ob-
antinutritional compounds such as condensed long, laterally compressed pod 6-15 cm x 1-2
tannins and trypsin inhibitors are lower than cm, bulging over the seeds, shortly hairy but
in other pulses. Suspensions of ground seed of glabrescent, yellow, 3-6-seeded. Seeds rectan-
white lupin have shown hypoglycaemic effects gular or square with rounded corners, laterally
in rats. compressed, 7-16 mm x 6-12 mm x 2-5 mm,
The major alkaloids of white lupin are lu- more or less smooth, white variably tinged
panine, 13-hydroxylupanine and sparteine. The salmon pink or mottled dark brown. Seedling
pharmacological effects of these alkaloids are with epigeal germination.
that they block ganglionic transmission, de- Lupinus comprises about 200 species, mostly
crease cardiac contractility and contract uter- American; only 12 species are native to the Old
ine smooth muscle. 'Sweet lupin' is defined as World. In tropical Africa 3 native or natural-
having less than 200 mg alkaloids/kg; it can be ized species occur and another 9 species have
consumed without special precautions. In bit- been introduced. Many Lupinus spp. are orna-
ter cultivars the alkaloids, which are water- mental garden plants, and 4 species are culti-
soluble, can be soaked out from seeds in run- vated on a larger scale as agricultural crops.
ning water. Processing techniques such as Lupinus albus represents a crop-weed complex
sprouting and fermentation into tempeh also with wide variability in wild and cultivated
strongly reduce the alkaloid content. Best con- types. Both types have been classified as sub-
trol is achieved by chemical extraction, which species although for the cultivated types a clas-
currently is not economically feasible. sification into cultivar groups and cultivars
When white lupin is fed as dried forage, lupi- would be more appropriate. The wild type is
nosis can occur. This disease is caused by the distinguished as subsp. graecus (Boiss. &
98 C E R E A L S AND PULSES

Spruner) Franco & P.Silva (synonym: Lupinus Management White lupin is propagated by
graecus Boiss. & Spruner) and can be found in seed. The 1000-seed weight ranges from 70 g in
south-eastern Europe and western Asia. The some Kenyan populations to more than 1 kg in
corolla is dark violet-blue, pods are 6—8cm x 1— modern seed cultivars. Seed can easily be
1.5 cm and shatter seeds at maturity, and stored for 2—4 years under normal conditions;
seeds are small, 7-10 mm x 6-8 mm x 2-3 mm, longer storage is possible at lower tempera-
mottled dark brown with impermeable seed tures. In areas with mild winters such as the
coat. The cultivated types are distinguished as Mediterranean region, seed is sown broadcast
subsp. albus (synonym: Lupinus termis or drilled from mid September to late October.
Forssk.), with corolla white, pods 9-15 cm x The seed rate is 50-180 kg/ha, the seeding
1.5-2 cm with non-shattering seeds at matur- depth 2.5—5cm. In Ethiopia seed is sown in the
ity, and seeds 10-14 mm x 8-12 mm x 3-5 main rainy season (July-September). White
mm, pinkish white or white with permeable lupin is often grown intercropped with cereals
seed coat. In Ethiopia 2 types of cultivated or forage legumes, or in rotation with cereals.
plants are found: a large-seeded type as grown Weed control is essential until a closed canopy
in Egypt and Sudan, but also a small-seeded is formed. White lupin is sensitive to P defi-
type with small leaves. In northern parts of the ciency, but its roots can make more P available
distribution area of white lupin, in South Af- through acidification of the rhizosphere, a
rica and in the Americas mostly sweet (low- property from which also associated crops
alkaloid) modern cultivars are grown, whereas benefit. Wheat intercropped with white lupin
in the Mediterranean region and eastern Africa has access to a larger pool of P, Mn and N than
bitter landraces prevail. sole-cropped wheat. Inoculation of the soil with
White lupin is mainly self-pollinating, but 5— Bradyrhizobium bacteria is beneficial, giving
10% outcrossing can occur. White lupin nodu- up to a 5-fold increase in yield and a higher
lates effectively with Bradyrhizobium bacteria. protein content of the seed. A well-known
Atmospheric nitrogen fixation rates up to 400 commercially available inoculant strain is the
kg N per ha have been observed in Europe and Australian WU425.
Australia. The major diseases of white lupin are root rot
Ecology Wild white lupin prefers disturbed and brown leaf spot caused by Pleiochaeta se-
sites and poor soils, where there is less compe- tosa, anthracnose (Colletotrichum acutatum),
tition from other species. White lupin is usu- resulting in early plant death through stem
ally grown at mean monthly temperatures dur- breakage, and rust (Uromyces lupinicolus).
ing the growing season of 15-25°C, the opti- Sources of resistance to anthracnose have been
mum being 18-24CC. Higher temperatures and found in Ethiopian landraces, but resistant
moisture stress hinder flowering and pod set- cultivars are not yet available. Bean Yellow
ting. White lupin is cold-tolerant, but tempera- Mosaic Virus (BYMV) is the major virus dis-
tures of—6 to -8°C are harmful at germination, ease; it is transmitted by aphids and by seed.
temperatures of - 3 to —5°C at flowering. A No sources of resistance have yet been identi-
rainfall of 400-1000 mm during the growing fied. White lupin is immune to Cucumber Mo-
period is optimal for yield. Lupin species are saic Virus (CMV), a major disease of other Lu-
drought-tolerant due to their deep roots, but pinus spp. Major pests are the bean seedling
are sensitive to moisture deficiency during the maggot (Delia platura, synonym: Phorbia pla-
reproductive period. tura) causing seedlings to wilt and die, beetle
White lupin is adapted to well-drained, mildly and moth larvae (e.g.Agriotes and Agrotis spp.
acid or neutral soils of light to medium texture, killing seedlings), slugs (attacking leaves),
with pH 4.5-7.5. Growth is hampered on heavy thrips (Frankliniella spp., attacking flower
clay and waterlogged soils, while calcareous or buds and leaves), mirid bugs (attacking young
alkaline soils induce chlorosis and reduce pods) and budworms (e.g. Helicoverpa armigera
growth, frequently precluding cultivation. The feeding on pod and seed). In Ethiopia harvest-
accepted maximum soil level of CaC03 is 3-5 ing is in December. Seed yields are 500-4000
g/100 g. Some cultivars of white lupin are more kg/ha.
tolerant to soil salinity and heavy soils than Genetic resources and breeding Major
most other crops. germplasm collections of white lupin are avail-
In Ethiopia white lupin is grown at 1500-3000 able in France (INRA, Station d'Amélioration
m altitude, on soils too poor for a good faba des Plantes Fourragères, Lusignan, 1400 ac-
bean crop. cessions), the United Kingdom (University of
 MACROTYLOMA 99

Reading, Reading, 1100 accessions), Australia MACROTYLOMAGEOCARPUM (Harms)

(Western Australian Department of Agricul- Maréchal & Baudet
ture, South Perth, 890 accessions) and Spain
(Servicio de Investigation y Desarrollo Tec- Protologue Bull. Jard. Bot. Belg. 47(1-2):
nológico, Guadajira, 690 accessions). In tropical 50 (1977).
Africa small collections are held in Ethiopia Family Papilionaceae (Leguminosae - Papil-
(International Livestock Research Institute ionoideae, Fabaceae)
(ILRI), Addis Ababa, Ethiopia, 25 accessions) Chromosome number 2n = 20, 22
and Kenya (National Genebank of Kenya, Crop Synonyms Kerstingiella geocarpa Harms
Plant Genetic Resources Centre, KARI, Ki- (1908).
kuyu, 20 accessions). Vernacular names Kersting's groundnut,
Major objectives in breeding of white lupin are geocarpa groundnut, ground bean (En).
to produce rapid-growing, alkaloid-free, dis- Lentille de terre, fève de Kandale, doï, dohi
ease-resistant (particularly against anthrac- (Fr).
nose), high-yielding, high-pH-tolerant, frost- Origin and geographic distribution The or-
tolerant dwarf cultivars, well adapted to local igin of Kersting's groundnut is not known, but
ecological conditions. It appears that bitter it may originate from northern Togo or central
cultivars tolerate cold and disease stress better Benin. Supposedly wild types of Kersting's
than sweet ones. The level of cross-pollination groundnut are found in Cameroon and the
may limit the relevance of sweet white lupin Central African Republic, but these may be
cultivars in regions where also bitter weedy or considered as representing a separate though
cultivated types are present, because pollen of related species. Kersting's groundnut is
the latter would reintroduce the bitter charac- cultivated in the West African savanna zone,
ter in farm-saved sowing seed. Sweet cultivars, from Senegal to Nigeria and Cameroon. It is
however, are a prerequisite for any further also grown in Mauritius and Fiji and has been
breeding advancement. Commercial cultivars grown in Tanzania. Kersting's groundnut is
are pure lines bred through pedigree selection. often said to be grown by elderly people only,
Some well-known cultivars of white lupin are: e.g. in Ghana, and thus gradually going out of
'Eldo', 'Kiev', 'Multolupa' and 'Ultra'. From production.
Ethiopia 'Bahar Dar' is known. Uses Kersting's groundnut is cultivated
Prospects White lupin is a promising an- primarily for its edible seeds. Mature seeds are
nual legume crop for human consumption, boiled with salt and eaten with palm oil or
green manuring and forage. The composition of groundnut oil, and accompanied with fer-
the seed and especially the high protein con- mented cassava flour, called 'gari', yams or
tent makes white lupin highly suitable for live- rice. They may also be boiled in soups and
stock diets as a protein-rich product in inten- served to guests as a sign of honour. Dry seeds
sive farming systems. The low level of antinu- are ground into flour used in making cakes and
tritional factors facilitates a direct on-farm use
of white lupin in self-sustained systems. Since
it often can grow on land unsuitable for other
crops (too saline, heavy, acid or poor), the de-
velopment of cultivars adapted to tropical Afri-
can conditions is highly recommended. Much
can be learned from the excellent results ob-
tained with Lupinus angustifolius in Australia.
Major references Cowling, Buirchell &
Tapia, 1998; Gladstones, Atkins & Hamblin
(Editors), 1998; Huyghe, 1997; van Santen et
al. (Editors), 2000; Westphal, 1974.
Other references al-Zaid et al., 1991;
Duke, 1981;Hanelt &Institute of Plant Genet-
ics and Crop Plant Research (Editors), 2001;
Haq, 1993;Hill, 1998;Kay, 1979; López-Bellido
& Fuentes, 1997; Thulin, 1989a; UC SAREP,
undated; USDA, 2005.
Authors P.C.M. Jansen Macrotyloma geocarpum - wild and planted
100 CEREALS AND PULSES

other dishes. In central Benin, for instance, nutritional factors, including tannins, haemag-
people eat a fried paste made from the seed glutinins and phytate. Boiling pre-soaked (12
alone ('ata') or with fermented maize paste hours at 27°C) seeds for 30 minutes reduces
('akassa'). Sometimes roasted seeds of black- tannin content by 98%, haemagglutinating
seeded types or fresh unshelled pods are boiled activity by 100%and phytate level by 70%.
with salt and eaten as snacks. Kersting's Description Annual herb with prostrate
groundnut seeds play an important role in rooting stems; stem pubescent or almost
traditional customs in West Africa, particularly glabrous, up to 10 cm long. Leaves alternate, 3-
in Togo, where they are used in funeral foliolate; stipules triangular-ovate, 2-7 mm
ceremonies of the Kabyé and Mauba people. long, pubescent; petiole erect, up to 25 cm long;
This seems to have contributed largely to the rachis c. 7 mm long; stipels linear-lanceolate,
survival of the crop in northern Togo. In many 2-5 mm long; petiolules hirsute, lateral ones 1-
areas consumption is limited to the male 2 mm long, terminal one 4-10 mm long;
members of the family, the headman in leaflets broadly ovate or obovate, 3-8 cm x 2 -
particular, and the seeds are a favourite dish of 5.5 cm, glabrous, 3-veined from the base.
voodoo priests. In the tradition of the Sisala Flowers in pairs or solitary in leaf axils, bi-
people of northern Ghana boiled seeds are the sexual, papilionaceous, almost sessile; bract-
only food served to surviving children during eoles lanceolate, (1—)3.5—4 mm long; calyx
the funeral oftheir mothers. pilose, tube 2-2.5 mm long, lobes linear-
The leaves of Kersting's groundnut are some- lanceolate, (2-)3.5-4 mm long; corolla white or
times eaten as a vegetable or in soups. In greenish-white, sometimes tinged with purple,
northern Ghana and central Benin the water standard 6-10 mm x 4.5-6 mm, wings 6-7 mm
in which the seeds have been boiled is taken x 1.5 mm, keel 5.5—6mm x 1 mm; stamens 10,
against diarrhoea. Powdered dry seed mixed 9 fused and 1 free; ovary superior, shortly
with water or local beer ('pita') is used as an stalked but stalk elongating during fruit
emetic in case of poisoning. Leaf decoctions act development, 1-celled, style slender, curved,
as a vermifuge. The Igbo of Nigeria use the stigma minute. Fruit an indéhiscent pod 0.5-
plant in the treatment of dysentery, venereal 2.5 cm x 0.5-1 cm, on a stalk up to 2 cm long,
diseases, fever and diabetes. In arid zones, the
vegetative parts serve as fodder after the
harvest.
Production and international trade Relia-
ble production statistics for Kersting's
groundnut are not available because it is of - "H' ',
little economic importance and mainly grown
for local consumption. Some trade exists
between neighbouring countries such as Togo,
Benin and Nigeria, but no statistics exist.
Because of the low yield and poor storage
capability the economic importance of Ker-
sting's groundnut has decreased considerably
in recent times.
Properties The composition of dried Ker-
sting's groundnut seeds per 100 g edible
portion is: water 9.7 g, energy 1457 kJ (348
kcal), protein 19.4 g, fat 1.1 g, carbohydrate
66.6 g, fibre 5.5 g, Ca 103 mg, P 392 mg, Fe
15.0 mg, thiamin 0.76 mg, riboflavin 0.19 mg,
niacin 2.3 mg and ascorbic acid 0 mg (Leung,
Busson & Jardin, 1968). The content of
essential amino acids per 100 g food is:
tryptophan 155 mg, lysine 1280 mg,
methionine 267 mg, phenylalanine 1125 mg,
threonine 738 mg, valine 1209 mg, leucine Macrotyloma geocarpum - 1, plant habit; 2,
1485 mg and isoleucine 871 mg (FAO, 1970). fruit; 3, seed.
Kersting's groundnut seeds contain anti- Redrawn and adapted byIskak Syamsudin
 MACROTYLOMA 101

(l-)2(-3)-seeded, constricted between the in 4-5 months, but it is also found on the
seeds, maturing on or below the soil surface. fringes of the humid tropics. Kersting's
Seeds oblong or oblong-ovoid, 5-10 mm x 4-7 groundnut tolerates poor, sandy soils, but
mm x 3—5 mm, whitish, red, brown or black, sandy loams are required for optimum yields.
sometimes striped, spotted or speckled. It is often found on slightly acid soils (pH 5).
Seedling with epigeal germination, with Propagation and planting Kersting's ground-
cotyledons falling off about 2-3 days after nut is propagated by seed. Seeds used for
emergence and 2—3 simple lanceolate primary planting are retained from the previous har-
leaves persisting until maturity. vest though some farmers may buy them
Other botanical information Macroty- locally. The 1000-seed weight is 50-150 g. In
loma comprises about 25 species, most of which West Africa Kersting's groundnut is sown from
are restricted to Africa. In Macrotyloma the beginning to the middle of the rainy
geocarpum 2 varieties have been distinguished: season. It is grown mostly in small fields or
— var. geocarpum: internodes short, petiole 8- backyards, in pure stands or intercropped with
25 cm long, terminal leaflet up to 7.5 cm x 5 yam, cowpea, cassava or other crops on mounds,
cm, pod (l-)2(-3)-seeded, seed c. 9 mm x 6 beds or ridges. When grown as a sole crop, it is
mm; only known from cultivation; often the first crop in a rotation, planted in
- var. tisserantii (Pellegr.) Maréchal & Baudet: rows 30-40 cm apart and 15cm within the row.
internodes long, petiole up to 1 cm long, Management Cultivation of Kersting's
terminal leaflet up to 3.5 cm x 2.5 cm, pod groundnut is traditional and management
l(-2)-seeded, seed c. 5 mm x 4 mm; found mainly consists of 2—3 manual weedings. The
wild in Cameroon and the Central African use ofinorganic fertilizers is not common.
Republic, and perhaps better considered a Diseases and pests In semi-arid regions
separate species (originally described as Kersting's groundnut is not subject to serious
Kerstingiella tisserantii Pellegr.), which is attacks from diseases or pests. In more humid
supported by the results of isozyme analysis regions fungal diseases (rust, mould) may
and possibly also chromosome number. occur. Stored seed is very liable to infestation
Genotypes are distinguished on the basis of by weevils (Piezotrachelus spp.) and beetles
seed colour. White types are best known and (Bruchidae).
used as food, whereas black types mainly serve Harvesting Kersting's groundnut is har-
as medicine or in cultural ceremonies, although vested when leaves start to turn yellow and
they are also used as food. wither. As the crop is harvested in the dry
Growth and development Germination of season, farmers generally dig up whole plants
Kersting's groundnut normally occurs within using a hoe and leave them in the field to dry
3—5 days after sowing. The seedling emerges for a few days, after which the pods are picked
with simple, opposite primary leaves; the first by hand, allowing easy separation of the pods.
3-foliolate leaves appear after 5-10 days. Often some seeds are left in the ground after
Flowering starts 30-65 days after sowing and harvesting and germinate with the return of
may continue until the plant dies. Self the rains, thus enabling Kersting's groundnut
pollination is the rule and 2 days after to persist in a semi-wild state.
fertilization a stalk is formed at the base of the Yield Dry seed yields of Kersting's ground-
ovary, carrying the ovary to the ground. This nut average 500 kg/ha.
mechanism is similar to that in groundnut, but Handling after harvest After harvest, the
different from that in bambara groundnut, pods of Kersting's groundnut are dried in the
where the peduncle grows to the ground. Pods sun to a moisture content of about 12% and
mature either on the soil surface or 1-2 cm stored in granaries or anywhere in the house.
under it. They reach maturity 40-60 days after They are shelled using a pestle and mortar or
flower opening. The duration of the crop cycle by beating with sticks. Usually the major part
is 90-180 days. Kersting's groundnut effect- of the production is sold. Seeds are mostly kept
ively nodulates with nitrogen-fixating bacteria in sealed containers. They are mixed with
ofthe Bradyrhizobium group. sand, pepper, ash or insecticide to ensure
Ecology Kersting's groundnut is found at longer storage.
altitudes up to 1600 m. It requires ample Genetic resources Work on the genetic
sunshine and an average temperature of 18- resources of Kersting's groundnut is relatively
34°C. It is grown successfully in semi-arid recent, and only a few small collections are
regions with an annual rainfall of 500-600 mm available. Twelve accessions collected in West
102 CEREALS AND PULSES

and Central Africa are kept in the gene bank of MACROTYLOMA UNIFLORUM (Lam.) Verde.
the International Institute of Tropical Agri-
culture, Ibadan, Nigeria. Other collections are Protologue Kew Bull. 24:322 (1970).
present in Guinea (Bureau des Ressources Family Papilionaceae (Leguminosae - Papil-
Phytogénétiques, Conakry, 8 accessions), Ghana ionoideae, Fabaceae)
(Plant Genetic Resources Centre, Bunso, 8 ac- Chromosome number 2n = 20, 22, 24
cessions), Togo (Institut de Recherches Agrono- Synonyms Dolichos uniflorus Lam. (1786),
miques Tropicales et des Cultures Vivrières, Dolichos biflorus auct. non L.
Lomé, 8 accessions) and Benin (Agricultural Vernacular names Horse gram, horse grain,
Research Centre of South Benin, Niaouli, 6 Madras gram (En).Kulthi, grain de cheval (Fr).
accessions). Little is known of the genetic Feijoeiro de lagartixa, favalinha, culita (Po).
diversity of Kersting's groundnut. In a recent Origin and geographic distribution Horse
survey of allozyme variation no diversity was gram is native to the Old World Tropics. It was
found within and among domesticated acces- probably domesticated in India, where its cul-
sions and within and among wild accessions, tivation is known since prehistoric times.
but the difference between domesticated and Nowadays horse gram is cultivated as a low-
wild accessions was much larger than that grade pulse crop in southern Asia, mainly from
found in other tropical legume species. India to Myanmar. It is also grown as a forage
Breeding No breeding programmes of Ker- and green manure in many tropical countries,
sting's groundnut are known to exist. especially in Australia and South-East Asia. In
Prospects Kersting's groundnut is a tropical Africa horse gram is recorded to occur
traditional crop of West Africa, and it has wild or naturalized in Central, East and
largely been replaced by more productive and Southern Africa. It has also been cultivated as
profitable crops, such as groundnut and a food crop and green manure in various tropi-
cowpea. The low yields, small size of the seeds, cal African countries, but it is unclear to what
amount of labour required for harvesting, and extent it is currently grown.
the liability to storage pests are the main Uses Mature whole or ground seeds of horse
causes for its decline. Kersting's groundnut has gram are eaten poached, boiled or fried.
not entirely disappeared due to its role in Sprouted seeds are widely consumed in India.
traditional ceremonies, but the fact that at In Myanmar the seeds are boiled, pounded
present it is mainly grown by elderly people with salt and fermented into a product similar
indicates that the decline will continue and to soya bean sauce.
that the prospects for this crop are bleak. Horse gram seeds are also fed to horses and
Major references Achigan Dako, Vodouhè cattle, usually after boiling. The stems, leaves
& Koukè, 2003; Amuti, 1980; Baudoin & and pod walls are used as fodder. Horse gram
Mergeai, 2001a; Burkill, 1995; Kay, 1979; is sown as a green manure or cover crop. In
Maréchal & Baudet, 1977; Mergeai, 1993; Indian traditional medicine horse gram seeds
Pasquet, Mergeai & Baudoin, 2002; Rehm, are used as a diuretic, astringent and tonic.
1989; Verdcourt, 1982. Properties The composition of whole horse
Other references Berhaut, 1976; Busson, gram seeds per 100 g edible portion is: water
1965; Dakora & Muofhe, 1997; Duke, 1981; 9.7 g, energy 1394 kJ (333 kcal), protein 22.5 g,
FAO, 1970; Gillett et al., 1971; Goli, 1997; fat 1.0 g, carbohydrate 60.5 g and fibre 4.7 g
Hepper, 1958; Hepper, 1963; ILDIS, 2002; (Leung, Busson & Jardin, 1968). The seeds
IPGRI, undated; Irvine, 1969; Leakey & Wills, contain antinutritional compounds such as
1977; Leung, Busson & Jardin, 1968; Obasi, lectins, trypsin inhibitors, phytates, tannins
1997; Purseglove, 1968; Rehm & Espig, 1991; and oxalic acid.
Schuster et al., 1998; Smartt, 1976; Tamini, Horse gram seeds have shown in-vivo anti-
1995; Verdcourt, 1980. hepatotoxic activity in rats. Lipid from the
Sources of illustration Verdcourt, 1982. seeds has shown in-vivo protective and healing
Authors E.G. Achigan Dako &S.R. Vodouhè activity against peptic ulcers in experiments
with rats. Extracts from the seeds have shown
in-vitro antilithic activity.
Botany Climbing herb with stems up to 60
cm tall, with a perennial fibrous rhizome; stem
annual, sparsely to densely covered with
spreading or appressed whitish hairs. Leaves
 MACROTYLOMA 103

ofwhich are restricted to Africa.

Within Macrotyloma uniflorum 4 varieties
have been distinguished:
- var. uniflorum: pods 6—8 mm wide; wild in
southern Asia and Namibia, widely culti-
vated in the tropics as a cover and forage
crop;
- var. stenocarpum (Brenan) Verde: pods 4 -
5.5 mm wide, shortly stiped and with more
or less smooth margins, leaflets pubescent;
occurring in Central, East and southern Af-
rica and in India, up to 1700 m altitude in
grassland, bushland and thicket, often on
sandy soils and in disturbed locations; culti-
vated in Australia and California (United
States);
- var. verrucosum Verde: pods 4-5.5 mm wide,
distinctly stiped and with obscurely to mark-
edly warted margins, leaflets pubescent; oc-
curring in East and southern Africa up to
550 m altitude in grassland and thicket;
- var. benadirianum (Chiov.) Verde: pods 4—
5.5 mm wide, shortly stiped and with slightly
warted margins, leaflets densely velvety; oc-
Macrotyloma uniflorum - 1, part of branch curring in East Africa (Somalia, Kenya) at
with inflorescence and young fruit; 2, fruits; 3, sea-level on sand dunes and thin soils on
seeds. coral rag.
Source: PROSEA Horse gram is self-pollinating. Total crop dura-
tion is usually 4-6 months. It effectively nodu-
alternate, 3-foliolate; stipules lanceolate, 4—10 lates with nitrogen-fixing bacteria of the Bra-
mm long, striate; petiole 1—7 cm long, rachis dyrhizobium group.
2.5-10 mm long; petiolules 1-2 mm long; leaf- Ecology Horse gram requires an average
lets ovate-rhombic, obovate or elliptical, 1—7(— temperature of 20—30°C and does not tolerate
8) cm x i_4(_8) C ni ; apex rounded to acute, frost. It is drought-resistant and can be grown
base rounded, lateral leaflets asymmetric, with rainfall as low as 380 mm. It is mostly
hairy to glabrescent on both surfaces. Inflores- grown in areas with less than 900 mm annual
cence an axillary (l-)2-3(-5)-flowered fascicle; rainfall. In higher rainfall areas it is grown on
bracts up to 3 mm long. Flowers bisexual, residual moisture in the dry season, e.g. after a
papilionaceous; pedicel 1—7 mm long; calyx rice crop. Most horse gram cultivars are short-
pubescent, tube 2 mm long, lobes triangular- day plants.
lanceolate, 3—8 mm long, long-acuminate, up- Horse gram grows on a wide range of soils with
per pair entirely fused; corolla with cream, pH 5-7.5, including poor soils. It does not tol-
yellow or greenish yellow standard, often with erate waterlogging.
a small purple blotch inside, obovate-oblong, 6— Management Horse gram is propagated by
12 mm x 4-7 mm, wings and keel greenish seed. The 1000-seed weight is 15-50 g. The
yellow, 5-10 mm long; stamens 10, 9 fused and crop is sown broadcast or in rows 20-90 cm
1 free; ovary superior, stiped, 1-celled. Fruit a apart, at a seed rate of 20-45 kg/ha. The sow-
linear-oblong pod 3—8cm x 4-8 mm, upcurved ing depth is 1-2.5 cm. In India horse gram is
towards apex, acuminate, densely hairy when usually sown as a sole crop, but sometimes it is
young, later more sparsely so, margins gla- intercropped, e.g. with finger millet, maize,
brous, smooth or warty, dehiscent, 5-10- chickpea, groundnut or castor. The main dis-
seeded. Seeds trapezoidal, oblong or rounded- eases on horse gram in India are horse gram
reniform, 3—8mm x 3-5 mm, pale to dark red- yellow mosaic virus (HgYMV), anthracnose
dish brown, speckled or mottled with black and (Colletotrichum lindemuthianum), leaf spot
orange-brown or all black. (Cercospora dolichi, synonym: Mycosphaerella
Macrotyloma comprises about 25 species, most cruenta), rust (Uromyces appendiculatum), root
104 CEREALS AND PULSES

rot {Pellicularia filamentosa, synonym: Thana- ICRISAT, undated; Jayaraj et al., 2000; Laskar
tephorus cucumeris) and dry root rot (Macro- et al., 1998; Leung, Busson & Jardin, 1968;
phomina phaseolina). Recorded pests include Mackinder et a l , 2001; Omokanye, 1996;
the gram caterpillar (Azazia rubricans, syno- Purseglove, 1968;Sudha et a l , 1995.
nym: Anticarsia irrorata) and the green pod- Sources of illustration Jansen, 1989c.
boring caterpillar (Etiella zinckenella). Horse Authors M. Brink
gram grown for the seeds is harvested when Based on PROSEA 1: Puises.
the pods begin to shrivel and the leaves begin
to dry and fall off. The plants are cut or up-
rooted, stacked, and dried in the sun for a MUCUNAGIGANTEA (Willd.) DC.
week, after which they are threshed using
sticks, stone rollers or oxen. Seed yields are Protologue Prodr. 2:405 (1825).
usually low (150—350 kg/ha in India) but much Family Papilionaceae (Leguminosae - Papil-
higher yields have been obtained with im- ionoideae, Fabaceae)
proved cultivars (900 kg/ha in India, 1100- Synonyms Mucuna quadrialata Baker (1871),
2200 kg/ha in Australia). In experiments in Mucuna longipedicellata Hauman (1955).
Nigeria in the 1990s seed yields of 700-1000 Vernacular names Sea bean, burny bean
kg/ha were obtained. When grown for fodder, (En). Liane cadoque, liane caiman, mort aux
horse gram can be harvested about 6 weeks rats (Fr).Mtera (Sw).
after sowing. Forage yields are 4-15 t dry mat- Origin and geographic distribution Mucu-
ter per ha. na gigantea is distributed in tropical Asia, Ja-
Genetic resources and breeding Germ- pan, Australia, Pacific Islands and Africa. In
plasm collections of horse gram are held in tropical Africa it is found from DR Congo to
Australia (Australian Tropical Crops & For- Kenya, Tanzania and Mozambique, also in
ages Genetic Resources Centre, Biloela, Madagascar and other Indian Ocean islands.
Queensland, 38 accessions) and Kenya (Na- Uses The seeds of Mucuna gigantea are con-
tional Genebank of Kenya, Crop Plant Genetic sidered edible in Kenya. In India boiled seeds
Resources Centre, KARI, Kikuyu, 21 acces- are sometimes eaten as a pulse, e.g. in the An-
sions). Cultivated horse gram is usually a mix- daman Islands. Aboriginals in Australia used
ture of several landraces with different seed to heat the seeds on hot stones or sand, remove
colours and periods of maturity. the peel, and grind them to flour, which was
Breeding activities are focused on yield poten- then mixed with water, wrapped in leaves and
tial, resistance to diseases and insensitivity to baked.
daylength. Improved cultivars have been de- Root decoctions of Mucuna gigantea are taken
veloped and released in India; a popular forage to treat gonorrhoea and schistosomiasis. In
and fodder-grain cultivar with indéhiscent pods India the bark is applied externally to treat
in Australia is 'Leichhardt'. In-vitro regenera- rheumatic complaints. Powdered seed is used
tion has been achieved by direct organogenesis as a purgative in Hawaii. The irritant hairs on
using shoot tip and cotyledonary node expiants, the outside of the pods are mentioned as being
and by somatic embryogenesis through cell used in criminal poisoning in Malaysia. In
suspension culture of callus induced on leaf Vietnam they are mixed with food to get rid of
expiants. rats.
Prospects It is unclear to what extent horse Properties Mucuna gigantea seeds contain
gram is presently grown in tropical Africa, and 1.7-2% L-dopa (levodopa; L-3,4-dihydroxy-
how frequently it is consumed as a pulse or phenylalanine), an amino acid which stimu-
used for other purposes. It seems an interest- lates the formation of the neurotransmitter
ing crop for dry areas in tropical Africa, but dopamine in the brain. Dopamine lessens tremor
more information is needed on the nutritional experienced in Parkinson's disease. However,
characteristics of the seed and on the accept- opinions differ on the side effects and efficacy
ability ofits taste for the African consumer. in the long run of L-dopa. Because of the pres-
Major references Gillett et al., 1971; Jan- ence of toxic compounds in the plant, it seems
sen, 1989c; Kay, 1979; Varisai Mohamed et al., advisable to eat the seed only after prolonged
2004;Verdcourt, 1982. soaking and boiling.
Other references Garimella, Jolly & Nara- Botany Large liana up to 30(-80) m long;
yanan, 2001; Hanelt & Institute of Plant Ge- stems initially covered with orange-brown bris-
netics and Crop Plant Research (Editors), 2001; tle hairs, glabrescent. Leaves alternate, 3-
 ORYZA 105

foliolate; stipules 3—5 mm x 1 mm, deciduous; from the wild. The presence of the intensely
petiole 4-15 cm long, rachis 1.5-3.5 cm long; irritant bristle hairs makes handling difficult.
stipels needle-shaped, 2-3 mm long, persistent; Genetic resources and breeding One ac-
petiolules c. 5 mm long; leaflets ovate or ellipti- cession from Kenya is kept in the National
cal, 4-15 cm x 2-9 cm, the lateral ones oblique, Genebank of Kenya, Crop Plant Genetic Re-
acuminate and markedly apiculate at apex, sources Centre, KARI, Kikuyu. In view of its
rounded at base, thinly appressed hairy when widespread distribution Mucuna gigantea is
young, soon glabrescent. Inflorescence an axil- not threatened by genetic erosion.
lary, pendulous false umbel 10-35 cm long, Prospects Because of the toxic compounds
with flowers on short lateral branchlets 5-10 in the seed necessitating long cooking and the
mm long; peduncle 4-22(-30) cm long. Flowers presence of irritant hairs on the pods it is
bisexual, papilionaceous; pedicel 1-2.5 cm long; unlikely that Mucuna gigantea will become a
calyx cup-shaped, 10-13 mm long, 2-lipped, more important food crop.
covered with fine grey hairs and long deciduous Major references Beentje, 1994; Dahal &
orange-brown bristle hairs, tube 7-11 mm long, van Valkenburg, 2003; Dick, 1994; du Puy et
lobes 2—3 mm long, the upper lip somewhat al., 2002; Gillett et al., 1971.
emarginate; corolla pale creamy-green, white Other references Eilittä et al., 2002;
or pale lilac, standard (2-)2.5-3.5 cm x (1.5-)2- Friedmann, 1994; ILDIS, 2002; Mackinder et
2.5 cm, round, with sparse orange-brown bris- al., 2001; Neuwinger, 2000; Polhill, 1990; Ra-
tle hairs, wings and keel (3-)3.5-4.5 cm long; jaram & Janardhanan, 1991; Wilmot-Dear,
stamens 10, 9 united and 1 free; ovary supe- 1984;Wilmot-Dear, 1991; Wilmot-Dear, 1992.
rior, 1-celled, style long, filiform, stigma small Authors M. Brink
and terminal. Fruit a stiped pod, oblong or
oblong-elliptical, 7-15 cm x 3-5.5(-6.5) cm x 1-
2 cm, each margin with 2 wings, densely cov- ORYZA BARTHII A.Chev.
ered with orange-brown bristle hairs at first,
becoming glabrous at maturity, l-4(-6)-seeded. Protologue Bull. Mus. natn. Hist, nat.,
Seeds 2.5-3 cm x 2-3 cm x 1-1.5 cm, discoid, Paris 16: 405 (1911).
dark brown or densely mottled with rust brown Family Poaceae (Gramineae)
or black, smooth, hilum extending around the Chromosome number 2n — 24
seed-margin for c.three-quarters of the circum- Synonyms Oryza breviligulata A.Chev. &
ference. Seedling with hypogeal germination; Roehr. (1914), Oryza stapfii Roshev. (1931).
first leaves scale-like or simple. Vernacular names Wild rice, self-sown rice,
Mucuna belongs to the tribe Phaseoleae and Mandinka rice (En). Riz sauvage annuel, riz de
comprises about 100 species distributed pan- marais, riz sauvage (Fr).
tropically. In tropical Africa about 10 species Origin and geographic distribution Oryza
are present. Several subspecies have been dis- barthii is distributed in tropical Africa from
tinguished within Mucuna gigantea, with Mauritania east to Ethiopia and south to Bot-
subsp. quadrialata (Baker) Verde, in Africa. swana and Zimbabwe.
However, Mucuna gigantea is very variable Uses The grains of Oryza barthii are some-
throughout its range and it seems not possible times collected if enough plants are available,
to retain subsp. quadrialata. and they serve as a famine food. They are
Initial growth of Mucuna gigantea is rapid: sometimes sold in markets. However, Oryza
seedlings may attain a height of more than 1m barthii is regarded mostly as a weed. Before
in 3 weeks. In Madagascar it flowers during flowering the plant provides good grazing for
the dry season. The flowers are much visited by livestock; after flowering the awns may cause
humming-birds. The seeds are dispersed by sea injury to the mouth.
currents. All green plant parts, including the Properties The grain of Oryza barthii has a
flowers, become black when bruised or dried. good flavour.
Ecology Mucuna gigantea is essentially a Botany Annual grass up to 150 cm tall,
littoral species found around the Indian Ocean, growing in tufts; stem (culm) erect or genicu-
but in tropical Africa it also occurs inland. It is lately ascending, with roots from the lower
found in coastal scrub, on riverbanks, and near nodes, spongy, striate, glabrous. Leaves alter-
water in woodland and forest edges, up to 1800 nate, simple and entire; leaf sheath striate,
m altitude. smooth; ligule 2—6(-9) mm long, truncate or
Management Mucuna gigantea is collected rounded; blade linear, 15—45 cm x 0.5-1.5 cm,
106 CEREALS AND PULSES

with acute apex, intense green, glabrous, Oryza barthii is probably as a source of resis-
smooth on the lower surface, slightly rough on tance to various diseases affecting Oryza sa-
the upper surface. Inflorescence a terminal tiva.
panicle 20-35 cm x 3-7.5 cm, rather dense, Major references Burkill, 1994; Hanelt &
erect or more rarely somewhat nodding, with Institute of Plant Genetics and Crop Plant Re-
erect or obliquely ascending branches. Spikelet search (Editors), 2001;Launert, 1971; National
oblong to narrowly oblong, 7-11 mm long (awn Research Council, 1996; Smith &Dilday, 2003.
excluded), deciduous, pale green to straw- Other references Abo, Sy &Alegbejo, 1998
coloured, 3-flowered but 2 lowest florets re- Akromah, 1987; Clayton, 1970; Clayton, 1972
duced to sterile lemmas 2.5-4.5 mm long; Engels, Hawkes & Worede (Editors), 1991
glumes reduced to a 2-lobed rim; lemma of fer- Gibbs Russell et al., 1990; Kaushal & Ravi,
tile floret slightly shorter than spikelet, boat- 1998; Lu, 1999; Phillips, 1995; Vaughan &
shaped, leathery, hairy, with 2 longitudinal Chang, 1992.
lateral grooves, with pink to purplish stiff awn Authors M. Brink
(4-)8-16(-19) cm long; palea about as long as
lemma but much narrower, with the apex
drawn out in a short blunt point; lodicules 2; ORYZA GLABERRIMA Steud.
stamens 6; ovary superior, with 2 plumose stig-
mas. Fruit a caryopsis (grain). Protologue Syn. pi. glumac. 1(1):3 (1853).
Oryza comprises about 20 wild species distrib- Family Poaceae (Gramineae)
uted throughout the tropics and subtropics, Chromosome number In - 24
and 2 cultivated species, Oryza sativa L. and Vernacular names African rice, red rice
Oryza glaberrima Steud. Oryza barthii is clas- (En). Riz africain, riz de Casamance (Fr).
sified in ser. Sativae, together with Oryza sa- Origin and geographic distribution Oryza
tiva, Oryza glaberrima and Oryza longistami- glaberrima was derived from the wild annual
nata A.Chev. & Roehr. Oryza barthii A.Chev. (synonym: Oryza bre-
Oryza barthii is predominantly inbreeding, viligulata A.Chev. & Roehr.). Oryza barthii
with an outcrossing rate of 5-20%. probably grew abundantly in lakes that existed
Ecology Oryza barthii grows in shallow in what is now the Sahara from 8000-4000 BC,
water in ponds and marshes, and as a weed in and it was harvested as a wild cereal. When
rice fields, up to 1500 m altitude. It may form the climate became drier, Oryza glaberrima,
pure stands, but is usually found scattered which had gradually developed from Oryza
with other aquatic grasses. It may become a barthii (probably around 1500 BC or later),
noxious weed and may act as a reservoir for was grown as a rainfed homegarden crop in
important rice diseases and pests. Oryza oases. When the population took refuge in the
barthii is a short-day plant. interior delta of the Niger river (around 1500
Management Oryza barthii is not normally BC) and became much larger, Oryza glaber-
cultivated, but the grain is collected from the rima was transformed into the current floating
wild. The grain shatters very easily, and the
panicles are usually collected before they are
mature. If ripe, the panicles are harvested over
a basket or calabash to collect falling grain.
Genetic resources and breeding Oryza
barthii has a relatively narrow genetic varia-
tion. It is considered a source of resistance to
various diseases affecting Oryza sativa, includ-
ing bacterial leaf blight (Xanthomonas oryzae
pv. oryzae), rice yellow mottle virus (RYMV)
and sheath blight (Thanatephorus cucumeris,
anamorph: Rhizoctonia solani).
Prospects Although Oryza barthii may
serve as a famine food during times of short-
age, it is probably more often considered a
weed of Oryza sativa than a valuable food
plant, and there seems to be no reason to jus-
tify its promotion. The greatest potential of Oryza glaberrima -planted
 ORYZA 107

rice crop. farming, African rice has been replaced by

African rice is now grown in a zone extending Asian rice, which is more productive, shatters
from the delta of the River Senegal in the west less easily and has a softer grain that is easier
to Lake Chad in the east. To the south-east its to mill. Small-scale farmers in West Africa,
range is bordered by the river basins of the however, often still prefer to grow African rice
Benue, Logone and Chari, but it has also been for its taste and culinary properties, its ability
recorded from the islands of Pemba and Zanzi- to withstand flooding, and its resistance to
bar (Tanzania). The areas of most intensive several diseases and pests.
cultivation ofAfrican rice are the floodplains of Description Annual grass up to 120 cm tall
northern Nigeria, the inland delta of the Niger (up to 5 m in some floating types), often tufted;
river in Mali, parts of Sierra Leone and the dryland types with simple culm often rooting at
hills on the Ghana-Togo border. African rice lower nodes, floating types often branching and
was probably introduced into the New World rooting at upper nodes too. Leaves alternate,
during the slave trade era, and it is still occa- simple; sheath terete, up to 25 cm long, with
sionally cultivated there, e.g. in Brazil, Guy- transverse veinlets; ligule c. 4 mm long, trun-
ana, El Salvador and Panama. cate, membranous; blade linear, flat, 20-25(-
Uses In parts of West Africa the grain of 30) cm x 6-9 mm, sagittate at base, rugose
African rice is a staple food, highly appreciated beneath. Inflorescence a terminal, ellipsoid,
for its taste and culinary qualities. It is also stiff and compact panicle up to 25 cm long,
used in traditional and ritual ceremonies, e.g. with ascendent racemose branches. Spikelets
in the Casamance region of southern Senegal. ellipsoid, c. 9 mm x 4 mm, more or less persis-
The finer parts of the bran and broken grains tent, 3-flowered but 2 lowest florets reduced to
are given as feed to chicken and other live- sterile lemmas separated from the lemma of
stock. In the Central African Republic the root the fertile, bisexual upper floret by a stipe;
is eaten raw to treat diarrhoea. glumes absent or strongly rudimentary; lemma
Production and international trade In sta- hispidulous, 5-veined, usually without apical
tistics on rice production in West Africa no awn; palea 3-veined; lodicules 2; stamens 6;
distinction is made between African rice and ovary superior, with 2 plumose stigmas. Fruit
Asian rice (Oryza sativa L.). It is estimated
that African rice is grown in less than 20% of
the total area allocated to rice in West Africa.
As a traditional food grain it is not traded in-
ternationally, but only within the region of
production.
Properties The composition of whole Afri-
can rice grain (hulled) per 100 g edible portion
is: water 11.3 g, energy 1524 kJ (364 kcal),
protein 7.4 g, fat 2.2 g, carbohydrate 77.7 g,
fibre 0.4 g, Ca 38 mg, P 294 mg, Fe 2.8 mg,
thiamin 0.34 mg and niacin 6.5 mg. Milled
African rice contains per 100 g fresh edible
portion: water 11.4 g, energy 1532 kJ (366
kcal), protein 6.3 g, fat 0.3 g, carbohydrate 81.6
g, fibre 0.2 g, Ca 22 mg, P 98 mg, Fe 1.7 mg,
thiamin 0.06 mg, niacin 2.0 mg and tryptophan
110 mg (Leung, Busson & Jardin, 1968). Afri-
can rice is superior to Asian rice in its content
of the important vitamin thiamin and in iron.
The degree of gelatinization depends on the
amylose content, which ranges from 14-30%,
and influences consistency of the rice in cook-
ing and thus consumer choice. Most cultivars of
African rice have red-skinned grain and some
are strongly scented. Oryza glaberrima - 1, plant habit; 2, inflores-
Adulterations and substitutes In most cence;3, spikelet.
regions of West Africa, at least in commercial Redrawn and adapted by W. Wessel-Brand
108 CEREALS AND PULSES

a laterally compressed caryopsis (grain) up to 9 propagated by seed. The weight of 1000 seeds
mm x 3 mm, often reddish, tightly enveloped is 20—27 g. Seed dormancy disappears a few
by lemma and palea. months after maturity; for experimental pur-
Other botanical information Oryza com- poses, dormancy can be broken by removing
prises about 20 wild species distributed the lemma and palea and about one-third of
throughout the tropics and subtropics, and 2 the albumen, allowing germination in 2-3
cultivated species, Oryza sativa and Oryza gla- days. Before sowing the soil may be prepared
berrima. Several classifications of Oryza have with a hoe or, as in Senegal, Gambia and
been made. Most recently the genus has been Guinea, with a long-handled spade, but soil
divided into 3 sections: sect. Pädia, sect. preparation is rarely practised. Seed is mostly
Brachyantha and sect. Oryza. Section Oryza is broadcast and transplanting is rarely prac-
subdivided into 3 series: ser. Latifoliae, ser. tised. For floating rice, seed is densely sown in
Australiensis and ser. Sativae. Oryza glaber- soil that has been recently weeded and that
rima, its direct ancestor Oryza barthii A.Chev. may or may not have been ploughed or hoed.
and the rhizomatous perennial Oryza longi- Cultivars are selected according to expected
staminata A.Chev. & Roehr. are classified in flood duration and generally have a growing
ser. Sativae, together with Oryza sativa. Mor- period of 4-6 months.
phologically, Oryza glaberrima can be distin- In West Africa from Senegal to northern Cam-
guished from Oryza sativa by its shorter ligule eroon, where rainfall generally exceeds 1000
and less-branched panicle. mm/year, African rice is mostly planted as an
Growth and development African rice upland crop, depending solely on rain and sur-
seedlings normally emerge in 4—5 days after face run-off. In some regions short-duration
sowing. The vegetative phase of African rice cultivars are grown that are adapted to annual
consists of a juvenile phase of about 3 weeks rainfall amounts as low as 700 mm. In Senegal
followed by a tillering phase of 3—4 weeks. and Gambia the crop is sown in moist loca-
Vegetative growth is rapid. Tillering, high leaf tions, often under palm trees, after simple soil
area index and high specific leaf area contrib- cultivation. This is locally called 'riz de pla-
ute to its high competitiveness against weeds. teau'. 'Riz de montagne' is grown throughout
However, culms tend to be weak and brittle, the forest zone covering western Côte d'Ivoire,
making African rice prone to lodging. African Liberia, the Fouta Djallon and eastern Guinea
rice is self-fertilizing. The duration of the crop mountains. It is grown in shifting cultivation,
varies from 3-6 months depending on cultivar often following logging, even on steep slopes.
and type of culture. Some cultivars selected for The undergrowth is cut and at the end of the
rainfed conditions are of very short duration, dry season fields are burned. Sowing is mostly
shorter than cultivars of Oryza sativa. Culti- carried out without any soil cultivation. Rice is
vars for deep water conditions tolerate flooding grown in pure stands or intercropped with
up to 2.5 m deep and culms may grow up to 5 other crops, e.g. maize. After 2-3 years, the
m long. Some shattering of seed occurs in many field is used to grow cash crops such as cacao or
cultivars. coffee, or left fallow. Farmers return after 10—
Ecology African rice grows well above 30°C, 20 years, or later, depending on the recovery of
but above 35°C spikelet fertility is noticeably the vegetation and the soil. In such fields, cul-
reduced. Temperatures below 25°C reduce tivars of shortest duration are grown and Afri-
growth and yield; temperatures below 20°C do can rice is only rarely replaced by Asian rice,
so markedly. African rice is grown from sea- e.g. in the forest zone of Guinea and western
level to 1700 m altitude. It is generally a short- Côte d'Ivoire.
day plant, but photosensitivity varies between Irrigated rice systems depend more on river
cultivars from day-neutral to strongly sensi- water than on rainfall and are found in areas
tive. with a much drier climate; the degree of control
African rice is grown on a wide range of soils. of irrigation is variable. Floodplain rice on hy-
Although preferring fertile alluvial soils, it dromorphic soils is found in Guinea, Côte
tolerates low soil fertility. Some cultivars can d'Ivoire, Mali, Burkina Faso and Nigeria.
produce higher yields than Asian rice on alka- Floating rice cultivars are very common in the
line and on phosphorus-deficient soils. They interior delta of the Niger river in Mali, and is
are also more tolerant to iron-toxicity. Floating also planted in Senegal, Gambia, Niger and
rice is planted on loam or clay soils. Nigeria. It grows sometimes very rapidly in
Propagation and planting African rice is length as the flood water rises, tolerating sub-
 ORYZA 109

mersion for several days. Cultivars grown have pollinates with Oryza glaberrima; the resulting
a crop duration of 4-5 months. red grains shatter more easily and have to be
Along the rivers in northern Senegal and in milled more tightly, resulting in more weight
Mali, in the northern part of the interior delta loss and higher costs. Under conditions of deep
of the Niger river south-west of Timbuktu, in a flooding, perennial wild rice (Oryza longista-
zone stretching from Dire and Goundam to the minata) is cut below the surface ofthe water in
series of lakes Faguibine, Gouber and Ka- order to kill it.
mango, rice is grown on floodplains after floods Harvesting The harvesting season for Afri-
have receded. In this cropping system, rice is can rice is October-December. Upland rice is
sown in moist soil and the crop development harvested first. Panicles are bundled and
relies on ground water ('riz de décrue'). Weeds stacked in elevated granaries under which a
are few. Both Oryza glaberrima and Oryza smoking fire is maintained to keep away stor-
sativa are grown and have a duration of 4-5 age insects. After manual or mechanical
months. Along the Atlantic coast, e.g. in Sierra threshing, grain can also be stored in bulk in
Leone, African rice is grown in mangrove bags. Floating rice is harvested in several
swamps. rounds mostly from canoes, which leads to con-
Management Weeding of African rice in siderable losses.
non-flooded areas is manual and often late. In Yield Yields of African rice obtained under
some regions, such as the Basse Casamance, traditional conditions rarely average more than
weed control is combined with land prepara- 1 t/ha. In experiments with deep water rice
tion: a first light irrigation favours the germi- cultivars carried out in Gao and Timbuktu
nation of weeds, which can subsequently be (Mali) from 1984-1987, yields of 1-4 t/ha were
eradicated. Mechanization and fertilizer appli- obtained.
cation are rarely practised. In floodplain and Handling after harvest The produce of
wet rice cultivation neither crop rotation nor African rice, whether stored before or after
fallow is practised, contrary to the practice for threshing, should be protected against pests,
upland rice. mainly insects and rodents. The paddy should
Diseases and pests The most important be dried well to reduce the moisture content to
and widespread disease of African rice is rice a maximum of 14%to achieve good storage and
blast (Pyricularia grisea; synonyms: Mag- a high milling yield. The grain ofAfrican rice is
naporthe grisea, Pyricularia oryzae). Rice yel- more brittle than that of Oryza sativa, making
low mosaic virus (RYMV) and soil parasites it more difficult to mill.
(nematodes) often cause large losses. There are Genetic resources IRD (Institut de Recher-
few control measures, but some cultivars are che pour le Développement, formerly ORSTOM)
resistant to such pathogens. In floodplain and and CIRAD (Centre de Coopération Interna-
wet rice systems the main problems are rizo- tionale en Recherche Agronomique pour le
phagous fish (Distichodus, Tilapia), while birds Développement) collected cultivated and re-
cause serious damage in all rice cropping sys- lated wild types of rice (both African and intro-
tems. Children armed with pebbles and slings duced) throughout their area of distribution.
offer some protection. Rodents, buffaloes, ele- Between 1974 and 1983, over 3700 samples
phants and hippopotamuses can all cause seri- were collected in Africa and Madagascar, of
ous damage. African rice gall midge (Orseolia which 20% are Oryza glaberrima and 12% re-
oryziphora), crickets and grasshoppers are also lated wild species. These collections are kept in
important pests, as are stem-borers that de- cold storage (4°C, 20% humidity) for medium-
stroy the apex of the plants and so prevent the term conservation and partly frozen at -20°C
formation of inflorescences. for long-term storage at IRD in Montpellier
Annual wild rice {Oryza barthii) is very com- (France). The collection is duplicated at CIRAD
mon in wet rice fields. It can be recognized by in France and at the International Rice Re-
its red awns but it is then too late to remove it. search Institute (IRRI), the Philippines. The
It is characterized by very strong shattering International Institute of Tropical Agriculture
and, as it often ripens before the cultivated (UTA), Ibadan, Nigeria keeps almost 2800 ac-
rice, it multiplies and spreads throughout the cessions, and the Africa Rice Center (WARDA),
rice field. It is sometimes harvested with the Bouaké, Côte d'Ivoire, almost 1900 accessions.
rice crop. If the seed is not cleaned carefully, Collections of Oryza glaberrima germplasm are
the field will be infested with wild rice within a also kept at the Bangladesh Rice Research
few years. Annual wild rice readily cross- Institute, Dhaka, Bangladesh (200 accessions)
110 CEREALS AND PULSES

and theUSDA-ARS National Small Grain Col- Cross-breeding of Oryza glaberrima and Oryza
lection, Aberdeen, Idaho, United States (174 sativa should continue to include programmes
accessions). African rice shows orthodox seed aiming at the transfer of genome fragments.
storage behaviour. Currently noin-situ conser- Such breeding programmes should be carried
vation programmes of rice of African origin out in association with a programme of in-situ
exist butthey would bedesirable. conservation of genetic resources of wild and
Breeding While the genetic variation in cultivated rice of African origin. For specific
Oryza glaberrima is small in comparison with objectives certain regions should be identified,
that of Oryza sativa, types with important e.g. Guinea for its diversity of rice cropping
characteristics have been identified: resistance systems, the regions of southern Chad/north-
to RYMV, rice blast (Pyricularia grisea), Afri- ern Cameroon andtheinterior delta ofthe Niger
can rice hispa (Trichispa sericea), the African river in Mali forthecontacts between wildand
rice gall midge (Orseolia oryziphora), and to cultivated types, andthe valley ofthe Ferlo in
several stem-borers and nematodes, including Senegal to study spontaneous populations of
Heterodera sacchari, Meloidogyne graminicola the annual Oryza barthii away from all rice
and Meloidogyne incognita. African rice shows cultivation. Improvement of African rice culti-
resistance tosalinity, drought andiron toxicity vation should aim at decreased lodging, in-
and it competes well with weeds. Various culti- creased yield, less seed scattering and de-
vars have shown partial resistance to and tol- creased brittleness ofthe grain.
erance of parasitic plants of the genus Striga. Major references Bezançon, 1994; Bren-
In general, hybrids between Oryza glaberrima ière, 1983;Jones et al., 1994;Jones et al., 1997;
and Oryza sativa are highly sterile in the Fi Linares, 2002; Lorieux, Ndjiondjop & Ghes-
and early generations. However, in a hybridi- quière, 2000; Lu, 1999; National Research
zation programme initiated in 1992,WARDA Council, 1996; Séré &Sy, 1997; Sumi & Kata-
succeeded in crossing thetwo species into sta- yama,1994.
ble and fertile progenies through backcrossing Other references Aluko et al., 2004; Bet-
and doubled haploid breeding. Interspecific tencourt & Konopka, 1990; Bezançon, 1995;
progenies, which are called 'New Rice for Af- Bouharmont, Olivier & Dumont de Chassart,
rica' (NERICA), are nowbeing grown by farm- 1985; Buddenhagen &Persley (Editors),1978;
ers in Africa. They are more productive than Burkill, 1994; Catling, 1992; Chang, 1995;
Oryza glaberrima, but retain favourable char- Guei, Adam &Traoré, 2002; Hanelt & Institute
acteristics such as competitiveness against of Plant Genetics and Crop Plant Research
weeds, resistance to diseases and pests, toler- (Editors), 2001; Heuer et a l , 2003; IPGRI, un-
ance to poor soils, andhigh grain quality.Few dated; Johnson et al., 1997;Leung, Busson &
genetic improvement programmes of Oryza Jardin, 1968; Nwilene et a l , 2002; Plowright et
glaberrima itself have been undertaken. a l , 1999;Purseglove, 1972;Rehm &Espig, 1991;
Extensive genetic linkage maps have been Ukwungwu, Williams & Okhidievbie, 1998;
made for rice, andIRD andWARDA are work- Watanabe et al., 2002.
ing together in a programme to systematically Sources of illustration National Research
integrate thegenome ofOryza glaberrrima into Council, 1996;Roshevitz, 1931.
that of Oryza sativa. Theobjective is to follow Authors G.Bezançon &S.Diallo
the introgression ofsmall genome fragments of
Oryza glaberrima into the genetic base of
Oryza sativa using molecular markers. ORYZA LONGISTAMINATA A.Chev. & Roehr.
Prospects For over 30 years it has been
predicted that African rice would disappear Protologue Compt. Rend. Acad. Sei., sér. 2,
under the pressure of widespread introduction Mec. Phys. Chim. Sei. Univers. Sei. Terre.159:
of improved cultivars of Oryza sativa, but this 561 (1914).
has not happened, although in Burkina Faso, Family Poaceae (Gramineae)
for example, a strong decline ofAfrican ricehas Chromosome number 2n= 24
been observed. The explanation for the resil- Synonyms Oryza barthii auct. nonA.Chev.
ience ofAfrican rice isthat itis highly appreci- Vernacular names Wild rice, redrice(En).
ated bythepeople ofWest Africa, who continue Riz sauvage vivace, rizvivace (Fr).
to grow African rice for its taste and culinary Origin and geographic distribution Oryza
properties, andthat itishighly adapted to par- longistaminata is distributed throughout tropi-
ticular growing conditions, e.g.asfloating rice. cal Africa (including Madagascar) and is also
 ORYZA i l l

found inSouth Africa. such as bacterial leaf blight (Xanthomonas

Uses Thegrains ofOryza longistaminata are oryzae pv. oryzae).
sometimes eaten and sold on local markets. Management Oryza longistaminata is
They serve as famine food, e.g. in Sudan and mostly collected from the wild and only occa-
Ethiopia. Dense stands provide good grazing sionally cultivated. The grains shatter easily,
for cattle. Thestraw isused for thatching. and it is common practice to harvest panicles
Botany Robust perennial grass up to 2.5m just before maturity or to shake ripe panicles
tall, with long, creeping, branched rhizomes; over a basket or calabash. The long, scabrid
stem (culm) up to 2.5cmor more in diameter, awns form a disincentive totouch thepanicle.
erect or ascending, with aerial roots from the Genetic resources and breeding As seed
lower nodes, glabrous. Leaves alternate, simple production of Oryza longistaminata is very
and entire; leaf sheath spongy, pale green to poor, in situ conservation is recommended.
brownish, smooth, glabrous; ligule (l-)1.5-5.5 Oryza longistaminata is considered a sourceof
cm long, acute, often split down the middle; resistance genes to various diseases affecting
blade linear, 10-45(-75) cmx0.5-1.5(^2.5)cm, cultivated Oryza sativa. Resistance to bacterial
acuminate, bright to dark green, glabrous, leaf blight has successfully been transferred.
smooth or slightly rough onthe lower surface, Oryza longistaminata is a host plant of rice
slightly rough on the upper surface. Inflores- yellow mottle virus (RYMV), an important dis-
cence a terminal panicle 16-40 cmx 2.5-8 cm, ease of Oryza sativa in Africa, but in general
dense, erect orslightly drooping, with obliquely Oryza longistaminata is more tolerant of it,
ascending to almost erect branches. Spikelet and some accessions are immune. Oryza longi-
asymmetrically elliptical-oblong, 7—12(-15) mm staminata is a potential source ofgenes forthe
long (awn excluded), deciduous, pale green to development ofperennial types ofOryza sativa,
brownish, 3-flowered but 2 lowest florets re- which would provide a permanent ground cover
duced to sterile lemmas (2-)2.5-4(-4.5) mm and reduce erosion.
long; glumes reduced to a membranous rim; Prospects Oryza longistaminata serves asa
lemma of fertile floret slightly shorter than famine food during times of shortage, but is
spikelet, boat-shaped, leathery, hairy, with also a noxious weed ofOryza sativa. The great-
pink or purplish, rather slender awn(2.5—)4— est potential of Oryza longistaminata is proba-
7.5(—8) cm long; palea slightly shorter than bly in Oryza sativa breeding as a source of
lemma and much narrower, acute or tapering genes conferring disease resistance and peren-
into a point; lodicules 2;stamens 6;ovary supe- nial habit.
rior, with 2plumose blackish stigmas. Fruit an Major references Burkill, 1994; Engels,
oblong caryopsis (grain) 7.5-8.5 mm long, gla- Hawkes & Worede (Editors), 1991;Hanelt &
brous, pale brown, glossy. Institute ofPlant Genetics and Crop PlantRe-
Oryza comprises about 20 wild species distrib- search (Editors), 2001;Launert, 1971; National
uted throughout the tropics and subtropics, Research Council, 1996.
and 2 cultivated species, Oryza sauva L.and Other references Abo, Sy&Alegbejo,1998;
Oryza glaberrima Steud. Oryza longistaminata Akromah, 1987;Clayton, 1970;Clayton, 1972;
is classified in ser. Sativae, together with Gibbs Russell et a l , 1990;Lu, 1999;Phillips,
Oryza sativa, Oryza glaberrima and Oryza 1995; Sacks, Roxas &Sta Cruz, 2003; Smith &
barthii A.Chev. Dilday, 2003; Vaughan &Chang, 1992.
Oryza longistaminata can be distinguished Authors M. Brink
from other wild Oryza spp. by its very long,
pointed ligule. Oryza longistaminata is partly
self-incompatible and allogamous. Often only ORYZA PUNCTATA Kotschy ex Steud.
few seeds are set and natural reproduction is
mainly byits rhizomes. Protologue Syn. pi. glumac. 1(1):3(1853).
Ecology Oryza longistaminata is found in Family Poaceae (Gramineae)
shallow or deep water in pans, pools, swamps, Chromosome number 2n=24,48
flood plains andriverbanks, upto 1800 m alti- Vernacular names Redrice, wadi rice (En).
tude. It often occurs in pure stands. Oryza Mchetez (Sw).
longistaminata is a noxious weed in wet-rice Origin and geographic distribution Oryza
cultivation; it suppresses cultivated rice and punctata is distributed in tropical Africa from
forms hybrids with it.It mayalso act as a res- Côte d'Ivoire to Sudan and southwards to An-
ervoir for important rice diseases and pests, gola, Zimbabwe, Mozambique and Madagascar.
112 CEREALS AND PULSES

It also occurs in South Africa and Thailand. Management Oryza punctata is collected
Uses The husked grains of Oryza punctata from the wild. The 1000-seed weight is about
are sometimes eaten as a famine food in Sudan 25 g. Husking requires vigorous pounding, re-
and Kenya. In Sudan they are consumed after sulting in the grain being seldom whole when
boiling with milk or water. eaten.
Properties Per 100 g dry matter the grain Genetic resources and breeding Oryza
of Oryza punctata from Sudan contains: crude punctata is considered a source of resistance to
protein 13.9 g, fat 4.0 g, soluble carbohydrate various diseases and pests affecting Oryza sa-
74.8 g, crude fibre 2.9 g, Ca 40 mg, Mg 270 mg, tiva, including bacterial leaf blight (Xanthomo-
P 550 mg, Fe 16.8 mg and Zn 3.9 mg. The es- nas oryzae pv. oryzae) and brown planthopper
sential amino acid composition per 100 g pro- (Nilaparvata lugens).
tein (16 g N) is: lysine 3.6 g, methionine 2.2 g, Prospects Although the grains of Oryza
phenylalanine 5.2 g, threonine 3.4 g, valine 5.9 punctata have a good nutritional quality, they
g, leucine 8.6 g and isoleucine 4.1 g (Salih & seem to be used as a famine food only, and the
Nour, 1992). plant is considered a noxious weed in rice cul-
Botany Annual or perennial grass 50-120(- tivation. Oryza punctata may be useful in
150) cm tall, growing in tufts; stem (culm) erect Oryza sativa breeding, although it is geneti-
or geniculately ascending, branched, striate, cally more distant than Oryza barthii A.Chev.
glabrous. Leaves alternate, simple and entire; and Oryza longistaminata A.Chev. & Roehr.
leaf sheath often spongy, distinctly striate; Major references Hanelt & Institute of
ligule 3—10 mm long, rounded, truncate or Plant Genetics and Crop Plant Research (Edi-
somewhat acute; blade linear, 15-45 cm x 0.5- tors), 2001; Kaushal & Ravi, 1998; Launert,
2.5 cm, acuminate, pale green or rarely glau- 1971; National Research Council, 1996; Salih &
cous, glabrous, usually slightly rough on both Nour, 1992.
surfaces. Inflorescence a terminal panicle 15— Other references Burkill, 1994; Clayton,
35 cm x 3-17 cm, loose, erect or somewhat 1970; Clayton, 1972; Gibbs Russell et al., 1990;
drooping, with spreading or ascending Lu, 1999; Mahmoud et al., 1995; Smith & Dil-
branches. Spikelet asymmetrically elliptical- day, 2003;Vaughan &Chang, 1992.
oblong or broadly oblong, (5-)5.5-6.5 mm long, Authors M. Brink
deciduous, greyish green or glaucous, 3-
flowered but 2 lowest florets reduced to sterile
lemmas 1-1.5 mm long; glumes reduced to a O R Y Z A S A T I V A L.
membranous, whitish narrow rim; lemma of
fertile floret slightly shorter than spikelet, Protologue Sp. pi. 1:333 (1753).
boat-shaped, leathery, hairy or rarely glabrous, Family Poaceae (Gramineae)
with pale yellow slender flexuous awn (1—)2— Chromosome number 2n = 12, 24, 36
7.5 cm long; palea slightly shorter than lemma Vernacular names Rice, paddy, Asian rice,
and much narrower, acute or tapering into a Asiatic rice (En). Riz, riz asiatique (Fr). Arroz
short point; lodicules 2; stamens 6; ovary supe- (Po). Mpunga (Sw).
rior, with 2 plumose blackish stigmas. Fruit an Origin and geographic distribution Oryza
oblong caryopsis (grain) 4-5 mm long, gla- sativa evolved in Asia, but the exact time and
brous, pale brown. place of its domestication are not known for
Oryza comprises about 20 wild species distrib- certain. Remains of rice in China have been
uted throughout the tropics and subtropics, dated to 6500 BC; the earliest archaeological
and 2 cultivated species, Oryza sativa L. and evidence from India goes back to 2500 BC.
Oryza glaberrima Steud. Oryza punctata is Oryza sativa was brought from Asia into tropi-
classified in ser. Latifoliae. cal Africa along different routes. Seamen-
Within Oryza punctata diploid (2re = 24) and farmers began sailing from Indonesia to Mada-
tetraploid (2n = 48) plants are known. Oryza gascar probably a few centuries BC and started
punctata can be crossed with Oryza sativa us- cultivating Oryza sativa there. Another impor-
ing embryo rescue techniques. tant contact between Africa and Asia at the
Ecology Oryza punctata is found in swampy dawn of the Christian era was the trade route
locations, on stream banks, in pond margins from Sri Lanka and India via Oman to Somalia
and pools, up to 1200 m altitude. It is a noxious and the islands Zanzibar and Kilwa off the
weed in rice cultivation and a potential seed coast of Tanzania. Most probably Oryza sativa
contaminant ofrice cultivars. migrated from Egypt, where it was introduced
 ORYZA 113

for paper, for the production of compost and

mushroom growing medium, for mulching
vegetable crops, for making ropes, sacks, mats
and hats, for roof thatching, and to make plas-
tering material (mixed with clay mud) for the
construction of houses, and for incorporation
into the soil or burning on the field as a way to
maintain/improve soil fertility.
Several traditional medicinal applications of
rice have been reported from tropical Africa:
leaf dressings are applied to ulcers and grain
decoctions are drunk to treat diarrhoea, as a
diuretic and as an emollient. Rice powder is
applied against itch in Senegal. In DR Congo a
decoction of the roots, leaves and husks is
taken against madness and beriberi.
Production and international trade Accord-
Oryza sativa -planted ing to FAO estimates the average annual world
production during 1999-2003 was 593 million t
about 800-900 AD, to West Africa. The final paddy (unhusked grain) from 153 million ha.
penetration of Oryza sativa into Africa was Asia accounts for 90% of the world production
along the slave trading routes from the East and area. During 1999-2003 tropical Africa
African coast and Zanzibar to DR Congo from produced on average 11.9 million t paddy (2%
about 1500 AD onwards. At the same time of world production) annually on 7.7 million ha
Oryza sativa was introduced into Senegal, (5% of world area); these data include African
Guinea Bissau and Sierra Leone by the Portu- rice (Oryza glaberrima Steud.), which occupies
guese on their return from expeditions to India. less than 20% of the rice area in West Africa.
Nowadays it is cultivated throughout the hu- The main producers are Nigeria (3.5 million t
mid tropics and in many subtropical and tem- from 2.9 million ha), Madagascar (2.6 million t
perate areas with a frost-free period longer from 1.2 million ha) and Côte d'Ivoire (1.1 mil-
than 130 days. lion t from 0.5 million ha). The annual world
Uses The rice grain is cooked by boiling or paddy production increased steadily from 241
steaming, and eaten mostly with pulses, vege- million t/year in 1961-1965 to 593 million
tables, fish or meat. Flour from rice is used for t/year in 1999-2003, and the harvested area
breakfast foods, baby foods, bread and cake from 121 to 153 million ha. In the same period
mixes and cosmetics. Starch made from broken the annual paddy production in tropical Africa
rice is used as laundry starch and in foods, increased from 3.6 to 11.9 million t/year, and
cosmetics and textile manufacture. Beers, the harvested area from 2.8 to 7.7 million ha.
wines and spirits are made from rice. Only 5% of the world's rice production enters
The husk or hull is used as fuel, bedding, ab- into international trade. Thailand is the
sorbent, packing material and as carrier for world's largest exporter of milled rice (26% of
vitamins and drugs; it is also made into build- world trade during 1998-2002) followed by
ing board. The charred hull is used for filtra- Vietnam, India, the United States, China and
tion of impurities in water, a medium for hy- Pakistan. All countries in tropical Africa are
droponics and manufacture of charcoal bri- net importers of milled rice and during 1998-
quettes. 2002 an average of 4.8 million t milled rice was
Rice bran or meal obtained in pearling and imported annually. This means that more than
polishing is a valuable livestock and poultry one third of the rice consumption in tropical
feed. Oil is extracted from the bran. Crude rice Africa is satisfied through imports. Main rice
bran oil is processed into solidified oil, stearic importers are Nigeria, Senegal and Côte
and oleic acids, glycerine and soap. Processed d'Ivoire. Per capita annual milled rice con-
bran oil is used for cooking, antirust and anti- sumption in tropical Africa varies tremen-
corrosive agents, textile and leather finishers, dously between 0.15 kg and 95 kg with an av-
and in medicine. erage of about 18 kg for the period 1998-2002.
Rice straw is used for animal feed and bedding, In Madagascar, Sierra Leone and Guinea Bis-
for the manufacture of straw boards and pulp sau it is the main source of energy.
114 CEREALS AND PULSES

Properties Raw brown rice contains per 100

g edible portion: water 13.9 g, energy 1518 kJ
(363 kcal), protein 6.7 g, fat 2.8 g, carbohydrate
81.3 g, dietary fibre 3.8 g, Ca 10 mg, Mg 110
mg, P 310 mg, Fe 1.4 mg, Zn 1.8 mg, thiamin
0.59 mg, riboflavin 0.07 mg, niacin 5.3 mg,
vitamin B-60.56 mg, folate 49 ug, ascorbic acid
0 mg. Raw polished rice contains per 100 g
edible portion: water 11.7 g, energy 1536 kJ
(367 kcal), protein 6.5 g, fat 1.0 g, carbohydrate
86.8 g, dietary fibre 2.2 g, Ca 4 mg, Mg 13 mg,
P 100 mg, Fe 0.5 mg, Zn 1.3 mg, thiamin 0.08
mg, riboflavin 0.02 mg, niacin 1.5 mg, vitamin
B6 0.30 mg, folate 20 (ig, ascorbic acid 0 mg
(Holland, Unwin & Buss, 1988). The essential
amino acid composition of raw polished rice per
100 g edible portion is: tryptophan 87 mg, ly-
sine 250 mg, methionine 140 mg, phenyla-
lanine 330 mg, threonine 230 mg, valine 390
mg, leucine 560 mg and isoleucine 260 mg
(Paul, Southgate & Russell, 1980). Milling and
polishing result in a loss of protein, fat, miner-
als (phosphorus and potassium) and vitamins
(thiamin, riboflavin and niacin). However,
these operations improve the storability and
reduce the cooking time. Oryza sativa - 1, plant base with roots;2, ligule
Rice grain endosperm may be waxy (glutinous) and auricles; 3, panicle with leaf;4, flowering
or non-waxy (non-glutinous) depending on the spikelet; 5, ovary with stigmas; 6, spikelet with
content of amylose and amylopectin. The mature grain.
higher the amylopectin content, the more glu- Source: PROSEA
tinous the product is. The endosperm also con-
tains sugar, fat, crude fibre, vitamins and inor- (up to 5 m long in some floating types), forming
ganic matter. The flavour of rice is variable small tufts; roots fibrous, arising from the base
and aromatic rice cultivars are highly appreci- of the shoots; stem (culm) erect or ascending
ated throughout the world. A major component from a geniculate base, terete, smooth, gla-
ofthe flavour is 2-acetyl-l-pyrroline. brous. Leaves alternate, simple; sheath
Rice bran contains: water 9.9%, gross energy coarsely striate, tight when young, later some-
1940 kJ (463 kcal) per 100 g, crude protein what loose, often somewhat spongy, green or
13.8%, crude fibre 7.8%, ether extract 16.4%. sometimes tinged with brown or purple,
After oil extraction, rice bran contains: water smooth, glabrous; ligule 1.5—3 cm long, trian-
9.8%, gross energy 1590 kJ (380 kcal) per 100 gular, acute, entire or split, membranous, usu-
g, crude protein 14.4%, crude fibre 9.3%, ether ally glabrous; auricles often present, falcate, 1-
extract 3.1%.The husk forms about 20% of the 5 mm long, hairy; blade linear, tapering to an
unhusked grain weight, and is very rich in acute point, 12-65 cm x 0.5-2 cm, bright green
silica. Rice straw contains approximately: wa- to glaucous, glabrous or puberulous, smooth on
ter 7.0%, protein 3.4%, fat 0.9%, carbohydrate the lower surface, slightly rough on the upper
47.8%, fibre 33.4% and ash 7.5%. It is nutri- surface, midrib usually distinct. Inflorescence a
tionally inferior to other cereal straws unless terminal panicle up to 50 cm long, erect, curved
ensiled. or drooping, with 50-500 spikelets; branches
Rice straw is not particularly suitable for pa- solitary or clustered, nearly erect to spreading.
permaking due to the high silica content (12— Spikelet solitary, asymmetrically oblong to
18%) and is used for this purpose mainly in elliptical-oblong, 7-11 mm x 2.5-3.5 mm, with
countries where wood is scarce, e.g. in India pedicel up to 4 mm long, 3-flowered but 2 low-
and China. The ultimate fibre cells are (0.4-) est florets reduced to sterile lemmas 2-3 mm
1.4(-3.4) mm long and (4-)9(-16) (im wide. long; glumes small; lemma of fertile floret 6-10
Description Annual grass up to 1.8 m tall mm long, boat-shaped, sometimes awned; palea
 ORYZA 115

about as long as lemma; lodicules 2; stamens 6; perature for germination is 30-32°C. Most cul-
ovary superior, with 2 plumose stigmas. Fruit tivars have a short dormancy or none at all,
a caryopsis (grain), ovoid, ellipsoid or cylindri- but in some it may last up to 4 months. Ten
cal, 5-7.5 mm x 2-3.5 mm, often whitish yel- days after germination the plant becomes in-
low or brown to brownish grey. dependent as the seed reserve is exhausted.
Other botanical information Oryza com- Tillering begins thereafter, although it may be
prises about 20 wild species distributed a week later in transplanted seedlings. In
throughout the tropics and subtropics, and 2 modern cultivars with an average maturation
cultivated species, Oryza sativa and Oryza gla- period, maximum tillering stage is attained
berrima. In the most recent classification around 45 days after transplanting and coin-
Oryza has been divided into 3 sections: sect. cides with panicle initiation. The duration of
Pädia, sect. Brachyantha and sect. Oryza. Sec- the vegetative stage ranges from 7 to more
tion Oryza is subdivided into 3 series: ser. Lati- than 120 days. The reproductive stage starts at
foliae, ser. Australiensis and ser. Sativae. panicle initiation, and the period from panicle
Oryza sativa is classified in ser. Sativae, to- initiation to flowering is around 35 days. Rice
gether with, among others, Oryza glaberrima, is almost 100% self-pollinating, but small
Oryza barthii A.Chev., and Oryza longistami- amounts of cross pollination by wind do occur.
nata A.Chev. & Roehr. Oryza glaberrima culti- It takes around 7 days to complete the anthesis
vars are grown only in Africa. Introgression of of all spikelets in a panicle, starting from the
characters from Oryza glaberrima, Oryza top and progressing downwards. The period
barthii and Oryza longistaminata may have from flowering to full ripeness of all the grains
added new dimensions to the variability of in a panicle is usually about 30 days. Low tem-
Oryza sativa. perature can delay maturity and high tempera-
Cultivated rice Oryza sativa is supposed to ture accelerates it. Floating rice has a long
have evolved from perennial types (Oryza rufi- maturation period of 7 months or more. Rice
pogon Griff.) to annual types (Oryza nivara roots can grow under low oxygen concentra-
S.D.Sharma & Shastri, sometimes included in tions. The roots are not typically aquatic as
Oryza rufipogon). There is a natural gene flow they are much branched and have a profusion
between these 3 species, and they form a large of root hairs; later, spongy tissue (aerenchyma)
species complex together with weedy forms of develops in the cortex.
rice (popularly called 'red rice' because of their Ecology Rice is grown as far north as 53°N
red endosperm). There are 2 major eco-geo- in Moho, northern China and as far south as
graphical cultivar groups of Oryza sativa: In- 35°S in New South Wales, Australia. It grows
dica Group, which mainly includes cultivars on dry or flooded soil and at elevations ranging
from the tropics, and Japonica Group, which from sea level to at least 2400 m. The average
includes cultivars from temperate/subtropical temperature during the growing season varies
areas. Traditional cultivars from Indica Group from 20-38°C. Night temperatures below 15°C
are tall, leafy, strongly tillering, and prone to can cause spikelet sterility. Temperatures
lodging; they respond poorly to fertilization, above 21°C at flowering are needed for anthe-
particularly to nitrogen, and are sensitive to sis and pollination. Upland rice requires an
photoperiod; they are hardy, resistant to dis- assured rainfall of at least 750 mm over a pe-
ease and tolerate unfavourable growing condi- riod of 3-4 months and does not tolerate desic-
tions; they will produce fair yields under condi- cation. Lowland rice tends to be concentrated
tions of low management. Modern Japonica in flat lowlands, river basins and deltas. The
Group cultivars are small, and are less tiller- average water requirement for irrigated rice is
ing, less leafy, resistant to lodging, insensitive 1200 mm per crop or 200 mm of rainfall per
to photoperiod and are early maturing. The month or an equivalent amount from irriga-
characteristics of the two cultivar groups have tion. Relative humidity within the crop canopy
become less distinct because of the interbreed- is high, since there is standing water in most
ing programmes in recent years. Rice may also rice crops. A low relative humidity above the
be classified according to the conditions under canopy during the dry season aggravated by
which it is grown, according to the size, shape strong winds can cause spikelet sterility. Tra-
and texture of the grain, or according to the ditional cultivars are generally photoperiod
period needed to mature. sensitive, and flower when daylengths are
Growth and development Rice seed ger- short (critical daylength of 12.5-14 hours).
minates in 24—48 hours. The optimum tem- Many modern cultivars are photoperiod insen-
116 CEREALS AND PULSES

sitive. - Upland rice, which may be subdivided into

The soils on which rice grows vary greatly: dryland rice, whereby moisture supply is en-
texture ranges from sand to clay, organic mat- tirely dependent on rainfall, and hydromor-
ter content from 1-50%, pH from 3-10, salt phic rice where the rooting zone is periodi-
content up to 1%, and nutrient availability cally saturated by a fluctuating water table,
from acute deficiencies to surplus. Rice does in addition to rainfall;
best in fertile heavy soils. The optimum pH for - Lowland rice, including mangrove swamp
flooded soil is 6.5-7.0. The often sandy texture rice along the coastal regions with tidal in-
of soils in tropical Africa is a constraint to pro- trusion, inland swamp rice on flat or V-
ductivity due to drought stress, low inherent shaped valley bottoms with varying degrees
soil fertility and leaching. Groundwater salin- of flooding, and rice on bunded fields under
ity problems occur in the dry Sahel zone where rainfed or irrigated conditions;
rice is grown under irrigation. In lowland - Deepwater rice, in which the rapid growth of
coastal West Africa rice productivity is affected the internodes keeps pace with the rising
by saline water intrusion. The majority of water up to 5 m or more, starting from 50 cm
mangrove swamp soils along the West African of standing water.
coast are furthermore potential or actual acid In upland rice cultivation the fields are nor-
sulphate soils. In West Africa iron toxicity in mally cleared through the slash-and-burn prac-
valley bottoms is most severe in areas where tice. Soil preparation is normally minimal. The
the adjacent uplands are strongly leached Ulti- rice is broadcast or dibbled when the rains
sols. Lowland rice and deep-water rice may be start. It is often grown as the first crop in rota-
subjected to both drought or complete submer- tion or intercropped with other crops such as
gence. In submerged soil the pH tends to be cassava, maize, sorghum, cowpea, groundnut
neutral, i.e. the pH of acid soils increases and other pulse crops.
whereas the pH of calcareous and sodic soils In lowland rainfed-rice areas the land is mostly
decreases. Ions of Fe, N and S are reduced, the prepared while it is wet and only in rare occa-
supply and availability of the elements N, P, Si sions when it is dry. The wetland tillage
and Mo improve, whereas the concentration of method consists of soaking the land until the
water-soluble Zn and Cu decreases. Toxic re- soil is saturated, ploughing to a depth of 10-20
duction products such as methane, organic cm using a plough drawn by oxen/small ma-
acids and hydrogen sulphide are formed. The chines or by using a hand hoe, preferably when
flooding of rice soils also creates a favourable there is a little water on the land, and harrow-
environment for anaerobic microbes and the ing, during which big clods of soil are broken
accompanying biochemical changes. As a re- and puddled with water. The important bene-
sult, the decomposition rate of organic matter fits of puddling include the apparent reduction
decreases. However, a thin surface layer gen- of moisture loss by percolation, better weed
erally remains oxidized and sustains aerobic control, and easy transplanting. In lowland rice
microbes. cultivation seedlings are mostly raised on wet
Propagation and planting Rice is propa- nursery beds and sometimes on dry nursery
gated by seed. The 1000-seed weight is 20-35 beds. Wet nursery beds are made in the pud-
g. The seed may either be broadcast or drilled dled or wet field. Normally farmers use 50-60
directly in the field, or seedlings may be grown kg of rice seeds to plant one ha. Seeds are pre-
in nurseries and transplanted. Direct seeding germinated and spread on the bed which is
is done in dry or puddled soil. In puddled soil kept constantly wet. Dry nursery beds are pre-
the (pre-germinated) seeds are broadcast. After pared near the water source before land prepa-
sowing the water level is kept at 0—5cm under ration. The seeds are sown and then covered
tropical conditions. In dry soil the seeds are with a thin layer of soil and watered until
sown just before or after land preparation. In saturation for uniform germination. Further
the latter case the seeds are then covered watering is applied as needed. In both cases
lightly with soil. The seeds are sown just before the seedlings are ready for transplanting 20—35
the rains begin and germination occurs after days after sowing. At transplanting heavy
heavy continuous rains. This method makes it tillering cultivars in fertile valley bottoms are
possible to have initial crop growth from early wider spaced (30 cm x 30 cm) than slightly
rains. tillering cultivars in upper, sandy fields (20 cm
In tropical Africa various rice-growing systems x 20 cm). The spacing in irrigated rice is nor-
are distinguished: mally 20 cm x 20 cm with 2—4 plants per hill
 ORYZA 117

(500,000-1,000,000 plants/ha). Rice is gener- initiation. Fertilizer is broadcast by hand. The

ally a sole crop under lowland conditions. Near most common mineral deficiencies in rice culti-
harvest, relay planting is rarely practised. In vation are of nitrogen and phosphorus, with
many parts of the tropics 2 or even 3 crops of potassium and sulphur in limited areas and
rice can be grown per year. There is a lack of sometimes zinc and silicon on peaty soils. Defi-
accurate data on the extent of different rice ciency of potassium is often associated with
systems in tropical Africa. The upland rice iron toxicity. Upland rice often suffers from
ecosystem, including hydromorphic rice, ac- sulphur deficiency. Zinc deficiency occurs regu-
counts for an estimated 50% of the total rice larly in rice areas because of the high pH and
area in tropical Africa; lowland rice cultivation, strong reduction of the soil. Influenced by re-
including mangrove swamp rice, inland swamp duction and poor internal drainage, several
rice and irrigated rice, accounts for 45% of the toxic elements such as iron, which inhibit the
total rice area; deep-water rice cultivation oc- uptake of phosphorus in the plant, may accu-
cupies the remaining 5%.Most rice is grown on mulate in the environment of the root. Often a
smallholdings of 0.5-2 ha. harmful excess of elements such as calcium is
Management The agronomy of rice is di- accompanied by a lack of other elements such
verse due to the differences in cultivation sys- as phosphorus, iron and zinc. Double cropping
tems. Growing of upland rice is usually rela- is inadvisable where physiological diseases
tively labour-extensive, but transplanting rice occur. Green manure and Azolla are rarely
by hand in puddled soil is a labour-intensive used in tropical Africa. However, the fast grow-
operation. Weeding is generally not necessary ing and actively nitrogen-fixing Sesbania ros-
in the first 2 weeks. Manual weeding is com- trata Bremek. & Oberm. is a promising green
mon practice, although chemical weed control manure crop. Nitrogen fixation also takes place
is also becoming popular in tropical Africa, in paddy soils by Azotobacter and blue green
especially in irrigated rice areas. Three timely algae (cyanobacteria). Organic fertilizers such
weedings are normally necessary in broadcast as farmyard manure and compost are not
rice. commonly applied to rice crops in tropical Af-
In the cultivation of lowland rice, the land is rica. Although soil conditions are normally
inundated from the time of planting until the improved by incorporating organic fertilizers,
approach of harvest. The water is supplied the result is not immediately apparent. Poor
either by flooding during the rainy season, by availability, transport problems and the high
growing the crop in naturally swampy land or amount of labour involved also discourage its
by controlled irrigation. The water level is kept use.
at a height of 5-15 cm to suppress weed growth The degree of mechanization is in general lim-
and to ensure water availability. Continuous ited in rice cultivation in tropical Africa. Occa-
flooding at a static 2.5-7.5 cm depth is best. sionally farmers use tractors or small two-
The fields may be drained temporarily to facili- wheel power tillers for land preparation and
tate weeding and fertilizing. At flowering the powered threshing machines during harvest.
water level is gradually reduced until the field For various reasons many rice fields are left
is almost dry at harvest. Generally 1.5-2 m of fallow in the dry season. In areas with suitable
water (rainfall plus irrigation) are required to climatic and soil conditions for dry-season cul-
produce a good crop. The period in which rice is tivation, rice may be rotated with crops such as
most sensitive to water shortage is from 20 other cereals, pulses and vegetables.
days before to 10 days after the beginning of Diseases and pests The most common and
flowering. severe disease of rice in tropical Africa is blast
Fertilizer application is limited in rice cultiva- (Pyricularia grisea, synonym: Pyricularia ory-
tion in tropical Africa. Only in irrigated rice zae). Although this disease is often related to
with controlled water supply and modern culti- drought stress and therefore especially severe
vars do farmers generally use significant in upland and drought-prone areas, it may also
amounts of mineral fertilizers. The amount of be severe elsewhere. Low light intensity, nutri-
fertilizer used is usually 60-120 kg N, 10-20 tional imbalances (especially K-deficiency),
kg P and 0-30 kg K per ha. Higher nitrogen excessive N-supply, and relatively low tem-
rates are used during the dry season when peratures (20-28°C) are further factors favour-
solar radiation is higher and increase in grain ing this disease. The blast fungus can infect
yield is larger. Generally, nitrogen fertilizer is rice leaves, nodes and floral parts, particularly
only topdressed, mostly before or at panicle the basal part of the panicle. Other important
118 CEREALS AND PULSES

diseases of rice in tropical Africa are bacterial nomic injury threshold. Cultural methods in-
leaf blight (Xanthomonas oryzae pv. oryzae), clude sanitation (the destruction of crop resi-
rice yellow mottle virus (RYMV, only found in dues, of alternative hosts including weeds and
Africa), brown spot (Cochliobolus miyabeanus), of habitats), tillage and flooding of fields, crop
leaf scald (Microdochium oryzae), sheath blight rotation, intercropping, proper timing of plant-
(Thanatephorus cucumeris), narrow brown leaf ing and harvest, use of trap crops, and proper
spot (Cercospora janseana) and sheath rot fertilizer and water management.
caused by Sarocladium oryzae. The use of re- Birds eat broadcast seeds, disturb young
sistant cultivars, the judicious application of N transplanted seedlings and eat rice grains;
fertilizer, adjusted planting time, crop rotation losses can be very high. Rodents attack rice at
and phytosanitary and quarantine measures all stages of growth and also stored grain, and
limit losses from rice diseases. Chemical con- losses due to rodents are often serious. Less
trol for blast and other rice diseases is hardly damage is caused by snails, crabs and shrimps.
used in tropical Africa. Parasitic weeds of the genus Striga may cause
Nematodes attack roots and young, unfurled serious losses in upland rice, e.g. Striga aspera
leaves and reduce rice production in certain (Willd.) Benth. and Striga hermonthica (Delile)
parts of tropical Africa. Most insect species Benth. in West Africa, and Striga asiatica (L.)
causing damage to rice in the field and to the Kuntze in the Indian Ocean Islands.
grain during storage in tropical Africa are in- Harvesting Grain should be harvested be-
digenous, and different from those found in fore it is fully mature (around 21—24% mois-
Asia. Internal stem feeders such as stem bor- ture), usually about 30 days after flowering, or
ers, the stalk-eyed fly and gall midge generally when 90% of the grains are firm and do not
cause the most severe damage. The most com- have a greenish tint. Wetting and drying cause
mon species of stem borers in tropical Africa grain cracking, cracks being formed more read-
are white stem borer (Maliarpha separatella), ily when the grain is quite hard. Harvesting by
pink stem borers (Sesamia spp.) and striped hand, the commonest method, is very labour-
stem borer (Chilo spp.). Damage results from intensive. In some areas a small knife is used,
larvae feeding within the stem, severing the but in many areas farmers use a sickle to cut
vascular system. Dead heart is the damage to the panicles plus some or all of the culms. Me-
the tiller before flowering. White head is the chanical harvesters are very rare in tropical
damage after flowering which causes the entire Africa. The harvested rice plants are either
panicle to dry. The damage from the stalk-eyed allowed to dry in the field or bundled for proc-
fly (mainly Diopsis macrophthalma) resembles essing in a selected area.
the dead heart damage from stem borers as it Yield Average rice yields are 1.4 t/ha in
generally attacks the rice plant at the early tropical Africa, 4.1 t/ha in Asia and 4.0 t/ha in
tillering stage. The feeding of the gall midge the world in general. Yields are generally
maggot (Orseolia oryzivora) stimulates the leaf higher during the dry season than during the
sheath to grow into a gall and tillers with galls wet season, and higher in lowland rice than in
do not bear panicles. Termites and mole crick- upland rice. The yield of upland rice varies
ets attack rice plants especially in rainfed up- between 0.5 and 1.5 t/ha in tropical Africa but
land rice. may reach 4 t/ha in Latin America. Rainfed
The most serious insect pests of stored rice are lowland rice is higher yielding than upland rice
the rice weevil (Sitophilus oryzae) and the but may suffer a drastic reduction in years
lesser grain borer (Rhyzopertha dominica). with drought or floods. In a rainfed bunded
These insects can completely destroy the grain. lowland rice area in Tanzania yields are 3-4
Insects can be controlled by chemical, cultural, t/ha in good years, but can drop to 0.5 t/ha in
and biological methods. In tropical Africa bad years. Yields of irrigated lowland rice in
farmers use insecticides but at far lower levels tropical Africa are generally 3-6 t/ha. Yields in
than in Asia. It is important to use various the deep-water rice areas are generally low
crop protection methods in an integrated pest (0.6-1.2 t/ha), but they are more stable than in
management (IPM) system for rice in tropical the upland rice areas of tropical Africa.
Africa that is sustainable, inexpensive, and Handling after harvest Threshing is gen-
environmentally safe. It should combine the erally done by hand, by beating the bundles on
use of resistant cultivars, cultural methods, a stone or drum, or by beating the panicles
biological control and, finally, chemical control with wooden sticks on a canvas. However, mo-
when pest damage threatens to exceed the eco- torized and pedal-driven threshing machines
 ORYZA 119

are becoming popular. Winnowing is usually ity to diseases and pests, and reaction to envi-
done by shaking and tossing the grain on a ronmental stresses, mineral deficiencies or
basket-work tray with a narrow rim. Some- toxicities. Large germplasm collections of
times hand-winnowing machines are used. Oryza sativa are also held in China (China
After winnowing, the grain is dried in the sun National Rice Research Institute, Huangzhou,
and is then ready for hulling or transport to 70,000 accessions) and India (National Bureau
the mill. Proper drying of the rice grains is of Plant Genetic Resources, New Delhi, 26,000
important to prevent germination and rapid accessions). Apart from at WARDA, in tropical
loss of quality. Optimum moisture content for Africa large collections are present in Nigeria
storage is 12.5%. Rice grain is mostly stored in (International Institute of Tropical Agriculture
sacks after drying. Increase in fat acidity dur- (UTA), Ibadan, 9400 accessions; National Ce-
ing improper storage reduces the eating qual- reals Research Institute, Badeggi, 3500 acces-
ity. Temperature and humidity during storage sions) and Madagascar (Département de Re-
affect rice quality. Rice for home consumption cherches Agronomiques de la République Mal-
is stored unhusked, as it is less susceptible to gache, Antananarivo, 2000 accessions). Collec-
deterioration. tion of wild rices is being emphasized for possi-
In rice milling the aim is to avoid breaking the ble new sources ofimportant genes.
kernels because whole kernels command a Breeding Rice grain yields in the tropics
higher price. There are different methods of have increased dramatically since the mid
milling. On milling, the grain gives approxi- 1960s with the introduction of 'IR8' and other
mately: husk 20%, whole kernels 50%, broken semi-dwarf cultivars, which do not lodge easily
kernels 16%, bran and meal 14%. The husked and allow high nitrogen fertilizer doses. In
or hulled rice is usually called brown rice, and tropical Africa these green revolution cultivars
this is then milled to remove the outer layers, are mainly used in irrigated rice with con-
after which it is polished to produce white rice. trolled water supply. Genetic improvement of
During milling and polishing some of the pro- rice in Africa was mainly focused on the upland
tein and much of the fat, minerals and vita- crop. This has led to the 'New Rice for Africa'
mins are removed, reducing the nutritional ('NERICA') cultivars, WARDA's major break-
value but increasing storability and reducing through in the early 1990s. 'NERICA' cultivars
cooking time. Parboiling (soaking, boiling and were the result of successful crossing of Oryza
drying) before milling improves the nutrient glaberrima with Oryza sativa. They combined
value of the grains but it is not common in higher tolerance to deep water, drought, weeds,
tropical Africa. blast and stalk-eyed fly from Oryza glaberrima
Genetic resources The exploration and col- with greater grain productivity and retention
lection of germplasm of African wild and culti- on the plant from Oryza sativa. 'NERICA' cul-
vated rice species was started in 1959 by Japa- tivars are proving to be popular with farmers,
nese researchers who were attracted by the not only because of their growth characteris-
great diversity. The earliest collections of rice tics, but also for their grain quality and nutri-
genetic resources in West Africa were built up tive value. They are further well suited to low-
at research stations at Rokupr, Sierra Leone input conditions. Breeding activities of WARDA
and Badeggi, Nigeria. Later on the French re- on lowland cultivars have led to the release of
search institutes ORSTOM and IRAT started cultivars with improved grain yield, resistance
collecting rice germplasm from francophone to blast and rice yellow mottle virus and toler-
countries and UTA, Ibadan, Nigeria, from ance to drought and iron toxicity. The im-
mainly anglophone countries. A combination of proved cultivar 'Sahel 108', released in 1994 by
these germplasm collections with almost WARDA, has a short life cycle enabling double-
15,000 accessions was then established by WAR- cropping in the irrigated rice systems in the
DA at Bouaké, Côte d'Ivoire. Most of these ac- Sahel. Wild Oryza species, such as Oryza
cessions are also available in the International barthii, Oryza longistaminata and Oryza punc-
Rice Germplasm Collection at the Interna- tata are useful sources of resistance to various
tional Rice Research Institute (IRRI), Los biotic and abiotic stresses. For instance, resis-
Banos, the Philippines where the largest Oryza tance to bacterial leaf blight has successfully
sativa collection is found with more than been transferred from Oryza longistaminata.
86,000 accessions, characterized on the basis of Biotechnology techniques used in rice breeding
about 80 traits. These traits not only include include plant tissue culture, molecular biology
morphological characters but also susceptibil- and genetic engineering. Two tissue culture
120 CEREALS ANDPULSES

techniques, embryo rescue and anther culture, tilizers with a higher efficiency and greater use
have already made important contributions. of nitrogen-fixing legumes, bacteria and blue-
Saturated genetic linkage maps based on mo- green algae. The applicability of methods of
lecular markers have been developed for rice, integrated soil fertility management (ISFM) in
using crosses between cultivars of Indica a certain locality can be best tested through
Group and Japonica Group, or between Oryza farmer field schools. An increased use of
sativa and Oryza longistaminata. These maps farmer field schools is also advocated for the
have made possible the identification of QTLs adoption of methods of integrated pest man-
for many useful traits, such as resistance to agement (IPM) by more rice farming house-
diseases and tolerance to drought. More than holds in tropical Africa. Further improvements
3000 molecular markers are available now, are expected from mechanization of rice farm-
making rice the best characterized crop. The ing, especially regarding land preparation,
project for sequencing the complete rice ge- weeding, harvesting, threshing and further
nome has recently been completed. Biotechnol- processing.
ogy's most novel contribution will probably be All these suggestions require research adjusted
in adding alien genes to the rice gene pool to the local conditions, a well-functioning ex-
through genetic engineering. One example is tension service, government support, and ac-
'Golden Rice', which is rice enriched with vita- tive participation of farming households. Some
min A. It is, however, still not clear if this ge- of the above topics are already being re-
netically modified rice will yield well, not be searched.
susceptible to diseases and pests and be palat- Major references Alam, John & Zan, 1985;
able. Several insecticidal toxin genes from Ba- Buddenhagen & Persley (Editors), 1978; Cat-
cillus thuringiensis (Bt) have been transferred ling, 1992; Grist, 1986; Lorieux, Ndjiondjop &
to rice and plants containing Bt genes have Ghesquière, 2000; Meertens, Ndege & Lupeja,
shown substantial resistance to stem borers 1999; Schalbroeck, 2001; Smith & Dilday,
and leaf folders. Recently, transgenic rice has 2003; Vergara & de Datta, 1996; Zan, John &
been obtained conferring resistance to sheath Alam, 1985.
blight. Genetic engineering is a relatively new Other references Abo, Sy &Alegbejo, 1998;
technology and one of the principal biosafety Burkill, 1994; Catling & Islam, 1999; Chang,
concerns is the spread of foreign genes by pol- 2000; Choudhury & Kennedy, 2004; de Datta,
len dispersal from transgenic rice to other rice 1981; de Vries & Toenniessen, 2001; Hanelt &
cultivars and wild rice species. Institute of Plant Genetics and Crop Plant Re-
Prospects At present, only an estimated 2% search (Editors), 2001;Holland, Unwin & Buss,
of the 200 million ha of wetlands in tropical 1988; Johnson et al., 1997; Jones et al., 1997;
Africa are used for lowland rice cultivation. Khush, 1997; Latham, 2004; Launert, 1971;
Therefore one of the biggest challenges for rice Lu, 1999; Neuwinger, 2000; Paul, Southgate &
development in tropical Africa is the utilization Russell, 1980; Sauvant, Perez & Tran, 2004;
of the large potential for expansion of lowland Vergara & Chang, 1983; Widjaja, Craske &
rice. The emphasis of genetic improvement Wootton, 1996.
should be directed to lowland rice ecosystems, Sources of illustration Vergara & de Dat-
which have a higher production potential than ta, 1996.
upland rice, for example the breeding of crosses Authors H.C.C. Meertens
of Oryza sativa and Oryza glaberrima for low- Based on PROSEA 10: Cereals.
land rice ecosystems. Any new types recom-
mended should be well adapted to the local
environment and methods of cultivation. For PANICUM KALAHARENSE Mez
that matter it is advisable that in the breeding
process greater use is made of farmer partici- Protologue Bot. Jahrb. Syst. 57: 187 (1921).
patory varietal selection (PVS) and farmer par- Family Poaceae (Gramineae)
ticipatory plant breeding (PPB). Breeding ac- Origin and geographic distribution Pani-
tivities for tropical Africa should include toler- cum kalaharense is distributed in Zambia,
ance of and adaptation to iron toxicity, salinity, Namibia, Botswana, Zimbabwe, Mozambique
alkalinity, acid sulphate soils, and relatively and South Africa.
extreme cool and hot temperatures. In tropical Uses The grain of Panicum kalaharense is
Africa there is still much room for increased eaten by the Wambo people in Namibia. Pani-
and integrated use of organic and mineral fer- cum kalaharense is also a pasture grass.
 PANICUM 121

Properties As a pasture grass Panicum P A N I C U M LAETUM K u n t h

kalaharense is recorded as being moderately
palatable when young. Protologue Révis. gramin. 2: 399, f. 113
Botany Robust, perennial grass up to 2.5 m (1831).
tall, tufted or with a short rootstock; stem Family Poaceae (Gramineae)
(culm) pubescent at base. Leaves alternate, Vernacular names Wild fonio, desert panic
simple and entire; leaf sheath densely hairy (En). Haze, fonio sauvage (Fr).
along margins in upper part; blade linear, 20- Origin and geographic distribution Pani-
50 cm x 2-8 mm, acuminate, flat or rolled, cum laetum is distributed from Mauritania,
tough, upper surface densely covered with Senegal and Gambia eastwards through the
short hairs. Inflorescence a broadly ovoid pani- southern Sahara and the Sahel to Eritrea; it is
cle 15-27 cm long, moderately to much also recorded from Tanzania.
branched. Spikelet ovoid, 3-4 mm long, 2- Uses Panicum laetum is one of the 'kreb'
flowered; lower glume broadly ovate, half to grasses, a group of grasses occurring in the
two-thirds the length of the spikelet, acute or Sahel and collected from the wild for human
acuminate, 3-5-veined, upper glume 5—9- consumption on a regular basis and especially
veined; lower floret male, lemma 7-9-veined, in times of food shortage; it is also regarded as
palea well-developed, upper floret female, a delicacy. The grain of Panicum laetum is
lemma and palea pale or dark, glossy; stamens crushed and made into porridge and cakes, and
3; ovary superior, stigmas 2. Fruit a caryopsis is sometimes collected on a large enough scale
(grain), ellipsoid, compressed. to be sold in local markets. Panicum laetum is
Panicum comprises about 470 species and is much appreciated by animals for grazing and is
mainly distributed in tropical and subtropical suitable for making hay or silage. It is consid-
regions, with some species extending to tem- ered to have potential for the restoration of
perate regions. Panicum kalaharense follows degraded pastures.
the C4-cycle photosynthetic pathway. Properties Whole grains ofPanicum laetum
Ecology Panicum kalaharense is considered collected in Mali contained per 100 g: water 3.3
to be drought resistant. It is locally common in g, energy 1580 k j (377 kcal), protein 9.5 g, fat
grassland and savanna habitats on sandy soils 4.8 g, carbohydrate 70.8 g, Ca 51 mg and Fe
in areas with an annual rainfall of 300-600 210 mg. The essential amino acid content of
mm, at 500-1200 m altitude. It is also found in whole grains per 16 g N was: tryptophan 1.3 g,
disturbed locations such as roadsides. lysine 2.0 g, methionine 2.6 g, phenylalanine
Management The grain of Panicum kala- 5.9 g, threonine 3.7 g, valine 6.0 g, leucine 11.3
harense is collected from the wild. g and isoleucine 4.7 g. Husked grains contained
Genetic resources and breeding A small per 100 g: water 1.9 g, energy 1630 kJ (389
collection of 3 accessions of Panicum kala- kcal), protein 12.4 g, fat 2.2 g, carbohydrate
harense is held at the International Livestock 82.1 g, ash 1.4 g, Ca 13 mg and Fe 24 mg. The
Research Institute (ILRI), Addis Ababa, Ethio- essential amino acid content of husked grains
pia. per 16 g N was: tryptophan 1.4 g, lysine 1.3 g,
Prospects Panicum kalaharense is an un- methionine 2.6 g, phenylalanine 6.3 g,
important local source of food and forage, and threonine 3.6 g, valine 6.0 g, leucine 12.2 g and
will probably remain so. Little is known about isoleucine 5.1 g (Beseth Nordeide, Holm &
this species and more information would be Oshaug, 1994). The most limiting amino acid is
useful, particularly on its food and fodder qual- lysine.
ity. Panicum laetum plants in mid-bloom in Niger
Major references Clayton, 1989; Gibbs Rus- contain crude protein 14.3%, crude fibre 28.8%,
sell et a l , 1990;Launert, 1970. crude fat 1.8%, nitrogen-free extracts 42.9%,
Other references Klaassen & Craven, 2003; Ca 0.30%, Mg 0.28% and P 0.42%.
Schulze et al., 1996. Botany Annual, tufted grass up to 75 cm
Authors M. Brink tall; stem (culm) slender, erect or geniculately
ascending, branched. Leaves alternate, simple
and entire; leaf sheath glabrous or bristly-
hairy; ligule short, fringed; blade linear-
lanceolate, flat, 5-25 cm x 5-12 mm, acumi-
nate, usually glabrous, margin smooth or bris-
tly hairy in lower part. Inflorescence an ovoid
122 CEREALS AND PULSES

panicle 6-20 cm long, much-branched, primary P A N I C U M MILIACEUM L.

branches ascending or spreading, branchlets
and pedicels slender. Spikelet narrowly ellip- Protologue Sp. pi. 1:58 (1753).
soid, 2.5-3 mm x 1.5 mm, acute, usually pale Family Poaceae (Gramineae)
green, 2-flowered; lower glume ovate, about % Chromosome number ïn = 36
the length of the spikelet, 5-7-veined, acute, Vernacular names Proso millet, common
upper glume elliptical, 7—11-veined, acute; millet, hog millet (En). Millet commun, kibi
lower floret sterile, lemma 9-11-veined, palea (Fr). Milho miudo, milho de canario (Po).
almost equally long, upper floret bisexual, Origin and geographic distribution Proso
lemma narrowly ovate, acute, pale, smooth, millet is of ancient cultivation. Its origin has
glossy; stamens 3; ovary superior, stigmas 2. not been ascertained, but it was probably do-
Fruit an ellipsoid caryopsis (grain) 1.5-2 mm mesticated in central and eastern Asia, where
long, compressed, yellowish. it has been cultivated for more than 5000
Panicum comprises about 470 species and is years. Proso millet has long been a major crop
mainly distributed in tropical and subtropical in northern China. In the Bronze Age it spread
regions, with some species extending to tem- widely in Europe, also to northern regions
perate regions. where the cold-susceptible foxtail millet (Se-
Ecology Panicum laetum is found in sea- taria italica (L.) P.Beauv.) could not be grown.
sonally moist locations in grassland, ditches, In Europe remains have been found in agricul-
and pond and river margins, often on black tural settlements dating back about 3000
clay soils. It is not particularly drought toler- years. Proso millet was the 'milium' of the Ro-
ant. In West Africa Panicum laetum often oc- mans and the true millet of history. It was in-
curs in very large, nearly pure stands. In Tan- troduced into North America after the arrival
zania it is found at 1000—1300 m altitude. of Columbus. In Europe and the United States
Management Panicum laetum is propa- its popularity as a cereal declined after the
gated by seed. The optimum temperature for large-scale introduction ofpotato and maize.
seed germination is 35°C. Scarification or re- Nowadays proso millet is cultivated for human
moval of the lemma and palea from the grain consumption mainly in eastern and central
greatly improves germination. In West Africa Asia, and to a lesser extent in eastern Europe
Panicum laetum is collected from the wild by (Russia, Danube region) and from western Asia
sweeping through the crop with a calabash, to Pakistan and India (Bihar, Andhra Pra-
bowl or tray when the ears are ready to shat- desh). It is occasionally grown in other parts of
ter. The grains ofPanicum laetum are favoured Europe and Asia and in North America, mainly
by quelea birds. as a source of feed for cage-birds and poultry,
Genetic resources and breeding A collec- and as fodder. In tropical Africa, it is cultivated
tion of 25 accessions of Panicum laetum is held in Ethiopia, eastern Kenya, Malawi, Botswana,
at the International Livestock Research Insti- Zimbabwe and Madagascar. It is also recorded
tute (ILRI), Addis Ababa, Ethiopia. In view of from Lesotho. Its importance in Kenya is said
its wide distribution and abundance Panicum
laetum is not threatened by genetic erosion.
Prospects Panicum laetum is of importance
in marginal areas and has potential for restor-
ing over-grazed pastures. The selection of im-
proved strains for grain and fodder production
is recommended.
Major references Beseth Nordeide, Holm &
Oshaug, 1994; Burkill, 1994; Naegele, 1977;
National Research Council, 1996; Phillips,
1995.
Other references Bartha, 1970; Clayton,
1972; Harlan, 1989b; Keith & Plowes, 1997; le
Grand, 1979; van der Hoek & Jansen, 1996a;
Veldkamp, 1996b;Veldkamp, Wijs &Zoetemeyer,
1989.
Authors M. Brink
Panicum miliaceum -planted
 PANICUM 123

to have declined since the 1950s following the results in volatile prices.
advance of maize cultivation. Proso millet has Properties Proso millet contains per 100 g
widely naturalized, and is sometimes a trou- edible portion: water 8.6 g, energy 1582 kJ (378
blesome weed, e.g. in the United States and kcal), protein 11.0 g, fat 4.2 g, carbohydrate
Russia. 72.9 g, dietary fibre 8.5 g, Ca 8 mg, Mg 114 mg,
Uses The husked grains of proso millet are P 285 mg, Fe 3.0 mg, Zn 1.7 mg, vitamin A 0
eaten whole, boiled like rice or after roasting. IU, thiamin 0.42 mg, riboflavin 0.29 mg, niacin
They are also cooked into porridge, or, after 4.7 mg, vitamin P>6 0.38 mg, folate 85 ug, ascor-
grinding, baked into flat bread or chapatti. bic acid 0 mg. The essential amino-acid compo-
Only the flour of gluten-rich types can be used sition per 100 g edible portion is: tryptophan
for leavened bread and cakes; the flour of other 119 mg, lysine 212 mg, methionine 221 mg,
types has to be mixed with wheat flour. In phenylalanine 580 mg, threonine 353 mg,
China, where proso millet flour is made into valine 578 mg, leucine 1400 mg and isoleucine
bread, cultivars with glutinous (waxy) en- 465 mg. The principal fatty acids are (per 100 g
dosperm are favoured; in Mongolia, where the edible portion): linoleic acid 2015 mg, oleic
grains are cooked like rice, non-glutinous culti- acid 739 mg, palmitic acid 528 mg, stearic acid
vars are grown. In Ethiopia, the grains are 145 mg and linolenic acid 118 mg (USDA,
fermented into a kind of beverage ('tella'). 2004). The grains have a relatively high indi-
Elsewhere they are used for making beer and gestible fibre content because the seeds are
brandy. The grain is a feed for animals, includ- enclosed in hulls which are difficult to remove
ing pigs, fowls and cage-birds. The plant is by conventional milling processes. The husked
used as a forage. The forage quality of the grain of proso millet has a slightly nutty fla-
straw is poor, and in India it is more often used vour.
for bedding for cattle. The straw is also made
into brooms. Starch from the grains has been
used for sizing textiles. Various medicinal uses
of proso millet have been recorded in Asia; the
seeds are used as a demulcent and as a treat-
ment for abscesses and boils, and stem and
root decoctions are taken against haematuria.
Production and international trade Pro-
duction statistics for proso millet are scarce
because they are usually lumped with those of
other millets. The average annual world pro-
duction of proso millet in 1981-1985 was esti-
mated at 4.9 million t, with the Soviet Union
(2.3 million t), China (1.6 million t) and India
(0.5 million t) as main producers. The annual
production in 1992—1994 was estimated at 4
million t. In tropical Africa the production of
proso millet is very low compared to that of
pearl millet (Pennisetum glaucum (L.) R.Br.)
and finger millet (Eleusine coracana (L.)
Gaertn.), but no statistics are available.
Compared to the total world trade in cereals,
the international trade in millets is insignifi-
cant. The world millet trade in 1999-2003
amounted to about 250,000 t/year. The share of
proso millet of the total recorded millet trade Panicum miliaceum - 1,upper part of flowering
has been estimated at about two-thirds. Most culm of plant type with loose inflorescence; 2,
proso millet traded internationally is imported inflorescence branch of plant type with loose
by the pet-food industry in industrialized coun- inflorescence; 3, upper part of flowering culm of
tries for use as bird feed. As millet yields are plant type with compact inflorescence; 4, inflo-
relatively low, prices on the world market are rescence branch of plant type with compact in-
generally higher than those of other cereals; florescence; 5, grains.
the small size of the international millet trade Source: PROSEA
124 CEREALS AND PULSES

Non-glutinous proso millet cultivars are con- vars and landraces, and 5 groups have been
sidered suitable for the diet of people with distinguished, mainly based on size and shape
coeliac disease. Experiments with rats indicate of the inflorescences. Within these groups, cul-
that proso millet protein may be useful as a tivars are mainly distinguished on the basis of
preventive food for certain types of hepatitis. grain colour (varying largely from white, yel-
Proso millet has been found to induce allergic low, brown, red, to almost black) and ecological
reactions in bird keepers. adaptation.
Description Erect annual grass up to 1.2(— Growth and development Proso millet
1.5) m tall, usually free-tillering and tufted, matures in 45-90 days. Emergence of the seed-
with a rather shallow root system; stem cylin- ling is usually in 4-8 days after sowing. During
drical, simple or sparingly branched, glabrous the vegetative phase, which is usually com-
to variously hairy. Leaves alternate, simple; pleted 16—20days after sowing, tillering occurs
leaf sheath variously hairy; ligule membra- and the inflorescence primordia are initiated.
nous, c. 1 mm long, ciliate; blade linear- From then it takes 20-25 days to flowering of
lanceolate, 10—42 cm x 0.5-2.5 cm, variously the main culm, but this period is somewhat
hairy, with 3-6 veins on each side of the mid- shorter at higher temperatures. It is accompa-
rib. Inflorescence a slender panicle 10-30(-45) nied by an increase in leaf area and rapid elon-
cm x 5-15 cm, open or compact, erect or droop- gation of stem internodes. The leaf number on
ing. Spikelets solitary, stalked, ovoid-ellipsoid, the main culm differs among cultivars, but
4-6 mm long, 2-flowered, glabrous; glumes each cultivar produces a fixed number of leaves
unequal, the upper as long as spikelet, many- before flowering. Flowering proceeds from top
veined; lower floret sterile, upper one bisexual to bottom. The flowers are normally self-
with thick broad (c. 2 mm) lemma and palea, 2 fertilized, but cross-fertilization frequently
lodicules, 3 stamens and superior ovary with 2 exceeds 10%. The period from flowering to
plumose stigmas. Fruit a caryopsis (grain), grain maturity has a duration of about 20-30
broadly ovoid, up to 3 mm x 2 mm, smooth, days, and is almost constant among cultivars.
variously coloured but often white, enclosed by At grain maturity the lower part of the inflo-
the persistent lemma and palea and shedding rescence as well as the stem and leaves are still
easily. green. Proso millet follows the Ci-cycle photo-
Other botanical information Panicum is synthetic pathway.
a large genus comprising about 470 species and Ecology Although proso millet is primarily a
is mainly distributed in tropical and subtropi- crop of temperate regions, it has a wide
cal regions, with some species extending to adaptability and can be grown in climates
temperate regions. Panicum miliaceum is a which are too hot and dry, and on soils which
complex species with wild and cultivated types. are too shallow and poor for successful cultiva-
In the literature the following two groups have tion of other cereals. It is cultivated further
been classified as subspecies: subsp. ruderale north than any other millet, the limit being the
(Kitag.) Tzvelev including all spontaneous June isotherm of 17°C and the July isotherm of
types, wild and weedy, and subsp. miliaceum 20°C. Cultivation as a grain crop occurs up to
comprising the cultivated types. The cultivated 3000 m altitude in the Himalayas. It suscepti-
types have sometimes been classified into a ble to frost. Proso millet has one of the lowest
cultivar group: Proso Millet Group. The wild water requirements of all cereals. An average
types have lax panicles, usually jointed annual rainfall of 200-450 mm is sufficient, of
spikelet stalks and narrow lemmas, whereas which 35-40% should fall during the growing
the cultivated ones have either lax or com- period. Most soils are suitable for proso millet,
pressed panicles, spikelet stalks without joints except coarse sand.
and wider lemmas. The true wild type is native Propagation and planting Proso millet is
to central China and is considered to be the propagated by seed. The 1000-seed weight is
ancestor of the cultivated types. In temperate (4.7-)5(-7.2) g. Proso millet seeds germinate
regions of Europe, Asia and the United States, well at temperatures of 10-45°C, with the
however, wild types occur that differ from the highest rate at temperatures between 35°C and
wild type in China and are most probably de- 40°C. The seed is either broadcast or drilled in
rivatives of cultivated types which have re- rows 20-25 cm apart and at a distance of 7.5
gained the ability of natural seed dispersal and cm in the row. This corresponds with a seed
spread as weeds. rate of 8-12 kg/ha. The recommended seed rate
Cultivated proso millet comprises many culti- for furrow planting in Kenya is 4 kg/ha, with a
 PANICUM 125

row distance of 30 cm and 10 cm between hoppers and termites. Birds and rats may de-
plants within the row. The seedbed should be stroy a considerable part ofthe harvest.
moist, firm and free of weeds. For optimal ger- Harvesting Proso millet is ready for harvest
mination seed should be soaked in water for 24 when the seed has a moisture content of 14—
hours and planted no deeper than 4 cm. In 15%. Delayed harvesting should be avoided, as
India the crop is sometimes grown from trans- the seed shatters easily if allowed to become
planted seedlings. Sowing early in the rainy too mature. Premature harvesting, on the
season is less important for proso millet than other hand, results in reduced yield and qual-
for cereals such as sorghum and pearl millet, ity. Plants are usually harvested by pulling
as yield reduction due to late sowing is rela- them up by the roots, and they are threshed
tively small. Proso millet is usually grown as a immediately to avoid grain loss. If proso millet
sole crop, but may be intercropped with other is harvested during the rainy season with high
cereals and with pulses. relative humidity, the grain must be dried to
In-vitro regeneration of proso millet is possible 14% moisture content. Households usually dry
on Murashige and Skoog medium, using ex- the grains over fire.
cised embryos, shoot tips and segments of Yield The average yield of proso millet un-
young inflorescences. der rainfed conditions is 400-700 kg/ha. With
Management The first weeks after sowing sufficient rain and fertile soils or under irriga-
are critical in proso millet cultivation, as initial tion and with application of fertilizers, yields of
growth is slow, thus making competition with over 2 t/ha have been obtained. The milling
weeds difficult. In Kenya the first weeding of recovery is 70-80%.
proso millet is recommended to take place at 2 - Handling after harvest Proso millet grain
3 weeks after emergence of the seedlings, and stores well for up to 5 years. Because of its
the second 2 weeks later. Little is known about small size it is hardly susceptible to insect at-
the fertilizer response of proso millet. In India tack. In India it is stored in granaries with clay
recommended fertilizer rates are 20-40 kg N, walls or clayjars; sometimes the grain is mixed
20 kg P and 0-20 kg K per ha. Proso millet is with ash or slightly baked before storage. It
usually grown as a rainfed crop, but in India it should be stored at 13% moisture content or
is sometimes irrigated. In Russia proso millet less.
is usually grown in rotation with a forage Genetic resources The largest germplasm
grass, wheat or barley. In Bangladesh the rota- collections of Panicum miliaceum are held in
tion may comprise a pulse, wheat, jute, rice, Russia (N.I. Vavilov All-Russian Scientific Re-
potato or a Brassica crop. search Institute of Plant Industry, St. Peters-
Diseases and pests Proso millet is relative- burg, about 9000 accessions), China (Institute
ly little affected by diseases and pests. The most of Crop Germplasm Resources (CAAS), Beijing,
important disease is smut (Sphacelotheca de- about 7500 accessions) and Ukraine (Institute
struens and Ustilago spp.). Control measures of Plant Production, Kharkiv, about 5000 ac-
include seed treatment with fungicide (copper cessions; Ustimovskaya Experimental Station
sulphate) and crop rotation. Other diseases for Plant Cultivation, Ustimovka, about 3500
recorded are anthracnose (Colletotrichum gra- accessions). In tropical Africa, accessions are
minicola), leaf blast (Pyricularia grisea), downy held in Kenya (National Genebank of Kenya,
mildew (Sclerospora graminicola), ergot Crop Plant Genetic Resources Centre, KARI,
(Claviceps spp.), rust (Puccinia and Uromyces Kikuyu). In China, proso millet germplasm is
spp.), leaf blight (Helminthosporium sp.) and being evaluated for resistance to smut, salt
foot rot (Sclerotium rolfsii). The bacterium tolerance and nutritional quality. Of 4200 ac-
Xanthomonas holcicola can cause melanopa- cessions described in China, 53% were non-
thy, a darkening ofthe endosperm. glutinous.
Proso millet can be damaged severely by mag- Breeding In Kenya some selection work
gots of the shootfly (Atherigona miliaceae), with local lines and lines obtained from
which attack the growing point. Infestation ICRISAT has been carried out. The recom-
usually begins in the seedling stage, but may mended cultivar in Kenya is 'KAT/PRO-1', de-
also occur in older plants. Tolerant lines have veloped by Kenya Agricultural Research Insti-
been identified in India. Other pests that are tute (KARI). It was selected from 'N40101', an
sometimes troublesome include stem borers introduction from the former Soviet Union,
(Chilo partellus, Chilo suppressalis and Sesa- received through ICRISAT. 'KAT/PRO-1' was
mia inferens), midges, bugs, army worms, grass- derived from single plants selected for erect
126 CEREALS AND PULSES

tillers, large inflorescences and large grains, lack of information on the crop. Measures re-
and repeated cycles of mass selection of the sulting in less tillering, e.g. narrow distances
progeny for the same traits and high yield po- between rows, give a more uniform crop matur-
tential. This cultivar is self-pollinated, has an ity and will reduce labour requirements for
open inflorescence and cream-coloured grains. bird scaring. The export market for millets in
It has the ability to stop growing when under general will remain small, as millet prices tend
severe drought stress, but it recovers quickly to be too high compared with those of other
and resumes growth when the source of the cereals. Further development of niche markets,
stress is removed. It can be grown up to 2000 e.g. waxy starch for Asian markets, may im-
m altitude, becomes about 80 cm tall, flowers prove the export potential of proso millet.
in 40—50 days, and matures in 65-80 days, Major references Cardenas, Nelson &
depending on altitude and season. The average Neild, 1984; FAO, 1995; Fröman & Persson,
yield of 'KAT/PRO-1' was 1400 kg/ha, which 1974; Hülse, Laing & Pearson, 1980; Ministry
was 50% higher than the mean of the local of Agriculture and Rural Development, 2002;
cultivars. M'Ragwa & Watson, 1994; Penninkhoff, 1984;
Breeding programmes in India and Russia aim Riley et al. (Editors), 1993; Seetharam, Riley &
at a higher productivity (drought resistance Harinarayana, 1990; van der Hoek & Jansen,
and earlier or late maturity), disease resistance 1996b.
(especially to smut) and grain quality (uniform Other references Bajaj, Sidhu & Dubey,
size and shape, yellow endosperm with high 1981; Baltensperger, 1996; Baltensperger,
carotenoid content). The main breeding method 2002; Bohle et a l , 2003; CSIR, 1966; de Wet,
employed in Russia is intraspecific hybridiza- 1995c; Douglas, 1974; Gibberd, 1996; Hanelt &
tion. The floral morphology of proso millet Institute of Plant Genetics and Crop Plant Re-
(small florets with tightly held lemma and search (Editors), 2001; Haq, 1989; ICRISAT &
palea) makes emasculation prior to anthesis FAO, 1996; Kashin et al., 1997; M'ragwa &
and artificial crossing difficult, but techniques Kanyenji, 1987; Nelson, 1984; Nishizawa et al.,
were developed in the United States and a 2002; Petr et al., 2003; Purseglove, 1972;
number of cultivars have been released since Seetharam, 1998; USDA, 2004; Veldkamp,
1984. In the United States breeding efforts 1996b.
include the development of cultivars with Sources of illustration van der Hoek &
higher yield, better harvestability and large Jansen, 1996b.
grain. In addition, germplasm with waxy Authors R.N. Kaume
starch characteristics (used for steam breads in Based on PROSEA 10: Cereals.
South-East Asia) is being developed, to expand
the export potential for proso millet in the
United States. PANICUM TURGIDUM Forssk.
Interspecific crossing of Panicum miliaceum
with some other Panicum species resulted in Protologue Fl. aegypt.-arab.: 18 (1775).
abnormal embryos, which could be rescued by Family Poaceae (Gramineae)
in-vitro ovary culture. Pollen sterility of the Chromosome number 2n = 18, 36, 54
hybrid progeny could partly be overcome by in- Vernacular names Desert grass, turgid
vitro propagation. panic grass (En).
Prospects The production of proso millet is Origin and geographic distribution Pani-
declining and the crop is being replaced in the cum turgidum is distributed from Mauritania
human diet by other cereals, especially rice, and Senegal eastwards through the Sahara
wheat and maize. However, it will continue to and Sahel to Sudan, Ethiopia, Eritrea and So-
be an important staple in semi-arid areas malia, and through northern Africa and west-
where hardly any other cereal can be grown. ern Asia to Pakistan and India.
Proso millet is considered a potentially useful Uses Panicum turgidum is one of the 'kreb'
quick-maturing crop for the drier regions of grasses, a group of grasses occurring in the
tropical Africa, to fill the hunger gap before the Sahel region and collected from the wild for
main cereals are harvested. Major constraints human consumption on a regular basis and
are low returns due to high labour require- especially in times of food shortage. Formerly
ments (mainly for bird scaring, but also for the grains of Panicum turgidum were gathered
planting and weeding) and low yields, limited in large amounts, but nowadays they are rarely
alternative uses, existing eating habits, and a harvested. The grains are mainly made into
 PANICUM 127

porridge. They may also be pounded and eaten dormant buds sprout rapidly after the onset of
without further preparation ('tebik'). the rainy season and the plants stay green over
In Djibouti the young shoots are eaten; they a very extended period, with flowering occur-
are said to be sweet. When green, Panicum ring towards the end of the rainy season and
turgidum is eaten by all livestock; when dry, during the early part of the dry season. The
only by camels and donkeys. The stems are seeds ofPanicum turgidum mature at different
used for thatching and for making mats, bas- times over an extended period, shatter easily
kets and cordage. The Tamachek people in and are often eaten by birds. Panicum tur-
Niger weave the straw as the weft, with thin gidum follows the C4-cycle photosynthetic path-
threads ofleather as the warp, into mats which way.
can only be rolled one way. Mats of Panicum Ecology Panicum turgidum is extremely
turgidum have been used as funeral shrouds. drought tolerant, growing in regions with an
In the Sahara the stems are used as fuel. Plant annual rainfall of 200-250 mm or sometimes
ash is mixed with tobacco for chewing and in even less. It occurs up to 3200 m altitude, in
southern Algeria the powder of ground stems is sandy deserts and semi-deserts, on dunes and
used as a wound-dressing. In Mauritania the seashores, and in sandy pockets in rocky out-
grains are credited with antidiabetic proper- crops. Panicum turgidum is an important plant
ties. Panicum turgidum has occasionally been of the Sahara and Arabian deserts, catching
used for dune fixation in arid areas. sand and forming hummocks, sometimes in
Properties The palatibility of the leaves of nearly pure stands. In Sudan it is dominant on
Panicum turgidum is low, but sufficient for grounds where locusts lay their eggs, and it
camels and donkeys, and, when young, for serves as food for young locusts. Panicum tur-
sheep and goats. Herdsmen in Niger say that gidum tolerates saline soils.
milk becomes foul-smelling 2-3 days after cows Management Panicum turgidum is not cul-
have grazed Panicum turgidum. The grains tivated as a cereal, but collected from the wild.
thresh free from the glumes but remain cov- It is sometimes protected from grazing until
ered by the tough lemma and palea. after seed harvesting, e.g. in southern Algeria
Botany Much-branched, glaucous, perennial and northern Mali. The panicles may be beaten
grass, forming rounded tussocks up to 1.5(-2) with a stick to obtain the grains. In Niger the
m tall and wide, with a thick rootstock and a panicles are rubbed between the hands. Pani-
fibrous root system up to 2 m deep and later- cum turgidum can be propagated by seed or by
ally spreading for up to 3.5 m; stem (culm) rootstock cuttings. Seeds do not germinate be-
erect or ascending, woody, rooting at the nodes. low 15°C and must be sown superficially. Ger-
Leaves alternate, simple and entire; blade lin- mination is best at 25—35°C. Transplanting of
ear-lanceolate, up to 20(-30) cm x 7 mm, often seedlings is possible.
much shorter than the sheath, flat, folded or Genetic resources and breeding A collec-
inrolled, stiff and pungent. Inflorescence a tion of 42 accessions of Panicum turgidum is
moderately branched, pyramidal panicle 2.5- held at the International Livestock Research
15(-30) cm x 5—9 cm, lax, primary branches Institute (ILRI), Addis Ababa, Ethiopia. High
distant, eventually spreading. Spikelet ovoid, grain-yielding types are particularly found in
3.5-4.5(—5) mm long, acute or acuminate, swol- the Middle East. Because of its wide distribu-
len, glabrous, often gaping at anthesis, 2- tion and abundance Panicum turgidum is not
flowered; glumes broadly ovate, acute to acu- threatened by genetic erosion.
minate, lower glume slightly shorter than the Prospects Panicum turgidum has value as
spikelet, 5—9-veined, upper glume 7-9-veined; a very drought-resistant grass suitable for
lower floret male, lemma 9-11-veined, palea sand-binding and for providing food and fodder.
well-developed, upper floret bisexual, lemma It is recommended to sample the existing
pale or yellowish, smooth, glossy; stamens 3; variation and to use the collected germplasm in
ovary superior, stigmas 2. Fruit a caryopsis a breeding programme aimed at developing
(grain) c. 2 mm long, reddish. superior cultivars. More information is needed
Panicum comprises about 470 species and is on the nutritional characteristics of the grain.
mainly distributed in tropical and subtropical Major references Burkill, 1994; Harlan,
regions, with some species extending to tem- 1989b; Kernick, 1992; Phillips, 1995; Williams
perate regions. & Farias, 1972.
Natural reproduction of Panicum turgidum is Other references Ahmad et al., 1994; Bog-
mainly vegetatively by stolons. In dry areas the dan, 1977; Clayton, 1972; Cope, 1995;Hanelt &
128 CEREALS AND PULSES

Institute of Plant Genetics and Crop Plant Re- it is consumed as a stiff porridge ('tô') or gruel
search (Editors), 2001; Haroun, 2000; Kernick, in Africa, or as flat unleavened bread ('cha-
1978; Kiambi, 1999; Naegele, 1977; National patti') in India. In Africa there are various
Research Council, 1996. other preparations such as couscous, rice-like
Authors M. Brink products, snacks of blends with pulses, and
fermented and non-fermented beverages. In
several Indian preparations parched seeds are
P E N N I S E T U M GLAUCUM (L.) R.Br. used. The stems are widely applied for fencing,
thatching and building, as fuel and as a poor-
Protologue Prodr.: 195 (1810). quality fodder. Split stems are used for bas-
Family Poaceae (Gramineae) ketry. A dye for leather and wood is obtained
Chromosome number 2n = 14 from red- and purple-flowered types. In African
Synonyms Pennisetum spicatum (L.) Körn. traditional medicine the grain has been applied
(1885), Pennisetum americanum (L.) Leeke to treat chest disorders, leprosy, blennorrhoea
(1907), Pennisetum typhoides (Burm.f.) Stapf & and poisonings, and the ground grain as an
C.E.Hubb. (1933). anthelmintic for children. A root decoction is
Vernacular names Pearl millet, bulrush drunk to treat jaundice; the vapour of inflores-
millet, cattail millet (En). Mil, mil à chandelle, cence extracts is inhaled for respiratory dis-
mil pénicillaire, petit mil (Fr). Milho zaburro, eases in children. In some areas the grains are
milho preto, milheto, massango liso (Po). used in rituals. Outside Africa and India pearl
Mwele (Sw). millet is mostly grown as a green fodder crop
Origin and geographic distribution Pearl for silage, hay making and grazing. Following
millet was domesticated in the Sahel 4000- the discovery that pearl millet can suppress
5000 years ago from Pennisetum violaceum root-lesion nematodes (Pratylenchus penetrans)
(Lam.) Rich. It spread to East Africa and from it is increasingly being used as an alternative
there to southern Africa, and, about 3000 years to soil fumigation in tobacco and potato crop-
ago, to the Indian subcontinent. It reached ping in Canada.
tropical America in the 18 th century and the Production and international trade Pro-
United States in the 19th century. Pearl millet duction statistics often combine data on all
is commonly grown as a grain crop in the millet species. Estimates based on total millet
semi-arid regions of West Africa and the driest production (FAO statistics) and relative impor-
parts of East and southern Africa and the In- tance of pearl millet in different countries indi-
dian subcontinent. It is also grown as a fodder cate an annual grain production of about 18
crop, e.g. in Brazil, the United States, South million t from a planted area of 26.5 million ha
Africa, and Australia. mostly in the dry regions ofAfrica (60%of area
Uses Pearl millet is the staple food for over and 58% of production) and the Indian subcon-
100 million people in parts of tropical Africa tinent (38% of area and 41% of production).
and India. Decorticated and pounded into flour Production statistics over the past 10 years
show a 20% increase in area planted in Africa,
with a 12%>increase in yield. Most of the area
increase is in Burkina Faso, Chad, Mali, Niger
and Nigeria, but yield levels increased only in
the latter two countries. During the same pe-
riod, the area planted to pearl millet in India
declined by 16%, but yield levels increased by
30%. Negligible quantities are traded interna-
tionally.
Properties Whole dried grain of pearl millet
contains per 100 g edible portion: water 12.0 g,
energy 1428 k j (341 kcal), protein 10.4 g, fat
4.0 g, carbohydrate 71.6 g, fibre 1.9 g, Ca 22
mg, P 286 mg, Fe 20.7 mg, ß-carotene traces,
thiamin 0.30 mg, riboflavin 0.22 mg, niacin 1.7
mg and ascorbic acid 3 mg (Leung, Busson &
Jardin, 1968). The content of essential amino
Pennisetum glaucum -planted acids per 100 g food is: tryptophan 189 mg,
 PENNISETUM 129

lysine 332 mg, methionine 239 mg, phenyla- obovate, 3-7 mm long, usually 2-flowered;
lanine 467 mg, threonine 374 mg, valine 535 glumes 2, lower one c. 1 mm long, upper one c.
mg, leucine 927 mg and isoleucine 397 mg 2.5 mm; lower floret male or sterile, upper one
(FAO, 1970). From a nutritional viewpoint bisexual; lemma ovate, pubescent on margins;
pearl millet is better than maize and sorghum. palea almost as long as lemma; lodicules ab-
Compared to that of other millets, the protein sent; stamens 3, anthers 2-5 mm long, tipped
ofpearl millet is rich in tryptophan. with brush-like bristles; ovary superior, obo-
Description Robust annual grass up to 4 m void, smooth, with 2 hairy stigmas, connate at
tall, with basal and nodal tillering; root system the base. Fruit a free-threshing caryopsis (grain),
extremely profuse, sometimes the nodes at globose to cylindrical or conical, 2.5-6.5 mm
ground level produce thick, strong prop roots; long, variously coloured, from white, pearl-col-
stem slender, 1-3 cm in diameter, solid, often oured or yellow to grey-blue or brown, occa-
densely villous below the panicle, nodes promi- sionally purple, hilum marked by a distinct
nent. Leaves alternate, simple; leaf sheath black dot at maturity.
often hairy; ligule short, membranous, with a Other botanical information Pennisetum
fringe of hairs; blade linear to linear- comprises about 80 species and occurs
lanceolate, up to 1.5 m x 8 cm, often pubescent, throughout the tropics. Pearl millet is not
margins minutely toothed, somewhat rough. closely related to most other Pennisetum spe-
Inflorescence a cylindrical, contracted, stiff and cies, although it hybridizes easily with ele-
compact panicle, suggesting a spike, up to 200 phant grass (Pennisetum purpureum Schu-
cm long; rachis cylindrical, bearing densely mach., a tetraploid with In = 28). Pennisetum
packed clusters of 1—5(—9)spikelets, subtended glaucum belongs to a complex of 3 taxa that
by a tuft (involucre) of up to 90 bristles about hybridize freely and are sometimes considered
as long as spikelets, but in some cultivars with as subspecies of Pennisetum glaucum. How-
a few stiff bristles up to 2 cm long. Spikelets ever, as long as the complicated taxonomy of
pearl millet has not been fully cleared up, it is
preferable to keep these taxa separate:
- Pennisetum glaucum: cultivated types, with
persistent, stiped involucres; the dense inflo-
rescences and non-shattering habit are con-
spicuous.
- Pennisetum sieberianum (Schltdl.) Stapf &
C.E.Hubb. (synonyms: Pennisetum steno-
stachyum (A.Braun & Bouché) Stapf & C E .
Hubb., Pennisetum dalzielii Stapf & CE.
Hubb., Pennisetum americanum (L.) Leeke
subsp. stenostachyum (A.Braun & Bouché)
Brunken): weedy types, resulting from intro-
gression between wild Pennisetum violaceum
and cultivated Pennisetum glaucum and
ranging in morphology between wild and cul-
tivated types; in the latter case they are
termed 'shibras' and they look much like
Pennisetum glaucum cultivars, but differ in
having deciduous shortly stiped involucres,
and spikelets which shatter before harvest;
bristles numerous, longer than spikelets;
widespread in the Sahel and also found,
though less frequently, in East and southern
Africa.
- Pennisetum violaceum (Lam.) Rich, ex Pers.
(synonyms: Pennisetum fallax (Fig. & De
Pennisetum glaucum - 1, plant habit; 2,part of Not.) Stapf & C.E.Hubb., Pennisetum ameri-
infructescence; 3, pair of spikelets surrounded canum (L.) Leeke subsp. monodii (Maire)
by involucre of bristles. Brunken): wild, variable type, with decidu-
Source: PROSEA ous sessile involucres which always contain a
130 CEREALS AND PULSES

single spikelet; bristles numerous, longer days, but mostly from 75-180 days. Time to
than spikelets; distributed from the West Af- flower initiation isthe main factor determining
rican Sahel region to Eritrea in very drylo- the life cycle of a cultivar. Floral initiation is
cations, independent of farming; sometimes weakly to strongly controlled by photoperiod,
harvested as a wild cereal in times of scar- with short days accelerating flower initiation.
city. Photoperiod response allows crop cycle length
Although many intermediate cultivars occur, 4 to be adjusted by time of planting, needed
cultivar groups (originally described as races) when rains begin late, to ensure that flowering
can be distinguished in cultivated Pennisetum and grain production occur at the same opti-
glaucum, based mainly on grain shape and mum time each year for a specific latitude. In
partly on distribution: short-duration, photoperiod-insensitive culti-
- Typhoides Group; grain obovoid, circular in vars the developmental stages (from germina-
cross-section, 2.5-5.5 mm x 1.5—3 mm x1— tion to flower initiation, to flowering and to
2.5 mm, inflorescence cylindrical or ellipsoi- maturity) areofapproximately equal duration.
dal, usually less than 0.5 m long; it is the Field establishment of pearl millet is affected
most primitive, the most variable and most by its relatively small seed size, especially in
widely distributed group, occurring all over crusting soils. Other factors that influence
the pearl millet range in Africa and India, stand establishment include high soil surface
and is probably ancestral for the other temperatures at emergence (as high as 50—
groups. 55°C), sand storms and early season moisture
- Globosum Group: grain globose, more than stress. During early development the roots
2.5 mm in diameter, inflorescence cylindri- grow more than the above-ground parts. Pearl
cal, often longer than 1 m; most common in millet produces an extensive and dense root
the Sahel region west of Nigeria. system, which may reach a depth of 1.2—1.6m,
- Leonis Group: grain oblanceolate in outline, exceptionally 3.5 m. Basal tillering occurs 2-6
circular in cross-section, 4-6.5 mm x 2-2.5 weeks after sowing, and when planted in
mm x 2-2.5 mm, apex acute, inflorescence widely spaced pockets up to 40 tillers may be
cylindrical; this is the smallest group and is produced, especially on long-season cultivars.
grown in Mauritania, Senegal and Sierra Secondary tillering from the upper nodes of
Leone. stems is a common response to drought, or to
- Nigritarum Group; grain obovoid but angu- damage to the stem or inflorescence. These
lar in cross-section, 3-5 mm x 1.5-2.5 mm x aerial tillers produce 2-3 leaves and a small
1.5-2 mm, inflorescence cylindrical; most inflorescence within 10-20 days; they maycon-
common in semi-arid regions from Nigeria to tribute 15%andoccasionally upto 50%of grain
Sudan. yield. It takes 15—20 days from inflorescence
Agronomically twomain groups ofcultivars are differentiation to flowering. Pearl millet has a
recognized in West Africa, based on growth protogynous breeding system, which encour-
duration : short-duration Gero (or Souna) cul- ages but does not enforce cross-pollination;
tivars and long-duration Maiwa (or Sanio) cul- 10% or more inbreeding may occur, depending
tivars. Gero cultivars are less photoperiod- on overlap in flowering between florets within
sensitive, are more widely grown and exhibit an inflorescence and among tillers. Pearl millet
more genetic diversity than Maiwa cultivars, in cultivars are therefore heterogeneous and het-
which flowering date is strongly controlled by erozygous random mating populations, which
daylength. Maiwa types are grown in regions exhibit substantial inbreeding depression.
where the rainy season is longer and where Heavy rainfall, low temperature and moisture
sorghum is the major cereal, but on poorer, stress reduce seed set. The grain-filling period
more drought-prone soils. Certain Maiwa mil- normally takes 22-25 days. The harvest index
lets are transplanted from nurseries into the of landraces is low (0.15-0.20), attaining 0.35
field andareknown as Dauro millet. in improved cultivars, and up to 0.45 in dwarf
The improved cultivars and dwarf single-cross hybrids.
hybrids grown in India are stronger tillering, Pearl millet is characterized by the C4 photo-
early (80days) tovery early (65days) maturing synthetic pathway. Vesicular-arbuscular my-
and less photoperiod-sensitive than African corrhizae (e.g.Gigaspora and Glomus spp.)and
cultivars. nitrogen-fixing bacteria (e.g. Azospirillium
Growth and development Pearl millet spp.) are commonly found associated with pearl
cultivars vary in time to maturity from 55—280 millet roots, which may assist with the uptake
 PENNISETUM 131

ofwater, N and P. for weeding. Pearl millet is highly responsive

Ecology In West Africa, from the oases of to increased soil fertility, but under traditional
the Sahara desert (under irrigation) to the rainfed farming conditions the application of
northern Sahel (characterized by 250 mm an- manure and chemical fertilizers is limited.
nual rainfall), pearl millet cultivars are grown Because of the depleted fertility status of most
that are photoperiod insensitive and mature in pearl millet soils some phosphorus and potas-
55-65 days. In the 250-400 mm rainfall zone, sium is needed for an optimal response to ni-
where very high temperatures are common, trogenous fertilizers. As fertilizers increase
especially at planting time, it is the dominant water use of the crop, plant populations and
cereal. The optimum temperature for germina- average seasonal water availability also need
tion of pearl millet seeds is 33-35°C; no germi- to be considered when making fertilizer rec-
nation occurs below 12CC. The optimum tem- ommendations. A pearl millet crop yielding
perature for tiller production and development about 3.1 t grain per ha in the West African
is 21-24°C, and for spikelet initiation and de- savanna was recorded as removing 132 kg N,
velopment about 25°C. Extreme high tempera- 28 kg P, 65 kg K and 31 kg Ca per ha from the
tures before anthesis reduce pollen viability, soil.
panicle size and spikelet density, thus reducing Diseases and pests Green ear caused by
yield. Pearl millet is tolerant of various soil downy mildew (Sclerospora graminicola), grain
conditions, especially of light and acid soils. Its smut (Tolyposporium penicillariae), rust (Puc-
large and dense root system allows it to grow cinia substriata var. penicillariae) and ergot
on soils with a low nutrient status. Pearl millet (Claviceps fusiformis) are important diseases of
does not tolerate waterlogging. Once estab- pearl millet, both in Africa and in Asia. Sources
lished, the crop is fairly tolerant of salinity. of resistance against all four have been identi-
Propagation and planting Propagation of fied and are being incorporated into new culti-
pearl millet is by seed, usually sown directly in vars, except for resistance to ergot which is
the field. Transplanting is carried out on a very polygenic and recessively inherited. Birds are
limited scale in India and West Africa (Dauro the major pest in pearl millet, especially
millet). The 1000-seed weight is 2.5-16 g. In Quelea spp. Bird scaring for several weeks be-
Africa short-duration cultivars are sown early, fore the harvest is essential. Farmers in West
after the first 20 mm rain of the season and Africa often do not harvest a larger area than
land preparation is limited to a light hoeing. they can protect from birds. Cultivars with
Land preparation for long-duration cultivars, long, hard bristles are less vulnerable than
which are sown later, is done more thoroughly. those without. Stem borer (Coniesta igne-
Pearl millet is usually sown directly in pockets fusalis), millet head miner (Heliocheilus al-
(hills) in rows at plant distances of 45 cm x 45 bipunctella) and millet midge (Geromyia penni-
cm to 200 cm x 200 cm depending on the crop- seti) are locally important. Other pests are
ping system (intercrop or sole crop). The pock- white grubs, grasshoppers, locusts, and various
ets are opened with a hoe or a stick, a pinch of Lepidoptera. Pearl millet is sometimes seri-
seeds is thrown in, and the hole is covered by ously attacked by adapted strains of the root
soil using the foot. At the first weeding the crop parasite Striga hermonthica (Delile) Benth. in
is thinned to 2 or 3 plants per pocket. Farmers West Africa.
tend to adjust plant density based on average Harvesting Pearl millet is harvested by
rainfall and soil fertility; it generally ranges hand, either by picking the panicles or by har-
from 20,000-50,000 plants per ha in pure vesting whole plants. In cultivars where tillers
stands. Seed rates vary accordingly from 2-5 ripen unevenly, several pickings are required.
kg per ha. Pearl millet is often intercropped Cultivars with long panicles are favoured for
with one to several crops, including cowpea, ease ofharvest, bundling and transport.
sorghum and groundnut. Yield Grain yields range from 250 kg/ha in
Management Pearl millet frequently needs the driest areas to 500-1500 kg/ha in the main
2—3weedings, which are done mostly by hand. production areas. Average yields in Africa and
With short-duration cultivars in Africa, weed- India are about 670 and 790 kg/ha, respec-
ing coincides with land preparation and plant- tively. Under optimal conditions hybrids may
ing of later crops. Manual weeding places se- reach grain yields of 5 t per ha in 85 days and
vere demands on available labour and limits yields of 8 t per ha have even been obtained. In
the area that can be managed properly. In a landraces the above-ground dry matter yield
few regions animal-drawn implements are used may be 3—10 t/ha. In hybrids bred specifically
132 CEREALS AND PULSES

for forage dry matter yields in a season range farmers. The discovery of cytoplasmic male
from 10-20 t/ha. sterility in 1958 in the forage breeding pro-
Handling after harvest A harvested pearl gramme at Tifton (Georgia, United States) led
millet crop is dried in the sun for a few days. In to the production of early-maturing, semi-dwarf
Africa whole panicles are commonly stored in grain hybrids in India, which covered 3 million
elevated granaries, built of mud or plant mate- ha by 1970. Despite the occurrence of disease
rials and covered with thatch. Sometimes they epidemics, some 50% of the Indian pearl millet
are stored in pits. Ash or neem (Azadirachta crop on family farms now consists of hybrids
indica A.Juss.) leaves may be put in layers to and improved cultivars, and yields have in-
reduce insect attack. Threshing is normally creased by 40% since 1965. Early breeding
done manually when grain is needed. If dry work in West Africa by the Institut de Recher-
and protected from insects, seed can be stored ches Agronomiques Tropicales et de Cultures
adequately at room temperature for several Vivrières (IRAT) and the East African Agricul-
years. Pearl millet flour, unless dry-milled and ture and Forestry Research Organization
well-packed, has poor storage quality because (EAAFRO) produced improved local cultivars,
ofrancidity due to the high oil content. but adoption was negligible. Since the early
Genetic resources Landraces of pearl mil- 1980s ICRISAT, working with a broader germ-
let have evolved over thousands of years plasm base and in conjunction with national
through natural and human selection. Selec- agricultural research programmes in West,
tion at different latitudes and in different agro- East and southern Africa, has produced better
climatic zones for crop duration, yield, adapta- cultivars. In 8 countries in West Africa 24 cul-
bility to nutrient-poor soils, resistance to tivars (11 for the Sahel zone and 13 for the
drought and diseases, and grain type has re- Sudan Zone) have been released and in East
sulted in local cultivars with a large range of and southern Africa 19 new cultivars are
morphological diversity and photoperiod sensi- available for 8 countries with adoption rates of
tivity. Continual introgression with wild and up to 50% (Namibia, Zimbabwe). Work at
weedy relatives in West Africa has further con- ICRISAT is focused on the identification of
tributed to the crop's genetic diversity. Genetic stable stress tolerance, wide adaptability and
variation is conserved and evaluated at the high yield potential. Sources of tolerance of the
Coastal Plains Experiment Station, Tifton major diseases have been identified and are
(Georgia, United States) and the International being incorporated into new cultivars. Two
Crops Research Institute for the Semi-Arid further cytoplasmic male sterility systems (A4
Tropics (ICRISAT) in Hyderabad (India), and A5), which have superior attributes both
where the world collection of over 24,000 en- for hybrid breeding and seed production, have
tries is housed. The International Plant Ge- been discovered. These allow access to different
netic Resources Institute (IPGRI) supports a heterotic patterns and more rapid development
programme, started in Burkina Faso, to im- of new hybrids, particularly topcross hybrids
prove the description and evaluation of mate- which are most suited to the higher disease
rial at the time and location of collection. One pressures in Africa. On average, single-cross
particular germplasm source, the Tniadi' culti- hybrids (male-sterile x inbred restorer) give
var from northern Togo and Ghana, has had a about 20% more grain yield than open-polli-
profound effect on pearl millet breeding. Selec- nated cultivars of similar maturity. Topcross
tions from it have been successful as cultivars hybrids (male-sterile x open-pollinated culti-
in northern India, Namibia and Botswana, and vars) benefit from the adaptability and durable
it has been extensively used in all breeding disease resistance of the open-pollinated type,
programmes. and give 10—15% more yield. The pearl millet
Breeding Both improved open-pollinated genome has now been well mapped, and
types and hybrids have been developed in pearl marker-assisted selection is being used to im-
millet ; cytoplasmic male sterility is available prove downy mildew and rust resistance.
for the commercial production of hybrid seed. Transgenic pearl millet plants with various
Cultivar breeding of pearl millet started in marker genes have been obtained using parti-
Africa in the 1950s and traditional cultivars cle bombardment.
are still widely used. Breeding work by the Forage breeding work, mainly in the United
Indian Council of Agricultural Research and States, Australia, Brazil and southern Africa,
ICRISAT has been most successful in develop- has also produced pearl millet hybrids. Inter-
ing cultivars that were rapidly adopted by specific hybrids between pearl millet and ele-
 PHASEOLUS 133

phant grass (Pennisetum purpureum) are substantially on existing land.

available; these are vigorous, variable, triploid Major references Anand Kumar & An-
and sterile, but selections are easily vegeta- drews, 1993; Andrews & Bramel-Cox, 1993;
tively propagated and used as a persistent per- Andrews & Kumar, 1992; Bidinger & Hash,
ennial forage by small farmers in South-East 2004; Brunken, de Wet & Harlan, 1977; Dendy
Asia, East and southern Africa and South (Editor), 1995; Khairwal et al., 1999; Oyen &
America. In the United States dwarf plant Andrews, 1996; Pearson (Editor), 1985; Renard
stature and synchronous maturity of tillers, in &Anand Kumar, 2001.
addition to earliness and tolerance of diseases Other references Andrews & Anand Ku-
have been successfully incorporated into pearl mar, 1996; Bationo et al., 1992; Bezançon,
millet to convert it to a new feed grain crop Renno &Anand Kumar, 1997;Bonamigo, 1999;
suitable for mechanical cultivation. Commer- Burkill, 1994; Clayton, 1989; Clayton &Renvo-
cial production of grain pearl millet for poultry ize, 1982; de Wet, 1995d; FAO, 1970; 2001;
feed has commenced in Georgia (United Hash, Schaffert & Peacock, 2002; Jagdale et
States). al., 2000; Leung, Busson & Jardin, 1968;
Prospects Pearl millet has great potential McDonough, Rooney & Serna-Saldivar, 2000;
because it has one of the highest rates of dry Monyo, 2002; National Research Council, 1996;
matter gain among the Ci-cycle cereals, a very O'Kennedy, Burger & Botha, 2004; Rachie &
flexible breeding system and a large amount of Majmudar, 1980; Rai et al., 2001; Stoop, 1986;
genetic variability in the primary gene pool yet Wilson, 2000.
to be used. The results of pearl millet breeding Sources of illustration Oyen & Andrews,
work in India, where gains from breeding have 1996.
been 1-2% per year over the last 35 years, Authors D.J. Andrews &K.A. Kumar
demonstrate what can be achieved in Africa. Based on PROSEA 10:Cereals.
Half of the millet area in India is now planted
to improved cultivars, including hybrids.
Proven breeding techniques and wider use of PHASEOLUSACUTIFOLIUS A.Gray
genetic resources will continue to produce bet-
ter cultivars. Protologue PI. wright. 1:43 (1852).
In Africa small and seasonably variable grain Family Papilionaceae (Leguminosae - Papil-
markets, lack of credit and bulk grain storage ionaceae, Fabaceae)
have constrained farmers from making the Chromosome number 2n = 22
monetary investments which will increase pro- Vernacular names Tepary bean, Texas
duction. However, the growing urban demand bean (En). Haricot tépari, tépari (Fr). Feijäo
for pearl millet flour, and food products like tepari (Po).
couscous allow farmer cooperatives in Niger, Origin and geographic distribution Tepary
Mali and Senegal to make contracts before bean is an ancient crop of the south-western
planting directly with urban grain processors, United States and northern Mexico. Recent
for the supply of grain meeting given stan-
dards, including grain type and colour, milling
quality, flavour and freedom from impurities.
In Senegal the cultivar is also specified. This
enables subsistence farmers to benefit from
seed of improved cultivars, and from purchased
inputs such as fertilizer. Better cultivars, pro-
duced by certified seed farmers, are being mar-
keted in Namibia, Nigeria and Senegal. Many
production technologies have been researched
and tested both for family farmers and larger-
scale pearl millet cultivation, including im-
proved cultivars, better crop management, soil
improvement and moisture conservation tech-
niques for the major pearl millet producing
regions in Africa. If reliable grain markets are
established, pearl millet production in Africa
can follow the example of India, and increase Phaseolus acutifolius -planted
134 CEREALS AND PULSES

studies indicate that the earliest remains of consumption, and no production and trade sta-
domesticated tepary bean, found in Tehuacân tistics are available.
Valley, Mexico, date from around 2300 years Properties Per 100 g edible portion the
ago. Isozyme analysis suggests that domestica- composition of dry tepary bean seeds is: water
tion took place in a single geographic region, 8.6 g, energy 1478 kJ (353 kcal), protein 19.3 g,
with the Mexican states of Jalisco and Sinaloa fat 1.2 g, carbohydrate 67.8 g, fibre 4.8 g, Ca
being potential candidates. Today, wild types 112 mg, P 310 mg, thiamin 0.33 mg, riboflavin
are distributed from the south-western United 0.12 mg, niacin 2.8 mg and ascorbic acid 0 mg
States (Arizona, New Mexico, Texas) to Gua- (Leung, Busson & Jardin, 1968). As with other
temala, with the core of their distribution in pulses, the seeds are low in the sulphur-
north-western Mexico. Tepary bean is also cul- containing amino acids methionine and cystine
tivated in the southern United States and Cen- (0.9-1.3% and 0.8-1.3% of total amino acids,
tral America. Cultivation of tepary bean de- respectively). With respect to antinutritional
creased strongly after World War II, but nowa- factors, such as trypsin inhibitors, flatulent
days the crop is regaining interest. Tepary oligosaccharides and phytic acid, tepary bean is
bean has been introduced and is cultivated in similar to cowpea and chickpea; the lectin ac-
Africa, Asia and Australia. It was introduced tivity is exceptionally high, but is readily re-
into francophone West Africa, Central Africa, duced by cooking, whereas cyanogenic gluco-
East Africa and Madagascar between the first sides have not been detected. Consumption of
and second World Wars, and is now grown raw tepary bean flour has been recorded to
there and as far south as Botswana. Tepary cause death in mice and rats within 3—4 days,
bean is also recorded as being grown in Mo- but soaking and cooking the seeds eliminated
rocco, Algeria, South Africa, Swaziland and toxicity completely. Tepary beans have a strong
Lesotho. flavour and odour and are less palatable than
Uses Tepary bean is mainly grown for its common beans. On storage the dry seeds be-
mature dry seeds, which are eaten after boil- come very hard and take a long time to cook.
ing, steaming, frying or baking. They are used Cultivars with white seeds have a more per-
in stews and soups, and mixed with whole- meable seedcoat than cultivars with black
grain maize. In Uganda the dry seeds are usu- seeds, resulting in a shorter cooking time.
ally boiled and then coarsely ground before Tepary bean hay contains 6.6% water, 9.9%
being added to soup. Occasionally it is eaten as protein, 1.9% fat, 43.1% N-free extract, 29.3%
a green bean or as bean sprouts. The leaves are fibre, and 9.2% ash. Pods and stems contain 8%
considered edible in Malawi, but are tougher water, 4.1% proteins, 0.5% fat, 43.6% N-free
than those of common bean (Phaseolus vulgaris extract, 37.0%fibre and 6.8% ash.
L.) and take longer to cook. Pods and stems Description Climbing, trailing or more or
remaining after removing the seed may be used less erect and bushy annual herb, with stems
for animal feed. In Botswana the seeds are a up to 4 m long; roots fibrous. Leaves alternate,
common supplementary feed for chickens. Te- 3-foliolate; stipules lanceolate, 2-3 mm long,
pary bean has occasionally been grown for fod- appressed to stem; petiole 2-10 cm long; stipels
der or green manure, e.g. in the United States. linear, up to 2 mm long; leaflets ovate to ovate-
It may be used as a cover crop and an intercrop lanceolate, 4-8 cm x 2-5 cm, acute, usually
in agroforestry systems. pubescent below. Inflorescence an axillary ra-
Production and international trade Tepary ceme, 2-5 flowered. Flowers bisexual, papil-
bean is mainly grown in Mexico and Arizona ionaceous; pedicel 3—7 mm long; calyx cam-
(United States). Large-scale commercial pro- panulate, 3—4 mm long, the upper 2 lobes
duction was tried in the early 1900s, but efforts united into one, the lower 3 triangular; corolla
were abandoned due to its unfavourable mor- white, pink or pale lilac, standard half-
phological characteristics compared to the reflexed, broad, emarginate, up to 1 cm long,
common bean, changes in eating habits and wings up to 1.5 cm long, keel narrow, coiled;
lack of information on its performance. Tepary stamens 10, 9 fused and 1free; ovary superior,
bean has recently gained importance in semi- c. 0.5 cm long, densely pubescent, style with a
arid parts of tropical Africa, e.g. in Sudan, thickened terminal coil, with collar of hairs
north-eastern Kenya, Uganda and Botswana, below the stigma. Fruit a compressed pod,
where most other grain legumes fail due to straight or slightly curved, 5—9cm x 0.5—1 cm,
drought and where short-duration crops are rimmed on margins, with short but distinct
often needed. Production is mainly for domestic beak, hairy when young, 2-9-seeded. Seeds
 PHASEOLUS 135

within 27—40 days. Self-pollination occurs be-

fore anthesis. In the tropics, short-duration
types may mature within 2 months, but most
types have a growth period of 70-90 days. In
cooler regions, such as coastal Algeria, the
growth period averages 120 days. The seeds of
many domesticated types of tepary bean are
shattered less easily than those ofwild types.
Tepary bean shows effective nodulation and
nitrogen fixation only with Bradyrhizobium
isolates. Hybrids of tepary bean and common
bean (the latter nodulating with Bradyrhizo-
bium but only fixing atmospheric nitrogen
when nodulated with Rhizobium leguminosa-
rum bv. phaseoli) can be divided in 2 groups:
one group only fixing nitrogen with Brady-
rhizobium, the other only with Rhizobium
leguminosarum bv. phaseoli.
Ecology Tepary bean is particularly suited
to arid regions as it is tolerant of drought, heat
and a dry atmosphere. Factors contributing to
the drought tolerance of tepary bean are sensi-
tive stomata, closing already at relatively high
water potentials, and a deep and extensive root
system. Tepary bean is found in regions with a
Phaseolus acutifolius - 1, flowering branch mean annual temperature of 17-26°C; the
with young fruit; 2,fruits; 3, seeds. minimum night temperature should not drop
Source: PROSEA below 8°C. It can grow in areas with an annual
rainfall ranging from 500-1700 mm, but where
globose to oblong, 4-7(-10) mm x 2-5(-7.5) annual rainfall exceeds 1000 mm, vegetative
mm, white, yellow, brown, purple, black or growth is usually excessive, at the expense of
variously speckled, dull. Seedling with epigeal seed yield. After flowering little or no rain is
germination; first pair ofleaves simple. needed. In most ofAfrica, tepary bean is grown
Other botanical information Phaseolus com- as a short-season crop, but in the more humid
prises about 50 species, most of them in the parts it is grown year-round. In Mexico and Ari-
Americas. Three varieties of Phaseolus acuti- zona, it is usually grown at medium altitudes.
folius have been distinguished, based on the Some tepary bean types require short days for
shape of leaflet and seeds. Var. acutifolius and flowering, but others seem day-neutral.
var. tenuifolius A.Gray comprise wild types Light, well-drained soils are preferred; reason-
from south-western United States and north- able yields can be obtained on poor sandy soils
western Mexico, whereas var. latifolius G.F. with pH 5-7. Tepary bean does not tolerate
Freeman comprises wild and cultivated types. waterlogging, and heavy clays are unsuitable.
Isozyme and AFPL analyses have shown no It is moderately tolerant of saline and alkaline
clear-cut differentiation between var. acuti- soils. The salt tolerance may not be physiologi-
folius and var. tenuifolius. cal but result from its ability to escape salinity
Growth and development Tepary bean due to its root system going deeper than that of
seeds absorb water easily; in moist soils the e.g. common bean.
testa wrinkles within 5 minutes, in water in 3 Propagation and planting Tepary bean is
minutes. This leads to quick germination. propagated by seed. The 1000-seed weight is
Seedling emergence is faster in white-seeded 100-220 g for cultivated and 15-50 g for wild
than in dark-seeded types. The seeds of domes- genotypes. Seeds are broadcast at a rate of 2 8 -
ticated types have no dormancy, which is a 34 kg/ha, or drilled in rows 60-90 cm apart
disadvantage in humid regions, where fallen with 10-45 cm between plants within the row.
seeds will germinate rapidly. The rate of ger- The sowing depth is 2.5-10 cm. Tepary bean is
mination increases with increasing tempera- sometimes sown on mounds, with 2-4 seeds
tures from 10°C to 35°C. Flowering occurs per mound. In Kenya tepary bean is sown at
136 CEREALS AND PULSES

15-20 kg/ha in a spacing of 60 cm x 30 cm. fabae) and the lesser corn stalk borer (Elasmo-
When grown for hay, seed rates are about 70 palpus lignosellus) has been observed. The
kg/ha. hard seed coat makes the seed resistant to
Tepary bean is grown as a sole crop or inter- storage pests such as bruchid beetles (Calloso-
cropped with cereals (sorghum, millet, maize), bruchus and Acanthoscelides spp.). In Uganda,
vegetables (Allium, Brassica, Capsicum, Cu- however, the rice weevil (Sitophilus oryzae) has
curbita spp.), or other pulses. In the United been recorded in stored seed.
States and Mexico tepary bean is sometimes Harvesting Pods on the same tepary bean
sown in unsorted admixtures with common plant do not usually mature simultaneously,
bean, thus providing greater yield stability and as pods may shatter if left to dry up in the
than common bean alone and higher potential field, they are normally harvested by hand as
yields than tepary bean alone. soon as they change colour, usually 2.5-3
Management Weeding of tepary bean is months after planting. Sometimes whole plants
essential, particularly during early growth. It are pulled up by hand. Normally the pods are
requires little weeding, however, when it is dried for a few days before they are threshed.
grown as an 'end-of-season' crop. Little is In Africa beating the dried pods or plants with
known about its nutrient requirements, and its sticks is common practice to thresh tepary
response to nitrogen and potassium fertilizers bean.
is not consistent. Irrigation may be applied, but Yield In Uganda average yields of tepary
is not usual. In intercropping the cultural prac- bean are 450-670 kg dry seeds per ha. In dry-
tices for the main crop are applied to tepary land farming in the United States yields are
bean. In Senegal and Mali tepary bean is 500-800 kg/ha, under irrigation 900-1700
grown as a kitchen garden vegetable. kg/ha. When grown for fodder, 5500-11,000
Diseases and pests Tepary bean is gener- kg/ha oven-dry hay can be obtained.
ally not affected by diseases in semi-arid re- Handling after harvest Unlike most other
gions, except during periods when humidity is pulses, tepary bean seed stores well and it
high. Within the species, variable levels of re- hardly needs storage pest control.
sistance exist against common bacterial (bean) Genetic resources The genetic basis of cul-
blight (Xanthomonas campestris pv. phaseoli), tivated tepary bean is narrower than that of
bean rust (Uromyces appendiculatus), Fusa- cultivated common bean and Lima bean
rium rot (Fusarium sp.), powdery mildew (Ery- (Phaseolus lunatus L.), and most genetic diver-
siphe polygoni), anthracnose (Colletotrichum sity for future improvement resides in the wild
lindemuthianum), angular leaf spot (Phaeoi- types. Collection of germplasm from the native
sariopsis griseola) and charcoal rot (Macro- area of tepary bean is recommended. Wild te-
phomina phaseolina). Tepary bean is suscepti- pary bean genepools have decreased due to
ble to white mould disease (Sclerotinia scle- habitat elimination and degradation, whereas
rotiorum) and also considered susceptible to domesticated tepary bean has suffered genetic
halo blight (Pseudomonas syringae pv. phaseo- erosion due to its shrinking area of cultivation.
licola); it is listed as a host of southern blight Exchange of genetic information between
(Sclerotium rolfsii) and Pythium rot (Pythium common bean and tepary bean is possible, but
aphanidermatum). Tepary bean is highly sus- only for the transfer of simple traits involving
ceptible to bean common mosaic virus (BCMV), only a few genes. Although common bean and
and also has shown susceptibility to alfalfa tepary bean have the same chromosome num-
mosaic virus (AMY), bean yellow mosaic virus ber and similar karyotypes their mitochondrial
(BYMV),beet curly top virus (BCTV), bean pod genomes, agroecological adaptations and mor-
mottle virus (BPMV) and bean golden mosaic phological characters are distinct, indicating
virus (BGMV). that they are fairly divergent species.
Leafhoppers (Empoasca kraemeri) and pod The largest germplasm collection of tepary
borers (Epinotia opposita) were found in water- bean (about 350 accessions) is held at CIAT
stressed trials. Several lines have shown resis- (Centro Internacional de Agricultura Tropical)
tance against the former, but the mechanism in Cali, Colombia. Another large collection is
was non-preference rather than antibiosis. The present at the USDA-ARS Western Regional
Mexican bean beetle (Epilachna varivestis) and Plant Introduction Station, Pullman, Washing-
the potato leafhopper (Empoasca fabae) have ton, United States (211 accessions). Smaller
been found to cause damage to tepary bean. collections are held in Australia (Australian
Some resistance to the black bean aphid (Aphis Tropical Crops & Forages Genetic Resources
 PHASEOLUS 137

Centre, Biloela; 70 accessions), Belgium (Na- to cause flatulence, the long cooking time, the
tional Botanical Garden of Belgium, Meise; 59 laboriousness of the harvest, and its strong
accessions), Mexico (Instituto Nacional de In- flavour and, according to some, objectionable
vestigaciones Forestales, Agricolas y Pecuarias odour. However, in northern Kenya and Nige-
(INIFAP), Col. San Rafael; 40 accessions) and ria, traditional dishes prepared with tepary
Guatemala (Centro Universitario de Sur Occi- bean instead of cowpea were found very ac-
dente (CUNSUROC), Universidad de San Car- ceptable. To promote tepary bean in Africa, the
los, Mazatenango; 31 accessions). In Africa, 29 selection of high-yielding cultivars, the devel-
accessions are held at ISRA (Institut Sénégal- opment of food products (protein supplements)
ais de Recherches Agricoles), Dakar, Senegal, with reduced odour, and the creation of a mar-
and 10 accessions at ILRI (International Live- keting infrastructure are a prerequisite.
stock Research Institute), Addis Ababa, Ethio- Major references CIAT, 2003; Dillen et al.,
pia. Most collections comprise wild as well as 1997; Hornetz, 1993; Jansen, 1989d; Kaplan &
cultivated types. Lynch, 1999; Kay, 1979; Miklas et a l , 1994;
Breeding Tepary bean is generally resistant National Academy of Sciences, 1979; Pratt &
to diseases and pests, tolerant to drought, heat Nabhan, 1988;Tinsley et al., 1985.
and salinity and has a short crop cycle. Pros- Other references Aganga et al., 2000; Bau-
pects for selection are favourable, as sufficient doin &Maquet, 1999;Debouck &Smartt, 1995;
variation in resistance to biotic and abiotic Freytag & Debouck, 2002; Garvin & Weeden,
stress factors exists within the species. Except 1994; Idouraine, Tinsley & Weber, 1989;
for some mass-selected populations, no im- Idouraine, Weber & Kohlhepp, 1995; Kaiser,
proved tepary bean cultivars have been re- 1981; Leung, Busson & Jardin, 1968; Lin &
leased to farmers. Rather, its favourable traits Markhart III, 1996; Markhart III, 1985;
have mainly been targeted to improve common Mogotsi, 1982; Munoz et a l , 2004; Nabhan &
bean instead of tepary bean itself. Resistance Felger, 1978; Purseglove, 1968; Schinkel &
to common bacterial (bean) blight, for instance, Gepts, 1988; Somasegaran, Hoben & Lewinson,
has been transferred into common bean 1991; Stanton, 1966; Thorn et al., 1983; White
through interspecific hybridization. In at- &Montes, 1993.
tempts to cross Phaseolus acutifolius with Sources of illustration Jansen, 1989d.
Phaseolus vulgaris, artificial cross-fertilization Authors K.K. Mogotsi
does not pose problems, but post-zygotic barri-
ers usually prevent normal embryo develop-
ment, and, as a rule, no viable hybrids are ob- PHASEOLUS COCCINEUS L.
tained. Embryo rescue through in-vitro culture
is normally required to complete hybridization Protologue Sp. pl. 2: 724 (1753).
successfully. Through recurrent backcrossing Family Papilionaceae (Leguminosae - Papil-
with alternating parents the hybrids become ionoideae, Fabaceae)
cross-fertile with both species. However, genes Chromosome number 2/t = 22
obtained from common bean tend to be pre- Vernacular names Scarlet runner bean,
dominant in these hybrids. Seeds have also runner bean, multiflora bean, case knife bean
been obtained from a cross involving a Phaseo- (En). Haricot d'Espagne (Fr). Feijào da Es-
lus acutifolius accession (NI 576) without in- panha, feijäo escarlata, feijäo trepador (Po).
vitro culture. Agrobacterium-mediated genetic Origin and geographic distribution Scarlet
transformation of this accession has been runner bean occurs wild from Mexico to Pa-
achieved, based on regeneration from callus. nama. It was probably domesticated in Mexico.
This has opened up the possibility of using Archaeological evidence indicates that scarlet
Phaseolus acutifolius to introduce transgenes runner bean was a domesticated crop in Mexico
into the economically more important Phaseo- around 900 AD. Nowadays scarlet runner bean
lus vulgaris. is cultivated in temperate countries and occa-
Prospects Tepary bean seems a very suit- sionally in highland areas of Central and South
able crop for resource-poor farmers in Africa, America, Africa (e.g. Ethiopia, Kenya, Uganda,
since its rapid germination, deep root system South Africa) and Asia. It is probably culti-
and short life cycle make it well adapted for vated in Madagascar and is recorded as being
production in arid or semi-arid regions. Rea- grown in the eastern part of tropical southern
sons for reluctance to adopt tepary bean as a Africa, although no specific countries are men-
food include the small seed size, the tendency tioned.
138 CEREALS AND PULSES

carbohydrate 3.2 g, fibre 2.6 g, Ca 33 mg, Mg

19 mg, P 34 mg, Fe 1.2 mg, Zn 0.2 mg, carotene
145 ng, thiamin 0.06 mg, riboflavin 0.03 mg,
niacin trace and ascorbic acid 18 mg (Holland,
Unwin & Buss, 1991). Many improved culti-
vars have substantial reduction in the fibrous
vascular strands of the pod sutures ('stringless
runner beans').
The tuberous root of scarlet runner bean is
edible, but it is fibrous and may contain toxic
compounds, which can be removed by soaking
or peeling and by discarding the cooking water.
Coccinin, a peptide isolated from the seed of
scarlet runner bean, has shown antifungal
activity against a range of fungi. It also inhib-
ited proliferation in leukaemia cell lines and
reduced the activity of HIV-1reverse transcrip-
Phaseolus coccineus - planted tase.
Description Perennial, climbing herb with
Uses In Central America the immature and stems up to 4(-7) m long or bushy annual herb
mature seeds are consumed, elsewhere mainly up to 60 cm tall; taproot tuberous. Leaves al-
the mature seeds, e.g. in Ethiopia. Preparation ternate, 3-foliolate; stipules triangular; petiole
is predominantly by boiling. In temperate re- (6-)8.5-10.5(-16) cm long, rachis (1.5-)2.5-4(-
gions the immature pods are most commonly 5) cm long; stipels c. 5 mm long; leaflets ovate-
eaten, sliced and cooked, as a vegetable. In rhombic, (5-)6.5-10.5(-12.5) cm x (3.5-)5-8.5(-
Central America the young shoots, leaves and 12.5) cm, base cuneate or truncate, apex acute,
inflorescences are sometimes used (boiled or thinly pubescent to glabrescent. Inflorescence
boiled and fried) as a vegetable while the tu-
berous roots are consumed boiled or chewed as
candy. A root decoction is taken against ma-
laria or applied to swollen eyes. In Central
America scarlet runner bean is grazed by live-
stock and dried into hay. It is grown as an or-
namental.
Production and international trade Accu-
rate production statistics for scarlet runner
bean are not available. Production is almost
exclusively for local use. Commercial produc-
tion of the pods is done in the United Kingdom
and Argentina and of the seeds of white-seeded
cultivars in South Africa. In Kenya scarlet run-
ner bean is mainly grown by smallholders.
Properties Per 100 g edible portion the
composition of dried scarlet runner bean seeds
is: water 12.5 g, energy 1415 kJ (338 kcal),
protein 20.3 g, fat 1.8 g, carbohydrate 62.0 g,
fibre 4.8 g, Ca 114 mg, P 354 mg, Fe 9.0 mg,
thiamin 0.50 mg, riboflavin 0.19 mg, niacin 2.3
mg and ascorbic acid 2 mg (Leung, Busson &
Jardin, 1968). The seeds contain antinutri-
tional factors such as trypsin inhibitors, and
must be cooked before being eaten to break
down these compounds.
Per 100 g edible portion, raw green pods (ends Phaseolus coccineus - 1, part of flowering
and sides trimmed) contain: water 91.2 g, en- branch; 2,fruit; 3, seed.
ergy 93 kJ (22 kcal), protein 1.6 g, fat 0.4 g, Source: PROSEA
 PHASEOLUS 139

an axillary or terminal raceme, many-flowered; be harvested 4-5 months after sowing. Bushy
peduncle (5-)ll-16.5(-25.5) cm long; rachis (2-) cultivars produce earlier and smaller crops
10-16(-39.5) cm long. Flowers bisexual, papil- than climbing cultivars. In Central America
ionaceous; pedicel 0.5-1.5 cm long; calyx cam- scarlet runner bean is sometimes grown as a
panulate, glabrescent, tube c. 3 mm long, the perennial: where stems die back during cooler
upper 2 lobes united, the lower 3 triangular, c. periods, the tuberous taproot remains viable
1 mm long; corolla scarlet, pink or white, stan- and produces new stems with returning
dard hood-shaped, circular or broadly obovate, warmth. In temperate regions scarlet runner
c. 17 mm x 17 mm, wings broadly obovate, c. 25 bean is grown as an annual. Scarlet runner
mm x 17 mm, keel coiled, c. 10 mm long; sta- bean fixes atmospheric nitrogen by symbiosis
mens 10, 9 fused and 1free; ovary superior, c. 6 with fast-growing Rhizobium bacteria.
mm long, finely pubescent, style coiled, with Ecology Scarlet runner bean is a crop for
collar of hairs below the stigma. Fruit a linear- temperate climates. In the tropics it is most
lanceolate, straight or slightly curved pod (4.5-) successful at altitudes of 1500-2000 m. In
9-13(-30) cm x 1.5-2.5 cm, laterally compress- Kenya it is grown at 1900-2600 m altitude, in
ed, beaked, glabrescent, rough with small Ethiopia up to about 2000 m. Scarlet runner
oblique ridges, (l-)3-5(-10)-seeded. Seeds el- bean is more tolerant of cool conditions than
lipsoid-oblong, 13-25 mm x 6-13(-16) mm, other Phaseolus species, but damage occurs at
black, white, cream or brown, often pink to temperatures below 5°C. At temperatures
purple mottled. Seedling with hypogeal germi- above 25°C fruit development is inhibited.
nation; first pair of leaves simple and opposite. Scarlet runner bean is extremely susceptible to
Other botanical information Phaseolus drought and requires a well-distributed rainfall
comprises about 50 species, most ofthem in the throughout the growing period. In Ethiopia it
Americas. Phaseolus coccineus is closely re- is successfully grown in areas with an average
lated to Phaseolus dumosus Macfad. (synonym: annual rainfall of 1500 mm. It needs a high
Phaseolus polyanthus Greenman; year-bean, relative humidity for seed set. Scarlet runner
sometimes also called runner bean) and Phase- bean comprises short-day and day-neutral
olus costaricensis Freytag & Debouck. Hybrids types.
between Phaseolus coccineus and these 2 spe- Scarlet runner bean is adapted to a wide range
cies have been obtained; natural hybridization of soils, but it prefers deep, well-drained,
also occurs. The 3 species can be crossed with loamy, light- to medium-textured soils, with
common bean (Phaseolus vulgaris L.), with the pH 6-7. Waterlogging is not tolerated.
latter as female parent, without embryo rescue, Propagation and planting Scarlet runner
although progenies may be partially sterile. bean is normally propagated by seed, but the
Where scarlet runner bean and common bean tuberous root with a piece of stem can also be
grow together, natural hybridization may oc- used. The 1000-seed weight is 700-3000 g. The
cur. Hybridization of scarlet runner bean with seedbed should be well prepared and weed free.
tepary bean (Phaseolus acutifolius A.Gray) is Normal planting densities are 50,000-75,000
also possible. plants/ha for climbing types and double those
Phaseolus coccineus is a variable species, and for bushy types, requiring about 75 kg and 150
levels of genetic variability are high, both in kg seed per ha, respectively. However, lower
wild and in cultivated populations. A white- densities have also been recorded. In Mauritius
seeded type of Phaseolus coccineus is known as scarlet runner bean is sown in rows 100 cm
'butter bean' in Kenya and South Africa, but apart with 30 cm within the row. The sowing
this name is normally applied to Phaseolus depth is 2.5-5 cm. In Central America scarlet
lunatus L. In Uganda, where the crop is grown runner bean is often intercropped with maize.
a high altitudes in Nakuru District, white- Management To obtain high-quality pods,
seeded cultivars are most common. scarlet runner bean is grown on trellises, poles,
Growth and development Scarlet runner fence lines or other support structures. How-
bean seeds germinate 10-14 days after sowing. ever, labour and material requirements are
Flowering starts 40-60 days after sowing. high and may impede cultivation. Climbing
Flowers open at sunrise and fade at sunset. types can yield without support if leading
Phaseolus coccineus is predominantly cross- shoots are pinched out to induce bushy growth.
pollinating. Harvesting of green pods starts Scarlet runner bean should be kept weed-free
around 3 months after sowing and can be eas- during the early growth stages and it is com-
ily sustained for 2-3 months. Mature seed can monly weeded once or twice. Tillage should be
140 CEREALS AND PULSES

shallow to avoid root damage. Supplementary Moderate levels of resistance to common bacte-
irrigation is beneficial. In Ethiopia scarlet run- rial blight (Xanthomonas campestris pv. phase-
ner bean is a garden crop. oli), Fusarium root rot (Fusarium solani f.sp.
Diseases and pests In the tropics scarlet phaseoli) and white mould (Sclerotinia scle-
runner bean is affected by anthracnose (Colle- rotiorum) have been transferred from scarlet
totrichum lindemuthianum) and Fusarium wilt runner bean to common bean. Scarlet runner
(Fusarium solani f.sp. phaseoli). The seed- bean is also considered as a potential source of
borne disease halo blight (Pseudomonas savas- resistance against other diseases of common
tanoi pv. phaseolicola, synonym: Pseudomonas bean, including anthracnose, Ascochyta blight
syringae pv. phaseolicola) has been isolated (Phoma exigua), angular leaf spot (Phaeoi-
from scarlet runner bean in South Africa. sariopsis griseola), powdery mildew (Erysiphe
Harvesting Green pods of scarlet runner polygoni) and rust (Uromyces appendiculatus).
bean are harvested when pod length reaches Considerable tolerance to bean flies (Ophio-
its maximum before the phase of rapid seed myia spp.) has been detected in scarlet runner
development. Picking is usually at 4—5 day bean, and tolerance has been transferred into
intervals. For dry seed production, plants are common bean. On the other hand, resistance to
pulled or cut when most pods are dry and then halo blight has been transferred from common
allowed to dry for a few days. Alternatively bean to scarlet runner bean.
pods may be handpicked in several rounds be- In vitro plant regeneration of scarlet runner
cause of asynchronous ripening. bean is possible using cotyledons, through di-
Yield Yields of green pods of 10 t/ha and of rect organogenesis as well as somatic embryo-
seeds of 1.5 t/ha are possible. The yield of dry genesis via callus.
mature seeds in Kenya has been estimated at Prospects Scarlet runner bean is a suitable
900-1100 kg/ha. pulse and vegetable crop for the humid high-
Handling after harvest After drying, scar- land tropics, although the need to provide sup-
let runner bean pods are threshed. port and the uneven maturation ofthe pods are
Genetic resources In Brazil 428 accessions serious drawbacks for commercial production.
are maintained by EMBRAPA/CENARGEN in Scarlet runner bean may have some potential
Brasilia. Large germplasm collections of scarlet at higher altitudes in tropical Africa, but more
runner bean are also maintained in the United information is needed on appropriate sowing
States (USDA-ARS Western Regional Plant and management practices. It is a potential
Introduction Station, Pullman, Washington, source of resistance to diseases and pests af-
478 accessions from throughout the world in- fecting common bean.
cluding Ethiopia and Kenya) and Mexico Major references Campion, 1995; Debouck
(Banco Nacional de Germoplasma Vegetal, & Smartt, 1995; Freytag & Debouck, 2002;
Universidad Autónoma Chapingo, Chapingo, Gepts (Editor), 1988; Kay, 1979; Singh, 2001;
311 accessions). In Africa 6 accessions are kept Smartt, 1989a; Suttie, 1969; Webster, Ross &
in South Africa (Division of Plant and Seed Sigourney, 1980;Westphal, 1974.
Control, Department of Agriculture, Pretoria) Other references Duke, 1981; du Puy et
and 1 in Ethiopia (International Livestock Re- al., 2002; Escalante et a l , 1994; FAO, 1989;
search Institute (ILRI), Addis Ababa). Fourie, 1998; Hidalgo & Beebe, 1997; Holland,
Breeding Breeding efforts for scarlet runner Unwin & Buss, 1991; Kaplan & Lynch, 1999;
bean have been directed to improvement of Knudsen (Editor), 2000; Leung, Busson & Jar-
culinary quality (stringlessness) and disease din, 1968; Liebenberg, 1995; Mahuku et al.,
resistance. Selection to improve cooking quality 2002a; Mahuku et al., 2002b; Nagl, Igna-
is promising since seed proteins of scarlet run- cimuthu & Becker, 1997; Ngai & Ng, 2004;
ner bean are more polymorphic than those of Schmit & Baudoin, 1992; Smartt, 1976; Sum-
common bean. For dry seed production, im- merfield & Roberts (Editors), 1985; Thulin,
provement of plant habit and shorter pods are 1989a; Yu, Stall &Vallejos, 1998.
appropriate objectives of selection. Cultivars Sources of illustration Smartt, 1989a.
with a determinate growth habit suitable for Authors M. Brink
mechanical harvesting CVenere' and 'Alarico') Based on PROSEA 1: Pulses.
have been developed in Italy, by crossing
Phaseolus coccineus with determinate Phaseo-
lus vulgaris cultivars and repeated back-
crossing with Phaseolus coccineus.
 PHASEOLUS 141

PHASEOLUS LUNATUS L. across the Pacific to the Philippines and from

there it spread to other parts of Asia, mainly
Protologue Sp. pi. 2: 724 (1753). Java and Myanmar (Burma), and to Mauritius.
Family Papilionaceae (Leguminosae - Papil- The slave trade introduced Lima bean from
ionoideae, Fabaceae) Brazil into Africa, particularly to the western
Chromosome number 2re=22 and central parts. Some large-seeded types
Vernacular names Lima bean, butter bean, from the Peruvian coast were distributed to
Madagascar bean (En). Haricot de Lima, pois south-western Madagascar and southern Cali-
du Cap, pois souche, pois savon (Fr). Feijào de fornia. Lima bean is now cultivated throughout
Lima, feijào favona, feijâo espadinho (Po). Mfi- tropical Africa and the rest of the tropics, and
wi (Sw). has frequently become naturalized.
Origin and geographic distribution Lima Uses Lima bean is cultivated primarily for
bean has a Neotropical origin with at least two its immature and dry seeds, which in tropical
centres of domestication: Central America Africa are usually eaten boiled, fried in oil or
(Mexico, Guatemala) for the small-seeded types baked. In Nigeria they are also cooked with
and South America (mainly Peru) for the large- maize, rice or yam and used in making special
seeded types. This distinction coincides with a kinds of soup and stew. The Yoruba people
classification into 2 types (Mesoamerican and process the seeds into porridges, puddings and
Andean) on the basis of morphological, ecologi- cakes. Immature green seeds, young pods and
cal, protein and molecular characters. Wild and leaves are eaten as a vegetable, e.g. in Ghana
cultivated forms of the same race are grouped and Malawi. In the United States, fresh and
together. The Andean wild populations have a dry Lima beans are processed on an industrial
very limited geographic distribution (Ecuador scale involving canning and freezing. Sprouts
and northern Peru). The Mesoamerican wild and young plants are cooked and eaten in
types extend from Mexico to Argentina through many Asian countries.
the east side of the Andes. Recent discoveries The seeds are sometimes used as fodder, but
have led to the proposition of 3 primary centres may lead to hydrogen cyanide poisoning when
of genetic diversity, 2 of which are also domes- used raw. The leaves and stems may be turned
tication centres: a centre of genetic diversity into hay or silage. Juice from the leaves is used
and domestication on the western slope of the in nasal instillations against headache and as
Andes in southern Ecuador and northern Peru; eardrops against otitis in Senegal and DR
a centre of genetic diversity and domestication Congo. In Nigeria the seeds are powdered and
in Central America; and a centre of genetic rubbed into small cuts on tumours and ab-
diversity in the region covering northern Peru, scesses to promote suppuration. In traditional
northern Colombia, northern Ecuador and Asian medicine the seeds and leaves are valued
western Venezuela. for their astringent qualities and used as a diet
In post-Columbian times, Lima bean spread against fever. Lima bean has been grown as a
throughout the Americas. Spaniards took seeds cover crop and for green manure.
Production and international trade Pro-
duction statistics for Lima bean from many
tropical regions are fragmentary and often
aggregated with other pulses. The United
States is the largest producer of Lima bean
with about 21,000 ha under cultivation (mainly
in California, Delaware, Maryland and Wis-
consin) and a production of (primarily fresh)
beans of about 70,000 t in 1995.Madagascar is
the second largest commercial producer with
an area cropped varying from 3000 to 19,000
ha (mainly in the flood plains of the semi-arid
coastal region in the south-western part) and a
production of dry seed of about 8000 t, almost
exclusively of large white-seeded types. Peru
comes third with a production of dry seed of
5000-5500 t from 5000-6000 ha. In other coun-
Phaseolus lunatus - planted tries, Lima beans are grown mostly in gardens
142 CEREALS AND PULSES

or as an intercrop, but there are no estimates mg, Fe 2.6 mg, vitamin A 65 IU, thiamin 0.15
of area or production. In Africa the area mg, riboflavin 0.10 mg, niacin 1.20 mg and
planted with Lima bean in the sub-humid and ascorbic acid 27.0 mg (Kay, 1979). Lima bean
humid tropics (especially Sierra Leone, Liberia, silage contains 27.3%dry matter, 3.3% protein,
Côte d'Ivoire, Ghana, Nigeria and DR Congo) 2.1% digestible protein and 14.2% digestible
has been estimated for the 1980s as 120,000- nutrients.
200,000 ha, with a total annual production of Description Climbing, trailing or more or
50,000-100,000 t. No trade statistics are avail- less bushy annual or perennial herb, with gla-
able. brous or pubescent stems up to 4.5(-8) m long;
Properties The composition of dried raw roots thin or swollen, up to 2 m deep. Leaves
seeds per 100 g edible portion is: water 11.6 g, alternate, 3-foliolate; stipules ovate to lanceo-
energy 1214 kJ (290 kcal), protein 19.1 g, fat late, 2-4 mm long; petiole 1.5-19 cm long, ra-
1.7 g, carbohydrate 52.9 g, dietary fibre 19.4 g, chis 0.5-5(-8) cm long; stipels 1-2 mm long;
Ca 85 mg, Mg 190 mg, P 320 mg, Fe 5.9 mg, Zn leaflets ovate, 3-19.5 cm x 1-11 cm, acute or
2.8 mg, carotene trace, thiamin 0.45 mg, ribo- acuminate, sparsely pubescent or glabrous.
flavin 0.13 mg, niacin 2.5 mg, vitamin B6 0.51 Inflorescence an axillary raceme or panicle up
mg and ascorbic acid trace (Holland, Unwin & to 15(—40) cm long, with many nodes, few-
Buss, 1991). The essential amino-acid composi- flowered to many-flowered. Flowers bisexual,
tion per 100 g raw Lima beans is: tryptophan papilionaceous; pedicel 5-10 mm long; calyx
180 mg, lysine 1440 mg, methionine 280 mg, campanulate, 2.5-3.5 mm long, puberulous, the
phenylalanine 1160 mg, threonine 800 mg, upper 2 lobes united, the lower 3 broadly tri-
valine 980 mg, leucine 1560 mg and isoleucine angular; corolla 7-10 mm wide, standard hood-
950 mg (Paul, Southgate & Russell, 1980). As shaped, 5-7 mm x 5-10 mm, white, pale green
in other pulses, the main limiting amino acids or rose-violet, wings spatulate to obovate, 7—10
are methionine and cystine. Antimetabolic fac- mm long, white or violet, keel sharply up
tors include protease inhibitors, lectins and
cyanogenic glucosides (linamarin or phaseolu-
natin). The latter are accompanied by an en-
zyme, linamarase, which can hydrolyze the
glucosides into a sugar and an aglycone, which
in turn is split into acetone and hydrogen cya-
nide (HCN). Hydrolysis occurs rapidly when
the soaked seeds are cooked in water; most of
the HCN then evaporates. Linamarin and li-
namarase are heat-sensitive but inactivated at
different temperatures: 140°C for the glucoside
and 80°C for the enzyme. If inactivation of the
enzymes takes place before complete hydroly-
sis, the residual glucoside may break down in
the human organism under the influence of
enzymes secreted by the intestinal microflora,
leading to poisoning. The HCN content is sig-
nificantly higher in wild types (2000-2400
ppm) than in cultivated ones (100-120 ppm).
Soaking Lima bean seeds in water overnight
easily eliminates the apparent toxicity, which
is explained by the release of HCN during the
process. In Nigeria, cooking time for dry seeds
is 1—1.5 hours. In Malawi, cooking times have
been recorded of 2-2.5 hours for dry, unsoaked
seeds and 1.5-2 hours for dry, soaked seeds.
Per 100 g edible portion, green pods contain 1.3
g protein, and green leaves 0.6 g. Immature
Lima bean seeds contain per 100 g edible por- Phaseolus lunatus - 1, flowering and fruiting
tion: water 66.3 g, protein 8.3 g, fat 0.7 g, car- branches; 2, flower; 3, seeds.
bohydrate 23.1 g, fibre 1.0 g, Ca 28 mg, P 111 Source: PROSEA
 PHASEOLUS 143

t u r n e d , w h i t e or pale green; s t a m e n s 10, 9 favouring self-pollination. However, cross-

fused a n d 1 free; ovary superior, c. 3 m m long, pollination often occurs too. P r e s s u r e on t h e
minutely hairy, style with a t e r m i n a l coil, with wings of fully-open flowers by visiting insects
collar of h a i r s below t h e stigma. F r u i t a n ob- forces t h e stigma a n d style to p r o t r u d e t h r o u g h
long pod (4.5-)5-10.5(-13) cm x l - 2 ( - 3 ) cm, t h e keel. T h e exposed stigma r e m a i n s receptive
compressed, generally curved, beaked, glabrous to pollen for several h o u r s . Bees visit t h e flow-
or pubescent, 2-4(-5)-seeded. Seeds kidney- ers for both pollen a n d nectar. Of buds, flowers
s h a p e d to rhomboid or globose, 8 - 1 1 m m x 6 - 7 a n d young pods, 75—85% a r e shed u n d e r field
mm, white, green, yellow, brown, red, purple, conditions. Early blooming inflorescences a r e
black or variously speckled, often with t r a n s - more productive t h a n later ones, a n d b a s a l
verse lines r a d i a t i n g from t h e h i l u m . Seedling nodes of t h e inflorescences a r e potentially more
with epigeal germination; first pair of leaves fruitful t h a n t e r m i n a l ones. F r u i t s e t t i n g pro-
simple a n d opposite. ceeds u n t i l a 'capacity set' is a t t a i n e d ; r e m a i n -
Other botanical information Phaseolus com- ing reproductive s t r u c t u r e s t h e n abscise. L i m a
prises about 50 species, most of t h e m in t h e b e a n can fix nitrogen by symbiosis w i t h Bra-
Americas. Wild a n d cultivated types of Phaseo- dyrhizobium bacteria.
lus lunatus h a v e been distinguished as var. E c o l o g y L i m a b e a n is p a r t i c u l a r l y well
Silvester B a u d e t a n d var. lunatus, respectively. suited to low-altitude h u m i d a n d s u b - h u m i d
W i t h i n t h e cultivated types some cultivar tropical climates, b u t it can be grown in a wide
groups h a v e been distinguished: Sieva Group r a n g e of ecological conditions. It is found in
with medium-sized flat seeds, Potato Group w a r m t e m p e r a t e zones as well as in arid a n d
with small globular seeds, a n d Big L i m a Group semi-arid tropical regions. L i m a b e a n is found
with large flat seeds. Wild types from t h e An- from sea-level u p to a l t i t u d e s h i g h e r t h a n 2000
des a p p e a r closest to t h e cultivated types. m. It comprises photoperiod-insensitive types
In M a l a w i types of Lima b e a n a r e distin- t h a t flower in daylengths up to 18 h o u r s , a n d
guished a n d n a m e d according to t h e size a n d short-day types t h a t require a d a y l e n g t h as
s h a p e of t h e seeds, e.g. ' m a y e m b a ' (large w h i t e short as 11-12.5 h o u r s to initiate flowers. Op-
or black speckled seeds, slightly bitter, seed t i m u m t e m p e r a t u r e s a r e 16-27°C; frost is not
coats moderately tough), 'butter' or ' M a d a g a s - tolerated. Average rainfall is 900—1500 m m per
car' (large flat w h i t e seeds, with a good flavour, year, b u t once established t h e crop tolerates as
seed coats soft a n d tasteless), 'moki' (small, little rainfall as 5 0 0 - 6 0 0 m m . Some types a r e
white, flat seeds, good flavour, seed coats mod- considered very drought r e s i s t a n t , due to their
erately tough), a n d 'pebugale' (seed s h a p e vari- deep, well-developed root system. Lima b e a n
able, colour pale pink, speckled red, slightly prefers well-aerated, well-drained soils w i t h
bitter w i t h tough seed coats). p H 6.0-6.8. However, some cultivars tolerate
G r o w t h a n d d e v e l o p m e n t G e r m i n a t i o n of acid soils with p H as low as 4.4.
L i m a b e a n seeds occurs 4—10 days after sow- P r o p a g a t i o n a n d p l a n t i n g P r o p a g a t i o n of
ing. Vegetative growth accelerates after one L i m a b e a n is by seed. Seed weight varies be-
m o n t h . Flowers a p p e a r 35—70 days a n d ripe t w e e n 30 g a n d 300 g per 100 seeds. B u s h types
pods 8 0 - 1 2 0 days after sowing w i t h short a r e u s u a l l y spaced 20—30 cm w i t h i n rows a n d
daylength. Cultivated L i m a b e a n h a s two dis- 6 0 - 1 0 0 cm b e t w e e n rows, while climbing types
tinct growth h a b i t s : a n i n d e t e r m i n a t e growth m a y be p l a n t e d on hills 9 0 - 2 0 0 cm a p a r t . L i m a
h a b i t (prostrate or climbing; w i t h axillary flow- b e a n m a y be p l a n t e d in groups of 3—4 p l a n t s ,
ering only) a n d a p s e u d o - d e t e r m i n a t e growth s e p a r a t e d by a t least 1 m. T h e n o r m a l seed
h a b i t (dwarf or b u s h p l a n t s ; with t e r m i n a l a n d r a t e varies b e t w e e n 5 5 - 8 0 kg/ha for small-
axillary flowering). T h e vegetative cycle of seeded cultivars a n d 1 3 0 - 1 6 0 kg/ha for large-
p s e u d o - d e t e r m i n a t e growth types is s h o r t e r seeded types. T h e p l a n t i n g density in south-
t h a n t h a t of i n d e t e r m i n a t e ones. T h e earliest w e s t e r n M a d a g a s c a r is (500-)2100(-4500)
b u s h cultivars m a t u r e within 90 days w h e r e a s pockets per ha, w i t h 3 - 5 seeds per pocket.
t h e climbing types r e q u i r e 6—9 m o n t h s . In In t h e more h u m i d tropics, L i m a b e a n is mostly
climbing types, flowering a n d fruiting m a y cultivated in h o m e g a r d e n s or intercropped
extend t h r o u g h o u t t h e wet season. T h e growth with cereals (maize, sorghum), root a n d t u b e r
h a b i t of p e r e n n i a l wild types is always inde- crops (yam, cassava) or other crops (e.g. ba-
terminate. n a n a , g r o u n d n u t , s u g a r cane). Sole cropping is
Pollen a n d stigma m a t u r e synchronously a n d more common in drier a r e a s (Madagascar,
in close proximity w i t h i n t h e u n o p e n e d bud, P e r u ) . In intercropping, seeds a r e often placed
144 CEREALS AND PULSES

in the same hill as the companion crop. Harvesting Green and mature pods of the
Management Weeding is necessary during climbing Lima bean types are usually picked
initial growth of Lima bean. In Africa it is usu- manually over an extended period (4-6 weeks).
ally done 1-3 times. In humid areas, climbing In drier areas (Madagascar), whole plants are
types are staked. Unstaked plants tend to have cut and left to dry in the field before the pods
lower yields because of poor foliage display, are removed and the stems are fed to livestock.
and lower seed quality because the pods may Mechanical picking is possible with erect culti-
be resting on the ground. In parts of West Af- vars maturing uniformly and setting pods well
rica, where Lima bean is intercropped with above the soil surface.
maize or sorghum, the cereals provide support Yield In the tropics, yields of dry seeds of
for climbing. Lima bean may also be planted Lima bean are 200-600 kg/ha in intercropping
after yam, with the stake of the previous yam and 1000-1500 kg/ha in sole cropping. Yields
crop providing support. In drier conditions in south-western Madagascar are (50-)400(—
(Madagascar, California and Peru), Lima bean 950) kg/ha. In experiments dry-seed yields in
may be left prostrate and irrigated 2-4 times pure stands have reached 2000—2500 kg/ha for
before maturity. In the south-western part of the bush types and 3000-4000 kg/ha for the
Madagascar Lima bean is planted on alluvial climbing types. In Madagascar yields of 15 t
soils in mounds or ridges as the flood waters green matter per ha have been obtained for use
from a nearby river recede, or is irrigated from as fodder.
a river. Fertilizer is not usually applied in Handling after harvest Pods of Lima bean
tropical areas. If fertilizer is applied, this is are usually threshed by hand, and seed is
often done at planting, in bands below and cleaned and sorted. Care should be taken with
adjacent to the seeds. Supplemental nitrogen threshing, as the seeds are brittle and easily
and phosphorus may be side-dressed at the damaged. In many tropical countries, seeds are
early bud stage and during fruit development. sometimes stored in jars or baskets, and cov-
Lima bean may be planted after a well- ered with a layer of sand or ash to protect them
fertilized crop, thus benefiting from residual against bruchid infestation.
fertilizer, especially phosphorus. Genetic resources There is a real risk of
Diseases and pests In the tropics the most loss of genetic diversity of Lima bean in pri-
serious diseases of Lima bean are web blight mary centres of diversity (Latin America) as
caused by Rhizoctonia solani, Fusarium root well as in secondary centres of cultivated types
rot caused by Fusarium solani, anthracnose (Africa and part of Asia). Over 2600 seed sam-
caused by Colletotrichum spp., downy mildew ples of Lima bean are available in the CIAT
caused by Phytophthora phaseoli, bacterial (Centro Internacional de Agricultura Tropical)
blight due to Xanthomonas campestris pv. collection at Cali (Colombia) with seeds coming
phaseoli and two viral diseases: Lima bean mainly from South and Central America, West
golden mosaic virus (LGMV) transmitted by Africa (mainly Ghana and Nigeria), East and
white flies (Bemisia sp.) and Lima bean green Central Africa, Madagascar, India, the Philip-
mottle virus (LBGrMV) transmitted by aphids. pines and Myanmar. The wild and weedy types
The use of fungicides is recommended to con- represent 3-5% of the total collection. Among
trol the fungal diseases. For bacterial blight the cultivated types, Sieva Group and Potato
the use of disease-free seed and crop rotation Group predominate, while accessions of the Big
are the recommended control measures. Lima Group come mostly from a few limited
Root-knot nematodes (mainly Meloidogyne areas, such as the South American Andean
incognita) can cause considerable yield reduc- region or the desert coast of Peru. According to
tion. Crop rotation with cereals can reduce the IPGRI other large Lima bean collections exist
nematode population in the soil. Mexican bean in Indonesia (Research and Development Cen-
beetle (Epilachna varivestis), aphids (mainly tre for Biology (RDCB), Bogor, 3850 acces-
Aphis craccivora), leafhopper (Empoasca doli- sions), the United States (Regional Plant In-
chi), flower thrips (Megalurothrips sjostedtî), troduction Station, Washington State Univer-
legume pod borers (Maruca vitrata, Cydia sp. sity, Pullman, Washington, 1060 accessions),
and Etiella sp.), and bruchids (Callosobruchus, Brazil (Empresa Brasileira de Pesquisa Ag-
Acanthoscelides and Zabrotes spp.) are serious ropecuâria (EMBRAPA ), Brasilia, 980 acces-
pests. Chemical control measures (e.g. with sions), the Philippines (National Plant Genetic
endosulfan) have been recommended to control Resources Laboratory, University of the Phil-
them. ippines Los Banos (UPLB), College, Laguna,
 PHASEOLUS 145

780 accessions) and Costa Rica (Escuela de of the wild taxa (e.g. resistance to Lima bean
Biologia, Universidad de Costa Rica (UCR), golden mosaic virus) has been observed in in-
San Pedro de Montes de Oca, 400 accessions). terspecific breeding material. Many commer-
In tropical Africa small Lima bean collections cial Lima bean cultivars have been developed
are present in Ghana (Plant Genetic Resources in the United States.
Centre, Crops Research Institute, Bunso, 40 Prospects Because of its high yield poten-
accessions; University of Ghana, Accra, 8 ac- tial, deep rooting and drought tolerance, Lima
cessions), Togo (Institut de Recherches bean has good prospects in tropical Africa. Pre-
Agronomiques Tropicales et des Cultures liminary investigations have shown the great
Vivrières, Lomé, 36 accessions; Direction de la potential and the large genetic diversity of
Recherche Agronomique (DRA), Lomé, 17 ac- Phaseolus lunatus germplasm. Some progress
cessions), Guinea (Programme de conservation in crop improvement has been made, mainly in
des ressources phytogénétiques, Institut de areas outside the region of origin (for example
Recherche, Conakry, 34 accessions), Senegal in temperate climates of the United States).
(Institut Sénégalais de Recherches Agricoles Much remains to be done in many regions of
(ISRA), Dakar, 23 accessions), Nigeria (Inter- the tropics, particularly to develop more stable
national Institute of Tropical Agriculture, and higher-yielding cultivars for the humid,
Ibadan, 15 accessions), Kenya (National Gene- sub-humid and semi-arid tropics. Breeding
bank of Kenya, Crop Plant Genetic Resources efforts should consider separately the two ma-
Centre, KARI, Kikuyu, 12 accessions) and jor growth habits. The climbing indeterminate
Ethiopia (International Livestock Research types usually give high but unstable dry-seed
Institute (ILRI), Addis Ababa, 2 accessions). yield and require an expensive system of stak-
The National Botanic Garden of Belgium at ing. In tropical regions, these types are mainly
Meise has been mandated by IPGRI as a base grown in intercropping systems with cereals or
repository collection of wild Phaseolus, which root and tuber crops. So far, few genotypes
includes wild accessions of Phaseolus lunatus suitable for intercropping have been bred,
and related species. which explains the poor performance of climb-
Breeding Evaluation and breeding based on ing Lima bean in such systems in spite of their
international collections were carried out at high potential. The bushy pseudo-determinate
UTA (Ibadan, Nigeria) between 1973 and 1980, types are more appropriate for sole cropping
and later at CIAT (Cali, Colombia) between and intensive production systems. However,
1980 and 1992, the general objectives being to results are discouraging, particularly in humid
increase dry seed yield, upgrade resistance to tropics, due to unfavourable plant architecture
pests and diseases and improve nutritional (profuse branching, pods within the leaf canopy
seed quality. Small-scale improvement pro- and severe lodging) and high susceptibility to
grammes of Lima bean using local collections diseases. The key factor to success would be to
are being conducted in Ghana, Nigeria, DR develop indeterminate bushy types with sev-
Congo, Zambia and Madagascar. The adopted eral traits of wide adaptation (such as deep
breeding methods are pure-line selection, bulk rooting, drought tolerance, disease resistance
and population improvement; the target crop- and high yield potential). Research priorities
ping systems include both sole cropping and should first be devoted to full exploitation of
intercropping systems. Erectness, resistance to the large genetic variation available in the
lodging and to web blight are prime criteria to primary genepool of Mesoamerican and An-
improve pseudo-determinate bush types. dean origin. Exploitation of the alien genepools
Earliness, photoperiod insensitivity, resistance and selection in the interspecific populations
to Lima bean golden mosaic virus and suitabil- should not be neglected when considering the
ity for intercropping are being sought in climb- challenge of high and stable yields in the hu-
ing types. Some promising types in the humid mid tropics.
tropics have been identified among the climb- Major references Baudoin, 1989; Baudoin,
ing forms. A large secondary genepool is avail- 1991; Baudoin, 2002; Baudoin & Mergeai,
able for improvement and the following wild 2001b; Burkill, 1995; Fofana, du Jardin & Bau-
species have been successfully crossed with doin, 2001; Freytag & Debouck, 2002; Kay,
Lima bean: Phaseolus jaliscanus Piper, Phaseo- 1979; Lyman, Baudoin &Hidalgo, 1985; Rollin,
lus maculatus Scheele, Phaseolus polystachyus 1997.
(L.) Britton, Sterns & Poggenb. and Phaseolus Other references Baudet, 1977; Baudoin,
salicifolius Piper. Introgression of useful genes 1988; Baudoin, 1993; Berhaut, 1976; Duke,
146 CEREALS AND PULSES

1981; Ezueh, 1977; Fofana et a l , 1999; Gillett century through Sofala (Mozambique), Zanzi-
et al., 1971; Hauman et al., 1954a; Hepper, bar and Mombasa, from where it was carried to
1958; Holland, Unwin & Buss, 1991; ILDIS, higher altitude areas of the interior by slave
2002; Kee, Glancey & Wootten, 1997; Maquet, trading caravans and merchants. Common
Vekemans & Baudoin, 1999; Maquet et al., bean became well established as a pulse crop in
1997; Paul, Southgate & Russell, 1980; Polhill, parts of Africa before the colonial era. Genetic
1990; Schmit et a l , 1993;Westphal, 1974; Wil- diversity of common bean and its pathogens
liamson, 1955. and linguistic evidence indicate that it became
Sources of illustration Baudoin, 1989. a major crop in Central African highland areas
Authors J.P. Baudoin (e.g. in Rwanda and Burundi) earlier than in
Based on PROSEA 1: Puises. other parts of Africa.
Nowadays, common bean is a crop of global
importance, especially in North and South
PHASEOLUSVULGARIS L. (common bean) America, Europe and Africa. The crop is of sig-
nificance in many African countries and most
Protologue Sp. pi. 2: 723 (1753). intensively grown in the Great Lakes areas of
Family Papilionaceae (Leguminosae - Papil- Central Africa. In tropical Africa common bean
ionoideae, Fabaceae) is a major food crop in urban as well as rural
Chromosome number 2n = 22 areas.
Vernacular names Common bean, haricot Uses The mature dry seeds of common bean
bean, kidney bean, navy bean (En). Haricot are eaten worldwide as a pulse and the imma-
commun, haricot (Fr). Feijäo, feijoeiro (Po). ture pods and seeds as a vegetable. In tropical
Mharagwe (Sw). Africa common bean is primarily produced and
Origin and geographic distribution Com- consumed as a pulse. The nutritional value of
mon bean originated in Central and South the seeds is recognized, but common bean is
America. Small-seeded and climbing ecotypes also important for adding diversity and flavour
are found in the wild in northern Argentina to carbohydrate-rich meals, such as those
and Central America. Common bean was inde- based on maize or banana. It is the major pro-
pendently domesticated in both Central Amer- tein source in various countries, e.g. in
ica (Mexico and Guatemala) and in the South Rwanda, Burundi and Kenya. Although com-
American Andes (mainly Peru). The resulting mon bean is sometimes recognized as the 'meat
gene pools are distinct. Archaeological evidence of the poor', it is also much appreciated by
indicates that common bean was a domesti- wealthier consumers. In tropical Africa com-
cated crop already in 6000 and 5000 BC in mon bean is most typically consumed boiled,
Peru and Mexico, respectively. Common bean often with seasoning and some oil added. It
was taken to other parts of the world since the may also be mashed or made into soup. In
16th century. Portuguese traders probably in- many parts of the world the dry seeds of com-
troduced common bean to Africa from the 16 th mon bean are canned, either alone or in tomato
sauce.
The leaves of common bean are sometimes
eaten as a vegetable, e.g. during the hunger
months of the year when not much food is
available, but relatively few cultivars have
leaves of sufficient tenderness. Crop residues
are often used as fodder. In Mali a powder of
carbonized seeds is applied to wounds.
In temperate regions of the world Phaseolus
vulgaris is mainly grown for the green imma-
ture pods (French bean), which are canned,
frozen or eaten fresh. In tropical Africa imma-
ture pods are mostly produced as a market
crop. A separate article in PROTA 2: 'Vegeta-
bles' deals with vegetable aspects of Phaseolus
vulgaris.
Production and international trade Reli-
Phaseolus vulgaris - planted able production statistics for dry common bean
 PHASEOLUS 147

are difficult to obtain, as its production is often flatulence-inducing compounds.

lumped together with that of other Phaseolus Common bean starch has shown cholesterol-
species. In 2000 the annual world production of lowering effects in rats, whereas lectins have
common bean was estimated by FAO to be 8.3 shown in-vitro inhibitory action against HIV-1
million t; the largest producer is Brazil. Africa reverse transcriptase. A decoction of the pods
produces about 2 million t annually on about has shown hypoglycaemic effects in rabbits.
3.5 million ha. Large producers (100,000- Seed coat compounds (methanol extracts, tan-
600,000 ha annually) in tropical Africa are DR nin fractions and pure flavonoids) have shown
Congo, Rwanda, Burundi, Ethiopia, Kenya, antioxidant activity.
Uganda, Tanzania, Malawi, Angola and Mo- Description Climbing, trailing or erect and
zambique; smaller producers (2,000-100,000 bushy annual herb, slightly pubescent; taproot
ha) are Cape Verde, Niger, Cameroon, Sudan, well developed, with many lateral and adventi-
Zambia and Zimbabwe. Most common bean tious roots; stem up to 3 m long, angular or
produced in tropical Africa is consumed by the nearly cylindrical. Leaves alternate, 3-foliolate;
producer, but 40% may be marketed to supply stipules triangular, small; petiole up to 15(-30)
urban areas and for export with a farm gate cm long, grooved above, distinctly thickened at
value of over US$ 200 million per year. Trade base, rachis (1.5-)2.5-3.5(—6) cm long; stipels
with neighbouring countries is important. For small; leaflets ovate, (5-)7.5-14(-20) cm x 5 -
example, significant amounts move from 10(—15) cm, lateral ones asymmetrical, central
Uganda to Kenya, Rwanda and Sudan, but one symmetrical, entire, slightly pubescent, 3-
Uganda may import from these countries when veined from the base. Inflorescence an axillary
localized deficits occur. An example of longer or terminal false raceme up to 15(-35) cm long,
distance trade of common bean is from Kivu in with flowers arranged along the rachis in pairs
eastern DR Congo to Kisangani and down the or solitary. Flowers bisexual, papilionaceous;
Congo River to Kinshasha. Some common bean pedicel up to 1 cm long, slender, with ovate
production is for specialized export markets, bracteoles; calyx campanulate, tube c. 3 mm
e.g. in Ethiopia for export to Europe and the long, lobes triangular, 2-3 mm long; corolla
Middle East, and in northern Tanzania for white to pale purple or red-purple, standard
export to Europe. very broadly obovate, hood-shaped, 1-1.5 cm
Properties A typical composition of common long, wings obovate, c. 2 cm long, keel sharply
bean per 100 g edible portion is: water 11.3 g, upturned, c. 1cm long; stamens 10, 9fused and
energy 1218 kJ (291 kcal), protein 21.4 g, fat
1.6 g, carbohydrate 49.7 g, dietary fibre 22.9 g,
Ca 180 mg, Mg 180 mg, P 310 mg, Fe 6.7 mg,
Zn 2.8 mg, carotene trace, thiamin 0.45 mg,
riboflavin 0.13 mg, niacin 2.5 mg, vitamin B6
0.56 mg, ascorbic acid trace (Holland, Unwin &
Buss, 1991). The essential amino acid composi-
tion per 100 g edible portion is: tryptophan 210
mg, lysine 1540 mg, methionine 240 mg,
phenylalanine 1130 mg, threonine 860 mg,
valine 990 mg, leucine 1640 mg and isoleucine
890 mg (Paul, Southgate & Russell, 1980).
Common bean is deficient in the essential
amino acids methionine and cystine. Composi-
tion alone is not a reliable indicator of its food
value as common bean is not very digestible.
Much of the phosphorus is phytate-bound and
the protein is only 55-65% digestible. Common
bean generally requires a long cooking time
and may have 'hard-to-cook' properties, which
may be due to genetic and environmental fac-
tors. Common bean contains antinutritional
compounds, such as lectins (haemagglutinins) Phaseolus vulgaris - 1, inflorescence; 2, fruiting
and trypsin inhibitors, but both are inactivated branch; 3, seeds.
by proper cooking. It also contains tannins and Source: PROSEA
148 CEREALS AND PULSES

1 free; ovary superior, c. 0.5 cm long, laterally climbing. Flowering in common bean generally
compressed, style upturned and spiralled, with starts 28-45 days after sowing. Self-fertil-
collar of fine hairs below the ellipsoid stigma. ization is the rule, but with 1-3% outcrossing.
Fruit a linear pod up to 20 cm long, straight or Immature pods for vegetable use can be har-
more commonly curved with a prominent beak, vested 25-30 days after flowering. The seed-
fleshy when immature, green or yellow, some- filling period may take 23-50 days. The length
times red, purple or with purplish stripes, (2-) of the crop cycle ranges from 60-90 days for
5—7(—12)-seeded. Seeds globose to kidney- determinate types and may be as long as 250-
shaped, ellipsoid or oblong, 0.5-1.5(-2) cm 300 days for indeterminate climbing types.
long, black, brown, yellow, red or white, some- Several Rhizobium species fix nitrogen with
times with speckled, flecked or saddled pat- Phaseolus vulgaris, including Rhizobium legu-
terns; hilum oblong to elliptical. Seedling with minosarum bv. phaseoli, Rhizobium etli and
epigeal germination; cotyledons oblong, thick; Rhizobium tropici. The nitrogen-fixing ability
first two leaves simple and opposite, subse- ofcommon bean is often considered poorer than
quent leaves alternate, 3-foliolate. that of other pulses such as cowpea, soya bean
Other botanical information Phaseolus com- and groundnut, although fixation rates up to
prises about 50 species, most of them in the 125kg of N per ha have been recorded.
Americas. Phaseolus vulgaris is closely enough Ecology In tropical Africa common bean is
related to some other Phaseolus species, e.g. well adapted to elevations of 1200-2200 m,
Phaseolus coccineus L. (runner bean) and with mean temperatures during the growing
Phaseolus acutifolius A.Gray (tepary bean), to season of 15-23°C. Still, 20% of the common
make interspecific hybridization possible. bean production in tropical Africa takes place
Andean types of Phaseolus vulgaris tend to at a mean temperature higher than 23°C. The
have larger seeds and leaves than the Central crop can withstand occasional daytime tem-
American types. All growth habits are found in peratures of 35°C, but this often results in
each gene pool, but determinate bush types flower abortion. Growth stops below 10°C and
and climbing types are more common in the the plant is killed by frost. At latitudes higher
Andean than in the Central American pool. In than 10° Phaseolus vulgaris may be grown at
tropical Africa some genetic diversity is found low altitudes during cooler months, generally
that is not found in the Americas. with irrigation, and usually for immature pod
Erect bush bean types are most common where harvest. Common bean production occurs with
mechanical harvest is practised but also in 250 mm mean rainfall during the growing sea-
smallholder agriculture. Climbing types are son but 65% of the production is estimated to
largely restricted to high altitude areas, espe- occur in areas with an average rainfall higher
cially in south-western Uganda, Rwanda, Bu- than 400 mm during the season. Occasional
rundi and eastern parts of DR Congo, but they water deficits severely reduce common bean
are also grown in northern and western Ma- yield. More important constraints than water
lawi, northern and southern Tanzania and deficits are diseases that are favoured by hu-
northern Zambia. Indeterminate trailing or mid conditions. Common bean genotypes vary
semi-climbing types are common in most bean for photoperiod sensitivity (short day plants or
growing areas and prevail under growing con- day neutral); photoperiod sensitivity is typi-
ditions that are marginal due to high tempera- cally greater in genotypes of Andean origin
tures, water deficits and low soil fertility. than in Meso-American ones.
Red, mottled, large-seeded cultivars are most Common bean prefers medium-textured, well-
common in tropical Africa, followed by culti- drained soils over 0.5 m deep. It is sensitive to
vars with red, small to medium-sized seed. soil acidity, including the associated alumin-
Other seed types may comprise 50% of the pro- ium and manganese toxicities. The optimum
duction. Black-seeded and white-seeded culti- pH is 6.0-7.5, but most common bean produc-
vars are not popular because of the colour of tion in tropical Africa is at soil pH 5-6 and 20%
the food preparations. takes place on soils with pH below 5. Common
Growth and development For seed germi- bean production in tropical Africa occurs
nation of common bean the soil must be mostly under conditions of P deficiency. Where
warmer than 12°C, with optimal emergence Phaseolus species have not grown previously,
occurring at soil temperatures of 22-30°C. symbiotic N-fixation may be inadequate to
Plant growth habits are broadly grouped into meet the N requirement ofthe plants.
determinate or indeterminate and bush or Propagation and planting Common bean
 PHASEOLUS 149

is normally propagated by seed, but vegetative annual or short-lived perennial crops. The ro-
propagation using stem cuttings is possible. tated crops are typically cereals, other pulses
The 1000-seed weight is 150-600 g. Common and root or tuber crops.
bean may be sown by broadcasting and row Diseases and pests Common bean is ex-
planting. Sole-crop sowing rates range from tremely susceptible to diseases and pests and
150,000-400,000 seeds per ha. With intercrop- more than 50% of the production in tropical
ping, sowing rates are less than for sole crop- Africa is estimated to be lost every year. The
ping. Indeterminate climbing common bean is seedborne fungal diseases angular leaf spot
sown 3—6 seeds per planting hole in rows 100- (Phaeoisariopsis griseola) and anthracnose
120 cm apart with 40—50cm spacing within the (Colletotrichum lindemuthianum), and the
row. Seeds are normally sown 3-4 cm deep, but bacterial diseases common bacterial blight
as deep as 7 cm if the soil surface is dry and (Xanthomonas campestris pv. phaseoli) and
not too heavy or prone to crusting. Often mix- halo blight (Pseudomonas savastanoi pv.
tures of different seed types are sown, e.g. in phaseolicola, synonym: Pseudomonas syringae
Rwanda, Tanzania and Malawi. In traditional pv. phaseolicola) are each among the top con-
agriculture the land is prepared by hand or straints of common bean production. Estimated
animal traction before sowing. Cultivation is total yield loss attributed to these diseases is
mostly on the flat, but sowing on hills or ridges more than 1 million t per year in sub-Saharan
may be practised where the soil is heavy or the Africa. Angular leaf spot and anthracnose are
groundwater table is high. sensitive to many fungicides, but smallholder
Only about 30% of the common bean produc- farmers generally do not use chemicals for
tion area in tropical Africa is planted as a sole common bean disease control. Cultivars vary in
crop. Intercropping with maize, banana and their reaction to these diseases. Pre- and post-
root or tuber crops is important with these emergence damping off caused by root rot com-
intercrop associations accounting for 40-50%, plexes (Pythium aphanidermatum, Rhizoctonia
10-20% and 10-20%, respectively, of the com- solani (group AG4) and Fusarium solani f.sp.
mon bean production area. Less common is phaseoli) is very important in areas with inten-
intercropping with sorghum, millet, pea, faba sive common bean production and low-fertility
bean, coffee and other crops. Climbing cultivars soils. Improving nutrient supply and sowing
are more often produced in sole cropping than resistant or tolerant cultivars are effective
non-climbing types, but the dense foliage in methods of reducing losses to root rot. Bean
sole cropping easily creates a humid environ- common mosaic virus (BCMV) is an aphid-
ment promoting diseases. Common bean is transmitted and seedborne virus and has been
sometimes grown as a relay crop on residual estimated to cause 180,000 t yield loss per year
moisture, e.g. in Malawi and southern Tanza- in sub-Saharan Africa. Resistance to BCMV is
nia. controlled by a single dominant gene, but this
Management For climbing cultivars of com- gene causes susceptibility to bean common
mon bean, 2 m high poles (usually straight mosaic necrosis virus (BCMNV; also known as
branches or stems ofbamboo orPennisetum) are black root) that is indigenous to Africa. These
placed after emergence to support the plants. closely related viruses each have more than
The crop is usually weeded once or twice, after one pathogenicity group. Resistance to all
which its canopy is sufficiently developed to groups can be achieved by deploying 2 or more
suppress weeds. Earthing-up is often done at recessive genes. Sowing of disease-free seed
about 3 weeks after sowing. This should be can be useful in control of seedborne diseases,
done carefully, because common bean is liable but such seed is scarce. Bean rust (Uromyces
to damage to the collar of the plant. Irrigation appendiculatus), Ascochyta blight (Phoma exi-
is uncommon except at higher latitudes with gua), powdery mildew (Erysiphe polygoni),
winter (dry season) production. Common bean floury leaf spot and web-blight (Rhizoctonia
is rarely fertilized in tropical Africa although N solani group AGI) together may cause yield
and P deficiencies are major constraints. Ade- losses of 600,000 t/year in sub-Saharan Africa.
quate P nutrition is important for symbiotic N- The most important insect pests are bean flies
fixation and there is often economical response or bean stem maggots (Ophiomyia spp.), espe-
to 20 kg N and 22 kg P per ha. Although much cially at early growth stages and when plants
of the common bean production is on acid soils, are stressed by water and nutrient deficits.
the use of lime to amend these soils is uncom- Bean flies can be controlled by treating seed
mon. The crop is grown in rotation with other with a systemic insecticide, such as imidaclo-
150 CEREALS AND PULSES

prid or endosulphan, either as a seed dust or as ers transport the common bean harvest from
a spray shortly after seedling emergence. In the field to their home to be spread on the
Africa cutworms (Agrotis spp.) and caterpillars ground and dried in the sun. After drying,
(Spodoptera spp.) may be a problem especially threshing may be by beating with long sticks,
in soils amended with farmyard manure, a driving over heaps of harvested plants with a
common practice by smallholder growers. tractor, or, less commonly, with a threshing
Thrips (Frankliniella occidentalis, Frank- machine. Before being stored, common bean
liniella schultzei and Megalurothrips sjostedti) seed is often dried in the sun to destroy
and pod borers (Helicoverpa armigera, Maruca bruchids and to reduce moisture content for
testulalis and Clavigralla spp.) cause 80,000— better storage. Prolonged drying can, however,
90,000 t and 130,000-140,000 t yield loss per induce a hard-to-cook condition. In some re-
year, respectively, in sub-Saharan Africa. gions seed is sorted to lots of single seed types,
Thrips, particularly Frankliniella occidentalis, while elsewhere complex mixtures of seed
are difficult to control as they are resistant to types are intentionally produced and con-
many commonly used pesticides. Pod borers sumed. Seeds may be stored with wood ash,
are easily controlled by Bacillus thuringiensis tobacco leaves or ash from bean stems.
products. Aphids (Aphis fabae and Aphis crac- Genetic resources Common bean is threat-
civora) are among the top 10 constraints to ened by genetic erosion due to non-traditional
common bean production and are worse under farming practices where relatively few geno-
dry conditions. Ootheca foliage beetle causes types are produced in pure stands, and, espe-
widespread damage in sub-Saharan Africa. cially in Latin America, to displacement of
Whitefly (Bemisia tabaci), a Madagascan bean common bean by more profitable crops. In-situ
leaf roller called 'cigarier' (Apoderus humeralis) conservation can be of importance especially in
and painted lady (Vanessa cardui, synonym: countries like Rwanda where many landraces
Pyrameis cardui) are of local importance. are found under diverse conditions and where
Bruchids (Zabrotes subfasciatus and Acan- they are often grown in complex mixtures of as
thoscelides obtectus) are major pests of stored many as 20 seed types.
common bean and have been attributed with The largest ex-situ collection of Phaseolus is at
being the sixth main cause of yield loss the International Center for Tropical Agricul-
(250,000 t/year) in sub-Saharan Africa. Pest ture (CLAT) near Cali, Colombia. It holds over
management typically involves the integration 40,000 accessions of which over 35,000 are of
of several low-cost practices including crop Phaseolus vulgaris. This was estimated to ac-
rotation, intercropping, sowing of resistant or count for 50-75% of the variability occurring in
tolerant cultivars, and insecticide use. the centres of diversification for domesticated
Harvesting Common bean may be har- types, but only less than 30% of diversity of
vested while most pods are still green but near wild types. Germplasm collections held in Af-
physiological maturity, for an early harvest of rica include: Bunda Agricultural College, Li-
a fresh, easy to cook pulse product, but most longwe, Malawi (6000 accessions), National
crops are harvested when mature. In tropical Genebank of Kenya, KARI, Kikuyu (3000 ac-
Africa harvesting is nearly all by hand. Non- cessions) and Institut des Sciences Agronomi-
climbing common bean plants are usually up- ques du Rwanda, Butare (3000 accessions).
rooted when most of the pods are dry, bundled, African national breeding programmes (e.g. in
and carried home. Pods of climbing types are Uganda) have smaller landrace collections.
normally harvested by hand as they mature, Breeding Common bean breeding programs
with repeated harvests over several weeks. in Africa and elsewhere have as their goal to
Yield Average common bean yields are improve yield potential; much of the progress is
about 1.5 t/ha in Europe and industrialized through improved tolerance or resistance to
countries of Asia, 1t/ha in North America and biotic and abiotic constraints. Improved resis-
0.7 t/ha worldwide. Average yields in tropical tance to diseases has been the main breeding
Africa are often around 0.6 t/ha. Under the goal and much success has been achieved, al-
best growing conditions, yields of 2.5 and 5 t/ha though resistance is often not durable due to
for non-climbing and climbing types, respec- genetic diversity and adaptive ability of the
tively, are achievable. Under irrigation in Ma- pathogens. Resistance to common bacterial
lawi, for instance, yields of 3.8 t/ha have been blight has been introduced from Phaseolus
obtained. coccineus and Phaseolus acutifolius. Improved
Handling after harvest Smallholder farm- insect resistance has been another major
 PlSUM 151

breeding goal. Wild Phaseolus species have mand is even likely to increase as population
been useful sources of genes, such as for resis- increases. Common bean production in tropical
tance to the bruchid Zabrotes subfasciatus. Africa is constrained by susceptibility to dis-
Breeding for tolerance to abiotic stresses has eases and pests. Breeding for resistance to or
gained in importance and lines with superior tolerance of diseases and pests has achieved
tolerance to acid soils, and others efficient in N considerable progress, but work remains to be
or P use, have been released in Africa. Promis- done as resistance is often not durable. Breed-
ing progress in breeding for drought tolerance ing for better tolerance to abiotic stresses, such
is being achieved where deep root systems are as aluminium toxicity, and for more efficient
combined with efficient transport of carbohy- use of inadequate soil water and nutrients, is
drates from leaves to seed under drought also necessary to improve production both on
stress. Progeny produced from crosses between marginal and on productive soils. Biotechno-
the Andean and Meso-American gene pools logical tools will play an increasingly important
typically is weak and of low productivity, but role in common bean breeding, e.g. the use of
breeders have developed superior lines and molecular mapping to locate resistance genes.
parents through inter-pool crosses which have Major references Abate & Ampofo, 1996;
superior traits from each pool. Allen, Buruchara & Smithson, 1998; Baudoin
Resource-constrained breeding programmes in et a l , 2001; Gepts & Debouck, 1991; Hidalgo,
Africa have benefited from regional and inter- 1991; Messiaen & Seif, 2004; Popelka, Terryn
national collaborative efforts, such as from & Higgins, 2004; Shellie-Dessert &Bliss, 1991;
germplasm generated at CIAT. Smartt, 1989b; Wortmann et a l , 1998.
Efficiency of breeding common bean is improv- Other references Beninger & Hosfield,
ing with increased use of molecular markers. 2003; Chacon S., Pickersgill & Debouck, 2005;
The 'Phaseomics' initiative facilitates collabo- Debouck & Smartt, 1995; Freytag & Debouck,
ration among research institutions in the de- 2002; Fukushima et al., 2001; Giller, 2001;
velopment of a cDNA library and sequencing of Gillett et a l , 1971; Graham & Ranalli, 1997;
the common bean genome. In-vitro regenera- Hanelt & Institute of Plant Genetics and Crop
tion of common bean for breeding purposes is Plant Research (Editors), 2001; Hidalgo &
possible using different expiants, including Beebe, 1997; Holland, Unwin & Buss, 1991;
shoot tips, petioles, seedlings, embryonic axes, Johnson, Pachico & Wortmann, 2003; Kay,
cotyledons, seedling nodes and meristematic 1979; Mackinder et al., 2001; Martinez Ro-
calli. No confirmed reports exist of stable mero, 2003; Paul, Southgate & Russell, 1980;
transgenic common bean plants based on Agro- Qi, Smithson & Summerfield, 1998; Roman-
bacterium tumefaciens systems, but transgenic Ramos, Flores-Saenz & Alarcon-Aguilar, 1995;
plants have been obtained by particle bom- Wang &Ng, 2000; Westphal, 1974.
bardment. However, the most efficient way to Sources of illustration Smartt, 1989b.
improve common bean with gene technology is Authors C S . Wortmann
probably to use Phaseolus acutifolius, which
can be routinely transformed using Agrobacte-
rium, and to cross the resulting transgenic PlSUMSATIVUM L.
plants with common bean using embryo rescue
techniques. Protologue Sp. pi. 2: 727 (1753).
Prospects Common bean is the most con- Family Papilionaceae (Leguminosae - Papil-
sumed pulse globally and a very important ionoideae, Fabaceae)
crop in tropical Africa, especially in Central, Chromosome number 2n = 14
East and southern Africa, both for its nutri- Vernacular names
tional value and its market potential. It is es- - Field pea, pea (En). Pois, pois sec (Fr). Ervil-
pecially important to smallholder farmers and ha (Po).Njengere, njegere (Sw).
women, who often are responsible for the com- - Garden pea, pea, petit-pois (En). Petit pois
mon bean crop. Regional trade of common bean (Fr). Ervilha (Po). Njengere, njegere (Sw).
is of economic significance for some countries - Sugar pea, pea pod, snap pea, snow pea,
and there is also some common bean produc- mange-tout (En). Pois mangetout, pois gour-
tion in tropical Africa for export to Europe and mand (Fr). Ervilha torta (Po).
the Middle East. There is no reason to expect Origin and geographic distribution The or-
that the importance of common bean will de- igin and progenitors of Pisum sativum are not
crease in the future; in tropical Africa the de- well known. The Mediterranean region, west-
152 CEREALS AND PULSES

seeds; garden pea, grown for the immature

green seeds; and sugar pea, grown for the im-
mature pods. The dry seeds of field pea are
first soaked in water to soften and are then
boiled and consumed as a pulse dish. Alterna-
tively, they are decorticated and split ('split
peas') before boiling. They are also consumed
roasted. The young pods of sugar pea are boiled
for a few minutes only, to preserve their crisp-
ness; after boiling they may be stir-fried before
consumption. The young seeds of garden pea
are also boiled for a few minutes. They are
commonly offered as canned or - in Western
countries — as deep frozen products. In Ethio-
pia the annual consumption per person of pea
seeds is estimated at 6-7 kg. Main dishes in-
clude 'shiro wot' (split pea seeds ground and
Pisum sativum -planted made into stew) and 'kik wot' (split pea seeds
boiled and made into stew). Snacks include
ern and central Asia, and Ethiopia have been 'eshet' (fresh green field pea seeds either eaten
indicated as centres of origin. Recently FAO raw or roasted), 'nifro' (boiled dry or fresh
designated Ethiopia and western Asia as cen- green pea seeds) and 'endushdush' (seeds
tres of diversity, with secondary centres in soaked first and then roasted). In local markets
southern Asia and the Mediterranean region. white- and cream-coloured seeds are preferred
Archaeological evidence of the use of pea dat- for 'kik'-making, and grey-coloured seeds for
ing from 8000 BC has been found in the Fertile 'shiro'-making. In Malawi and some Asian
Crescent. The first cultivation of pea appears countries, the young shoots are used as a leafy
to have been in western Asia, from where it vegetable. In Western countries dry, mature
spread to Europe, China and India. In classical pea seeds are extensively used as animal feed.
times Greek and Roman authors mentioned its The haulms or straw after threshing are used
cultivation as a pulse and fodder crop. Pea was as forage, hay, silage and green manure. Apart
already well known in the mountain regions of from being an important source of food and
Central and East Africa before the arrival of feed, pea plays a role in soil fertility restoration
the Europeans and was a well-established and as a suitable rotation crop that fixes atmos-
important food crop in Rwanda and south- pheric nitrogen.
western Uganda by 1860. The use of the edible The seeds of pea are claimed to have beneficial
pods was first described in the Netherlands effects on many types of skin complaints; face
and France during the 16th century, whereas masks made from crushed seeds are used to
the use of immature seeds as a vegetable began treat acne and wrinkled skins.
in Europe a century later. Production and international trade FAO
At present, Pisum sativum is found in all tem- estimated the annual world dry pea seed pro-
perate countries and in most tropical high- duction in 1999-2003 at about 10.5 million t
lands. Field pea is extensively grown in the from 6.2 million ha. The main producers are
highlands of eastern Central Africa and East Canada (2.1 million t/year from 1.1 million ha),
Africa (notably Ethiopia), and in southern Af- France (1.9 million t/year from 400,000 ha),
rica. In parts of Rwanda and Uganda it is the China (1.1 million t/year from 900,000 ha) and
main pulse crop. Field pea is hardly grown in the Russian Federation (1.1 million t/year from
West Africa. In Africa garden pea and sugar 700,000 ha). The annual production in tropical
pea are mostly considered exotic products. Africa for this period was about 310,000 t from
They are regionally of some importance, sugar 470,000 ha. Here, the main producers are
pea more in Francophone countries, garden pea Ethiopia (135,000 t/year from 184,000 ha), DR
more in Anglophone countries. Imported Congo (65,000 t/year from 96,000 ha), Burundi
canned garden pea seeds are available every- (32,000 t/year from 49,000 ha), Tanzania
where in food shops. (28,000 t/year from 63,000 ha), Uganda (18,000
Uses Three main types of pea cultivars can t/year from 29,000 ha) and Rwanda (14,000
be distinguished: field pea, grown for the dry t/year from 30,000 ha). The annual world pro-
 PISUM 153

duction of green pea seeds in 1999-2003 was Raw garden pea seeds, immature taken from
about 8.7 million t from 1.0 million ha, the the pods (refuse 63%) contain per 100 g edible
main producers being India (3.4 million t/year portion: water 74.6 g, energy 348 kJ (83 kcal),
from 300,000 ha), China (1.5 million t/year protein 6.9 g, fat 1.5 g, carbohydrate 11.3 g
from 190,000 ha) and the United States (1.0 (starch 7.0 g), fibre 4.7 g, Ca 21 mg, Mg 34 mg,
million t/year from 96,000 ha). In tropical Af- P 130 mg, Fe 2.8 mg, Zn 1.1 mg, carotene 300
rica about 30,000 t green pea seed was pro- (Xg,thiamin 0.75 mg, riboflavin 0.02 mg, niacin
duced annually from 6400 ha, mainly in Kenya 2.5 mg, folate 62 u,g, ascorbic acid 24 mg.
(23,000 t/year from 5600 ha). Raw sugar pea pods, with the ends trimmed
Statistics on the international trade in pea (refuse 8%) contain per 100 g edible portion:
seed are generally scanty, as they are mostly water 88.7 g, energy 134 kJ (32 kcal), protein
aggregated in 'pulse crops' as a whole. The 3.6 g, fat 0.2 g, carbohydrate 4.2 g (starch 0.8
main exporting countries are Canada, Austra- g), fibre 4.2 g, Ca 44 mg, Mg 28 mg, P 62 mg,
lia, France and China. Canada focuses on the Fe 0.8 mg, Zn 0.5 mg, carotene 695 p.g, thiamin
European stock feed market and in recent 0.2 mg, riboflavin 0.15 mg, niacin 0.6 mg, folate
years on the food market in India. Australia 10 |lg, ascorbic acid 54 mg (Holland, Unwin &
focuses on the food markets and the domestic Buss, 1991).
feed market. The top importers for pea feed or Description Annual, climbing, glabrous
food are Spain, Bangladesh, Belgium, India, herb up to 2(-3) m tall (up to 1.3 m for sugar
China, United States, Colombia, United Arab pea types); taproot well developed, up to 1.2 m
Emirates and Malaysia. Almost all the produc- long, with many lateral roots; stem terete, with
tion in Ethiopia is consumed locally. Most no or few basal branches, internodes hollow.
sugar pea pods produced in the world are sold Leaves alternate, pinnate, with l-3(-4) pairs of
in local markets. Western countries import leaflets and ending in a usually branched ten-
large quantities of sugar pea pods from devel- dril; stipules leaf-like, up to 8(-10) cm x 4 cm;
oping tropical countries because locally pro- petiole (2-)4-6(-7.5) cm long; leaflets shortly
duced ones are available for only a short time stalked, ovate to elliptical, 1.5-8 cm x 0.5-4
ofthe year and because of the high labour costs cm, entire to toothed, sometimes converted into
of picking. Kenya exports yearly 4500 t sugar
pea pods to the European Union. Garden pea
seeds are mostly exported as canned or frozen
products from Western countries, e.g. the
United States and France, but statistical data
are not available.
Properties Whole mature dried seeds of
field pea contain per 100 g edible portion: wa-
ter 13.3 g, energy 1269 kJ (303 kcal), protein
21.6 g, fat 2.4 g, carbohydrate 52.0 g (starch
47.6 g), fibre 15.0 g, Ca 61 mg, Mg 120 mg, P
300 mg, Fe 4.7 mg, Zn 3.7 mg, carotene 245 ug,
thiamin 0.6 mg, riboflavin 0.3 mg, niacin 3.0
mg, vitamin Be 0.13 mg, ascorbic acid trace
(Holland, Unwin & Buss, 1991). The content of
essential amino acids per 100 g food is: trypto-
phan 210 mg, lysine 1620 mg, methionine 210
mg, phenylalanine 1000 mg, threonine 860 mg,
valine 1000 mg, leucine 1480 mg and isoleucine
930 mg (Paul, Southgate & Russell, 1980). The
composition of wrinkled pea seeds is different
from rounded ones; they have less starch (27-
37 g) and more fat (5 g) and sugars. Antinutri-
tional factors in pea seeds include trypsin in-
hibitors, haemagglutinins (lectins), tannins,
oligosaccharides and phytate. Cultivars with a Pisum sativum- 1, shoot with flower; 2,part of
darker seed coat contain more tannin, which shoot with fruit; 3,seed.
tends to decrease their digestibility. Source: PROSEA
154 CEREALS AND PULSES

tendrils. Inflorescence an axillary, 1-3-flow- of the flowering period is 2—3 weeks in culti-
ered raceme. Flowers bisexual, papilionaceous; vars for mechanical harvesting, up to one
calyx with tube 4-8 mm long, lobes as long or month in garden cultivars. For field peas the
longer than tube; corolla white to purple, stan- period from emergence to dry seed harvest
dard 1-3 cm x 2.5—4.5 cm, wings a little ranges from 3—6months depending on cultivar
shorter than standard, keel much shorter; and environment. Most field pea cultivars
stamens 10, 9 united and 1 free; ovary supe- grown in Africa have an indeterminate growth
rior, 1-celled, style curved, longitudinally habit. In a 2-season experiment with 63 geno-
grooved. Fruit an oblong-ovate pod 3.5-15 cm x types in Ethiopia at 3000 m altitude, the period
1-2.5 cm, pendant, 2-11-seeded. Seeds globose, to flowering and maturity ranged from 80—104
sometimes wrinkled, 5-8 mm in diameter, days and 149-163 days, respectively. Pea flow-
varying in colour from uniform yellow (sugar ers are self-pollinated, with usually less than
pea), green (crinkled garden pea) to purple or 1% outcrossing. Pea is nodulated by Rhizobium
spotted or cream-white, sometimes with black leguminosarum.
hilum. Seedling with hypogeal germination; Ecology Pea requires a relatively cool cli-
cotyledons remaining within testa; first 2 mate, with average temperatures between 7—
leaves simple. 24°C, and with optimum yields at average
Other botanical information Pisum com- temperatures of 13—21°C, although maximum
prises a few species and is related to Lathyrus, rates of development and vegetative growth are
Lens and Vicia, from which it can be distin- reached at considerably higher temperatures.
guished by its terete stems, very large stipules It can be grown at elevations above 1000 m
and longitudinally grooved style. near the equator, or at lower elevations (even
Pisum sativum has long been studied by ge- in coastal areas) during the cool season at lati-
neticists; Knight did his crossing experiments tudes between 15—20°.Young plants can with-
on it in 1787, and it was the subject of the pio- stand frost if progressively hardened by lower-
neering work of Gregor Mendel in the 19thcen- ing temperatures. Pisum sativum is grown in
tury. Within Pisum sativum several varieties areas with an annual rainfall as low as 400
or subspecies have been distinguished. A clas- mm, but the optimum is 800—1000 mm/year. It
sification in cultivar groups is more appropri- is slightly susceptible to daylength, with long
ate. Sativum Group is cultivated worldwide, days promoting flowering. In most tropical
including tropical Africa. Abyssinicum Group circumstances it can be considered day-neutral.
(Abyssinian pea) is cultivated in the northern In Ethiopia rainfed field pea is grown at 1800—
(Tigray and Wollo) and south-eastern (Arsi) 3000 m altitude, because it suffers from dis-
parts of Ethiopia; it is also grown in Yemen. eases and drought at lower altitudes and from
The latter differs in having leaves with only frost at higher altitudes. It is mostly grown in
one pair of leaflets (Sativum Group: 2-3 pairs), the main rainy season (June-December). In
and smaller, red-purple flowers. It has slightly Uganda pea plants grow best at altitudes above
glossy seeds with a black hilum; these may 1800 m, and in Kenya optimum yields are ob-
mature earlier. Other cultivar groups, varieties tained at 2100-2700 m altitude. Pea grows on
or subspecies occur outside Africa; 2 of these a wide range of soil types with moderate fertil-
represent wild populations from southern ity levels, well drained and with pH 5.5—7.0,
Europe and western Asia. although some cultivars tolerate a pH up to
Purple coloured flowers are associated with 7.5. It is seriously affected by soil acidity, alu-
bitter tasting green seeds. For this reason minium toxicity and waterlogging.
nearly all garden pea cultivars are white- Propagation and planting Pea is propa-
flowered, while most field pea cultivars are gated by seed. The 1000-seed weight ranges
purple-flowered and sugar pea cultivars can from 100 g to 500 g. Sugar pea is sown in dou-
have white or purple flowers. ble rows 10 cm apart with 60 cm (30-80 cm)
Growth and development Pea seeds ger- between the double rows. Within the rows the
minate at ambient temperatures of between4— seed of small cultivars is sown 3—5 cm apart,
24°C, with 13-18°C being optimal. In sugar for taller cultivars up to 10 cm apart. Garden
pea cultivars flowers appear between the 6 th pea is sown rather densely, with plant densi-
and 12th nodes according to cultivar earliness, ties up to 80 plants per m2. The seed should be
normally 5—7 weeks after emergence. At opti- sown 4-7 cm deep. Per ha 60-200 kg of seed is
mum temperatures, pods are ready for harvest- required, with the highest rates for garden pea.
ing 12 days later. For garden pea the duration Field pea is mostly broadcast in Africa. Even
 PlSUM 155

though it does not require a fine seedbed, 2-3 150 kg K and 30 kg Mg per ha. Irrigation is
ploughings with animal-drawn ploughs or one necessary in dry conditions, e.g. 10 mm twice a
disc ploughing followed by two disc harrowings week.
may be beneficial. Timely sowing is essential Diseases and pests Ascochyta blight is a
for optimum yields, since late-sown crops are disease complex caused by Ascochyta pisi, My-
often affected by low moisture availability and cosphaerella pinodes (Ascochyta pinodes), and
heavy aphid infestation at medium altitudes Phoma medicaginis (Ascochyta pinodella); it is
and by frost at high altitudes. In Ethiopia field widespread throughout the world. It is fa-
pea is produced either as a sole crop or in voured by frequent rains and high humidity.
mixed cropping with other crops, e.g. faba bean Moderate levels of resistance have been de-
(Vicia faba L.). In the latter case, faba bean tected in landraces and in the related Pisum
provides physical support and good aeration to fulvum Sibth. & Sm., which occurs wild in
field pea, whereas field pea suppresses weed western Asia. Powdery mildew caused by Ery-
growth. In Ethiopia mixed cropping of field pea siphe pisi is widespread and important wher-
with faba bean significantly slows down the ever pea is grown. Resistant cultivars have
rate of Ascochyta blight development and re- been developed. In sugar pea a recessive resis-
sults in higher yields than pure stands. In the tance gene is present in the cultivar 'Manoa
Kilimanjaro region of Tanzania pea is grown Sugar' bred in Hawaii. Bacterial blight (Pseu-
during the cool season in association with crops domonas syringae pv. pisi) is common where
such as coffee, banana, tomato and maize. The pea is grown intensively and humidity is high.
same practice is found in parts of Kigezi Dis- Downy mildew (Peronospora viciae) may de-
trict of Uganda. In Malawi (Ntheu District) it velop at high altitudes where temperatures are
is also grown during the cool season, mostly as between 1°C and 18°C. As Ascochyta blight,
a garden crop in mixed stands with other crops, powdery mildew, bacterial blight and downy
notably wheat. In the tropics, e.g. Rwanda and mildew are seedborne, the use of certified dis-
south-western Uganda, field pea is often the ease-free seed is essential. If own seed is to be
first crop after a fallow period. In temperate used, it may be treated with a systemic fungi-
areas sugar pea is sown either in autumn or in cide to control Ascochyta blight and powdery
early spring. mildew. In addition, wide row spacing, eradica-
Management Sugar pea plants are nor- tion of weeds, surface irrigation and rotations
mally supported. The stems are not twining, of three years or longer help to manage bacte-
but grasp the support with their tendrils. They rial blight and other diseases. Aphanomyces
do not need vertical poles, but the poles can be root rot (Aphanomyces euteiches) is a major
crossed, or the plants are supported by wire root pathogen of pea worldwide. It is extremely
mesh, horizontal wires, vertical lattices or nets, difficult to control, as no effective fungicides
depending on the potential height of the culti- are available. The development of resis-
var grown. Garden pea is seldom supported, tance/tolerance to this disease will be neces-
field pea not at all. Weeds should be rigorously sary for effective control. Another important
controlled. The critical period of weed competi- soilborne disease is Fusarium wilt caused by
tion is 3—8weeks after emergence. Both annual Fusarium oxysporum f.sp. pisi, but cultivars
and perennial grasses affect field pea. Weeds resistant to this disease are available. Aphid-
can be controlled by hand weeding where la- transmitted virus diseases include bean yellow
bour is cheap, whereas chemical weed control mosaic virus (BYMV), pea seedborne mosaic
is more practical in large-scale production. virus (PSbMV), pea leaf-roll (BLRV -bean leaf-
Early land preparation can encourage weed roll luteovirus) and pea enation mosaic virus
seeds to germinate so that they can be de- (PEMV). Recent sugar pea cultivars bred in
stroyed in subsequent cultivation. southern France are relatively tolerant to se-
Field pea normally needs no fertilizer N as the vere infestation by these viruses (e.g. 'Super-
amount present in the soil and fixed by the mangetout' compared to the traditional 'Ca-
plant is sufficient. The total uptake of a crop rouby de Maussane'). The pea cyst nematode
yielding 5-6 t of seed per ha is 30-35 kg/ha P (Heterodera goettingiana) can cause consider-
and 200—250 kg/ha K. Young sugar pea and able crop loss; control measures are crop rota-
garden pea respond well to a starter dose of N tion and the use of chemicals.
fertilizer, even when nodulation occurs. An Insect pests attacking pea include cutworms
indicative fertilizer recommendation on light (Agrotis spp.), aphids (including the pea aphid
medium-rich alkaline soils is 40 kg N, 50 kg P, Acyrthosiphon pisum, a vector of many virus
156 CEREALS AND PULSES

diseases, which has become a major pest in or reduces attack by moulds and insects. The
Ethiopia and Uganda), bollworms (Heliothis seed should be stored in a dry and cool place,
armigera and Spodoptera exigua) and the pea free of pests and protected from absorbing
weevil (Bruchus pisorum). Bruchids (Calloso- moisture from the surroundings. In tropical
bruchus spp.) are a major storage pest of field Africa, e.g. in Ethiopia, pea seed is not stored
pea, e.g. in Ethiopia. for more than one season because of insect
The parasitic weed Orobanche crenata Forssk. damage, particularly by bruchids. Small-scale
causes crop losses in pea in the Mediterranean farmers do not commonly use insecticides. Bins
region. made of earth (smeared with cow dung) or
To control insect pests and diseases, integrated wooden materials (sealed with mud) are the
pest management (IPM) is recommended: use most commonly used storage structures in
of resistant/tolerant cultivars; use of certified tropical Africa. Sugar pea pods can be kept for
disease-free seed or seed treatment of own only 2—3days at temperatures of 20—25°C, but
seed; keeping fields weed-free; appropriate for more than 15 days at 2.5-5°C in perforated
fertilizing and irrigation; growing pea for seed plastic bags or crates covered with perforated
in semi-arid and/or arid areas; regular moni- plastic sheets. Garden pea seeds may be kept
toring ofthe crop; and judicious use ofbiocides. for 1—3 weeks at temperatures of 0-4°C and a
Harvesting Sugar pea pods and garden pea relative humidity of 88-92%.
seeds are ready for harvesting 8-12 weeks Genetic resources A large genetic diversity
from sowing, field pea seeds one month later. has been found in Pisum sativum collections
Pods of sugar pea are hand-picked every sec- from both Africa (e.g. Ethiopia) and Asia (e.g.
ond day during a 15-20 day period. Garden pea India). Genetic erosion in field pea is probably
seeds are either handpicked or — in large scale less than in cereals, because of less progress in
production for canning - machine-harvested. cultivar development and hence less replace-
Late harvesting of field pea may result in ment of landraces by a few new cultivars.
shedding and rotting of pods and shattering of Many germplasm collections of pea are held all
the seeds. Therefore, harvesting should be done over the world. The world collection of cultivars
at the appropriate stage: when the leaves begin and mutant forms of Pisum sativum is housed
to yellow, the lower pods begin to wrinkle, and at the Nordic Gene Bank, Alnarp, Sweden
the seed moisture content is reduced to 16- (about 2700 accessions). Emphasis in the col-
18%. In most parts of Africa where the time of lection is on lines with multiple disease resis-
harvest more or less coincides with the start of tance, wild and primitive types, lines carrying
the dry season, it is easy to achieve low mois- structural mutations, breeding lines and culti-
ture contents while the crop is still in the field. vars of special interest. Large Pisum sativum
Most field pea cultivars have an indeterminate collections are held in Australia (Australian
growth habit and the pods do not mature si- Temperate Field Crops Collection, Horsham,
multaneously. Therefore, the harvested plants Victoria, 6300 accessions), the Russian Federa-
should be dried before threshing. In most parts tion (N.I. Vavilov All-Russian Scientific Re-
of Africa (e.g. Ethiopia), harvesting of field pea search Institute of Plant Industry, St. Peters-
is done with sickles, the crop is transported to burg, 6200 accessions), Italy (CNR - Istituto di
threshing ground and stacked for a few days to Genetica Végétale, Bari, 4100 accessions), the
dry in the sun. The stack is then spread on the United States (Western Regional Plant Intro-
ground and threshed usually by beating with duction Station, Pullman, 3500 accessions;
sticks or by trampling with animals. Horticultural Sciences Department, NY State
Yield Yields offield pea range from less than Agricultural Experiment Station, Geneva, 2500
1 t/ha in Africa and South America to over 4 accessions), China (Institute of Crop Germ-
t/ha in Europe. The average world yield is plasm Resources (CAAS), Beijing, 3400 acces-
around 1.7 t/ha. Under good growing conditions sions), and the United Kingdom (John Innes
sugar pea yields of up to 8 t/ha edible pods per Centre, Department of Applied Genetics, Nor-
ha can be obtained. Garden pea may produce wich, 2700 accessions). The largest collection of
4-7 t/ha young seeds. Pisum sativum germplasm in Africa is located
Handling after harvest The initial seed at the Institute of Biodiversity Conservation,
moisture content of field pea must be reduced Addis Ababa, Ethiopia, with over 1600 acces-
to the required level of about 12% before stor- sions.
age. Optimum moisture content reduces the Breeding All commercial cultivars of Pisum
deterioration rate during storage and prevents sativum are pure lines. The main breeding
 PlSUM 157

objectives in temperate regions are colour and crease yields by higher plant densities and to
quality for fresh product markets and canning, make fruit picking easier. Well-known culti-
mechanization and cold tolerance. Breeding in vars of sugar pea in Africa are 'Sugar Snap',
most parts of the tropics has an improved seed 'Carouby de Maussane', 'Oregon Sugar Pod',
yield as a first priority through the develop- 'Shield' and 'Sugar Queen'. Some cultivars of
ment of productive cultivars tolerant/resistant garden pea are 'Alderman', 'Télévision' and
to different stress factors and suitable for dif- 'Green Feast'. Many growers use their own
ferent agro-ecological conditions. Some pro- seed originating from old introductions.
gress has been made. In addition to improved A consensus genetic linkage map has been de-
yield potential, sources of resistance to pow- veloped for Pisum sativum based on various
dery mildew have been identified. Attempts to linkage maps. Quantitative trait loci associated
transfer resistance to Ascochyta blight from a with, among others, seed yield, seed protein
wild type have not been successful because of concentration, early maturity, lodging resis-
complications due to polygenic inheritance and tance, plant height and resistance to various
linkage with other traits. The presence of biotic stresses (including Ascochyta blight,
physiological races of the pathogens is another Aphanomyces root rot and Orobanche crenata)
problem. Manipulation of morphological traits have been identified. Procedures for direct as
has resulted in determinate types with even well as indirect, callus-mediated somatic em-
maturity, suitable for mechanization and semi- bryogenesis of pea have been developed for
leafless types with reduced lodging. A peculiar breeding purposes. Transgenic plants have
mutant character, 'Afila', with tendrils in the been produced using Agrobacterium-based
place of leaflets has been introduced in com- transformation vectors, e.g. to increase resis-
mercial dwarf field pea cultivars. Breeding tance to Callosobruchus chinensis, Calloso-
efforts during the past three decades in Africa bruchus maculatus and Bruchus pisorum by
have resulted in the release of a number of incorporating oc-amylase-inhibiting capacity
cultivars (obtained by introduction, hybridiza- from Phaseolus vulgaris.
tion and local selection), but most farmers still Prospects Field pea will remain important
use their own farm-saved seed of local culti- in Central and East Africa, as well as in tem-
vars; well-known cultivars are 'Mitali' and perate areas. It is a major and cheap source of
'Miseriseri'. In Ethiopia more than 15 culti- protein, fixes atmospheric nitrogen, and plays
vars, with superior yield potential, seed size, an important role in farming systems by break-
seed colour and disease resistance, have been ing cereal monoculture. A drawback is its sus-
released for different agro-ecological condi- ceptibility to diseases, which can best be coun-
tions. These cultivars include 'Holetta' (from teracted by the development of resistant culti-
local collection), 'Tegegnech' (introduced from vars. As a potential export crop, it might repre-
Burundi), 'Hassabe' and 'Markos' (introduced sent a special opportunity in the years to come
from the International Center for Agricultural and the major pea-producing countries of tropi-
Research in the Dry Areas, ICARDA) and 'Adi', cal Africa could benefit from African (Morocco
'Milky' and 'Wolmera' (obtained by hybridiza- and Sudan), European (Netherlands, France
tion of adapted local cultivars with introduc- and Greece), Middle Eastern (Israel and
tions from the United States and ICARDA). Yemen) and Asian (India and Pakistan) mar-
For sugar pea breeding the most urgent objec- kets. Sugar pea and garden pea will become
tive is powdery mildew resistance (available in gradually more important in city markets in
'Manoa Sugar') and to a lesser extent As- tropical Africa. Sugar pea is often considered a
cochyta resistance (from green pea cultivars). tastier vegetable than French bean and it could
The 'edible pod' character (absence of 'parch- be interesting to develop its production for the
ment' in the pod walls) is induced by two reces- domestic African market and for export. Gar-
sive genes. A mutation inducing thickening of den pea could be produced locally on a larger
this wall of up to 3 mm was recently introduced scale to replace imports in canned form.
in American cultivars, giving rise to the 'sugar Major references Cousin, 1992; Davies,
snap pea'. The 'sugar snap' character will be 1989; Ellis & Poyser, 2002; Kay, 1979; Knight
interesting if it appears attractive to consum- (Editor), 2000; Kraft & Pfleger, 2001; Nadol-
ers. It might also be interesting to introduce ska-Orczyk & Orczyk, 2000; Telaye et al. (Edi-
more new characters into sugar pea, e.g. true tors), 1994;Thulin, 1989a; Westphal, 1974.
dwarfs which could be grown without support, Other references Aburjaj &Natsheh, 2003;
or climbing semi-leafless types in order to in- AVRDC, 1992; FAO, 1998; Griga, 2002; Hanelt
158 CEREALS AND PULSES

& Institute of Plant Genetics and Crop Plant from weeds. Rye has been spread to all conti-
Research (Editors), 2001; Hebblethwaite, nents, especially to areas with temperate grow-
Heath & Dawkins, 1985; Holland, Unwin & ing conditions. Occasionally it is cultivated at
Buss, 1991; INRA, 2000; Kalloo, 1993; Kraft, high elevations in the tropics and subtropics.
Larsen & Inglis, 1998; Makasheva, 1983; Mes- In tropical Africa rye is cultivated in the high-
siaen et a l , 1991; Olivier & Annandale, 1998; lands of East Africa and it has been grown suc-
Paul, Southgate &Russell, 1980;Pilet-Nayel et cessfully in Malawi. In Ethiopia rye is spar-
a l , 2002; Pope, Polhill & Martins (Editors), ingly grown in the highlands of Arsi, where it
2003; Rubatzky &Yamaguchi, 1997; Schroeder was introduced through Swedish projects in
et al., 1995; Valderrama et a l , 2004; Wroth, the 1960s. Rye has been grown experimentally
1998. in Zambia and Mozambique, but apparently
Sources of illustration Davies, 1989. with little success. In Nigeria it has been tried
Authors C.-M. Messiaen, A.A. Seif, M. Jarso as a fodder plant in the 1980s. Rye is also
& G. Keneni grown in Morocco, Algeria, Egypt and South
Africa.
Uses Rye grain is used as a food for humans,
SECALE CEREALE L. but on a worldwide scale it is more important
as animal feed. The grain is processed into
Protologue Sp. pi. 1: 84 (1753). bread, cakes, crackers etc. For making bread,
Family Poaceae (Gramineae) whole or broken grain can be used; for making
Chromosome number 2n - 14 cake, the grain needs to be milled. Rye flour is
Vernacular names Rye (En). Seigle (Fr). often mixed with wheat flour. In Africa rye
Centeio (Po). flour is considered to make good porridge with
Origin and geographic distribution The an equal amount of maize flour; if used alone,
centre of origin of rye is not known exactly, but it is considered too sweet. Rye grain can be
its current centre of diversity is in the moun- sprouted to make malt for beer; several alco-
tainous areas of Afghanistan, Iran and the holic beverages are prepared by distilling
Middle East. Probably from there, rye was malted rye grains, e.g. rye whiskey in North
spread to the surrounding areas in Asia, north- America and vodka in Poland and Russia. Rye
ern Africa and later, just like wheat, to Russia, flour is used as filler for thickening soups and
central and western Europe, where it is culti- sauces.
vated under temperate climatic conditions. Rye Rye grain is used as a fodder, especially in pig
is a typical 'secondary crop': it was primarily a husbandry. Starch from the grain is industri-
weed in wheat and barley fields, later adopted ally used in the production of glue, matches,
as a crop. It may have been domesticated be- gum for sizing paper, and plastics. Rye straw is
fore 3000-4000 BC. Rye grains dating back to harvested for feed (cattle), litter (in livestock
6000 BC have been found in Turkey, but it is sheds), thatching, mulching material, indus-
not known if these were from crop plants or trial use (paper/cardboard), packing material
(nursery plants, cheese) and fuel. Immature
rye is harvested as a whole crop forage and it is
grown as a green manure or cover crop. In
Europe and India rye is sometimes grown as a
host plant for ergot (Clauiceps purpurea),
which is used medicinally, e.g. against mi-
graine. Rye pollen extracts are registered and
commercially available as a medicine against
benign prostatic hyperplasia in western
Europe, Japan, Korea and Argentina. In
Europe rye is under investigation as a biomass
energy crop.
Production and international trade Accord-
ing to FAO statistics, the total world rye pro-
duction in 1999-2003 amounted to 20 million
t/year from 9 million ha. The main producers
are the Russian Federation (5.6 million t/year
Secale cereale -planted from 3.2 million ha), Poland (4.2 million t/year
 SECALE 159

from 1.9 million ha) and Germany (3.9 million dihydroxy-l,4(2H)-benzoxazin-3-one and its
t/year from 0.7 million ha). No production sta- decomposition product 2(3H)-benzoxazolinone.
tistics are available for tropical Africa. World Weed-control effects of a rye mulch remain for
rye exports in 1998-2002 amounted to about 30-75 days after the rye is killed.
1.8 million t/year. The main exporter was Description Annual tufted grass up to 1.5(-
Germany (1.1 million t/year); the main import- 3) m tall, often blue-green; stem (culm) erect,
ers were Japan (360,000 t/year), the Russian slender, hollow except at nodes, glabrous but
Federation (170,000 t/year), South Korea pubescent near the spike, producing tillers and
(170,000 t/year) and China (140,000 t/year). roots at base; root system extensive, penetrat-
Properties Rye contains per 100 g edible ing to 2 m depth. Leaves alternate, simple; leaf
portion: water 11.0 g, energy 1402 kJ (335 sheath long and loose, with small auricles; lig-
kcal), protein 14.8 g, fat 2.5 g, carbohydrate ule short, jagged; blade linear-lanceolate, 10—
69.8 g, dietary fibre 14.6 g, Ca 33 mg, Mg 121 20 cm x 1-2 cm, smooth or slightly scabrous.
mg, P 374 mg, Fe 2.7 mg, Zn 3.7 mg, vitamin A Inflorescence a terminal spike 7-15 cm long,
11 IU, thiamin 0.32 mg, riboflavin 0.25 mg, curved, much awned, with spikelets alternat-
niacin 4.3 mg, vitamin Be 0.29 mg, folate 60|Xg ing and closely inserted on a long zigzag rachis.
and ascorbic acid 0 mg. The essential amino- Spikelets 2-flowered, with bisexual florets;
acid composition per 100 g edible portion is: glumes subulate, 1-veined, up to 1 cm long;
tryptophan 154 mg, lysine 605 mg, methionine lemma lanceolate, up to 2 cm long, tapering
248 mg, phenylalanine 674 mg, threonine 532 into a 2-8 cm long awn, 3(-5)-veined, keel
mg, valine 747 mg, leucine 980 mg and isoleu- prominently set with stiff bristles; palea about
cine 549 mg (USDA, 2004). Rye cultivars com- as long as lemma, awnless, scabrid on the keel;
paratively rich in lysine are known. The main stamens 3; ovary superior, with 2 plumose
fatty acids are (per 100 g edible portion): li- stigmas. Fruit a caryopsis (grain), oblongoid,
noleic acid 958 mg, oleic acid 280 mg, palmitic 4.5—10 mm x 1.5-3.5 mm, narrowly grooved,
acid 271 mg and linolenic acid 147 mg. Due to short-pointed, pale brown, glabrous.
the limited gluten content, bread made from Other botanical information Secale com-
rye flour has a compact structure; rye grain or prises 3 species, and is distributed from east-
grit is usually combined with wheat flour to
improve the volume and texture of the bread.
Because rye is not gluten-free, it is not suitable
for inclusion in the diet of people with coeliac
disease.
Rye starch has a high water-absorbing capac-
ity, making it suitable for use in adhesives. The
feed value of rye grain is lower than that of
other cereal grains, due to decreased feed in-
take, of which the causes are unclear. There-
fore rye is used in mixtures with other grains.
Rye straw is not very suitable as fodder be-
cause it is tough and fibrous. In a study on
forage quality in the United States, the crude
protein content of whole rye plants declined
from 27.8% in the vegetative stage through
24.2% in the booting stage to 13.4% in the
heading stage; the in-vitro dry matter digesti-
bility in these three stages was 79%, 81% and
70%, respectively.
Although rye pollen extracts are used to treat
benign prostatic hyperplasia, results from long-
term studies are not available and a meta-
study did not show sufficiently strong evidence.
Rye, its residues and aqueous extracts have Secale cereale - 1, plant habit; 2, flowering
allelopathic properties, enhancing the suitabil- spikelet; 3, floret without lemma; 4, fruiting
ity of rye for use as a weed-suppressing cover spikelet; 5, grains.
crop. The main allelopathic compounds are 2,4- Source: PROSEA
160 CEREALS AND PULSES

ern Europe to central Asia, with 1 species also endure frost down to -25°C. Tillering, shoot
occurring in South Africa. Only Secale cereale growth and flower initiation require rather low
is cultivated. Secale strictum (C.Presl) C.Presl temperatures (10—15°C); for adequate growth
subsp. africanum (Stapf) K.Hammer (synonym: during reproductive development the mean
Secale africanum Stapf) is only found in a sin- daily temperature must not exceed 20°C. Rye is
gle locality in South Africa. It is recordedly tolerant to drought. Flowering is favoured by
eaten as a cereal. It is liked by livestock and dry and sunny weather. Continuous rain, high
birds and is considered a potential pasture humidity and low temperatures hamper polli-
plant. nation, causing incomplete grain set. Winter
In the literature 2 subspecies have been distin- rye is a long-day plant; the reproductive devel-
guished within Secale cereale: subsp. cereale opment is stimulated by daylength increasing
(comprising the cultivated types, with a tough from 14 to 20 hours. Therefore, winter rye is
rachis) and subsp. ancestrale Zhuk. (compris- mainly grown between 40-65°N. Cultivars of
ing the wild and weedy types, with a more or spring rye are occasionally grown at high ele-
less fragile rachis, mainly found in western vations in subtropical and tropical areas. They
Asia). However, more subspecies have also are less sensitive to daylength and do not need
been distinguished. Within cultivated rye there vernalization. Their flowering and seed set are
are many landraces (usually with long culms satisfactory at a daylength of 12-13 hours. Rye
and small grains) and cultivars. can be grown on most well-aerated soil types
Hybrids of rye and wheat called triticale with a pH from 5-7.5; it is mainly grown on
(xTriticosecale) have been developed and these light, sandy and peaty soils.
show a mix of characteristics from the parents, Propagation and planting Rye is propa-
combining the hardiness of rye with the high gated by seed. The 1000-seed weight is 20-40
yield and quality of wheat. Triticale is pres- g. The optimal planting time for winter rye
ently grown only locally in tropical Africa, e.g. usually ranges from mid-September until mid-
in Ethiopia, Kenya, Tanzania and Madagascar, October in Europe. Seed can be broadcast by
and also in northern Africa and South Africa. hand but needs to be covered to achieve ade-
As a new food crop, it fell short of expectations, quate germination. Better conditions are cre-
but it is becoming increasingly popular as a ated by drilling seed mechanically at a uniform
forage crop. depth of 2-4 cm in rows 10-25 cm apart. Seed
Growth and development Rye germinates rates range from 100-150 kg/ha to obtain an
within 4 days at a soil temperature of 4-5°C, optimal density of 200-300 plants/m 2 . Spring
and more rapidly at higher temperatures. At rye needs to be planted as early as possible, if
the appearance of the fourth leaf, tillers and necessary even during winter, if soil conditions
roots are formed to anchor the plant. Shoot are suitable for preparing a seed-bed. Spring
initiation ceases as the plant enters the repro- rye tillers poorly, so requires a higher seed rate
ductive stage. Then, stem elongation starts and (150-200 kg/ha) than winter rye.
initiation and differentiation of the inflores- Management Rye competes strongly with
cence take place. In each spike 40-45 spikelets weeds, but they can cause problems at harvest.
are initiated, 30-35 of which bear 1-2 grains, They can be controlled mechanically by har-
resulting in 45-55 grains per spike. Flowering rowing or hoeing, or by herbicides during the
lasts 3-5 days for a spike and 8-12 days for a tillering stage. Lodging can cause considerable
rye crop. Rye is cross-pollinated by wind. The damage. The amount of fertilizer required is
post-floral period for grain-filling is 4-5 weeks. largely related to the expected yield; about 20
The period from sowing to harvesting varies kg N, 4 kg P and 13kg K are removed from the
from 4-10 months. The duration of growth is soil per t grain yield. About 75-80% of the N
largely dependent on temperature during re- and P is removed with the grains, whereas 75%
productive development. In temperate regions of the K remains in the straw. N is often the
so-called winter rye is planted in autumn to most yield-limiting nutrient. For yields over 5
receive sufficient cold and short days to induce t/ha a split N-application is preferred.
vernalization and reproductive growth; spring Diseases and pests Rye is considered rela-
rye is planted in early spring and can be har- tively tolerant to diseases. Nevertheless, after
vested after 4-6 months. germination snow mould (Fusarium nivale) can
Ecology Rye is a crop of temperate climates; cause considerable plant losses and brown or
in the tropics it is grown at high altitudes, e.g. leaf rust (Puccinia recondita f.sp. secalis) can
at 3000-3900 m in Ethiopia. Seedlings can severely damage leaves and stems. The most
 SECALE 161

conspicuous disease is ergot (Claviceps pur- before or during storage. After drying in the
purea), which infects the grain especially when field, straw is usually baled and stored in
grain set is poor; it produces alkaloid- barns or stacks for later use.
containing sclerotia. Grains with ergot are Genetic resources Large germplasm collec-
toxic, causing gangrenous or convulsive ergo- tions of rye are kept in the Russian Federation
tism, and can make a rye stock unsuitable for (N.I. Vavilov All Russian Scientific Research
human and animal consumption. No sources of Institute of Plant Industry, St. Petersburg,
resistance to ergot have yet been identified in 2635 accessions), Germany (Institute for Plant
rye. Other diseases include eyespot (Pseudocer- Genetics and Crop Plant Research (IPK), Gat-
cosporella herpotrichoides), sharp eyespot ersleben, 1990 accessions), the United States
(Rhizoctonia solani), powdery mildew (Erysiphe (USDA-ARS National Small Grains Germ-
graminis), stem rust (Puccinia graminis), plasm Research Facility, Aberdeen, Idaho,
glume blotch (Septoria nodorum) and leaf 1823 accessions) and Poland (Plant Breeding
blotch (Rynchosporium secalis). Most fungal and Acclimatization Institute (IHAR), Radzikow,
diseases can be controlled by fungicides, but Blonie, 1366 accessions; Botanical Garden of
damage by snow mould, sharp eyespot and the Polish Academy of Sciences, Warsaw, 1362
ergot can only be restricted by using healthy accessions). The only rye germplasm collections
and disinfected seed. Resistance, e.g. to leaf in Africa recorded by IPGRI are in South Africa
rust and stem rust, is found in several rye cul- (Division of Plant and Seed Control, Depart-
tivars, and resistance to these diseases has ment of Agriculture Technical Service, Preto-
been transferred from rye to wheat through ria, 178 accessions; Small Grain Institute,
intergeneric crosses. Damage by viruses is of Bethlehem, 52 accessions).
minor importance. The nematode Ditylenchus Breeding Rye breeding programmes have
dipsaci can affect rye, but it is not common. given priority to winter types, and aspects as
Insect pests are not important in rye cultiva- winter hardiness, straw stiffness, disease resis-
tion. tance and resistance to sprouting in the ear
Harvesting Time of harvest of rye is mid- have received much attention. These breeding
summer in Europe when the moisture content efforts have resulted in a considerable increase
of the grain is below 15%.The crop can be har- in grain yield and yield stability, shorter
vested by hand; the method of harvesting, plants, reduced lodging and enhanced harvest
threshing, collecting and storing can be similar index. Well-known cultivars include 'Petkus',
to that used for sorghum and millets. For com- 'Pearl', 'Steel' and 'King II'. Efforts to exploit
bine harvesting, it is best to wait until the heterosis for enhancing grain yield have re-
moisture content has dropped below 16%. sulted in hybrids that have entered commercial
However, to prevent loss of quality due to production with high-input management. Hy-
sprouting in the ear, the crop may be harvested brids outyield conventional cultivars by 10-
at a higher moisture content (18-20%), espe- 20%, but they demand more inputs (seed, crop
cially if wet weather conditions prevail and protection). Tetraploid cultivars have been
delay ripening. Then subsequent drying will be developed, with more vigorous growth and lar-
required, in sheaves in the field or mechani- ger grains. Secale cereale has been crossed with
cally during storage. Secale strictum with the objective of improving
Yield Rye yields vary widely, from less than winter hardiness and resistance to drought and
1 t/ha in Africa, Latin America and Australia diseases. Perennial rye cultivars, intended for
to over 5 t/ha in some western European coun- use as fodder, have also been developed by
tries. The world average yield is about 2 t/ha. crossing the two species. For the production of
Handling after harvest Low moisture con- ergot, male-sterile lines are used, facilitating
tent of the rye grain and low storage tempera- the infection by the fungus.
tures are desirable for long-term storage. The Rye is considered one of the most recalcitrant
moisture content of the grain should be less plants for tissue culture and genetic transfor-
than 13% if rye is to be stored for 6 months mation. However, systems for the stable ge-
(without ventilation) at 15°C. If the stock is netic transformation of rye using Agrobacte-
regularly ventilated, a moisture content of 14- rium tumefaciens or biolistic methods have
15% may be acceptable. In temperate regions, been developed. It is possible to obtain large
such low moisture contents are often not numbers of genetically identical plants by in-
reached at harvesting, and grain needs to be vitro regeneration using immature inflorescen-
dried by warm air. Cleaning is commonly done ces as expiants. Genetic linkage maps of rye,
162 CEREALS AND PULSES

on the basis of various marker types (RFLPs,

AFLPs, RAPDs and microsatellite markers)
have also been constructed. Genes conferring
resistance to leaf rust have been identified.
Prospects Rye may be inferior in several
ways to the predominant world cereals (wheat,
rice and maize), but it will continue to be an
important crop because of its winter hardiness,
tolerance to drought, ability to grow on poor
soils, and consumer demand for baked products
with the unique flavour of rye. There is consid-
erable scope for improving yields. The applica-
tion ofhigh quality seed, new (hybrid) cultivars
and advanced management practices can in-
crease yield levels in the short term. The pros-
pects of rye in tropical Africa seem limited. It
has been tried in various countries, but its cul-
tivation has not become important. Setaria italica - planted
Major references Darwinkel, 1996; Dar-
winkel, 1999; Frederiksen & Petersen, 1998; evolved from the wild Setaria viridis (L.)
Froman &Persson, 1974; Hanelt &Institute of P.Beauv. (green foxtail millet), and it was most
Plant Genetics and Crop Plant Research (Edi- probably first domesticated in the highlands of
tors), 2001; Khlestkina et al., 2004; Kuip & central China, from where it spread to India
Ponte (Editors), 2000; Popelka & Altpeter, and Europe soon thereafter. Evidence for this
2003; Popelka, Xu & Altpeter, 2003; Smartt & origin, however, is not conclusive and its do-
Simmonds (Editors), 1995. mestication may have taken place anywhere in
Other references Acharrya, Mir & Moyer, the area extending from Europe to Japan, per-
2004; Allkämper, 1984; Barnes & Putnam, haps even several times independently. Foxtail
1987; Bosworth, Hoveland & Buchanan, 1986; millet was the 'panicum' of the Romans. At
Burgos & Talbert, 2000; Gibbs Russell et al., present foxtail millet is cultivated all over the
1990; Launert, 1971; Maikhuri, Nautiyal & world. In tropical Africa it is cultivated to a
Khali, 1991; Masiunas et al., 1997;Musa, 1985; limited extent in upland areas in East Africa
Nwankiti, 1984; Phillips, 1995; Roux et al., and occasionally recorded elsewhere, e.g. in
2004; Raemaekers (Editor), 2001; Scholz & Cameroon and southern Africa (Malawi, Zim-
Eilerbrock, 2002; USDA, 2004; Vazquez & Li- babwe, Mozambique). In these areas it also
nacero, 1995; Weston, 1996; Williamson, 1955; occurs as an escape. Foxtail millet is also
Wilt et al., 2000. grown in South Africa and Lesotho.
Sources of illustration Darwinkel, 1996. Uses The husked grain of foxtail millet is
Authors M. Brink used as food in Asia, south-eastern Europe and
Based on PROSEA 10: Cereals. Africa. It is most important in China and In-
dia. The grain may be cooked and eaten like
rice, either entire or broken. It can be ground
SETARIA ITALICA (L.) P.Beauv. and made into unleavened bread or, when
mixed with wheat flour, into leavened bread.
Protologue Ess. Agrostogr.: 51, 170, 178 The flour is also made into cakes, porridges
(1812). and puddings. In northern China foxtail millet
Family Poaceae (Gramineae) forms part ofthe staple diet; it is usually mixed
Chromosome number 2n = 18 with pulses and cooked, or the flour is mixed
Vernacular names Foxtail millet, Italian with that of other cereals in the preparation of
millet, German millet (En). Panis, millet des bread and noodles. It is considered a nutritious
oiseaux, millet d'Italie (Fr). Painço, milho food and is often recommended for the elderly
painço, milho painço de Itâlia (Po). Kimanga and for pregnant women. Since the 1990s it
(Sw). has been used in China for the industrial
Origin and geographic distribution Foxtail preparation of mini crisp chips, millet crisp
millet is an old crop, grown since 5000 BC in rolls and flour for baby foods. Foxtail millet is
China and 3000 BC in Europe. It probably used in the preparation of beer and alcohol,
 SETARIA 163

especially in Russia and Myanmar, and for lateral branches short, bearing 6-12 spikelets.
vinegar and wine in China. Sprouted seeds are Spikelets almost sessile, subtended by 1-3
eaten as a vegetable, e.g. in China. bristles up to 1.5 cm long, elliptical, usually
In Europe and the United States foxtail millet about half as long as the bristles, 2-flowered;
is primarily grown as bird feed. It is an impor- lower glume small and 3-veined, upper glume
tant fodder crop ('moha'); in the United States slightly shorter than spikelet, 5-veined; lower
and Europe it is grown for hay and silage, and floret sterile, upper one bisexual with 5-veined
in China the straw is an important fodder. The lemma and palea, 2 lodicules, 3 stamens and
straw is also used for thatching and bedding, superior ovary with 2 plumose stigmas. Fruit a
e.g. in India. The bran serves as animal feed caryopsis (grain), broadly ovoid, up to 2 mm
and can be used for oil extraction. Foxtail mil- long, pale yellow to orange, red, brown or
let is credited with diuretic, astringent and black, tightly enclosed by lemma and palea.
emollient properties and is used to treat rheu- Other botanical information Setaria com-
matism. It can be sown in contour strips for prises about 100 species distributed in the
erosion control. tropics, subtropics and temperate regions. Fox-
Production and international trade Pro- tail millet is the most economically valuable
duction statistics for foxtail millet are scarce species of the genus. Several wild Setaria spe-
because they are usually lumped with those of cies are harvested for their seeds, e.g. Setaria
other millets. The annual world production of finita Launert in Namibia. Setaria sphacelata
foxtail millet in the early 1990s was estimated (Schumach.) Stapf &C.E.Hubb. ex M.B.Moss is
at 5 million t (18% of total millet production), cultivated as a forage throughout the tropics
with China being the main producer. In tropi- and its grains are gathered as a famine food in
cal Africa the production of foxtail millet is Africa. The grains of Setaria pumila (Poir.)
much lower than that of pearl millet (Pennise- Roem. & Schult, are also eaten as a famine
tum glaucum (L.) R.Br.) and finger millet food, e.g. in Mali, Burkina Faso, Sudan and
(Eleusine coracana (L.) Gaertn.), but no statis- Ethiopia, but it is more important as a forage.
tics are available. In India and China foxtail Setaria verticillata (L.) P.Beauv. is a forage
millet is mainly grown for home consumption. plant, but also collected as a famine food, e.g.
Properties The composition of foxtail millet
grain per 100 g edible portion is: water 12 g,
energy 1470 k j (351 kcal), protein 11.2 g, fat
4.0 g, carbohydrate 63.2 g, crude fibre 6.7 g, Ca
31 mg, Fe 2.8 mg, thiamin 0.6 mg, riboflavin
0.1 mg and niacin 3.2 mg (FAO, 1995). The
essential amino-acid composition per 100 g
grain is: tryptophan 103 mg, lysine 233 mg,
methionine 296 mg, phenylalanine 708 mg,
threonine 328 mg, valine 728 mg, leucine 1764
mg and isoleucine 803 mg (FAO, 1970). The
starch granules are spherical, angular or poly-
hedral with a diameter of 6-17 |im. Most fox-
tail cultivars are non-glutinous and are thus
suitable for the diet of people with coeliac dis-
ease. The bran contains about 9%oil.
Description Erect annual grass up to 150(-
175) cm tall, tufted, often variously tinged with
purple; root system dense, with thin wiry ad-
ventitious roots; stem erect, tillering at base,
sometimes branched. Leaves alternate, simple;
leaf sheath 10-15(-25) cm long, glabrous or
slightly hairy; ligule short, fimbriate; blade
linear, 15-30(-50) cm x 0.5-2.5(-4) cm, acumi-
nate at apex, midrib prominent, slightly rough. Setaria italica - 1, upper part of plant; 2,
Inflorescence a spike-like panicle 5—30cm x 1- sheath mouth with ligule; 3, flowering spikelet
2(-5) cm, erect or pendulous, continuous or with bristles; 4,fruiting spikelet.
interrupted at base; rachis ribbed and hairy; Source: PROSEA
164 CEREALS AND PULSES

in Niger, Sudan and Namibia. drought. Flowering is normally accelerated by

Setaria italica is a 'crop-weed complex', i.e. short days, but day-neutral cultivars exist.
with wild and cultivated types. These types Foxtail millet prefers fertile soils with a pH of
show no crossing barriers and isozyme analysis about 6.5, but can be grown successfully on a
and molecular studies have confirmed their wide range of soils, from light sands to heavy
similarity. The wild types are considered to clays, and even yields reasonably well on poor
represent Setaria viridis (green foxtail millet), or marginal soils. It does not tolerate waterlog-
the cultivated ones Setaria italica (foxtail mil- ging.
let). Propagation and planting Foxtail millet
Green foxtail millet occurs worldwide as a is propagated by seed. The 1000-seed weight is
variable, annual weed, especially common in about 2 g. Dormancy is common in freshly har-
temperate regions. It differs from foxtail millet vested seed. The recommended seed rate for
in its completely caducous spikelets, upper sole cropping in Kenya is 4 kg/ha, with a dis-
glume about as long as the spikelet and more tance of 30 cm between rows and 10 cm within
roughly papillose lemma. It is sometimes con- the row. In China and India it is sown at a seed
sidered a subspecies of Setaria italica: subsp. rate of 5-15 kg/ha when grown in pure stands,
viridis (L.) Thell. It is also known as green with plant densities of 300,000-1.5 million
bristle grass, and is one of the world's most plants/ha. It is either broadcast or drilled in
noxious weeds, but it is sometimes used as rows 20-60 cm apart, with 5-20 cm within the
fodder or for medicinal purposes. row, and thinning may be practised. The usual
Foxtail millet is very variable and numerous sowing depth is 3-6 cm and a fine, firm seed-
cultivars exist, differing in time to maturity, bed is required. Foxtail millet is grown as a
plant height, size, habit and structure of inflo- sole crop or intercropped, e.g. with finger mil-
rescence, number, colour and length of bristles, let, cotton, sorghum or pigeon pea in India.
and colour of grain. Primitive cultivars have Management In Kenya the first weeding of
numerous, strongly branched stems (like green foxtail millet is recommended at 2-3 weeks
foxtail millet), while advanced cultivars pro- after emergence of the seedlings, and the sec-
duce a single stem with a large, solitary inflo- ond one 2 weeks later. In India foxtail millet is
rescence. usually weeded once at about 3 weeks after
Growth and development Foxtail millet sowing. Foxtail millet responds well to manur-
generally starts flowering at about 60 days ing, but generally only irrigated crops are ma-
after sowing, and flowering lasts for 10-15 nured. It is usually grown as a rainfed crop,
days. Flowering proceeds from the top of the but it may also be grown under irrigation, e.g.
panicle downward. The flowers open late at in India. Crop rotation of foxtail millet with
night or early in the morning, and close soon finger millet and sorghum is common in India.
after opening. Foxtail millet is largely self- Sometimes it is grown as a catch crop when
pollinating with an average outcrossing rate of paddy rice has failed.
4%; natural hybrids between wild and culti- Diseases and pests The most serious dis-
vated types occur. Total crop duration is 80- eases of foxtail millet are blast {Pyricularia
120 days, although some cultivars only need 60 setariae), downy mildew (Sclerospora gramini-
days to mature. Foxtail millet has largely lost cola), leaf rust (Uromyces setariae-italiae) and
the ability of natural seed dispersal, and shows smut (Ustilago crameri). Downy mildew and
a tendency toward uniform plant maturity. smut can be controlled by treating the seed.
Foxtail millet follows the C4-cycle photosyn- Important insect pests of foxtail millet are
thetic pathway. shoot flies (Atherigona spp.), crickets, borers
Ecology Foxtail millet is primarily a crop of and caterpillars. Foxtail millet is highly sus-
subtropical and temperate regions; in the trop- ceptible to bird attack in the field, and mice
ics it is grown up to 2000(-3300) m altitude. It and rats also damage the crop. In stored grain,
does not tolerate frost. In China and India it is seed smut (Sorosporium bullatum) and kernel
mainly grown in areas with an annual rainfall smut (Ustilago paradoxa) may cause consider-
of 400-800 mm. Foxtail millet is not particu- able losses in addition to the common cereal
larly drought-resistant, but its short crop cycle storage insects.
makes it suitable for low-rainfall areas and it Harvesting Foxtail millet is harvested
can be grown in semi-arid regions with rainfall manually by cutting off the panicles and
less than 125 mm in the 3-4 months of growth. threshing them. Mechanical harvesting with a
It is, however, susceptible to long periods of combine or binder is possible. In southern In-
 SORGHUM 165

dia whole plants may be cut and threshed by ers) and flowering behaviour of foxtail millet,
trampling by cattle or by passing a stone roller artificial cross-pollination is difficult, but an
over the plants. When grown for fodder, foxtail effective procedure for artificial hybridization
millet should be harvested before flowering. of foxtail millet has been developed in the
Yield The average annual yield of rainfed United States. High levels of heterosis for
foxtail millet is 800-900 kg/ha of grain and grain yield (67%) and panicle length (68%)
2500 kg/ha of straw. Improved cultivars in have been found.
China yield 1800 kg/ha of grain in regions with Prospects On a worldwide scale foxtail mil-
less than 900 mm annual rainfall. Much higher let has lost its importance as a food crop in
grain yields can be obtained with irrigation (in competition with major cereals such as wheat,
China experimental yields have reached 11 rice, maize and sorghum. However, because of
t/ha). As forage it may yield 15-20 t green mat- its short crop cycle and the fact that it can be
ter per ha or 3.5 t hay. grown on a wide range of soil types it may re-
Handling after harvest Foxtail millet main a useful crop in Asia on poor agricultural
should be dried thoroughly before storage. The land in regions with low rainfall or a short
grain is usually husked just before processing growing season. The prospects for foxtail millet
because husked grains are readily infested in tropical Africa seem limited, but it may gain
with insects. Husking can be done with a stone importance as a niche crop in dry regions at
roller or with rice milling machinery. In China medium to high altitudes.
mini crisp chips are made by cooking husked Major references de Wet, Oestry-Stidd &
grains, pressing the product to 1mm thickness, Cubero, 1979; FAO,undated; Hanelt & Institute
drying, frying in oil and flavouring. Crispy rolls of Plant Genetics and Crop Plant Research
are prepared from husked grains which are (Editors), 2001; Oduori, 1993; Prasada Rao &
soaked in water, ground and, after addition of de Wet, 1997; Prasada Rao et al., 1987; Purse-
sugar, toasted between 2 iron plates and glove, 1972; Rahayu & Jansen, 1996; Riley et
formed into rolls. al. (Editors), 1993;Seetharam, Riley & Harina-
Genetic resources Large collections of fox- rayana, 1990.
tail millet germplasm are kept by the Institute Other references Benabdelmouna et al.,
of Crop Germplasm Resources (CAAS), Beijing, 2001; Benabdelmouna, Abirached-Darmency &
China (25,380 accessions), the International Darmency, 2001; Burkill, 1994; Clayton, 1989;
Crops Research Institute for the Semi-Arid CSIR, 1972; de Wet, 1995b; FAO, 1970; FAO,
Tropics (ICRISAT), Patancheru, India (1528 1995; Gibbs Russell et al., 1990; Hülse, Laing
accessions) and the All India Coordinated Mi- & Pearson, 1980; ICRISAT & FAO, 1996;
nor Millet Project, University of Agricultural Klaassen & Craven, 2003; le Thierry d'En-
Sciences, Bangalore, India (1300 accessions). In nequin et a l , 2000; Li et al., 1998; Malm &
Africa a collection of 451 foxtail millet acces- Rachie, 1971; Ministry of Agriculture and Ru-
sions is kept at the National Dryland Farming ral Development, 2002; Petr et al., 2003; Siles,
Research Station, Machakos, Kenya. Resis- Baltensperger & Nelson, 2001; Siles et al.,
tance to blast and rust has been identified in 2004; Wanous, 1990.
germplasm collections. Sources of illustration Hanelt & Institute
Breeding Foxtail millet breeding is mainly of Plant Genetics and Crop Plant Research
carried out in China and India. Major breeding (Editors), 2001;Rahayu &Jansen, 1996.
objectives are developing high-yielding culti- Authors M. Brink
vars which produce protein-rich seed and are Based on PROSEA 10: Cereals.
resistant to diseases, pests and lodging, and
adapted to local ecological circumstances. In
China, for example, cultivars with a short S O R G H U M BICOLOR (L.) Moench
growing cycle and a high drought and cold tol-
erance have been developed; these can be Protologue Methodus: 207 (1794).
grown in the summer season after winter Family Poaceae (Gramineae)
wheat. The recommended cultivar in Kenya is Chromosome number 2n = 20
'KAT/FOX-1'; it matures in 3-4 months. Tech- Vernacular names Sorghum, sorgo, guinea
niques applied in foxtail millet breeding in- corn, great millet, durra (En). Sorgho, gros mil,
clude selection, hybridization (using male- sorgho rouge (dye cultivars), sorgho des teintu-
sterile lines) and radiation-induced mutations. riers (dye cultivars) (Fr). Sorgo, milho miudo,
Due to the floral morphology (very small flow- massambala (Po).Mtama (Sw).
166 CEREALS AND PULSES

ing. Sorghum flour is used to make thick or

thin porridge, pancake, dumplings or couscous,
opaque and cloudy beers and non-alcoholic
fermented beverages. In Africa sorghum grain
is germinated, dried and ground to form malt,
which is used as a substratum for fermentation
in local beer production. White grain is gener-
ally preferred for cooking while red and brown
grains are normally used for beer making.
Where bird pressure is high, e.g. around Lake
Victoria, red and brown types rich in tannin
may be grown for food instead of white types.
In China sorghum is extensively distilled to
make a popular spirit and vinegar. Sorghum
grain is a significant component of cattle, pig
and chicken feeds in the United States, Central
and South America, Australia and China, and
Sorghum bicolour - planted is becoming important in chicken feed in India.
It requires grinding, rolling, flaking or steam-
Origin and geographic distribution The ing to maximize its nutritional value.
greatest diversity in both cultivated and wild Several non-edible sorghum cultivars are ex-
types of Sorghum is found in north-eastern clusively grown for the red dye present in the
tropical Africa. The crop may have been domes- leaf sheaths and sometimes also in adjacent
ticated in that region, possibly Ethiopia. Vari- stem parts. In Africa this dye is used particu-
ous hypotheses have been put forward as to larly for goat-skin leather (e.g. in Nigeria), but
when the crop was domesticated, from as early also for mats, textiles, strips of palm leaves
as 5000-3000 BC to around 1000 BC, but the and grasses used in basketry and weaving,
latter period is more widely accepted now. ornamental calabashes, wool (e.g. in Sudan), as
From north-eastern Africa sorghum was dis- a body paint and to colour cheese and lick-
tributed all over Africa and along shipping and stones for cattle (e.g. in Benin). A similar dye
trade routes through the Middle East to India. can be extracted from the grain refuse (glumes
From India it is believed to have been carried and grain wall) of several red sorghum culti-
to China along the silk route and through vars grown for food or for beer-making. In Ni-
coastal shipping to South-East Asia. From geria the red sorghum dyes were traditionally
West Africa sorghum was taken to the Ameri- used by the Bunu, Aworo, Igbira and Okpella
cas through the slave trade. It was introduced people for a fabric called 'abata', used as a fu-
into the United States for commercial cultiva- neral hanging, decorated with patterns made
tion from North Africa, South Africa and India by thick threads added to the weft of the fabric.
at the end of the 19th century. It was subse- The fabrics in which the dominant colours were
quently introduced into South America and derived from sorghum were known as 'ifala'.
Australia. It is now widely cultivated in drier Sorghum is also used to provide the violet col-
areas ofAfrica, Asia, the Americas, Europe and ours decorating the masks worn during certain
Australia between latitudes of up to 50°N in dances by Yoruba people in southern Benin
North America and Russia and 40°S in Argen- and in south-western Nigeria. In Côte d'Ivoire
tina. Sorghum types exclusively cultivated for sorghum and other tannin-rich dyes are used
the dye in the leaf sheaths can be found from in combination with mud to create the patterns
Senegal to Sudan. of the painted cloths produced in the Korhogo
Uses Sorghum is an important staple food, region. The dye was formerly exported to Mo-
particularly in semi-arid tropical regions of rocco where it was used in the leather industry.
Africa and Asia, and an important feed grain In China sorghum types with red panicles and
and fodder crop in the Americas and Australia. leaf sheaths were also used for dyeing. In the
In the simplest food preparations, the whole 19th century red sorghums were exported to
grain is boiled (to produce a food resembling Europe where the dye was known as 'carmin de
rice), roasted (usually at the dough stage), or sorgho'. It was extracted by squeezing out the
popped (like maize). More often the grain is juice, which was then fermented. Used with
ground or pounded into flour, often after hull- wool or silk mordanted with tin or chrome, the
 SORGHUM 167

result was a colourfast red-brown that was million ha), Sudan (3.4 million t/year from 5.3
once known as 'rouge badois'. 'Durra red', a million ha), Argentina (3.0 million t/year from
similar product, was imported from India into 630,000ha), China (3.0million t/year from 840,000
the United Kingdom where the dye was known ha), Australia (1.9 million t/year from 690,000
as 'Hansen brown' or 'Meyer brown'. Recently ha), Ethiopia (1.4 million t/year from 1.2 mil-
the use of sorghum dye in hair dying products lionha) and Burkina Faso (1.3million t/year from
has been patented. 1.4 million ha). In sub-Saharan Africa annual
The stems of sweet sorghum types are chewed production increased from around 10 million t
like sugar cane and, mainly in the United from 13 million ha in the early 1960s to about
States, a sweet syrup is pressed from them. In 20 million t from 25 million ha in the early
North America and eastern Europe special 2000s.
types with very long, fibrous and few-seeded Almost all sorghum traded on international
inflorescences, known as 'broomcorn', are markets is for use as livestock feed. Average
grown to make brooms. Sorghum plant resi- world exports of sorghum in 1998-2002
dues are used extensively as material for roof- amounted to 6.3 million t/year, almost all from
ing, fencing, weaving and as fuel. The stems the United States (5.6 million t/year). The
can be used for the production of fibre board. main importers are Mexico and Japan. In
Danish scientists have made good panelling tropical Africa most sorghum is grown for
using stem chips of sorghum. The stover re- home consumption (except for beer production).
maining after harvesting the grain is cut and In southern and eastern Africa malting sor-
fed to cattle, sheep and goats, or may be ghum for beer brewing has developed into a
grazed. Some farmers grind harvested stover large-scale commercial industry, using about
and mix it with sorghum bran or salt to feed 150,000 t of sorghum grain annually. In
livestock. Sorghum is also grown for forage, Uganda commercial production of lager beer
either for direct feeding to ruminants or for using sorghum instead of barley is becoming a
preservation as hay or silage. Sorghum flour is great success (annual requirement of sorghum
used to produce an adhesive in the manufac- is 3000 t) and is very promising for other Afri-
ture of plywood. Sweet sorghum is suitable for can countries. In Nigeria sorghum malting has
the production of alcohol, while the bagasse is become a major industry for lager and stout
a suitable source of paper pulp for the produc- beer brewing and for malt beverages, using
tion of kraft paper, newsprint and fibre board. about 15,000 t of sorghum annually. In South
Sorghum has various applications in African Africa an instant breakfast cereal is made from
traditional medicine: seed extracts are drunk sorghum that is similar in quality but much
to treat hepatitis, and decoctions of twigs with cheaper than wheat or maize products. Annual
lemon against jaundice; leaves and panicles are production is 12,000 t and is increasing stead-
included in plant mixtures for decoctions iiy.
against anaemia. The Salka people in northern In West Africa small tied bundles of 4-6 leaf
Nigeria use sorghum in arrow-poisons. The red sheaths of sorghum dye cultivars are offered
pigment is said to have antimicrobial and anti- for sale on local markets (in the 1990s the price
fungal properties and is also used as a cure for was about 150 CFA). In 1993 in Burkina Faso,
anaemia in traditional medicine. the red pigment was successfully extracted
Production and international trade Sor- chemically from sorghum leaf sheaths and of-
ghum grain is the fifth most important cereal fered for sale as dry powder on the world mar-
in the world after wheat, rice, maize and bar- ket.
ley. In Africa it comes second after maize in Properties The composition of sorghum
terms of production. According to FAO esti- grain per 100 g edible portion is: water 9.2 g,
mates, the average world production of sor- energy 1418 kJ (339 kcal), protein 11.3 g, fat
ghum grain in 1999-2003 amounted to 57.7 3.3 g, carbohydrate 74.6 g, Ca 28 mg, P 287
million t/year from 42.6 million ha. The pro- mg, Fe 4.4 mg, vitamin A 0 IU, thiamin 0.24
duction in sub-Saharan Africa was 19.0 mil- mg, riboflavin 0.14 mg, niacin 2.9 mg and
lion t/year from 22.8 million ha. The main pro- ascorbic acid 0 mg. The essential amino acid
ducing countries are the United States (12.0 composition per 100 g edible portion is: trypto-
million t/year in 1999-2003 from 3.2 million phan 124 mg, lysine 229 mg, methionine 169
ha), India (7.6 million t/year from 9.8 million mg, phenylalanine 546 mg, threonine 346 mg,
ha), Nigeria (7.6 million t/year from 6.9 mil- valine 561 mg, leucine 1491 mg and isoleucine
lion ha), Mexico (6.0 million t/year from 1.9 433 mg. The principal fatty acids are per 100 g
168 CEKEALS AND PULSES

edible portion: linoleic acid 1305 mg, oleic acid gus Bipolaris maydis in the production of api-
964 mg and palmitic acid 407 mg (USDA, geninidin in these cultivars deserves further
2004). Sorghum grain is first limiting in lysine, research. Used without a mordant, the dye
then in methionine and threonine. Much of the obtained from sorghum gives a dark red that is
protein in sorghum is prolamine (39-73%), fairly colourfast and still much used in eastern
which is poorly digestible. As a result, maxi- Africa, particularly Sudan and Ethiopia, for
mum available protein in sorghum grain is dyeing leather, cotton and the grasses and
usually 8-9%. The tannin content of sorghum reeds used for woven matting. Black colours
also affects its nutritional value. High- and are obtained with natron salt and iron mor-
low-tannin sorghum types are distinguished. dants. From red sorghum grain the pigments
High-tannin sorghum types (sometimes called apigenin, quercimeritrin, kaempferol gluco-
'brown sorghums', although the grain may also sides, apigenidin glucosides, apigeninidin,
be white, yellow or red) have less nutritional luteolinidin and 7-O-methyl-luteolin-glucoside
value but have agronomic advantages, includ- have been isolated. From the stem of red sor-
ing resistance to birds, insects, fungi and de- ghum cultivars the constituents of the red dye
creased sprouting in the panicle. Sorghum were the anthocyanidin apigeninidin (17%) and
types without a pigmented grain wall ('white the flavonoids luteolin (9%) and apigenin (4%).
sorghums') do not contain condensed tannins The anaemia curing property of the red pig-
and have a nutritional value similar to that of ment has been confirmed in tests with rats.
maize. Decortication, parboiling, malting or Description Annual grass up to 5 m tall,
steeping in alkali solutions significantly reduce with one to many tillers, originating from the
the tannin content of sorghum grain. In gen- base or stem nodes; roots concentrated in the
eral the endosperm accounts for 82—84%of the top 90 cm of the soil but sometimes extending
grain weight, the germ for 9-10% and the grain to twice that depth, spreading laterally up to
wall for 6-8%. The starch granules in the en- 1.5 m; stem (culm) solid, usually erect. Leaves
dosperm have a diameter of (4-)15(-25) |im. alternate, simple; leaf sheath 15—35 cm long,
The starch normally contains 70-80% amy- often with a waxy bloom, with band of short
lopectin and 20-30% amylose, although some white hairs at base near attachment, reddish
types contain 100% amylopectin and others up in dye cultivars, auricled; ligule short, c. 2 mm
to 62% amylose. The gelatinization tempera- long, ciliate on upper free edge; blade lanceo-
ture ranges from 68-75°C. Sorghum grain does late to linear-lanceolate, 30-135 cm x 1.5-13
not contain gluten and cannot be used for leav- cm, initially erect, later curving, margins flat
ened products unless mixed with wheat. or wavy. Inflorescence a terminal panicle up to
The composition of the green plant varies ac- 60 cm long; rachis short or long, with primary,
cording to age and cultivar but it normally con- secondary and sometimes tertiary branches,
tains 78-86 g of water per 100 g of fresh mate- with spikelets in pairs and in groups ofthree at
rial. On a dry basis it contains per 100 g: pro- the ends of branches. Spikelet sessile and bi-
tein 12 g, carbohydrate 40-50 g and fibre 20- sexual or pedicelled and male or sterile, with 2
30 g. The glycoside dhurrin occurs in the aerial florets; sessile spikelet 3-10 mm long, with
parts of most sorghum. Dhurrin is hydrolyzed glumes approximately equal in length, lower
to hydrocyanic acid (HCN), which is highly glume 6-18-veined, usually with a coarse keel-
toxic and can kill grazing animals. It is particu- like vein on each side, upper glume usually
larly concentrated in the young leaves and till- narrower and more pointed, with central keel
ers and in plants that are suffering from for part of its length, lower floret consisting of
drought. HCN content usually declines with a lemma only, upper floret bisexual, with
age, reaching non-toxic levels 45-50 days after lemma cleft at apex, with or without kneed and
planting, and HCN is destroyed when the fod- twisted awn, palea, when present, small and
der is made into hay or silage. thin, lodicules 2, stamens 3; ovary superior, 1-
The red pigment in sorghum dye cultivars is celled with 2 long styles ending in feathery
composed of anthocyanic compounds, particu- stigmas; pedicelled spikelet persistent or de-
larly rich (95%) in the stable apigeninidin chlo- ciduous, smaller and narrower than sessile
ride (3-deoxyanthocyanidin) and tannins of the spikelet, often consisting of only two glumes,
condensed proanthocyanidins group (producing sometimes with lower floret consisting of
phlobaphen reds). The red pigment in the sor- lemma only and upper floret with lemma, 2
ghum leaf sheath makes up to over 20% of the lodicules and 3 stamens. Fruit a caryopsis
dry weight. The role of the non-pathogenic fun- (grain), usually partially covered by glumes, 4—
 SORGHUM 169

groups would, however, be more appropriate.

The 5basic races are:
- Bicolor: the most primitive cultivated sor-
ghum, characterized by open inflorescences
and long clasping glumes that enclose the
usually small grain at maturity. Cultivars
are grown in Africa and Asia, some for their
sweet stems to make syrup or molasses, oth-
ers for their bitter grains used to flavour sor-
ghum beer, but they are rarely important.
They are frequently found in wet conditions.
- Caudatum: characterized by turtle-backed
grains that are flat on one side and curved on
the other; the panicle shape is variable and
the glumes are usually much shorter than
the grain. Cultivars are widely grown in
north-eastern Nigeria, Chad, Sudan and
Uganda. The types used for dyeing also be-
long here and are known as 'karan dafi' by
the Hausa people in Nigeria.
- Durra: characterized by compact inflorescen-
ces, characteristically flattened sessile
spikelets, and creased lower glumes; the
grain is often spherical. Cultivars are widely
grown along the fringes of the southern Sa-
hara, western Asia and parts of India. The
Sorghum bicolor - panicles and spikelets of the durra type is predominant in Ethiopia and in
5 basic races: 1, bicolor;2, caudatum; 3, durra; the Nile valley in Sudan and Egypt. It is the
4,guinea; 5, kafir. most specialized and highly evolved of all
Source: PROSEA races and many useful genes are found in
this type. Durra cultivars range in maturity
8 mm in diameter, rounded and bluntly from long to short-season. Most of them are
pointed. drought resistant.
Other botanical information Sorghum com- - Guinea: characterized by usually large, open
prises 20—30 species. Sorghum bicolor belongs inflorescences with branches often pendulous
to section Sorghum, together with the 2 peren- at maturity; the grain is typically flattened
nial species Sorghum halepense (L.) Pers. and and twisted obliquely between long gaping
Sorghum propinquum (Kunth) Hitchc. At pre- glumes at maturity. Guinea sorghum occurs
sent, Sorghum bicolor is mostly considered as primarily in West Africa, but it is also grown
an extremely variable crop-weed complex, along the East African rift from Malawi to
comprising wild, weedy and cultivated annual Swaziland and it has also spread to India
types (classified as subspecies) which are fully and the coastal areas of South-East Asia.
interfertile. The cultivated types are classified Many subgroups can be distinguished, e.g.
as subsp. bicolor (synonyms: Sorghum ankolib with cultivars especially adapted to high or
Stapf, Sorghum caudatum Stapf, Sorghum low rainfall regimes. In the past the grain
cernuum Host, Sorghum dochna (Forssk.) was often used as ship's provisions because it
Snowden, Sorghum durra (Forssk.) Stapf, Sor- stored well.
ghum membranaceum Chiov., Sorghum nigri- - Kafir: characterized by relatively compact
cans (Ruiz & Pav.) Snowden, Sorghum subgla- panicles that are often cylindrical in shape,
brescens (Steud.) Schweinf. & Asch., Sorghum elliptical sessile spikelets and tightly clasp-
vulgare Pers.) and they are subclassified into ing glumes that are usually much shorter
different races on the basis of grain shape, than the grain. Kafir sorghum is an impor-
glume shape and panicle type. Five basic races tant staple across the eastern and southern
and 10—15hybrid combinations of 2 or more of savanna from Tanzania to South Africa.
these races are recognized and grouped into Kafir landraces tend to be insensitive to pho-
subsp. bicolor. A classification into cultivar toperiod and most commercially important
170 CEREALS AND PULSES

male-sterile lines are derived from kafir type expanded and light interception is approaching
sorghum. its maximum; lower leaves have begun to se-
Hybrid races exhibit various combinations and nesce. During the boot stage, the developing
intermediate forms of the characteristics of the panicle has almost reached its full size and is
5 basic races. Durra-bicolor is found mainly in clearly visible in the leaf sheath; leaf expansion
Ethiopia, Yemen and India, guinea-caudatum is complete. The peduncle grows rapidly and
is a major sorghum grown in Nigeria and Su- the panicle emerges from the leaf sheath.
dan, and guinea-kafir is grown in East Africa Flowering follows soon after panicle emer-
and India. Kafir-caudatum is widely grown in gence, with the interval largely determined by
the United States and almost all of the modern temperature. Individual panicles start flower-
North American hybrid grain cultivars are of ing from the tip downwards and flowering may
this type. Guinea-caudatum with yellow en- extend over 4-9 days. Sorghum is predomi-
dosperm and large seed size is used in breeding nantly self-pollinating; cross-pollination may
programmes in the United States. range from 0-50%, but is on average about 5 -
The wild representatives are classified as 6%. Grain filling occurs rapidly between flow-
subsp. verticilliflorum (Steud.) Piper (syno- ering and the soft dough stage, with about half
nyms: Sorghum arundinaceum (Desv.) Stapf, the total dry weight accumulating in this pe-
Sorghum bicolor (L.) Moench subsp. arundina- riod. Lower leaves continue to senesce and die.
ceum (Desv.) de Wet & J.R.Harlan): tufted an- By the hard dough stage, grain dry weight has
nual or short-lived perennial, with slender to reached about three-quarters of its final level.
stout culms up to 4 m tall; leaf blade linear- At physiological maturity, determined by the
lanceolate, up to 75 cm x 7cm; panicles usually appearance of a dark layer at the hilum (where
large, somewhat contracted to loose, up to 60 the grain is attached to the panicle), maximum
cm x 25 cm, branches obliquely ascending, dry weight has been achieved. Moisture con-
spreading or pendulous. Wild types extend tent of the grain is usually between 25—35% at
across the African savanna and have been in- this stage. The time taken between flowering
troduced into tropical Australia, parts of India and maturity depends on environmental condi-
and the New World. tions but normally represents about one-third
The weedy plants are usually considered as of the duration of the crop cycle. Further dry-
hybrids between subsp. bicolor and subsp. ver- ing of the grain takes place between physio-
ticilliflorum, and named subsp. drummondii logical maturity and harvest, which usually
(Steud.) de Wet (synonyms: Sorghum x-drum- occurs when grain moisture content has fallen
mondii (Steud.) Millsp. & Chase, Sorghum below 20%. Leaves may senesce rapidly or stay
aterrimum Stapf, Sorghum sudanense (Piper) green with further growth if conditions are
Stapf); they occur in Africa wherever cultivated favourable. Early maturing sorghum cultivars
sorghum and its wild relatives are sympatric take only 100 days or less, whereas long-
because they cross freely. These weedy plants duration sorghum requires 5-7 months. Sor-
occur in recently abandoned fields and field ghum follows the C4-cycle photosynthetic
margins as a very persistent weed; stem up to pathway.
4 m tall; leaf blade lanceolate, up to 50 cm x 6 Ecology Sorghum is primarily a plant of
cm; panicle usually rather contracted, up to 30 hot, semi-arid tropical environments that are
cm x 15 cm, often with pendulous branches. A too dry for maize. It is particularly adapted to
well-known forage grass, 'Sudan grass', belongs drought due to a number of morphological and
to this complex. physiological characteristics, including an ex-
Growth and development The optimum tensive root system, waxy bloom on leaves that
temperature for sorghum seed germination is reduces water loss, and the ability to stop
27-35°C. Seedling emergence takes 3-10 days. growth in periods of drought and resume it
Panicle initiation takes place after approxi- when the stress is relieved. A rainfall of 500-
mately one third of the growth cycle. By this 800 mm evenly distributed over the cropping
stage the total number of leaves (7-24) has season is normally adequate for cultivars ma-
been determined and about one-third of total turing in 3-4 months. Sorghum tolerates wa-
leaf area has developed. Rapid leaf develop- terlogging and can also be grown in areas of
ment, stem elongation and internode expansion high rainfall. It tolerates a wide range of tem-
follow panicle initiation. Rapid growth of the peratures and is also grown widely in temper-
panicle also occurs. By the time the flag leaf is ate regions and at altitudes up to 2300 m in the
visible, all but the final 3 to 4 leaves are fully tropics. The optimum temperature is 25-31°C,
 SORGHUM m

but temperatures as low as 21"C will not dra- more than once as a ratoon crop, e.g. in loca-
matically affect growth and yield. Sterility can tions with a bimodal rainfall pattern. Sorghum
occur when night temperatures fall below 12- is often grown in intercropping systems with
15°C during the flowering period. Sorghum is maize, pearl millet, cowpea, common bean,
susceptible to frost, but to a lesser extent than groundnut and bambara groundnut; in India
maize and light night-frosts during ripening also with pigeonpea.
cause little damage. Sorghum is a short-day Dye cultivars are never grown in large quanti-
plant with a wide range of reactions to photo- ties. Farmers usually grow a few plants in or
period. Some tropical cultivars fail to flower or around their normal sorghum field or near the
to set seed at high latitudes. In the United house.
States, Australia and India the existence of Management Sorghum does not compete
mild photoperiod-sensitive to virtually insensi- well with weeds during the early stages of
tive cultivars has been recorded. growth, and it is recommended that weeding be
Sorghum is well suited to grow on heavy Verti- done early during the seedling stage. In tropi-
sols commonly found in the tropics, where its cal Africa weeding is commonly done once or
tolerance of waterlogging is often required, but twice with a hoe but sometimes animal-drawn
is equally suited to light sandy soils. The best or tractor-drawn cultivators are used. Where
growth is achieved on loams and sandy loams. couch grass (Cynodon dactylon (L.) Pers.) is a
Sorghum tolerates a range of soil pH from 5.0- problem more frequent weeding is necessary.
8.5 and is more tolerant of salinity than maize. Sorghum may be weeded by a combination of
It is adapted to poor soils and can produce inter-row cultivation with animal-drawn im-
grain on soils where many other crops would plements and hand weeding within rows.
fail. Chemical weed control is almost non-existent
In the floodplains of the Senegal and Niger among small farmers. Thinning can be carried
rivers and in parts of Chad and Cameroon sor- out at the same time as hand weeding, or at
ghum is sown in the early dry season when the intervals during the crop cycle, particularly
water recedes, and the crop survives on resid- where thinnings are used to feed livestock.
ual moisture ('culture de décrue'). Subsistence farmers rarely apply fertilizer, but
Propagation and planting Sorghum is application of farmyard manure or ash is com-
normally grown from seed. The 1000-grain mon. In South Africa and the United States
weight is 13-40 g. Seed dormancy is not com- high doses of fertilizers are used in the produc-
mon in cultivated sorghum. A fine seedbed is tion of sorghum. In tropical Africa sorghum is
preferable but is often not achieved. The seed grown mainly as a rainfed crop, but it is grown
is usually sown directly into a furrow following under irrigation in Sudan. It is grown in rota-
a plough, but can also be broadcast and har- tions with maize, pearl millet, finger millet,
rowed into the soil. Optimum plant spacing cotton and other crops. It is often planted late
depends on soil type and availability of mois- in the rotation, as it tolerates low soil fertility.
ture. In low-rainfall areas a population of Under certain conditions decomposing roots of
20,000 plants/ha is normal, in high-rainfall sorghum have an allelopathic effect on the sub-
areas 60,000 plants/ha. For favourable condi- sequent crop, including sorghum.
tions, spacings of 45-75 cm between rows and Diseases and pests Common seed and
15-25 cm within the row, resulting in 80,000- seedling rot diseases in sorghum are caused by
180,000 pockets per ha, are normal; for drier or soil- and seed-borne Aspergillus, Fusarium,
less fertile conditions rows 1m apart, or broad- Pythium, Rhizoctonia and Rhizopus spp. They
casting at 6 kg seed per ha. Aplanting depth of are controlled by treatment of the seed with
2.5-5 cm is common, and up to 25 seeds may be fungicides. Anthracnose (Colletotrichum gra-
sown per pocket. Occasionally, seedlings are minicola) is common in hot and humid parts of
grown in a nursery and transplanted into the Africa. Control measures include the use of
field early in the dry season, e.g. on the flood- resistant cultivars and crop rotation. Downy
plains round Lake Chad in Africa ('sorgho repi- mildew (Peronosclerospora sorghi) may cause
qué'). Sweet sorghum in the United States is serious yield losses; the use of resistant culti-
also sometimes transplanted. Sorghum can vars and seed treatment are recommended.
also be propagated vegetatively by splitting Smuts (Sporisorium spp.) are important pani-
tillers from established plants and transplant- cle diseases. Loose and covered kernel smut are
ing them, a practice that is often used by small controlled by seed treatment with fungicides;
farmers to fill gaps. Sorghum may be harvested head smut and long smut by using resistant
172 CEREALS AND PULSES

cultivars and cultural practices such as crop (Sitophilus oryzae), flour beetle (Tribolium
rotation and removal of infected panicles. castaneum) and the grain moth (Sitotroga ce-
Grain mould is caused by a complex of fungal realella). Damage can be minimized by drying
pathogens (predominantly Cochliobolus luna- grain adequately before storage. Cultivars with
tus (synonym: Curvularia lunata), Fusarium hard grain also suffer less damage.
spp. and Phoma sorghina) that infect the grain The parasitic weed Striga (especially Striga
during development and can lead to severe hermonthica (Del.) Benth., but also Striga asi-
discoloration and loss of quality. It is most se- atica (L.) Kuntze, Striga densiflora Benth. and
vere in seasons when rains continue through Striga forbesii Benth.) has become a major
the grain maturity stage and delay the harvest. constraint to sorghum cultivation, particularly
Control measures include adjustment of the in Africa, where severe infestations can lead to
sowing date to avoid maturation during wet grain losses of 100% and land being aban-
weather, and the use ofresistant cultivars. doned. Striga can be controlled by cultural
Important pests of sorghum in tropical Africa methods such as rotation with trap crops or
are shoot fly (Atherigona soccata) and stem with crops that are not susceptible (e.g.
borers (particularly Busseola fusca, Chilo par- groundnut, cotton or sunflower), rigorous re-
tellus and Sesamia calamistis). Shoot fly larvae moval of the weeds before flowering and appli-
attack shoots of seedlings and tillers, and cause cation of nitrogen fertilizer and herbicides. A
'dead hearts'. Stem borers cause damage in all few sorghum cultivars that are resistant or
crop stages. Damage by both shoot fly and stem tolerant to Striga have been identified.
borers can be reduced by early, non-staggered Harvesting Sorghum is usually harvested
planting and seed or soil treatment with insec- when the grain moisture content has fallen
ticides. Resistance to shoot fly is associated below 20%, and the grain has become hard.
with low yield. Foliage pests include army Harvesting is done by hand using a knife to cut
worms (Spodoptera and Mythimna spp.); they the panicles, which are temporarily stored in
are controlled by contact insecticides. Larvae of sacks before being taken to the threshing floor
the sorghum midge (Stenodiplosis sorghicola, for further drying to a moisture content of 12—
synonym: Contarinia sorghicola) feed on the 13%. Alternatively, the whole plant is cut or
young grains in the panicle. Damage can be pulled up and the panicle removed later. Com-
limited by sowing early-maturing cultivars and bine harvesting is possible, but many small
avoiding staggered planting. Head bugs (Eu- farmers cannot afford to buy the machinery. In
rystylus and Calocoris spp.) suck on developing South Africa combine harvesting is more com-
grains, resulting in yield loss, grain deforma- mon.
tion and discoloration and infection by moulds. For dye production, leaf sheaths are harvested
Guinea type sorghum is generally less affected. when the plant comes to maturity, about 4-6
In practice, control methods of diseases and months after sowing. They can be used imme-
pests are mainly preventative or cultural, in- diately or dried and stored.
cluding selection of optimum planting dates, Rainfed forage sorghum is usually cut only
seed treatment and crop rotation. Early sowing once, soon after flowering. Forage sorghum
is particularly important as a mechanism to crops grown under more favourable conditions,
avoid large insect populations at times when often with irrigation and high levels of fertil-
plants are most susceptible to damage. High izer, can be harvested and then left to regrow
levels of host plant resistance are available for (ratoon).
sorghum midge, but only low levels of resis- Broomcorn is harvested by hand as mechanical
tance for the other pests. Chemical control of harvesters are not available. Sweet sorghum is
diseases and insect pests is rarely practised in harvested when the seed is in the soft dough
tropical Africa. stage when the sugar content of the stalk is
Birds, especially Quelea quelea, cause impor- highest.
tant yield losses. Control measures include the Yield Average sorghum grain yields on
choice of suitable planting dates, timely har- farmers' fields in Africa are as low as 0.5-0.9
vesting, bird scaring and the destruction of t/ha because sorghum is often grown in mar-
roosting and nesting sites. Brown sorghum is ginal areas under traditional farming practices
less preferred by birds than the tannin-free (low inputs, traditional landraces). Under fa-
white sorghum. vourable conditions sorghum can produce grain
Sorghum is very susceptible to damage by yields up to 13 t/ha. In South Africa, with in-
storage pests, the main ones being rice weevil tensive agricultural practices and improved
 SORGHUM 173

cultivars, average commercial yield was 2.3 shaken. Alternatively, the dye liquid is painted
t/ha in 2001. In China, where sorghum is on the tanned surface with the fingers or a
grown with high levels of inputs, yield aver- brush. The hide is then rinsed in cold water
ages 3.6 t/ha and in the United States 3.8 t/ha. acidulated with lime juice or tamarind pulp.
Forage yields from single-cut cultivars and After the hide has been dried, the process is
hybrids can reach 20 t/ha of dry matter. Multi- completed by rubbing the hide with a smooth
cut cultivars and hybrids usually give only stone on a wooden block. It is estimated that
slightly higher total yields but produce better 1.25 1 of dye bath is sufficient for about 6 skins
quality forage. Sweet sorghum yields about of medium size. Another recipe uses about 30
1000 1 syrup per ha in the United States. Aver- leaf-sheaths of sorghum, about half a spoonful
age broomcorn yields are 300-600 kg/ha, of soda, a handful of'sant' pods (Acacia nilotica
enough to make 350-800 brooms. (L.) Willd. ex Delile) or 2 handfuls of chips of
Handling after harvest The harvested mangrove bark, 2 spoonfuls of palm oil and 1.5
grain of sorghum is usually sun-dried, often in 1 of water. These are all mixed together and
the panicle. Panicles, particularly those to be boiled, the juice of 5 or 6 limes added, and the
retained for seed, may be stored hanging from liquid is left to simmer for 2 hours. It is then
the ceiling of kitchens over cooking fires where ready for application on the skin by brushing
the smoke helps to deter insect attack. Alterna- or rubbing.
tively, the heads may be threshed after drying To obtain a dye of constant high quality, a
and the grain stored in granaries, above or laboratory extraction technique has been de-
below ground, designed to prevent insect at- signed in Burkina Faso. Sorghum leaf sheaths
tack. are crushed into fine particles, a solvent is
Traditional food preparation of sorghum is added in an acid or basic medium (both give
quite varied. The whole grain may be ground similar results) and a red liquid is produced.
into flour or decorticated before grinding to By addition of an acid the dyestuff is precipi-
either a fine particle product or flour which is tated and is centrifuged off. The end product is
then used in various food products. To prepare a fine, burgundy-red powder with an apigen-
porridge, water is boiled and sorghum flour is inidin concentration of 50-60%, ready for use
gradually added until the desired consistency as a dye. Pure apigeninidin can be obtained by
of the paste is reached. Regular stirring is further processing ofthe powder.
needed to mix the contents thoroughly. An- Forage sorghum can be fed to livestock while
other simple form of sorghum food preparation still green or can be stored in various ways for
is to boil the grain before or after decorticating. later use. The forage is often dried and stacked
To make beer, sorghum grain is germinated, or can be made into silage. Stover left after
dried, pounded into flour and mixed with water harvest of grain is often grazed by animals.
and left to ferment in a warm place for some Genetic resources A major collection of
days. To make the non-fermented drink sorghum germplasm is maintained and dis-
'mageu' in Botswana and South Africa, milled tributed to interested researchers by the Inter-
sorghum malt is mixed with water and kept at national Crops Research Institute for the Semi-
room temperature for 2-3 days. Occasional Arid Tropics (ICRISAT), Patancheru, India.
stirring may be necessary. The collection extends to over 36,000 acces-
In a traditional method of dyeing hides with sions from all the major sorghum-growing re-
sorghum dye in West Africa, a watery extract gions of the world (90 countries). Large germ-
of wood ashes, preferably from the wood of plasm collections of sorghum are also held in
Anogeissus leiocarpa (DC.) Guill. & Perr., is the United States (Southern Regional Plant
prepared and allowed to stand for 3-4 hours. Introduction Station, Griffin, Georgia, 30,100
The major active compound of the lye is potas- accessions; National Seed Storage Laboratory,
sium- or sodium carbonate. The red leaf Fort Collins, Colorado, 10,500 accessions) and
sheaths are pulverized and placed in a large China (Institute of Crop Germplasm Resources
vessel in which the dyeing is carried out. From (CAAS), Beijing, 15,300 accessions). In tropical
time to time a little lye is added and diluted Africa large germplasm collections of sorghum
with plain water as desired, obtaining a crim- are held in Zimbabwe (SADC/ICRISAT Sor-
son liquid. The tanned hide that has been ghum and Millet Improvement Program, Ma-
dressed with oil is folded with the tanned side topos, 12,340 accessions), Ethiopia (Institute of
outwards, the hide is immersed for about two Biodiversity Conservation (IBC), Addis Ababa,
minutes in the dye bath, wrung out and 7260 accessions), Kenya (National Genebank of
174 CEREALS AND PULSES

Kenya, Crop Plant Genetic Resources Centre, formation, but the efficiency is generally low,
KARI, Kikuyu, 3410 accessions) and Uganda especially with the former technique.
(Serere Agricultural and Animal Production Prospects Sorghum is a hardy, drought-
Research Institute, Serere, 2635 accessions). tolerant crop with a high potential yield, which
Breeding The main objectives in sorghum plays an important role in tropical Africa and
breeding include high grain yield, white grain elsewhere, especially as source of food and fod-
for human consumption with improved nutri- der, but also for a range of other uses, includ-
tional value and processing quality, and red or ing as a source of dye. Sorghum has lost part of
brown grain for feed purposes and brewing. In its traditional area in tropical Africa to maize,
many countries the emphasis is on producing which yields better in more favourable envi-
cultivars which combine high grain yield with ronments, is less liable to bird damage and
high stover yields because of the importance of easier to process. It is to be expected, however,
the residues as animal feed. Incorporation of that sorghum will remain an important food
resistance to major yield-limiting diseases and security crop in less favourable environments
pests, and tolerance of abiotic stresses are also in tropical Africa. Important problems in sor-
of high priority. Resistance to grain moulds ghum cultivation to be addressed by research
and other diseases as well as to insect pests and breeding activities are the large yield
such as head bugs and sorghum midge has losses caused by parasitic weeds (especially
been identified. High-yielding improved culti- Striga hermonthica), anthracnose, downy mil-
vars of sorghum are available in most of the dew, grain moulds, sorghum midge and stem
main producing countries. These include culti- borers. Improved sorghum cultivars are not
vars and hybrids produced using cytoplasmic widely grown in tropical Africa, and the im-
male sterility. Compared to traditional land- provement of seed supply systems should ac-
races they have a weak photoperiodic response company sorghum improvement programmes
and they are less hardy, less tall, with a lower in this region. Demand for sorghum for non-
grain quality but a higher yield potential. traditional uses is likely to increase. In particu-
Striga-resistant cultivars have been released in lar, the use of sorghum as a feed grain, already
Africa and India, e.g. 'Framida' in Ghana and well established in many industrialized coun-
Burkina Faso. Cultivars resistant to grain tries, is likely to become more common in de-
mould have also been released. Special culti- veloping countries. However, sorghum faces
vars with high biomass production and good strong competition from maize in the interna-
forage quality are bred for animal feed. Modern tional feed grain market. Similarly, as in-
sorghum cultivars predominate in the Ameri- creased affluence results in increased demand
cas, China and Australia, but in Africa they for meat and dairy products, the use of sor-
occupy probably less than 10% of the area un- ghum as a forage crop in intensive production
der sorghum. In India about 50% of the sor- systems in many tropical regions is likely to
ghum area is sown to modern cultivars and expand. The use of sorghum as a raw material
50% to traditional landraces. for industrial processes will also increase. Re-
The sorghum genome is relatively small (about search should focus on innovations that are
760 Mbp) compared to that of maize (about likely to reduce the costs of production of sor-
2500 Mbp), and construction of a physical ge- ghum. This should include research to increase
nome map is in progress. Several genetic link- yield levels of available cultivars, and to im-
age maps have been developed, mainly based prove agronomic practices. Emphasis should be
on RFLP markers. Various genes have been placed on enhancing resistance to the main
tagged, e.g. genes associated with head smut biotic and abiotic stresses and on production of
resistance, leaf blight resistance and shatter- cultivars richer in high quality proteins.
ing. Many QTLs have been mapped, including Sorghum dye may profit from the trend of in-
those associated with plant height, tillering, creasing use of natural colourants in foods and
seed size, drought resistance and rust resis- cosmetics. Rising harvesting costs of broomcorn
tance. In-vitro plant regeneration has been in North America and Europe may offer possi-
achieved from calli derived from young leaf bilities for expanding this commodity in Africa.
bases, shoot apices, immature inflorescences Major references Chantereau et al., 1997;
and immature embryos. Protocols have been de Vries & Toenniessen, 2001; de Wet, 1978;
developed for the production of stably trans- Doggett, 1988; Murty & Renard, 2001; Rooney
formed sorghum plants using microprojectile & Serna-Saldivar, 2000; Smith & Frederiksen,
bombardment orAgrobacterium-mediated trans- 2000; Stenhouse & Tippayaruk, 1996; Sten-
 SPOROBOLUS 175

house et a l , 1997;Taylor, 2003. short tertiary branchlets. Spikelet 1.5-2.5 mm

Other references Balole, 2001; Bellemare, long, dark green, 1-flowered; lower glume nar-
1993; Burkill, 1994; Byth (Editor), 1993; Dal- rowly oblong to lanceolate, 0.5-1.5 mm long,
ziel, 1926; Gao et al., 2005; Harlan & de Wet, veinless, upper glume narrowly ovate, 1.5-2
1972; Kouda-Bonafos et al., 1994; Ministry of mm long, 1-veined; lemma narrowly ovate, as
Agriculture and Rural Development, 2002; long as the spikelet or almost so, 1-veined;
National Research Council, 1996; Neuwinger, palea similar to lemma, but 2-veined; stamens
2000; Ogwumike, 2002; Pale et al., 1997; Phil- 3, c. 1mm long; ovary superior, with 2 plumose
lips, 1995; Reddy, Ramesh & Reddy, 2004; Rey stigmas. Fruit a caryopsis (grain), obovoid, c.
et a l , 1993; Sanders, Ahmed & Nell, 2000; 0.5 mm long, truncate, tetragonal in section.
Seshu Reddy, 1991; USDA, 2004; Westphal, Sporobolus comprises about 160 species and
1981. occurs in the tropics and subtropics, extending
Sources of illustration Stenhouse & Tip- into warm temperate regions. It may resemble
payaruk, 1996. Eragrostis, which differs in its 2-many-
Authors T.V. Balole &G.M. Legwaila flowered spikelets (1-flowered in Sporobolus)
Based on PROSEA 10: Cereals. and 3-veined lemma (1-veined in Sporobolus).
The species of Sporobolus are often difficult to
identify because they intergrade to such an
SPOROBOLUS FIMBRIATUS (Trin.) Nees extent that their limits are often not sharply
defined. This is also the case for the variable
Protologue Fl.Afr. austral, ill.: 156 (1841). Sporobolus fimbriatus. Sporobolus fimbriatus
Family Poaceae (Gramineae) follows the C4-cycle photosynthetic pathway.
Chromosome number 2n - 18, 36, 54 Ecology Sporobolus fimbriatus is commonly
Vernacular names Dropseed, perennial found up to 2000 m altitude in open woodland
dropseed grass, fringed dropseed (En). and grassland, often in shallow rainwater
Origin and geographic distribution Sporo- pans, sometimes on rocky hillsides, also in dis-
bolus fimbriatus is found wild and occasionally turbed or shady locations.
cultivated from Sudan and Somalia south- Management The grain of Sporobolus fim-
wards to South Africa. It has been introduced briatus is mostly collected from the wild. In
elsewhere, e.g. into the United States. experiments in South Africa ungrazed planted
Uses In southern Africa the grains of Sporo- pasture of Sporobolus fimbriatus produced 3.3
bolus fimbriatus are eaten during times of food t dry matter per ha per year, and grazed pas-
shortage; they may be ground to prepare a por- ture 2.7 t dry matter per ha per year.
ridge. Sporobolus fimbriatus is a good pasture Genetic resources and breeding A collec-
grass and is browsed by stock, e.g. sheep and tion of 47 accessions of Sporobolus fimbriatus
cattle. It has been planted for soil stabilization. (46 from South Africa and 1 from Botswana) is
Properties In South Africa the crude pro- held in the United States (USDA-ARS Western
tein content of Sporobolus fimbriatus ranges Regional Plant Introduction Station, Pullman,
from 14%in spring to 10%in autumn, and the Washington). In Africa germplasm collections
digestibility from 70% in spring to 63% in au- are held in Kenya (National Genebank of
tumn. The plant may contain hydrocyanic acid, Kenya, Crop Plant Genetic Resources Centre,
but poisoning is seldom a problem. KARI, Kikuyu, 21 accessions), South Africa
Botany Perennial, tufted grass up to 1.7 m (Grassland Research Centre, Department of
tall, with a short rhizome; stem (culm) 2-3 mm Agricultural Development, Pretoria, 4 acces-
in diameter at the base, erect, usually un- sions) and Ethiopia (International Livestock
branched. Leaves mostly basal, simple; basal Research Institute (ILRI), Addis Ababa, 1 ac-
leaf sheath papery, glabrous or hairy along the cession). In view of its wide distribution and
margins, terete to strongly compressed and common occurrence Sporobolus fimbriatus is
keeled, persistent; ligule ciliate; leaf blade lin- not threatened by genetic erosion.
ear, 10-30(-60) cm x 2-7.5(-14) mm, tapering Prospects The present role of Sporobolus
to a filiform apex, flat, folded or involute, the fimbriatus seems limited to being a local source
white midrib prominent above, rough on the of food during times of shortage and of fodder.
surfaces. Inflorescence a panicle 15-65 cm It is unlikely to increase in importance in the
long, linear to lanceolate, the branches not in future.
whorls, 2-12 cm long, smooth or somewhat Major references Clayton, Phillips & Ren-
rough, with the spikelets on the secondary or voize, 1974; Cope, 1995; Cope, 1999; Gibbs Rus-
176 CEREALSANDPULSES

sell et al., 1990;Phillips, 1995. spikelets on the panicle branches and the par-
Other references Ben-Shahar, 1991; du tially or complete sterile lowermost panicle
Pisani & Knight, 1988; Hanelt & Institute of branch whorl.
Plant Genetics and Crop Plant Research (Edi- Ecology Sporobolus panicoides is locally
tors), 2001; Klaassen & Craven, 2003; Myre, common in sunny or lightly shaded locations,
1972; Sânchez-Monge y Parellada, 1981; Shar- up to 2100 m altitude, in woodland on sandy
ma &Sharma, 1979;van der Westhuizen et al., soils, in granite sandveld and on rocky hill-
2001; van Wyk & Gericke, 2000; Watt & sides, often at roadsides or in other disturbed
Breyer-Brandwijk, 1962. localities.
Authors M. Brink Management The grains of Sporobolus
panicoides are only collected from the wild.
Genetic resources and breeding One ac-
S P O R O B O L U S PANICOIDES A.Rich. cession of Sporobolus panicoides is kept at the
National Genebank of Kenya, Crop Plant Ge-
Protologue Tent. fl. abyss. 2: 399 (1850). netic Resources Centre, KARI, Kikuyu. Sporo-
Family Poaceae (Gramineae) bolus panicoides is not threatened by genetic
Vernacular names Famine grass (En). erosion as it is widespread and locally common.
Origin and geographic distribution Sporo- Prospects The present role of Sporobolus
bolus panicoides is found in East and southern panicoides is very limited, being a local source
Africa from Sudan and Ethiopia southwards to of food during times of shortage. It is not prob-
South Africa, and in tropical Arabia. able that it will become more important in the
Uses The grains of Sporobolus panicoides future.
are eaten during times offood shortage. Major references Clayton, Phillips & Ren-
Botany Annual, slender grass up to 1m tall; voize, 1974; Cope, 1995; Cope, 1999; Gibbs Rus-
stem (culm) erect, solitary or tufted. Leaves sellet al., 1990;Phillips, 1995.
simple; leaf sheath papery, glabrous, but hairy Other references Fröman &Persson, 1974;
near the margins, slightly compressed; blade IPGRI, undated; Klaassen & Craven, 2003;
linear, 5-30 cm x 2-6 mm, attenuate at apex, Mackie, 1976; Shava &Mapaura, 2002.
flat or involute, pale green, glabrous or Authors M. Brink
sparsely hairy above. Inflorescence a narrowly
ellipsoid panicle 4-22 cm long, the branches in
a succession of whorls, with 1-4 spikelets per TRITICUMAESTIVUM L.
branch. Spikelet 2-3.5 mm long, pallid with
purple tinge above, 1-flowered; lower glume1— Protologue Sp. pi. 1: 85 (1753).
1.5 mm long, rarely minute, narrowly ovate to Family Poaceae (Gramineae)
lanceolate, obtuse to acute at apex, veinless, Chromosome number 2re= 42
glabrous, upper glume as long as the spikelet, Synonyms Triticum vulgare Vill. (1787).
elliptical-oblong to ovate, acute at apex, 1- Vernacular names Bread wheat, common
veined, glabrous; lemma a little shorter than wheat, wheat (En). Blé tendre, blé, froment
spikelet, elliptical-ovate, 1-veined; palea 2- (Fr). Trigo mole, trigo (Po). Ngano (Sw).
veined; stamens 3, 1—1.5 mm long; ovary supe- Origin and geographic distribution Bread
rior, with 2 plumose stigmas. Fruit a caryopsis wheat arose in the corridor extending from
(grain), 1-2 mm in diameter, oblong-globose, Armenia in Transcaucasia to the south-west
bright brown or orange. coastal areas of the Caspian Sea in Iran. Hy-
Sporobolus comprises about 160 species and bridization of a wild Aegilops species (Aegilops
occurs in the tropics and subtropics, extending tauschii Coss., with the D-genome) with em-
into warm temperate regions. It resembles mer, an old type of cultivated wheat belonging
Eragrostis, which differs in its 2-many- to Triticum turgidum L., gave rise to the
flowered spikelets (1-flowered in Sporobolus) hexaploid wheats, but it is unknown whether
and 3-veined lemma (1-veined in Sporobolus). bread wheat or spelt wheat (Triticum spelta L.)
The species of Sporobolus are often difficult to appeared first. The earliest archaeological
identify because they intergrade to such an finds of spelt wheat are from the southern
extent that their limits are often not sharply Caspian area and are dated at around 5000
defined. However, Sporobolus panicoides is BC. Finds of bread wheat are difficult to dis-
easily recognized by its comparatively large, tinguish from durum wheat (Triticum tur-
brightly coloured grain, the sparsity of gidum), but one thinks that those found in the
 TRITICUM 177

as a snack and during social gatherings as 'ni-

tro' (boiled whole grain often mixed with
pulses), 'kollo' (roasted grain) and 'dabo-kollo'
(ground and seasoned dough, shaped and deep
fried).
Industrial uses of wheat products centre on the
production of glues, alcohol, oil and gluten.
By-products of flour milling, particularly the
bran, are used almost entirely to feed livestock,
poultry or prawns. Wheat germ (from wheat
embryos) is sold as a human food supplement.
Straw is fed to ruminants or used for bedding
material, thatching, wickerwork, newsprint,
cardboard, packing material, fuel and as sub-
strate for mushroom production. In many dry
parts ofthe world it is chopped and mixed with
clay to produce building material.
Triticum aestivum -planted Production and international trade Accord-
ing to FAO estimates, the average world pro-
Caucasus, on the anatolian plateau (Turkey), duction of wheat grain (bread wheat and du-
in Central Europe and in Central Asia from the rum wheat together) in 1999—2003 amounted
fifth millennium onwards belong to bread to 576 million t/year from 209 million ha.
wheat. The D-genome in fact conferred to bread Worldwide, bread wheat constitutes more than
wheat and spelt wheat the adaptation to cold 90% of the area under the cultivated wheats.
winters and humid summers, allowing them to The main wheat producing countries are China
conquer temperate Eurasia, whereas the Medi- (96.8 million t/year from 25.2 million ha), India
terranean remained the area of emmer and (71.0 million t/year from 26.4 million ha), the
durum wheat. By the third millennium BC, United States (56.9 million t/year from 20.6
bread wheat had reached China. In 1529, the million ha), the Russian Federation (39.4 mil-
Spanish took it to the New World. Bread wheat lion t/year from 21.7 million ha) and France
was introduced into tropical Africa by Arab (35.1 million t/year from 5.0 million ha). Wheat
traders, missionaries and colonial settlers. It is production in tropical Africa in 1999-2003 was
not known exactly when it reached Ethiopia. It 2.5 million t/year from 1.6 million ha, the main
was brought from northern Africa to West Af- producing countries being Ethiopia (1.4 million
rica, where it was already known around 1000 t/year from 1.1 million ha), Kenya (272,000
AD. In the early 20 th century it was introduced t/year from 137,000 ha), Sudan (254,000 t/year
into Kenya and eastern DR Congo. from 124,000 ha), Zimbabwe (237,000 t/year
Bread wheat today is grown in almost all parts from 43,000 ha), Zambia (87,000 t/year from
of the world. In tropical Africa, it is mainly 13,000 ha), Tanzania (82,000 t/year from
produced in Nigeria, Sudan, Ethiopia, Kenya, 60,000 ha) and Nigeria (75,000 t/year from
Tanzania, Zambia and Zimbabwe. 53,000 ha). In Ethiopia close to 50% of the
Uses Bread wheat flour is made into numer- wheat production consists of bread wheat, the
ous products including bread (leavened or flat; other 50%of durum wheat. From 1961-1965 to
baked, steamed or deep fried), pastries, crack- 1999-2003 the world production of wheat in-
ers, biscuits, pretzels, noodles, farina, break- creased from 248 to 576 million t/year, whereas
fast foods, baby foods and food thickeners. It is the harvested area remained stable at around
also used as a brewing ingredient in certain 210 million ha. In the same period the wheat
beverages (white beer). Leavened breads are production in tropical Africa increased from
the most popular use of wheat in almost all 960,000 to 2.5 million t/year, and the harvested
parts of the world. Increased bread consump- area from 1.2 to 1.6 million ha.
tion is often linked to increasing urbanization Average world export of wheat amounted to
and higher per capita income. Bread wheat 115 million t/year in 1998-2002, the main ex-
utilization has also been adapted to local cui- porters being the United States (26.7 million
sine. In Ethiopia, for instance, the flour is used t/year), Canada (16.5 million t/year), Australia
to prepare 'injera' (pancake-like unleavened (15.9 million t/year), France (15.9 million
bread), porridge and soup. The grain is eaten t/year) and Argentina (10.0 million t/year).
178 CEREALS AND PULSES

Main importers are Italy, Brazil, Japan and as gluten, comprising 78-85% of the total
Iran, each importing more than 5 million wheat endosperm protein. Gluten is composed
t/year. All countries in tropical Africa are net mainly of glutenin (polymeric) and gliadin
importers. The main importer in tropical Africa (monomeric ) proteins. Glutenins confer elastic-
is Nigeria (1.9 million t/year in 1998-2002), ity and dough strength, while gliadins confer
followed by Ethiopia (770,000 t/year), Sudan mainly viscous flow and extensibility to the
(710,000 t/year) and Kenya (570,000 t/year). gluten complex. Wheat flour contains roughly
The share of food aid in wheat imports is as equal amounts of glutenins and gliadins, and
high as 80%for some countries. their imbalance may influence its visco-elastic
Properties The composition of wheat grain properties.
is 7-8% coat material, 90% endosperm and 2 - Description Annual, tufted grass up to 150
3% embryo. The embryo mainly comprises oil cm tall, with 2-5(-40) tillers; stem (culm) cy-
and protein, and little starch. The endosperm lindrical, smooth, hollow except at nodes.
is starchy, and is surrounded by the aleurone Leaves distichously alternate, simple and en-
layer which is rich in proteins. When a wheat tire; leaf sheath rounded, auricled; ligule mem-
grain is milled, the outer layers and embryo branous; blade linear, 15-40 cm x 1—2 cm, par-
are separated from the endosperm. The pulver- allel-veined, flat, glabrous or pubescent. Inflo-
ized endosperm becomes wheat flour, while the rescence a terminal, distichous spike 4—18 cm
other parts form the bran. The endosperm var- long, with sessile spikelets borne solitary on
ies both in hardness and vitreousness: hard zigzag rachis. Spikelet 10-15 mm long, later-
bread wheat grain high in gluten protein tends ally compressed, 3-9-flowered, with bisexual
to be vitreous and low-protein soft wheat grain florets, but 1-2 uppermost ones usually rudi-
tends to be opaque. Hard bread wheat grain is mentary, sometimes only 1 of the florets bisex-
best suited for bread making while the soft ual; glumes almost equal, oblong, shorter than
wheat grain is best for cookies, cakes and pas- spikelet, thinly leathery, keeled towards the
tries. Flour colour varies from white to slightly tip, apiculate to awned; lemma rounded on
yellow. back but keeled towards the tip, leathery,
Bread wheat grain (hard red spring type) con-
tains per 100 g edible portion: water 12.8 g,
energy 1377 kJ (329 kcal), protein 15.4 g, fat
1.9 g, carbohydrate 68.0 g, dietary fibre 12.2 g,
Ca 25 mg, Mg 124 mg, P 332 mg, Fe 3.6 mg, Zn
2.8 mg, vitamin A 9 IU, thiamin 0.50 mg, ribo-
flavin 0.11 mg, niacin 5.7 mg, vitamin BÖ 0.34
mg, folate 43 ug and ascorbic acid 0 mg. The
essential amino-acid composition per 100 g
edible portion is: tryptophan 195 mg, lysine
404 mg, methionine 230 mg, phenylalanine 724
mg, threonine 433 mg, valine 679 mg, leucine
1038 mg and isoleucine 541 mg. The principal
fatty acids are per 100 g edible portion: linoleic
acid 727 mg, palmitic acid 283 mg and oleic
acid 236 mg. Soft, white bread wheat grain
contains per 100 g edible portion: water 10.4 g,
energy 1423 kJ (340 kcal), protein 10.7 g, fat
2.0 g, carbohydrate 75.4 g, dietary fibre 12.7 g,
Ca 34 mg, Mg 90 mg, P 402 mg, Fe 5.4 mg, Zn
3.5 mg, vitamin A 9 IU, thiamin 0.41 mg, ribo-
flavin 0.11 mg, niacin 4.8 mg, vitamin B6 0.38
mg, folate 41 |ig and ascorbic acid 0 mg (USDA,
2005). Bread wheat grain is deficient in the
amino acids lysine and threonine, and some-
what in isoleucine and valine. It is a good Triticum aestivum - 1, lower part of plant; 2,
source of B-group vitamins and minerals. ligule and auricles; 3, inflorescence; 4, spikelet;
Wheat grain possesses a unique viscoelastic 5, floret (lemma andpalea removed); 6, grains.
and insoluble storage protein complex known Source: PROSEA
 TRITICUM 179

awned or blunt; palea 2-keeled, hairy on the tics from the parents, combining the hardiness
keels; lodicules 2, ciliate; stamens 3; ovary su- of rye with the high yield and quality of wheat.
perior, tipped by a small fleshy hairy append- Triticale is presently grown only locally in
age and with 2 plumose stigmas. Fruit an el- tropical Africa, e.g. in Ethiopia, Kenya, Tanza-
lipsoid caryopsis (grain), at one side with a nia and Madagascar, and also in northern Af-
central groove, reddish brown to yellow or rica and South Africa. As a new food crop, it
white. fell short of expectations, but it is becoming
Other botanical information Triticum is a increasingly popular as a forage crop.
classic example of allopolyploidy consisting of Growth and development Germination of
diploid (2/i - 14), tetraploid (2n = 28) and wheat occurs at temperatures of 4-37°C, the
hexaploid (2re = 42) species. Selection at the optimum being 12-25°C. The radicle emerges
diploid and tetraploid levels has proceeded first and the coleoptile emerges 4-6 days after
from wild species with hulled grain and brittle germination. The primary roots may remain
rachis to the free-threshing species with tough functional for life unless destroyed by disease
rachis; hexaploid wheats are not known in the or mechanical injury, but they constitute only a
wild, they appeared in cultivation. The classifi- small portion of the total root system. The first
cation of the genus Triticum and other related true leaf of the seedling emerges from the col-
genera within the tribe Triticeae was strongly eoptile. Secondary roots start to develop about
debated. Polyploidy and biphyletic genome two weeks after seedling emergence. They arise
differentiation (B vs. G genome) are isolating from the basal nodes and form the permanent
mechanisms offering adequate species borders. root system, which spreads out and may pene-
In this approach, Triticum comprises only 5-6 trate as deep as 2 m, but normally no more
species, including the diploid Triticum mono- than 1 m. Leaf and tiller production increase
coccum L. (einkorn, grown sporadically in rapidly soon after crop emergence. The dura-
southern Europe and western Asia), the tion of the vegetative stage may vary from 20-
tetraploid Triticum turgidum L. and the hexa- 150 days depending on temperature and the
ploid Triticum aestivum L. (comprising all cul- cultivar's vernalization and daylength re-
tivated hexaploids). Spelt wheat (Triticum sponse. For floral induction, spring types usu-
spelta L.) is sometimes separated from Triti- ally require temperatures between 7°C and
cum aestivum. It is a hexaploid, not free- 18°C for 5-15 days, while winter types require
threshing wheat, with only 2—3 florets per temperatures between 0°C and 7°C for 30-60
spikelet, cultivated in small quantities in days. Flowering begins at the middle third of
Europe, Africa and on the plateau of western the spike and continues towards the basal and
Iran. It can be cultivated under extreme cir- apical parts in 3-5 days. All spike-bearing till-
cumstances, not demanding fertile soils, being ers eventually flower almost simultaneously.
relatively disease resistant, and having good Wheat is normally self-pollinated; cross-
taste, food and baking qualities. Before 1850 it pollination is 1-4%. Pollen is largely shed
was a very important wheat in Europe, declin- within the floret. Stigmas remain receptive for
ing afterwards, especially because it has to be 4-13 days. Pollen is viable for up to 30 minutes
hulled before milling, but is now gaining in only. Grains in the centre of the spike and in
popularity in organic wheat cultivation. the proximal florets tend to be larger than the
Commercially, wheat is classified into distinct other ones. Physiological maturity is reached
categories of grain hardness (soft, medium- when the flag leaf (uppermost leaf) and spikes
hard, and hard) and colour (red, white and turn yellow and the moisture content of the
amber). Based on growing habit, bread wheat fully formed grain has dropped to 25—35%.The
is divided into two subclasses, spring or winter, complete crop cycle of bread wheat varies from
but facultative types exist. These subclasses in 50-200 days in tropical Africa.
turn may also be divided into grades, which are Ecology Bread wheat can be grown from
generally used to adjust prices, based mainly within the Arctic Circle to near the equator,
on grain soundness (effects of rain, heat, frost, but it is most successful between 30-60°N and
insect and mould damage), cleanliness, grain 27-40°S. Optimum temperatures for develop-
protein content and cc-amylase activity. In ment are 10-24°C, with minima of 3-4°C and
tropical Africa mostly spring wheats are grown. maxima of 30-32°C. An average temperature
Hybrids of wheats (tetraploid or hexaploid) and of about 18°C is optimal for yield. Tempera-
rye called triticale (xTriticosecale) have been tures above 35°C stop photosynthesis and
developed and these show a mix of characteris- growth, and at 40°C the heat kills the crop.
180 CEREALS AND PULSES

Wheat does not grow well under very warm In tropical Africa bread wheat is produced
conditions with high relative humidity, and in mainly under rainfed conditions, except in Ma-
the tropics it is best grown at higher elevations lawi, Zambia and Zimbabwe where it is grown
(1200-3000 m) or in the cooler months of the as an irrigated (flood and sprinkler) 'winter'
year. Bread wheat requires at least 250 mm season crop. In Nigeria wheat production is
water during the growing season for a good restricted to the river basin irrigation schemes
crop; it can be grown in areas that receive 250- of its northern states. Irrigation has great po-
750 mm rain annually. The sensitivity to tential to increase wheat production in Sudan
daylength differs among genotypes, but most and Somalia. Care must be taken not to
are quantitative long-day plants; they flower over-irrigate since wheat is sensitive to early
earlier at long daylengths, but they do not re- waterlogging. Irrigation timing is based either
quire a particular daylength to induce flower- on pre-defined crop stages or on estimates of
ing. soil moisture depletion.
Soils best suited for bread wheat production The mean nutrient removal per 1 t/ha of grain
are well aerated, well drained, and deep, with is 40-43 kg N, 5-8 kg P, 25-35 kg K, 2-4 kg S,
0.5% or more organic matter. Optimum soil pH 3-4 kg Ca, 3-3.5 kg Mg, and smaller amounts
ranges between 5.5 and 7.5. Wheat is sensitive of micronutrients. The exact values depend on
to soil salinity. the available nutrients and water in the soil,
Propagation and planting Bread wheat is the temperature, and the cultivar. Average
propagated by seed. The 1000-seed weight is fertilizer rates in tropical Africa range from 9
30-50 g. It is advisable to use certified seed kg N and 10 kg P on rainfed wheat in Ethiopia
that has been treated with fungicides against to 180 kg N, 84 kg P and 50 kg K on irrigated
soil- and seed-borne diseases, but this is rarely wheat in Zimbabwe. Commercial fertilizer ap-
practised in tropical Africa. Wheat is sown by plication ranges from less than 1%of the wheat
hand or machine. When broadcast, the seed is area in Burundi to 100% in Kenya and Zim-
incorporated in the soil using an animal-drawn babwe. Organic manure and compost are not
plough or machine-drawn disc. The seed may commonly used on wheat, except in Rwanda.
also be dibbled directly into a furrow behind a Boron deficiency, resulting in grain set failure,
plough and covered, or machine-planted in can be observed on certain soils; boron is ap-
rows. Common seed rates are 150-200 kg/ha plied to irrigated wheat in Zambia, Zimbabwe
for broadcasting and 75-120 kg/ha for row- and Madagascar. Copper is applied to most
planting. The optimum spacing is 10-25 cm rainfed wheat in Kenya, and manganese is
between rows, but it may extend up to 35 cm. needed in certain areas of Tanzania. Soil acid-
The sowing depth is 2-5(-12) cm, with deeper ity can be a constraint, e.g. in wheat production
planting required in dry conditions. At a sow- areas at lower elevations in Zambia. Liming
ing depth beyond 10-12 cm seedling emergence might raise the pH, but its economic returns
is poor. When using a no-till planting machine, are poor for rainfed wheat.
sowing can be done straight into the stubble of Wheat is best rotated with non-grass crops,
the previous crop. For rainfed wheat, the seed particularly with pulses. In the highland areas
can be dry-sown, before the start of the rainy of East Africa wheat is grown continuously or
season, or when the soil is moist. Bread wheat in rotation with other cereals, pulses or rape-
is usually grown in sole cropping. seed (Brassica oilseed crops). In other regions
Management Uniform crop stand and early double cropping systems are common, with
vigour discourage weed growth in bread wheat. irrigated wheat grown in the cool dry season
In this respect tillering allows the crop to com- and crops such as cotton, sorghum, maize, soya
pensate for poor stands and variable weather bean and groundnut in the hot rainy season. In
conditions. Yield losses due to weeds are Zimbabwe, for instance, double cropping of
caused by early competition in the first 4-5 irrigated wheat and rainfed soya bean is widely
weeks. Hand weeding, tillage practices, stubble adopted, with the same machinery for sowing
management, pre-sowing irrigation, proper and harvesting used for both crops.
crop rotation and herbicides may control In tropical Africa wheat is produced in farming
weeds. Herbicide use in tropical Africa ranges systems ranging from small scale, labour-
from little to none in many countries (e.g. Su- intensive, rainfed systems, e.g. in Kenya and
dan, Rwanda, Burundi, Madagascar) to almost southern Tanzania, to highly mechanized
complete coverage in Kenya, Tanzania, Zambia schemes and farms, e.g. in Nigeria, Sudan,
and Zimbabwe. northern and central Tanzania and Zimbabwe.
 TEITICUM 181

Diseases and pests Bread wheat is affected spikes, or during combine harvesting. In most
by several diseases and pests. In tropical Africa parts of tropical Africa wheat stubble is grazed
stripe rust or yellow rust (Puccinia striiformis), by livestock.
spread by air-borne uredospores, and Septoria Yield Yields of bread wheat in tropical Af-
blotches, particularly Septoria leaf blotch (Sep- rica vary from 400 kg/ha in Somalia and 700
toria tritici, synonym: Mycosphaerella gramini- kg/ha in Angola to 5 t/ha in Zambia and 6.3
cola), are the major diseases in the highlands. t/ha in Zimbabwe. The mean yield of wheat in
Stem rust or black rust (Puccinia graminis) tropical Africa is estimated at about 1.5 t/ha.
can be very damaging in Ethiopia, Kenya and Lower yields are due to high temperature, high
some parts of Sudan; like stripe rust it is humidity, disease pressure and the low levels
spread by air-borne uredospores. Other dis- of fertilizer applied. Maximum recorded grain
eases important in some years are common yields of irrigated winter and spring wheats
bunt (Tilletia spp.), loose smut (Ustilago tritici, are 14 and 9.5 t/ha, respectively; the absolute
synonym: Ustilago nuda f.sp. tritici), barley maximum yield, based on genetic potential, is
yellow dwarf virus (BYDV) and bacterial leaf estimated at 20 t/ha.
streak or black chaff (Xanthomonas translu- Handling after harvest Threshed grain of
cens). The use of resistant cultivars is the most bread wheat is winnowed, cleaned and pre-
effective control measure against these dis- pared for store or market. Seeds should be
eases. However, resistance breakdown is very dried to a moisture content of 13—14% for safe
frequent for stripe rust. Fungicide application storage. High temperatures and moist condi-
to control stripe rust occurs in Kenya, Uganda tions may result in spoilage. Regular re-drying
and Tanzania. may be necessary to maintain seed viability, if
The most important insect pests in tropical the seed is not stored in an airtight container.
Africa are aphids, which may also transmit Genetic resources The International Maize
viruses. The African migratory locust (Locusta and Wheat Improvement Center (CIMMYT),
migratoria) is a periodic pest that causes crop Mexico City, Mexico (60,400 accessions) and
damage in northern and eastern Ethiopia. The the International Center for Agricultural Re-
Hessian fly (Mayetiola destructor) has long search in the Dry Areas (ICARDA), Aleppo,
been an important pest in regions adjacent to Syria (9700 accessions) maintain extensive
the Mediterranean Sea in northern Africa, germplasm collections of Triticum aestivum.
southern Europe and western Asia. Pest con- Large germplasm collections are also held in
trol with commercial insecticides in tropical the United States (USDA-ARS National Small
Africa is rare, except in Sudan, Zambia and Grains Germplasm Research Facility, Aber-
Zimbabwe for aphids. Birds (especially Quelea deen, Idaho, 42,000 accessions), China (Insti-
quelea) are especially important in irrigated tute of Crop Germplasm Resources (CAAS),
wheat. Beijing, 35,900 accessions), and the Russian
Important storage insects, e.g. in Ethiopia, Federation (N.I. Vavilov All-Russian Scientific
include Sitophilus spp. on whole grains, and Research Institute of Plant Industry, St. Pe-
Tribolium spp. and Ephestia cautella (syno- tersburg, 25,900 accessions). In tropical Africa
nym: Cadra cautella, flower moth) on wheat the Institute of Biodiversity Conservation (Ad-
flour. Clean storage conditions and maintain- dis Ababa, Ethiopia) has the largest collection
ing grain moisture and temperature at suffi- of bread wheat (3400 accessions). Wheat is a
ciently low levels inhibit insect activity and priority crop for collection and conservation.
development. Rodents, mainly the black rat More collection needs to be done of its wild and
(Rattus rattus), also damage stored seeds. weedy relatives in regions where they are na-
Harvesting In tropical Africa bread wheat tive, of landraces in areas where they have not
is usually harvested with sickles or knives, and been collected before, and of new or obsolete
on large-scale farms with combines. A crop improved cultivars with specific traits from
harvested at physiological maturity (grain breeding programmes around the world for
moisture content 25-35%) must be dried thor- future improvement work.
oughly before threshing. Wet weather at har- Breeding CIMMYT and ICARDA have large
vest time can cause serious losses in grain breeding programmes and, upon request, have
quality because the grain sprouts readily. the international mandate to disseminate
Sickle-harvested wheat plants are stacked or bread wheat germplasm to national pro-
spread out to dry in the sun. Threshing is done grammes. In tropical Africa, Ethiopia and
by trampling animals, by beating bagged Kenya have strong public sector breeding pro-
182 CEREALS AND PULSES

grammes. In Zimbabwe, there is private sector come, there is likely to be a concomitant de-
wheat research, and to some extent in Kenya mand for traditional and new convenient, proc-
and Zambia too. High grain yield and disease essed wheat-based products. No tropical Afri-
resistance, mainly to stripe rust and Septoria, can countries are 100% self-sufficient for wheat
are the major objectives. Major breeding meth- and the region is confronted by rapidly increas-
ods used in tropical Africa are conventional. A ing wheat imports. In many of these countries
number of high-yielding cultivars, mostly wheat production is constrained by limited
spring types derived from CIMMYT germ- usage of high-yielding cultivars, fertilizer,
plasm, have been released in tropical African other inputs and irrigation. Increases in wheat
countries. In 1995 their estimated usage production may come from area expansion to
ranged from 5% in Malawi to 100% in Zambia non-traditional areas, coupled with social and
and Zimbabwe. economic incentives, and further increases in
Bread wheat is one of the crops that benefited yield by agronomic research and breeding.
most from transfer of genes from other species, Since the 1990s, area expansion ofbread wheat
such as Aegilops, Hordeum and Secale spp., by has been observed in Sudan, Ethiopia, Kenya,
artificial hybridization, mainly to increase re- Tanzania and Zambia. Research to improve
sistance to diseases, especially rusts. Develop- wheat yields at a global scale includes further
ments in molecular genetics and genetic engi- mixing of germplasm through wide hybridiza-
neering of wheat have been slower than in ce- tion and synthetic hexaploids, biotechnology
reals such as rice and maize, due to its ploidy tools, hybrid wheat, and basic studies on wheat
level, size and complexity of its genome, the physiology and host-plant relationships of vari-
low level of polymorphism and relatively ineffi- ous diseases and pests. Tolerances to drought,
cient transformation systems. Consequently, heat, aluminium soil acidity and waterlogging
far fewer maps exist in wheat and few QTL are some of the abiotic factors that require
(quantitative trait loci) studies have been re- continued research attention.
ported. On the other hand, the hexaploid na- Major references CIMMYT, 1985; Curtis,
ture of bread wheat and its amenity to cytoge- Rajaram &Gomez Macpherson (Editors), 2002;
netic manipulation have offered unique tools Heisey & Lantican, 1999; Heyene (Editor),
for molecular genetic studies. These include 2002; Klatt (Editor), 1988; Payne, Tanner &
the uses of aneuploid stocks to assign molecu- Abdalla, 1996; Saunders & Hettel (Editors),
lar markers to specific chromosome arms, of 1994; Tanner & Raemaekers, 2001;van Ginkel
chromosomal deletion stocks for physical map- &Villareal, 1996;Wiese, 1987.
ping and of chromosome substitution lines to Other references Ageeb et al. (Editors),
map genes of known chromosomal location. 1996; Bowden, 1959; Braun et al. (Editors),
The development of improved chemical hybrid- 1997; Byerlee & Moya, 1993; Dvorak et al.,
izing agents, which allows breeders to sur- 1998; Edwards, 1997; Feldman, Lupton &Mil-
mount the problems associated with cytoplas- ler, 1995; Gebre-Mariam, Tanner & Hulluka
mic male sterile systems, has considerably (Editors), 1991; Hanson, Borlaug & Anderson,
increased the progress towards the develop- 1982; Hu et al., 2003; Jordaan, 1999; Khairal-
ment of economically acceptable hybrid wheat lah et a l , 2001; Phillips, 1995; Pickett, 1993;
cultivars. Recently, an efficient Agrobacterium- Quisenberry & Reitz (Editors), 1967; Roelfs,
mediated transformation system has been de- Singh & Saari, 1992; Simmonds & Rajaram
veloped for the large-scale production of trans- (Editors), 1988; USDA, 2005; Walker & Boxall,
genic wheat plants. Private companies have 1974; Zhou et al., 2003.
developed transgenic herbicide-resistant bread Sources of illustration van Ginkel & Vil-
wheat cultivars, but these have not yet been lareal, 1996.
produced commercially. Authors G. Belay
Prospects Since bread wheat is the most Based on PROSEA 10: Cereals.
important food grain source for humans, the
need to continuously increase its production
cannot be overemphasized. Bread consumption
from wheat in tropical Africa is low and varies
from country to country; wheat consumption
ranges from 2.5 kg/person/year in Uganda to
43.3 kg/person/year in Sudan. However, with
the increasing trends of urbanization and in-
 TRITICUM 183

TRITICUM TURGIDUM L. BC. In tropical Africa durum wheat is pre-

dominantly grown in Ethiopia and to some
Protologue Sp. pi. 1:86 (1753). extent in Eritrea and Angola. In other coun-
Family Poaceae (Gramineae) tries, e.g. Sudan and Tanzania, it has been
Chromosome number 2n — 28 grown experimentally. Durum wheat is also
Synonyms Triticum dicocconSchrank (1789), widely grown in northern Africa (from Morocco
Triticum durum Desf. (1798). to Egypt), Mediterranean Europe (Italy, south-
Vernacular names Durum wheat, maca- ern France), Turkey, the Middle East (Syria,
roni wheat (En). Blé dur (Fr). Trigo duro, trigo Jordan, Iraq), Russia, Asia (Iran, Afghanistan,
rijo (Po). India, China), North America (Canada and the
Origin and geographic distribution Hybrid- United States) and Argentina.
ization between the diploids Triticum urartu Uses Throughout the world durum wheat is
Tumanian ex Gandylian (A-genome) and the mainly ground to semolina (coarse flour) that is
yet unconfirmed B-genome donor (possibly a made into various pasta products (macaroni,
species of Aegilops section Sitopsis), followed spaghetti, noodles) and traditional flat bread
by chromosome doubling gave rise to the first (little leavened). In tropical Africa durum
wild tetraploid wheat. Remains of primitive wheat utilization has been adapted to the local
types of cultivated Triticum turgidum (emmer cuisine. In Ethiopia it is used mainly to make
wheat, which has hulled grain) were discovered 'kitta' (unleavened bread), 'injera' (flat pan-
at several archaeological sites in Syria and cake-like unleavened bread), and homemade
dated at around 8000 BC. Emmer wheat be- alcoholic and non-alcoholic drinks. Durum
came the predominant cultivated wheat in the wheat is also preferred for preparation of
Fertile Crescent (southern Turkey, northern 'kinchie' (crushed kernels, cooked with milk or
Iraq and adjacent regions of Iran and Syria, as water and mixed with spiced butter), which is
well the Joran valley) and spread into much of often served for breakfast. The grain is eaten
Asia, northern Africa and Europe. It remained as a snack and during social gatherings as 'ni-
the main wheat for several thousands of years. fro' (boiled whole grain often mixed with
Free-threshing types, such as durum wheat, pulses), 'kollo' (roasted grain), and 'dabo-kollo'
arose by accumulation of mutations and subse- (ground and seasoned dough, shaped and deep
quent selection from the primitive emmer fried). In northern Africa and the Middle East
wheat. Around the beginning of the Christian durum wheat is preferred for making couscous;
era durum wheat had replaced emmer in most its granules result from the agglomeration of
of the wheat growing areas of the Old World. semolina particles. In the Middle East it is also
Durum wheat is now commercially the most durum wheat which is used for making 'bul-
important type of Triticum turgidum. It is not ghur', i.e. a wheat which is parboiled, dried and
certain when and how durum wheat reached then crushed.
tropical Africa, but it might have reached the The straw of durum wheat is fed to animals
northern highlands of Ethiopia around 3000 and used as bedding material for animals and
for thatching.
Production and international trade Durum
wheat and bread wheat statistics are usually
combined and therefore individual and reliable
statistics on durum wheat are difficult to ob-
tain. According to FAO estimates, the average
world production of wheat grain (durum wheat
and bread wheat together) in 1999-2003
amounted to 576 million t/year from 209 mil-
lion ha. Worldwide, durum wheat constitutes
less than 10% of the area under the cultivated
wheats. Major durum wheat producers are
northern Africa, where it covers nearly 50% of
the total wheat area, the United States, Can-
ada and the Russian Federation. The main
durum wheat-producing country in tropical
Africa is Ethiopia. Close to 50% of its total
Triticum turgidum -planted wheat production (1.4 million t/year in 1999—
184 CEREALS AND PULSES

2003) is durum wheat. In Ethiopia durum perior, tipped by a small fleshy hairy append-
wheat production is mainly for subsistence. age and with 2 plumose stigmas. Fruit an el-
Ethiopia also produced emmer wheat, but this lipsoid caryopsis (grain), at one side with a
crop is disappearing. central groove.
P r o p e r t i e s Durum wheat grain contains per Other botanical information Triticum is a
100 g edible portion: water 10.9 g, energy 1418 classic example of allopolyploidy consisting of
kJ (339 kcal), protein 13.7 g, fat 2.5 g, carbohy- diploid (2/i = 14), tetraploid (2re = 28) and
drate 71.1 g, Ca 34 mg, Mg 144 mg, P 508 mg, hexaploid (2n = 42) species. Selection at the
Fe 3.5 mg, Zn 4.2 mg, vitamin A 0 IU, thiamin diploid and tetraploid levels has proceeded
0.42 mg, riboflavin 0.12 mg, niacin 6.7 mg, from wild species with hulled grain and brittle
vitamin Ek 0.42 mg, folate 43 ug and ascorbic rachis to the free-threshing species with tough
acid 0 mg. The essential amino-acid composi- rachis; hexaploid wheats are not known in the
tion per 100 g edible portion is: tryptophan 176 wild, they appeared in cultivation. The classifi-
mg, lysine 303 mg, methionine 221 mg, cation of the genus Triticum and other related
phenylalanine 681 mg, threonine 366 mg, genera within the tribe Triticeae was strongly
valine 594 mg, leucine 934 mg and isoleucine debated. Polyploidy and biphyletic genome
533 mg. The principal fatty acids are per 100 g differentiation (B vs. G genome) are isolating
edible portion: linoleic acid 930 mg, palmitic mechanisms offering adequate species borders.
acid 422 mg and oleic acid 335 mg (USDA, In this approach, Triticum comprises only 5—6
2005). Durum wheat grain is deficient in the species, including the diploid Triticum mono-
amino acids lysine and threonine, and some- coccum L. (grown sporadically in southern
what in isoleucine and valine. It is a good Europe and western Asia), the tetraploid Triti-
source of B-group vitamins and minerals. Du- cum turgidum L., and the hexaploid Triticum
rum wheat grain is vitreous, amber in colour, aestivum L.
and is the hardest of all wheats. The physical Some tetraploid cultivated wheats are some-
and chemical characteristics of durum wheat times specifically distinguished from Triticum
gluten provide greater stability of the dough turgidum. Triticum aethiopicum Jakubz. is a
and make it specially suited for pasta products.
For preparation of pasta, the grain is milled
only as far as the semolina stage; a finely
ground flour is not required. In the process of
cooking, pasta products of good quality do not
disintegrate or become soft, mushy, starchy or
sticky. Protein should have a minimum level of
12%. Durum wheat is not suitable for making
cakes and leavened bread because of its high
gluten content and dough strength.
Description Annual, often strongly tufted
grass up to 170 cm tall; stem (culm) cylindrical,
smooth, hollow except at nodes. Leaves distich-
ously alternate, simple and entire; leaf sheath
rounded, auricled; ligule membranous; blade
linear, 15-40 cm x 1-2 cm, parallel-veined,
flat, slightly hairy. Inflorescence a terminal,
dense, distichous spike 4-12 cm long, with ses-
sile spikelets borne solitary on zigzag, hairy,
tough rachis. Spikelet 10-15 mm long, laterally
compressed, 4-7-flowered, with bisexual flo-
rets, but the 1—3 uppermost ones usually rudi-
mentary; glumes almost equal, oblong, shorter
than to almost as long as spikelet, thinly leath-
ery, 5—11-veined, strongly keeled throughout,
apiculate to awned; lemma rounded on back Triticum turgidum - 1, inflorescences; 2,
but keeled towards the tip, leathery, with an spikelet; 3, grains.
awn 8-20 cm long; palea 2-keeled, hairy on the Redrawn and adapted by Achmad Satiri Nur-
keels; lodicules 2, ciliate; stamens 3; ovary su- haman
 TRITICUM 185

special type of free-threshing wheat, a tradi- north-western highlands at 1800-2800 m alti-

tional cereal crop in Ethiopia and the southern tude during the main rainy season ('meher')
part of the Arabian Peninsula. Its spikes are between August and December. Highly rust-
loose to dense, its glumes are usually awned resistant cultivars are needed to grow durum
and its grain mostly purple. Triticum dicoccon wheat below 1900 m in Ethiopia. High tem-
Schrank (emmer wheat) is the oldest cultivated peratures and low humidity improve grain
tetraploid wheat, domesticated in the area of quality, and durum wheat is susceptible to low
Palestine, south-western Syria and north- temperatures and severe frosts. The minimum
western Jordan. It has disarticulating spikes amount of water required for an acceptable
with 2-grained spikelets and hulled grains, not crop is 250 mm.
easy to decorticate. At present it is still culti- Soils best suited for durum wheat are well aer-
vated in Ethiopia, Iran, Turkey, Transcauca- ated, well drained and deep, with 0.5% or more
sia, former Yugoslavia, the Czech Republic, organic matter. Optimum soil pH is 5.5-7.5.
Slovakia and India. Triticum durum Desf. is Durum wheat is sensitive to soil salinity. In
the free-threshing durum wheat or macaroni Ethiopia durum wheat is preferentially grown
wheat, that appeared in the Mediterranean, on heavy black clay soils (Vertisols); farmers
and is cultivated in regions with a hot dry cli- usually delay planting and use surface drain-
mate; it has its greatest diversity in Ethiopia. age systems (furrows) to avoid waterlogging. N
It has slender spikes and comparatively long and micronutrient deficiencies can be limiting
glumes. Triticum polonicum is the free- on Vertisols. Durum wheat can also be grown
threshing Galicia wheat (erroneously named on light soils (Andosols), but here short, stiff
'Polish wheat' by Linnaeus), occasionally culti- and disease resistant cultivars are required.
vated in the same areas as the true durum Propagation and planting Durum wheat
wheat. It has much longer glumes (2.5—3 cm). is propagated by seed. The 1000-seed weight is
In Ethiopia it is found only in mixture with 30—40 g. Durum wheat can be sown by hand or
other wheats. The free-threshing rivet wheat machine; in Ethiopia it is usually broadcast.
or cone wheat (Triticum turgidum L. sensu Dormancy can be a problem in introduced cul-
stricto) is cultivated in northern Africa, south- tivars, but not in the local Ethiopian landraces.
ern and central Europe and Asia. It has stout Seeding rate is commonly 100-150(-175)
spikes nearly square in section and compara- kg/ha, the higher rates being necessary on
tively short glumes. It is also grown in Ethio- heavy clay soils where stand establishment is
pia, usually in mixtures. usually poor on flat seedbeds. It is advisable to
Most durum wheat cultivars are spring or use certified seed that has been treated with
semi-winter types. Only a few winter types are fungicides against soil- and seed-borne dis-
known. eases, but this is not practised in tropical Af-
Growth and development Germination of rica. In Ethiopia an oxen-drawn implement
wheat occurs at temperatures of 4-37°C, the ('maresha') is used to till the land before sow-
optimum being 12-25°C. The coleoptile ing, with 2-3 ploughings made before planting.
emerges 4-6 days after germination. Flowering In Ethiopia planting dates vary from mid-July
begins at the middle third of the spike, then to early September.
rapidly progressing both upward and down- Management Weed competition during
ward. Durum wheat is predominantly self- tillering of the durum wheat crop, usually in
pollinated; in Ethiopia cross-pollination rates the first 10-50 days after sowing, is most det-
up to 4.3% have been recorded. Physiological rimental to grain yield. Uniform crop stand
maturity is reached when the moisture content and early vigour discourage weed growth.
of the fully formed grain has dropped to 2 5 - Competition occurring later in the crop cycle
35%. The complete crop cycle of durum wheat can affect grain numbers and grain weight, but
is 90-115 days in Ethiopia. usually has smaller effects on grain yield.
Ecology Durum wheat is better suited to Weeds can be controlled by hand weeding,
regions with a low average annual rainfall proper crop rotation, pre-seeding irrigation,
than bread wheat, e.g. in the Middle East, machine cultivation, or application of chemical
northern Africa and parts of Mediterranean herbicides. In tropical Africa hand weeding
Europe. In the tropics durum wheat is best remains the most common means of weed con-
grown at higher elevations or in the cooler trol. Blanket fertilizer recommendation rates
months of the year. In Ethiopia durum wheat for durum wheat in Ethiopia are 41 kg N and
is mostly produced in the central, northern and 26 kg P per ha; additionally 23 kg/ha N can be
186 CEREALS AND PULSES

top-dressed under heavy rain conditions at wheat are separated from the residues by win-
early growth stages. However, farmers in nowing. The clean seeds are stored, sold or
Ethiopia do not usually give priority to durum processed for home consumption. Harvested
wheat when applying commercial fertilizer. durum wheat grain should be dried to moisture
Diseases and pests The most important content of 13—14% for safe storage. High tem-
diseases of durum wheat in tropical Africa are peratures and moist conditions may result in
stem rust (Puccinia graminis) and leaf rust spoilage. Regular re-drying may be necessary
(Puccinia recondita f.sp. tritici, synonym: Puc- to maintain seed viability, if the seed is not
cinia triticina). The use of resistant cultivars is stored in an airtight container.
the most effective control measure against Genetic r e s o u r c e s The International Cen-
these diseases. In cooler regions, stripe rust or ter for Agricultural Research in the Dry Areas
yellow rust (Puccinia striiformis) limits durum (ICARDA), Aleppo, Syria (21,010 accessions)
wheat production, e.g. in the Arsi highlands of and the International Maize and Wheat Im-
Ethiopia. provement Center (CIMMYT), Mexico City,
The most important insect pests in tropical Mexico (7880 accessions) maintain large germ-
Africa include aphids (which may also transmit plasm collections of Triticum turgidum. Large
viruses), and grasshoppers. The African migra- germplasm collections are also held in the
tory locust (Locusta migratoria) is a periodic United States (USDA-ARS National Small
pest that causes crop damage in northern and Grains Germplasm Research Facility, Aber-
eastern Ethiopia. The Hessian fly (Mayetiola deen, Idaho, 42,030 accessions), the Russian
destructor) has long been an important pest in Federation (N.I. Vavilov All-Russian Scientific
regions adjacent to the Mediterranean Sea in Research Institute of Plant Industry, St. Pe-
northern Africa, southern Europe and western tersburg, 5580 accessions) and Australia (Aus-
Asia. Control of insect pests with commercial tralian Winter Cereals Collection, Agricultural
insecticides in tropical Africa is rare. Impor- Research Centre, Tamworth, New South
tant storage insects in Ethiopia include Sito- Wales, 5520 accessions). In tropical Africa the
philus spp. on whole grains, and Tribolium Institute of Biodiversity Conservation (Addis
spp. and Ephestia cautella (synonym: Cadra Ababa, Ethiopia) has the largest collection of
cautella, flower moth) on wheat flour. Rodents, Triticum turgidum (12,500 accessions). Since
mainly the black rat (Rattus rattus), also dam- Ethiopia is an important centre of diversity of
age stored seeds. durum wheat, several studies since the 1970s
Harvesting In tropical Africa durum wheat have dealt with the magnitude and structure of
is usually harvested with sickles and rarely by Ethiopian durum wheat germplasm using
machine. A crop harvested at physiological morphological, protein, cytological and molecu-
maturity (grain moisture content 25-35%) lar markers. While the area of durum wheat
must be dried thoroughly before threshing. Wet has decreased since the 1970s, no drastic
weather at harvest time can cause serious changes in the overall diversity are evident.
losses in grain quality because the grain Breeding CIMMYT and ICARDA have the
sprouts readily. Plants are stacked or spread international mandate to disseminate durum
out to dry in the sun. Threshing is mostly done wheat germplasm to national programmes. In
by trampling animals. tropical Africa a strong breeding programme
Yield Durum wheat yields on farmers' fields has been underway in Ethiopia since 1976.
in Ethiopia vary from 800 kg/ha to 2.5 t/ha; High grain yield and disease resistance, mainly
mean yield is estimated at less than 1 t/ha. to stem and leaf rusts, have been the major
Yields tend to be rather low due to the low ap- objectives, and recently industrial quality has
plication of improved cultivars and optimal been included. Genotype x environment inter-
production practices, and low levels of fertilizer actions in Ethiopia are very high and therefore
applied. Yield progress in durum wheat has emphasis has shifted from wide to specific ad-
generally been lower than that in bread wheat. aptation. Major breeding methods are conven-
However, durum wheat grain yields of 5-6 t/ha tional, and include selection from indigenous
can be obtained with irrigation and the use of landraces and introductions from CIMMYT
improved cultivars and better production prac- and ICARDA, and hybridization. Greater suc-
tices. Straw yields are equally important in cess was achieved from the international intro-
Ethiopia and range from 9-15 t/ha. ductions than from the landrace selections.
Handling after harvest In tropical Africa, More than 16 durum wheat cultivars have
e.g. in Ethiopia, threshed grains of durum been officially approved, but their area does
 TYLOSEMA 187

not exceed 10%of the total durum wheat area. 1991; Morris & Sears, 1967; Scarascia
'Boohai', 'Foka', 'Kilinto' and 'Yerer' are among Mugnozza (Editor), 1973; Srivastava, 1984;
the most widely sown cultivars. Tanner & Raemaekers, 2001; Tesemma & Be-
Linkage-maps of durum wheat have been de- lay, 1991;van Ginkel &Villareal, 1996; Wiese,
veloped and important QTLs (quantitative 1987.
trait loci) for grain quality traits have been Other references Alamerew et al., 2004;
identified. The developments in wheat molecu- Belay, 1997; Belay, Tesemma & Mituku, 1993;
lar genetics and genetic engineering have been Belay et al., 1997; Bowden, 1959; Curtis, Ra-
relatively slow, especially when compared to jaram & Gomez Macpherson (Editors), 2002;
other cereals such as rice and maize, due to its Elouafi & Nachit, 2004; Eticha et al., 2005;
ploidy level, size and complexity of its genome, Feldman, Lupton & Miller, 1995; Gashawbeza
the low level of polymorphism and relatively et al., 2003; Jauhar, 2003; Mac Key, 1966; Mo-
inefficient transformation systems. Breeding of hamed, 1999; Payne, Tanner & Abdalla, 1996;
durum wheat is less advanced than of bread Perrino et al., 1996; Phillips, 1995; Tarekegn,
wheat; in addition it has benefited less in wide 1994; Tsegaye, 1996; USDA, 2005; Walker &
hybridization and alien gene transfers. Durum Boxall, 1974.
wheat is an important component species for Sources of illustration Landwehr, 1976;
bread wheat breeding through formation of Vaughan &Geissler, 1997.
synthetic hexaploids, and for the production Authors G. Belay
and development of triticale ÇxTriticosecale),
the hybrid ofwheat and rye.
Prospects In tropical Africa, Ethiopia has TYLOSEMAESCULENTUM(Burch.) A.Schreib.
the greatest potential for durum wheat because
of its favourable growing environments in the Protologue Mitt. Bot. Staatssamml. Mün-
cool dry highlands and tradition of growing the chen 3:611 (1960).
crop. There is an increasing demand for quality Family Caesalpiniaceae (Leguminosae - Cae-
durum wheat grain by the local pasta indus- salpinioideae)
tries, which is usually met through import. Synonyms Bauhinia esculenta Burch. (1824).
Breeding programmes have developed cultivars Vernacular names Marama bean, morama
that meet the quality demand by the industry, bean, gemsbok bean, camel's foot (En). Ma-
but in the absence of premium price over the rama (Fr).
higher-yielding bread wheat cultivars, farmers Origin and geographic distribution Mara-
are losing interest in growing durum wheat. ma bean is native to the Kalahari desert and
The future trends of durum wheat production, neighbouring sandy regions in Angola, Na-
as a result of unfavourable market prices, mibia, Botswana and South Africa, but it also
therefore may seem discouraging. On the other occurs in Zambia and Mozambique. Experi-
hand, large-scale commercial farmers are en- mental cultivation in Kenya, South Africa,
tering into durum wheat production to supply Australia, Israel and the United States (Texas)
the industry, some even replacing bread wheat
due mainly to price competition from imported
flour. Adaptive research is needed to develop
durum wheat that reliably produces 2-3 t/ha in
farmers' fields. More progress is also required
in agronomic research, identification of suit-
able production areas and in establishing an
attractive pricing and marketing structure for
farmers. The crucial factor is a stable and
long-term commitment from the government,
the farmers, the private sector (including seed
producers) and national research programmes.
With this functional partnership in place,
Ethiopia could even export quality durum
wheat.
Major references Bechere, Kebede & Be-
lay, 2001; Bechere, Tesemma & Mitiku, 1994;
Gebre-Mariam, Tanner & Hulluka (Editors), Tylosema esculentum - wild
188 CEREALS AND PULSES

has been successful. 2.1 g, fat 0.1 g and carbohydrate 4.4 g. Young
Uses Marama bean is an important part of tubers have a sweet and pleasant taste and the
the diet of the Khoisan people in the Kalahari, texture has been described as similar to that of
where subsistence agriculture is marginal due artichoke. The tubers are reddish when dried.
to drought and low soil fertility, and it is a deli- Description Perennial herb or shrub, with
cacy among other peoples in southern Africa. tuberous root; stems prostrate and trailing, up
The seeds are eaten boiled or roasted. They to 6 m long, herbaceous or lower parts woody,
may be boiled with maize meal or ground into rusty-hairy, with axillary forked tendrils 1-4
flour to prepare a porridge or a coffee- or cocoa- cm long. Leaves alternate, simple; stipules 3-5
like drink. Roasted seeds are sometimes sold mm x 2-3 mm; petiole 1.5-3.5cm long;blade 2-
locally but only on a small scale. Marama lobed for more than half its length, glabrous or
beans have a pleasant sweet flavour when pubescent beneath; lobes reniform, 3.5-5 cm x
boiled or roasted, comparable to roasted 5-6.5 cm. Inflorescence a lateral raceme up to
cashew nuts or almonds, although bitter types 16 cm long; peduncle 2-4 cm long. Flowers
are known. The roasted seeds have sometimes bisexual, zygomorphic, 5-merous, heterosty-
been used by Europeans in southern Africa as lous; pedicel 2-4.5 cm long; sepals free but up-
a culinary substitute for almonds. Immature per 2 fused, 8-12 mm x 2-3 mm, rusty-hairy;
seeds and stems may be eaten cooked as a petals unequal, 4 larger ones 1.5—2.5 cm x l_
vegetable or in soups. The seed oil is used in 1.5 cm and tapering into a basal claw, the up-
Botswana for cooking and for making butter. per one smaller, yellow turning reddish with
Young tubers are eaten baked, boiled or age; stamens 2, free, with filaments 6-12 mm
roasted, as a vegetable dish. Tubers older than long, staminodes 8, with filaments 3-6 mm
2 years become fibrous and bitter and are usu- long; ovary superior, 5-6 mm long, 1-celled,
ally not eaten, but they are an important emer- style elongate, stigma small. Fruit an ovoid to
gency source of water for humans and animals. oblong pod 3.5-6 cm x 3-4 cm, flattened,
The pods and tubers are recorded to be eaten woody, l-2(-6)-seeded, constricted between the
by animals, but it is not clear whether the foli- seeds. Seeds ovoid to globose, 1.3-2.5 cm x 1.2-
age is browsed, as contradictory reports exist. 1.5 cm, reddish to brownish black.
Marama bean may have potential as a ground
cover or ornamental.
Properties Mature, shelled marama bean
seeds contain per 100 g: water 3.9 g, energy
2660 kJ (635 kcal), protein 31.8 g, fat 42.2 g
and carbohydrate 18.9 g (Bower et al., 1988).
The protein content of marama bean is compa-
rable to that of soya bean, and the oil content is
twice as high as that of soya bean and compa-
rable to that of groundnut. The essential
amino-acid composition per 100 g food is: tryp-
tophan 219 mg, lysine 1119 mg, methionine
257 mg, phenylalanine 874 mg, threonine 822
mg, valine 1149 mg, leucine 1774 mg and iso-
leucine 1119 mg (FAO, 1970). The seeds have a
relatively high trypsin inhibitor activity, which
can be remedied by cooking. The seed oil is
golden-yellow, with a nutty odour and a pleas-
ant, although slightly bitter flavour, and has
been described as similar to almond oil in con-
sistency and taste. Its principal fatty acids are
oleic acid (48-49%), linoleic acid (19-26%),
palmitic acid (12-14%), stearic acid (7-10%)
and arachidic acid (3%). Per 100 g dry weight
the defatted seed meal contains: energy 194 kJ Tylosema esculentum - 1, part of flowering
(46 kcal), protein 55.0 g, available starch 13.0 g stem; 2,fruit; 3, seed.
and fibre 1.6 g. Per 100 g, the tubers of a 5- Redrawn and adapted by Achmad Satiri Nur-
month-old plant contain: water 92.1 g, protein haman
 TYLOSEMA 189

Other botanical information Tylosema com- digging when they weigh about 1kg.
prises 5 species and occurs in southern and Yield In the Kalahari young tubers of
eastern Africa. Some taxonomists do not con- marama bean of 1 year old and about 1 kg in
sider Tylosema a separate genus, but include it weight are preferred. Tubers may reach 10 kg
in Bauhinia. Tylosema fassoglense (Schweinf.) after a few years and tuber weights of up to
Torre & Hillc, which also has edible seeds and 300 kg have been reported; a tuber weighing
tubers, has longer leaf stalks and more shal- 277 kg contained 224 1water. Information on
lowly lobed leaves. seed yields of marama bean is not available.
Growth and development In field experi- Handling after harvest Raw seeds of ma-
ments in Kenya marama bean seeds started to rama bean store well and remain edible for
germinate 9-10 days after planting. Once ger- years. Dry storage is preferable. Oil can be
minated the seedlings develop rapidly. extracted from the seeds by conventional press-
Marama bean has been recorded as not flower- ing or solvent extraction. To obtain water from
ing until the 3 rd or 4 th year after planting, but the tuber, the skin is scraped away and a hole
in experiments in Texas plants started flower- is made. The flesh in and around the hole is
ing after 2 years, and fruits and seeds were mashed with a wooden stick until the consis-
formed after 3.5 years. In its native area tency is porridge-like. This porridge is put into
marama bean flowers from October to March. a piece of fabric and the water is squeezed out
It is predominantly outcrossing and may be with both hands. The water can also be ex-
self-incompatible; it is pollinated by insects. In tracted from the tubers by pounding pieces of it
cultivation fruit and seed set tend to be low. In in a container.
southern Africa the stems die back during the Genetic resources Marama bean is consid-
dry and cool period (May-August), but the tu- ered neither rare nor threatened. No substan-
ber remains viable and produces new stems tial germplasm collections are known to exist.
when the temperature rises. Marama bean The International Livestock Research Institute
does not form root nodules and relies on soil (ILRI), Addis Ababa, Ethiopia, the National
nitrogen. Its drought-adaptive mechanisms Genebank, Muguga, Kenya, and the Plant Ge-
include closure of leaves, the maintenance of netic Resources Unit of the Agricultural Re-
green-leaf area under drought by early search Council, Pretoria, South Africa, have 1
stomatal closure, and the use of moisture re- accession each. Marama bean shows orthodox
serves in the tuber (which shrinks greatly in seed storage behaviour.
dry years). Marama bean plants have long Breeding Programmes for selection and
trailing stems that creep along the ground and breeding work of marama bean are recorded to
avoid the effects of the strong destructive be underway in the United States, Australia
windstorms ofthe Kalahari. and Israel. RAPD analysis of 3 populations
Ecology Marama bean occurs naturally in from various parts of Botswana has shown that
an extreme environment with high tempera- a considerable amount of genetic variation ex-
tures (typical daily maximum of 37°C in the ists within marama bean, most of it within
growing season), low rainfall (100—900 mm) rather than between populations. Sufficient
and long periods of drought. It is found on genetic variation for breeding may be found by
sandy and limestone (including dolomite) soils, sampling 30-40 plants from 1or 2 populations.
but not on soils developed over granite or ba- Prospects Marama bean is regarded as hav-
salt. Marama bean is found in grassland and ing considerable potential as a crop for arid
wooded grassland vegetation. It occurs in local- and semi-arid regions, and it is being investi-
ized patches. gated in Australia, Israel and the United
Propagation and planting Propagation of States (Texas). It has potential for its roasted
marama bean is by seed. Germination is some- seeds and as a source of oil. However, before
times said to be improved by scarification. large-scale cultivation can be promoted, more
Soaking will kill the seed and it should not be information is needed on its ecological re-
sown in waterlogged soils. The 1000-seed quirements, adaptability to cultivation and
weight is 2-3 kg. Preliminary results under agronomy. Furthermore, genetic improvement
laboratory conditions show that vegetative and germplasm collection need attention and
propagation using sprouts is possible. research should be carried out on the presence
Harvesting The seeds of marama bean are of toxic constituents or antinutritional factors
collected in its native area from the wild and in the seeds and tubers.
by hand. The tubers are harvested by hand- Major references Bower et al., 1988; Da-
190 CEREALS AND PULSES

kora, Lawlor & Sibuga, 1999; Keegan & van and bracelets.
Staden, 1981; Ladizinsky & Smartt, 2000; In traditional African medicine root decoctions
Monaghan & Halloran, 1996; National Acad- of Tylosema fassoglense are taken to treat gas-
emy of Sciences, 1979; Powell, 1987; Ross, trointestinal problems in various countries.
1977; van Wyk & Gericke, 2000; Wickens, They are also used against anaemia, fever and
1998. pneumonia, and to heal the uterus after child-
Other references Brummitt & Ross, 1976; birth. The pulverized tuber is taken for the
Chandel & Singh, 1984; FAO, 1970; Francis & treatment of venereal diseases. The leaf sap is
Campbell, 2003; Graham & Vance, 2003; Hao applied to treat inflammations of the middle
Gang et a l , 2003; Hartley, Tshamekeng & ear. Infusions of powdered flowers are drunk
Thomas, 2002; Hornetz, 1993; ILDIS, 2002; against jaundice and hypertension. A decoction
IPGRI, undated; Keith &Renew, 1975; Ketsha- of the roots and flowers is drunk to treat impo-
jwang, Holmback &Yeboah, 1998; Leger, 1997; tence. Children are encouraged to eat the pods,
Lock, 1989; Mitchell et al., 2003; Schreiber, because these are thought to be good for the
1967; USDA, ARS & National Genetic Re- stomach. In veterinary medicine root decoc-
sources Program, 2001; Victor, undated; Viet- tions of Tylosema fassoglense are administered
meyer, 1978;Watt &Breyer-Brandwijk, 1962. as a galactagogue to cows before calving, and
Sources of illustration Powell, 1987. as a drench for a retained placenta.
Authors L.J.G.van der Maesen Properties The composition of seeds of Ty-
losema fassoglense per 100 g edible portion is:
water 7.5 g, energy 1888 kJ (451 kcal), protein
TYLOSEMAFASSOGLENSE (Schweinf.) Torre 43.5 g, fat 32.6 g, carbohydrate 14.6 g, fibre 4.2
& Hillc. g, Ca 80 mg, P 200 mg and Fe 40 mg (Malaisse
& Parent, 1985). Seeds collected in DR Congo
Protologue Bol. Soc. Brot., ser. 2, 29: 38 and Burundi yielded 24-30 g oil per 100 g, with
(1955). as principal fatty acids linoleic acid (36—43%),
Family Caesalpiniaceae (Leguminosae - Cae- oleic acid (33-35%), palmitic acid (12-16%),
salpinioideae) stearic acid (3-5%), behenic acid (3-5%) and
Chromosome number In — 52 arachidic acid (2-4%). Per 100 g the defatted
Synonyms Bauhinia fassoglensis Schweinf. seedcake meal contains 59 g protein, with a
(1868), Bauhinia kirkii Oliv. (1871). very high level of tyrosine (7-9 g per 100 g dry
Vernacular names Sprawling bauhinia, weight) and relatively high proportions of ly-
creeping bauhinia (En). Bauhinia rampant sine and proline (3-4 g and 4-5 g per 100 g dry
(Fr). weight, respectively). The seedcake meal con-
Origin and geographic distribution Tylo- tains substantial amounts of trypsin inhibitors
sema fassoglense occurs wild from Sudan and (295 TUI/mg) and phytate (3.5 g per 100 g dry
Ethiopia southwards to Namibia, Mozambique weight), but cyanogenic glycosides have not
and South Africa. been detected. For human or animal consump-
Uses The seeds of Tylosema fassoglense are tion of the seedcake, removal or inactivation of
frequently eaten, for instance in DR Congo, the trypsin inhibitors is recommended. Re-
Ethiopia, Tanzania, Malawi and South Africa. cently a cyanoglucoside (lithospermoside) has
Immature and mature seeds can be eaten raw, been isolated from the roots. In the rainy sea-
but they are usually cooked or roasted. The son the tuberous root may contain 86% water.
pods are also eaten raw or cooked. The seeds Botany Perennial herb or shrub, with tu-
are a coffee substitute. berous root; stem prostrate and trailing or
The leaves and young branches of Tylosema climbing, up to 6 m long, herbaceous or woody
fassoglense are grazed. In Kenya the Masai and below, young parts rusty-tomentose or rusty-
Kipsigis people make rope from the stems and hairy, with axillary forked tendrils (2-)3—6.5(—
plaited items from the young stems or from the 9.5) mm long. Leaves alternate, simple; stip-
fibres. The fibre is also suitable for making ules 2-4 mm x 2 mm, persistent; petiole (2-)3-
cloth. Water is obtained from the tuber in arid 10(-20) cm long; blade bilobed for up to one-
regions, and the tuber is sometimes made into third (sometimes up to half) its length, (5-)7-
porridge. Sap from the shoots can be used as 13(-20) cm x (4-)8-15(-24) cm, base deeply
potable water. The roots are used to produce a cordate, lobes ovate to obovate, sometimes
brown dye. In Ethiopia the seeds, after being rounded, subglabrous to densely rusty pubes-
hardened over a fire, are strung into necklaces cent beneath. Inflorescence a lateral raceme 5 -
 UROCHLOA 191

45 cm long; peduncle (2-)4-12(-18) cm long. a l , 2005; Dubois et al., 1995; Dubois et a l ,

Flowers bisexual, zygomorphic, 5-merous, het- 1994; Ross, 1977.
erostylous; pedicel (1.5—)2-4.5(-6) cm long; Other references Fort, Jolad & Nelson,
sepals l-1.5(-2.5) cm x 3—4 mm, with upper 2 2001; Grobbelaar & Clarke, 1975; Huxham et
completely united and the other 3 free; petals al., 1998; Lock, 1989; Malaisse & Parent, 1985;
unequal, 4 (larger ones) obovate-circular, (1.5—) Neuwinger, 2000; Tabuti, Lye &Dhillion, 2003;
2-4(-4.5) cm x 1-3 cm and tapering into a Thulin, 1989a; van Wyk & Gericke, 2000; Watt
basal claw, the upper one much smaller, yel- & Breyer-Brandwijk, 1962.
low, sometimes fading to pink; stamens 2, free, Authors M. Brink
with filaments 10-18 mm long, staminodes 8,
with filaments 3-6 mm long; ovary superior, 5 -
6 mm long, 1-celled, pubescent, style elongate, UROCHLOAMOSAMBICENSIS (Hack.) Dandy
stigma small. Fruit an obovoid to oblong-ovoid
pod 5-12 cm x 3-7.5 cm, flattened, woody, 1-2- Protologue Journ. Bot. 69:54 (1931).
seeded. Seeds ellipsoid to globose, somewhat Family Poaceae (Gramineae)
compressed, 1.5—3 cm x 1-2 cm, chestnut- Chromosome number 2n — 28, 30, 42
brown to black. Vernacular names Sabi grass, common
Tylosema comprises 5 species and occurs in urochloa, bushveld signal grass (En).
southern and eastern Africa. Some taxonomists Origin and geographic distribution Uro-
do not consider Tylosema a separate genus, but chloa mosambicensis is distributed from Kenya
include it in Bauhinia. Tylosema fassoglense is southwards to South Africa; it has been intro-
extremely variable, especially in its indumen- duced as a pasture grass into many other
tum, leaf size and inflorescence size. tropical countries, including Ghana and Mada-
Growth of Tylosema fassoglense is rapid, with gascar. It was introduced into Australia in the
the shoots growing up to 5 cm per day. In early 1900s and has become an important
southern Africa Tylosema fassoglense flowers grass for the northern Australian beef indus-
from October to March. Regeneration after fire try.
is rapid. Tylosema fassoglense does not form Uses In southern Africa the grain of Uro-
root nodules and relies on soil nitrogen. chloa mosambicensis is commonly used as a
Ecology Tylosema fassoglense occurs up to cereal; the ground grain is made into porridge.
2100 m altitude in woodland and grassland, Urochloa mosambicensis is a useful, drought-
sometimes in cultivated areas. It grows well on resistant, palatable pasture grass also suitable
poor, sandy soils, but is also found on rocky or for hay making. It is planted as a pasture grass
clay soils. It is moderately tolerant to flooding in East Africa, southern Africa, Madagascar,
and drought. India, Sri Lanka and Australia. In South Af-
Management Tylosema fassoglense is col- rica it is sown to improve overgrazed pastures.
lected from the wild. Fresh tuber weights up to In India it is used against soil erosion. In Aus-
78 kg have been recorded. To prepare porridge tralia it plays a role in mine site rehabilitation.
from the tuber, it is scraped clean, then grated, Properties Young green leaves of Urochloa
crushed or pounded, and ground into a fine mosambicensis typically contain up to 2.5% N,
meal which is cooked. 0.2% P and are 65-70% digestible. In the late
Genetic resources and breeding No sub- wet season these values are 1.2%, 0.15% and
stantial germplasm collections of Tylosema 55-60%, respectively. Dry leaves and stems are
fassoglense are known to exist. The Plant Ge- much lower in quality and typically contain
netic Resources Unit of the Agricultural Re- 0.5% N and 0.2% P. Information on the nutri-
search Council, Pretoria, South Africa, has 1 tional characteristics of the grain is not avail-
accession. Tylosema fassoglense is considered able.
neither rare nor threatened. Botany Tufted or stoloniferous perennial
Prospects Tylosema fassoglense has inter- grass up to 1.5 m tall; stem (culm) ascending,
esting properties, such as tolerance of low soil sometimes rooting at the lower nodes. Leaves
fertility and drought, seeds with high levels of alternate, simple and entire; leaf sheath silky
protein and fat, and tuberous roots storing pubescent; ligule a ciliate membrane; blade
water. Therefore, research into the potential of linear, 2-30 cm x 3-20 mm, pale to bright
this plant and its possible cultivation is cer- green, more or less hairy. Inflorescence com-
tainly justified. posed of 2-20 racemes borne on a central axis
Major references Brenan, 1967; Castro et 3-15 cm long; racemes (l-)2-9(—14) cm long,
192 CEREALS AND PULSES

bearing solitary spikelets on a narrowly winged can be broken by hammer-milling, destroying

rachis. Spikelet ovate, 2.5-5.5 mm x 1.5-3 mm, the hard lemma. In India Urochloa mosambi-
glabrous or hairy, acuminate, 2-flowered with censis is also propagated vegetatively using
lower floret male and upper bisexual; lower rooted cuttings. In pastures a seed rate of 4
glume elliptical-oblong, shorter than spikelet, kg/ha is recommended, or 2 kg/ha when grown
3-veined, shiny, upper glume as long as the intercropped with other pasture plants. Uro-
spikelet, 5-veined with cross-veins, granulöse chloa mosambicensis does well in intercropping
to rugulose, with a mucro; lemma acuminate, with leguminous pasture plants and is com-
leathery, 5-veined, with a mucro, palea shorter monly grown together with Stylosanthes spp.
than lemma; stamens 3; ovary superior, with 2 To obtain the grain, the inflorescences are
plumose stigmas. Fruit a strongly flattened picked when still slightly green and spread out
caryopsis (grain), pale buff or cream. in the sun to dry. When dry, the grains are
Urochloa comprises about 12 species distrib- easily rubbed from the stalks; they are ground.
uted in the Old World tropics, mainly in Africa. Grain yields of 100-300 kg/ha per year have
It is distinguished from the related Brachiaria been recorded from Australia. In pastures dry
by the shape and orientation of the spikelets matter yields of 1—8t/ha per year are produced.
but the boundary between the two genera is Genetic resources and breeding The larg-
unclear due to a number of intermediate spe- est germplasm collections of Urochloa mosam-
cies. It has been proposed that Brachiaria be bicensis are held in Australia (Australian Trop-
nearly completely reduced to Urochloa, which ical Crops & Forages Genetic Resources Cen-
would increase the size of Urochloa to about tre, Biloela, Queensland, 73 accessions, mainly
120 species, with a pantropical distribution. from African countries; CSIRO Townsville Di-
Within Urochloa the species are sometimes vision of Tropical Crops and Pastures, Towns-
difficult to separate. Urochloa mosambicensis ville, Queensland, 63 accessions). In Africa 18
is the perennial counterpart of the annual accessions are held in South Africa (Grassland
Urochloa trichopus (Höchst.) Stapf, which does Research Centre, Department of Agricultural
not possess dormant buds at the base. The Development, Pretoria), 7 accessions in Ethio-
grain of Urochloa brachyura (Hack.) Stapf, pia (International Livestock Research Institute
distributed in East and southern Africa, is (ILRI), Addis Ababa) and 7 accessions in Kenya
eaten in Namibia; the plant is also grazed by (National Genebank of Kenya, Crop Plant Ge-
animals. netic Resources Centre, KARI, Kikuyu). In
Seeds of Urochloa mosambicensis germinate view of its wide distribution and abundance,
early in the wet season and vegetative growth Urochloa mosambicensis is not threatened by
continues until soil water is exhausted. Flower- genetic erosion.
ing starts 3-4 weeks after the start ofthe rainy The collection held in Biloela has been investi-
season and continues until growth ceases. Seed gated for a range of morphological and agro-
matures in 3-4 weeks. Leaves live for 5-25 nomical attributes, and considerable variation
weeks depending mainly on water supply. was found in time to maturity, stolon develop-
Plants are often short-lived (3-4 years). ment, plant height and yield. Cultivars of
Urochloa mosambicensis is an obligate apo- Urochloa mosambicensis have been registered
mict. It follows the C4photosynthetic pathway. in Australia, e.g. 'Nixon' and 'Saraji'.
Ecology In its natural habitat Urochloa Prospects Urochloa mosambicensis is a
mosambicensis occurs up to 1600 m altitude in useful wild cereal in southern Africa, but it has
regions with an average annual rainfall of400— more potential as a pasture grass for semi-arid
1200(—1600) mm, in savanna woodland and tropical regions. Investigations are needed to
open grassland, often in disturbed or over- assess the nutritional quality of the grains.
grazed locations (e.g. fallow land, roadsides). It Major references Burkill, 1994; Clayton,
grows in a wide range of soils, but prefers 1989; Hanelt &Institute of Plant Genetics and
lighter, more fertile soils. In northern Austra- Crop Plant Research (Editors), 2001; Mclvor,
lia it becomes dominant after fires. 1992; Pengelly &Eagles, 1999.
Management The grains of Urochloa mo- Other references Bogdan, 1977; Clayton &
sambicensis are mostly collected from the wild, Renvoize, 1982; FAO, undated; Gibbs Russell
but sometimes plants are grown in gardens et al., 1990; Mackay, 1974; Prakash & Uniyal,
alongside maize. The 1000-seed weight is 1-1.7 1980; van Wyk & Gericke, 2000; Veldkamp,
g. Fresh seed has dormancy, which breaks 1996a.
down after 9-12 months storage. Dormancy Authors M. Brink
 UROCHLOA 193

U E O C H L O A TRICHOPUS (Höchst.) Stapf but the boundary between the two genera is
unclear due to a number of intermediate spe-
Protologue Oliv., Fl. trop. Afr. 9(4):589 cies. It has been proposed that Brachiaria be
(1920). nearly completely reduced to Urochloa, which
Family Poaceae (Gramineae) would increase the size of Urochloa to about
Chromosome number 2n=14, 28 120 species, with a pantropical distribution.
Origin and geographic distribution Uro- Within Urochloa the species are sometimes
chloa trichopus is distributed in the more dry difficult to separate. Urochloa trichopus isthe
regions throughout tropical Africa. It also oc- annual counterpart of the perennial Urochloa
curs in Yemen and has been introduced into mosambicensis (Hack.) Dandy, which possesses
Brazil andIndia. dormant buds at thebase.
Uses The grain of Urochloa trichopus is Ecology Urochloa trichopus occurs fromsea-
sometimes gathered for food, e.g. in Kordofan level up to 1500 m altitude in semi-arid cli-
(Sudan), Tanzania, Botswana and Zimbabwe. mates, in grassland and savanna woodland;
In Botswana itis ground into flour, whichmay also in disturbed locations and as an arable
be cooked with water, milk or melon juice or weed.
made into cake;itisalso used formaking beer. Management Urochloa trichopus is col-
Urochloa trichopus is valued as a fodder in lected from thewild. In Botswana stored grain
semi-arid regions; in Brazil and India it is a is attacked by weevils, ants and rats, but it
forage grass. generally stores well. In Botswana the grain is
Properties The fodder value of Urochloa considered difficult to thresh and pound.
trichopus plants in the Sahel is:crude protein Urochloa trichopus is considered a weed in
10.7%, crude fibre 28.5%, crude fat 1.4%, nitro- Ethiopia.
gen-free extractives 45.2%, P 0.19%, K 4.69%, Genetic resources and breeding TheIn-
Ca 0.38%, Mg 0.37% and Na 0.02%. In Bot- ternational Livestock Research Institute
swana the crude protein content of Urochloa (ILRI), Addis Ababa, Ethiopia, holds 5 acces-
trichopus ranges from 6.2%in the dry season sions of Urochloa trichopus (3from Ethiopia; 2
(July) to 10.4% in the rainy season (January), from Mali). Three accessions from Ethiopia are
and the dry matter digestibility ranges from held at Centro Internacional de Agricultura
41% in July to 57%inJanuary. Information on Tropical (CIAT), Cali, Colombia; 2 accessions
the nutritional characteristics of the grain is from Tanzania in the Australian Tropical
not available. Crops & Forages Genetic Resources Centre,
Botany Coarse, tufted annual grass up to Biloela, Queensland. In view ofits wide distri-
1.7 m tall; stem (culm) geniculately ascending, bution, Urochloa trichopus isnotthreatened by
often rooting at the lower nodes. Leaves alter- genetic erosion.
nate, simple andentire; leaf sheath glabrous to Prospects Urochloa trichopus is a useful
slightly pubescent; ligule a ciliate membrane; source of food and fodder in semi-arid regions
blade linear, 5-30 cm x 5-20 mm, acuminate, of tropical Africa, butis unlikely to increase in
glabrous orhairy. Inflorescence composed of3 - importance. For use as a cereal, the small
20 racemes borne on a central axis 4—20cm grain size and difficulty in processing are con-
long; racemes 1-14 cm long, bearing solitary sidered serious limitations. Its role as a pas-
spikelets ona narrowly winged rachis. Spikelet ture grass will probably remain modest com-
ovate, 2.5-5.5 mm long, glabrous or less often pared to that ofits perennial and more persis-
hairy, acuminate, 2-flowered with lower floret tent counterpart Urochloa mosambicensis.
male and upper bisexual; lower glume ellipti- Major references Burkill, 1994; Clayton &
cal-oblong, slightly shorter than spikelet, 3- Renvoize, 1982; Gibbs Russell et al., 1990; Mo-
veined, upper glume aslong asthespikelet, 5(- diakgotla et a l , 1999;Phillips, 1995.
7)-veined with cross-veins; lemma acuminate, Other references Bartha, 1970; Clayton,
leathery, 5-veined, with a mucro; palea shorter 1972; Clayton, 1989; Cope, 1995; Hanelt &
than lemma; stamens 3;ovary superior, with 2 Institute ofPlant Genetics and Crop Plant Re-
plumose stigmas. Fruit a strongly flattened search (Editors), 2001;Longhi-Wagner &deOli-
caryopsis (grain). veira, 2002; Pratchett, 1983; Trouin, 1970;
Urochloa comprises about 12 species distrib- Veldkamp, 1996a.
uted intheOldWorld tropics, mainly in Africa. Authors M. Brink
It is distinguished from the related Brachiaria
by the shape and orientation of the spikelets
194 CEREALS AND PULSES

VATOVAEAPSEUDOLABLAB(Harms) J.B.Gillett up to 8-seeded. Seeds almost globose to irregu-

larly ellipsoid or squarish, 4.5-7 mm x 4.5-6.5
Protologue Kew Bull. 20(1): 104 (1966). mm x 2.5-3 mm, brown, sometimes speckled
Family Papilionaceae (Leguminosae - Papil- with black.
ionoideae, Fabaceae) Vatovaea comprises a single species. Although
Origin and geographic distribution Vato- Vatovaea pseudolablab becomes woody, plants
vaea pseudolablab is found wild in the drier may already flower when still quite herba-
parts of Sudan, Ethiopia, Somalia, Kenya, ceous; they are basically self-pollinating.
Uganda and Tanzania; also in Yemen and Ecology Vatovaea pseudolablab is found up
Oman. to 1500 m altitude in dry grassland or bush-
Uses The seeds of Vatovaea pseudolablab land in regions with an annual rainfall of 250-
are eaten either raw or after boiling or roast- 600 mm, often along lava or drainage lines,
ing. Immature pods, flowers and leaves are occasionally in seasonally wet grassland on
eaten raw or cooked as a vegetable. The tuber- clay.
ous and juicy roots are edible and are con- Management Vatovaea pseudolablab is com-
sumed raw or after boiling or roasting. They monly collected from the wild, but only occa-
are sometimes eaten as a snack, especially sionally sown, e.g. in Kenya. In Ethiopia it is
after roasting; they are also used as emergency semi-domesticated by Konso farmers, who keep
food and as a source of water. Flour made from it in their fields intercropped with other food
the roots is mixed with sorghum flour to pre- plants. Vatovaea pseudolablab can be propa-
pare a stiff porridge. It is normally stored and gated by seed. The tuberous roots can be dug
used in lean periods. Farmers grow and con- out any time ofthe year; they are best gathered
sume Vatovaea pseudolablab commonly, but when the foliage has died back. Flour is pro-
during food shortages more people rely on it for duced from the roots by peeling, chopping, dry-
their daily food. The plant is eaten by cattle, ing and grinding.
goats, sheep, camels and donkeys. The root Genetic resources and breeding Vatovaea
fibres are made into rope, hats and fly whisks. pseudolablab populations are dwindling in
Properties The tuberous roots of Vatovaea East Africa and in the Arabian Peninsula be-
pseudolablab are fibrous and contain much cause it is a popular food and fodder. Its ge-
juice; they have a pleasant, sweet taste even netic pool is likely to shrink fast if no action is
when eaten raw. taken. Two accessions of Vatovaea pseudolab-
Botany Liana or shrub up to 1.5(—3) m tall; lab are kept in Ethiopia at the International
stem branched, glabrous to sparsely pubescent; Livestock Research Institute (ILRI), Addis
roots tuberous. Leaves alternate, 3-foliolate; Ababa.
stipules oblong, c. 3.5 mm x 1.5 mm; petiole up Prospects Vatovaea pseudolablab is a use-
to 6 cm long, ribbed, rachis up to 2 cm long; ful multipurpose plant for dry regions and its
stipels small; petiolules 1-2 mm long; leaflets potential seems worthwhile exploiting. It is
ovate to narrowly ovate-rhomboid, up to 8 cm x recommended to start collecting and evaluating
6.5 cm, sometimes slightly 3-lobed, glabrous to germplasm and to test accessions for their per-
sparsely pubescent. Inflorescence an axillary formance in the field. Promising material
false raceme up to 50 cm long, pubescent, should be multiplied further. Furthermore,
many-flowered; peduncle 6-21 cm long; bracts investigations should be carried out on the
up to 2 mm long. Flowers bisexual, papiliona- agronomy of the plant and its nutritional prop-
ceous; pedicel c. 3 mm long; calyx c. 5 mm long, erties.
5-lobed, 2-lipped, the lower 3 lobes rounded- Major references Beentje, 1994; Gillett et
triangular, the upper 2 lobes united; corolla a l , 1971; Maundu, Ngugi & Kabuye, 1999;
greenish purple, standard 1-2 cm x 1.5—2 cm, Schippers, 2000;Thulin, 1993.
emarginate, with 2 appendages near the base, Other references African Studies Center,
wings with a long narrow spur, keel incurved; undated; Gillett, 1966; Huxham et al., 1998;
stamens 10, 9 fused and 1free; ovary superior, ILDIS, 2002; IPGRI, undated; Maundu, 1997;
linear, 1-celled, style long, incurved, usually Morgan, 1981;Thulin, 1989a; Thulin, 1989b.
hairy inside towards the apex and with a re- Authors M. Brink
flexed appendage above the stigma. Fruit a
linear-oblong pod 4.5-6 cm x 0.5-1 cm, curved,
flattened, widening towards the apex, dehis-
cent, at first silky pubescent, later glabrescent,
 VICIA 195

VICIA FABA L. immature pods are eaten. In Ethiopia and Eri-

trea main dishes include 'shiro wot' (hulled
Protologue Sp. pi. 2: 737 (1753). seeds ground and made into sauce), 'kik wot'
Family Papilionaceae (Leguminosae - Papil- (hulled and split seeds boiled and made into
ionoideae, Fabaceae) sauce), and 'ful' (hulled and boiled seeds, spiced
Chromosome number 2n — 12 and minced with butter). Snacks include 'eshet'
Vernacular names Faba bean, broad bean, (fresh green seeds eaten raw or roasted), 'kolo'
horse bean, field bean, tick bean (En). Fève, (roasted dry seeds), 'nifro' (boiled dry or fresh
fève de(s) marais, fèverole, fèvette, gourgane green seeds), 'gunkul' (seeds soaked, sprouted
(Fr). Faveira, fava (Po). and roasted), 'ashuk' (seeds roasted first and
Origin and geographic distribution Faba then soaked), and 'endushdush' (seeds soaked
bean is only known in cultivation. Its centre of first and then roasted). Throughout the Arab
origin and domestication is probably in western world faba bean seeds are consumed minced
Asia, from where it spread into Europe, Africa with onion, garlic and herbs, and eaten for
and central Asia. Ethiopia and Afghanistan are breakfast as 'ful medames'. Stewed seeds are
considered secondary centres of diversity. Faba eaten at any time of the day; seeds made into a
bean domestication occurred between 7000 and paste are often used as a sandwich filling.
4000 BC, and by the 3 rd millennium BC it was Mature seeds and vegetative parts of faba bean
widely distributed throughout the Mediterra- serve as concentrate, hay and silage for domes-
nean region. The evolution of the large-seeded tic animals. The straw is used as fuel for cook-
type was much later (around 500 AD), and may ing in Ethiopia. The stems and leaves are used
have occurred in western Asia and in the Medi- as green manure, e.g. in China. In China seeds
terranean region. Faba bean was probably not and vegetative parts have a wide range of me-
grown to any extent in Japan and China before dicinal applications. In Europe the inside of the
1200 AD, nor in the Americas before the arri- green pods is rubbed on warts to remove them.
val of the Spaniards. Nowadays, faba bean is Production and international trade Accord-
widely grown in temperate and subtropical ing to FAO statistics the world production of
regions and at higher altitudes in the tropics. dry faba bean seeds in 1999—2003 amounted to
In tropical Africa it is mainly found in East 3.9 million t/year from 2.6 million ha. The main
Africa, especially in Sudan and Ethiopia. producing countries are China (1.9 million
Uses Faba bean is grown as a field crop for t/year from 1.2 million ha), Ethiopia (405,000
the dry mature seeds and as a garden crop for t/year from 370,000 ha), Egypt (396,000 t/year
the immature seeds or pods. In Ethiopia, Medi- from 134,000 ha) and Australia (242,000 t/year
terranean countries, the Middle East and from 164,000 ha). The annual production in
China the dry mature seeds are widely used as sub-Sahara Africa in 1999-2003 was estimated
food, and in many countries the green imma- at 510,000 t, almost entirely from Ethiopia
ture seeds are boiled or eaten raw as vegetable. (405,000 t) and Sudan (100,000 t). The annual
In Mediterranean countries and in India whole world production of dry faba bean seeds de-
clined from about 5 million t (from 5 million
ha) in the early 1960s to about 4 million t (from
2.7 million ha) in the early 2000s. The reduc-
tion in area under cultivation in China from
about 3.5 million ha in the early 1960s to about
1.25 million ha in the early 2000s accounted for
the largest share of the reduction in produc-
tion. In contrast, the annual production in sub-
Sahara Africa increased during the same pe-
riod from 230,000 t (250,000 ha) to 540,000 t
(450,000 ha). The world production of green
faba bean seeds in 1998-2003 was estimated at
940,000 t/year from 2.6 million ha, with Algeria
(118,000 t/year), China (114,000 t/year) and
Morocco (112,000 t/year) as the largest produc-
ers; the production of green faba bean seeds in
tropical Africa is negligible.
Vicia faba -planted World exports of dry faba bean seeds in 1998-
196 CEREALS AND PULSES

2002 amounted to 475,000 t. The main export-

ing countries were Australia (201,000 t), the
United Kingdom (114,000 t), China (63,000 t)
and France (53,000 t). The main importers in
this period were Egypt (197,000 t), Italy
(169,000 t) and Spain (52,000 t). The exports
from African countries are negligible.
Properties The composition of raw mature
faba bean seeds per 100 g edible portion is:
water 11.0 g, energy 1425kJ (340 kcal), protein
26.1 g, fat 1.5 g, carbohydrate 58.3 g, dietary
fibre 25.0 g, Ca 103 mg, Mg 192 mg, P 421 mg,
Fe 6.7 mg, Zn 3.1 mg, vitamin A 53IU, thiamin
0.56 mg, riboflavin 0.33 mg, niacin 2.8 mg,
vitamin Ik 0.37 mg, folate 423 (ig and ascorbic
acid 1.4 mg. The essential amino acid composi-
tion per 100 g edible portion is: tryptophan 247
mg, lysine 1671 mg, methionine 213 mg,
phenylalanine 1103 mg, threonine 928 mg,
valine 1161mg, leucine 1964 mg and isoleucine
1053 mg. The principal fatty acids per 100 g
edible portion are: linoleic acid 581 mg, oleic
acid 297 mg and palmitic acid 204 mg (USDA,
2004).
In certain persons genetically predisposed,
living mainly in the Mediterranean area, con- Vicia faba - 1, flowering and fruiting branch; 2,
sumption of faba bean seeds, particularly im- seeds.
mature ones, and even the inhalation of pollen, Source: PROSEA
results in 'favism', a kind of haemolytic anae-
mia resulting from the accumulation of ß- varying in shape, toothed; leaflets ovate to el-
glycosidase (vicine and convicine) and their liptical, (3-)4-8(-10) cm x l-2(-4) cm, entire.
aglycones in individuals deficient in the en- Inflorescence an axillary, sessile, short raceme,
zyme glucose-6-phosphate dehydrogenase in 1-6-flowered. Flowers bisexual, papilionaceous,
their red blood cells. Soaking before cooking almost sessile; calyx campanulate, 5-lobed,
inactivates the toxic compounds. Other antinu- tube c. 7 mm long, lobes almost equal, nar-
tritional factors in faba bean seeds include rowly triangular, 2—8 mm long; corolla white,
trypsin inhibitors, lectins (haemagglutinins), marked by a dark brown blotch, fragrant,
tannins, oligosaccharides and phytate. Faba standard broadly ovate, c. 2.5 cm x 1.5 cm,
bean seeds have lipid-lowering effects in hu- approaching the keel, wings oblong-ovate, c. 2.5
mans and rats. Proteins isolated from the seed cm x 0.5 cm, keel c. 1.5 x 0.5 cm; stamens 10, 9
have shown antioxidative activity, whereas the united and 1free, c. 15 mm long, anthers ellip-
lectin agglutinin may slow the progression of soid to ovoid, about 1 mm long, dark brown;
colon cancer. Faba bean straw is a good feed ovary superior, sessile or nearly so, very slen-
with high protein content (5-20%) and digesti- der, compressed, c. 1.5 cm long, style abruptly
bility (50% of the dry matter). The high tannin upturned, c. 3 mm long, with a tuft of hairs
content of the seeds (up to 9%) results in a bit- near the glandular-papillate stigma. Fruit a
ter taste when they are fed raw to animals, but narrowly oblong, cylindrical to flattened pod,
cultivars have been developed with low tannin (3-)5-10(-30) cm x l-1.5(-3) cm, bulging over
content (1%) and high digestibility. the seeds, sparsely pubescent when mature,
Description Erect, robust, stiff, glabrous, beaked, 2-6-seeded. Seeds ovoid to oblong,
annual herb up to 2 m tall; stem stout, square, compressed, 1—3 cm in diameter, brown, red-
hollow with one or more basal branches; tap- dish or green; hilum narrowly oblong. Seedling
root well-developed, with strong lateral roots. with hypogeal germination.
Leaves alternate, paripinnate, with 2-6 leaf- Other botanical information Vicia com-
lets, without tendril, but rachis ending in a prises about 120 species, mainly in the tem-
short acumen; stipules conspicuous, widely perate regions of the northern hemisphere and
 VICIA 197

South America, with a few species in Africa. mediate between self- and cross-pollinating.
Vicia faba is unique in the genus: it has larger Cross-pollination rates up to 92% have been
but fewer chromosomes and the greatest recorded, but they are mostly between 20% and
amount of DNA content (around 13,000 Mbp). 50%. Insects facilitate cross-pollination. The
No other Vicia could successfully be crossed duration of the growth cycle varies from 3
with Vicia faba despite many attempts. Mor- months (Sudan, Canada) to 11 months (north-
phometry and seed-protein electrophoresis western Europe). In Ethiopia the growth cycle
studies have shown marked differences be- is 3-7 months. Faba bean is effectively nodu-
tween Vicia faba and wild relatives {Vicia nar- lated by Rhizobium leguminosarum.
bonensis L., Vicia galilaea Plitmann & Zohary Ecology Faba bean is grown in temperate
and Vicia hyaeniscyamus Mouterde). regions, as a winter crop in the subtropics, and
The infraspecific taxonomy of Vicia faba is as a high-altitude crop in the tropics. It is not
confusing. Several varieties have been distin- suited to the lowland tropics, where it may
guished, based on the shape and size of the flower well but usually does not produce pods.
seeds. Cultivars with small and rounded seeds A mean daily temperature around 13°C is op-
are often called tick bean, those with interme- timal for growth. In Ethiopia faba bean is
diate seed size horse bean, and those with grown at 1300-3800 m altitude, but mostly at
large and flat seeds broad bean. However, 2000—2500 m. Rust is the major production
there is no discontinuity in seed size between constraint below 1800 m, and frost above 3000
the groups, and they can be freely crossed. Ar- m. Faba bean requires an annual rainfall of
bitrarily, small-seeded types have been recog- 700-1000 mm, of which more than 60% during
nized as those with a 1000-seed weight of less the growing period. Long photoperiods reduce
than 700 g, medium-seeded types with 700- the time to flowering and the position of the
1200 g, and large-seeded types with more than first flowering node, e.g. in northern European
1200 g. cultivars, but under field conditions daylength-
Vicia sativa L. is widely cultivated as a forage. neutrality is often observed. Faba bean prefers
Its seeds, young stems and leaves are recorded well-drained, almost neutral soils (pH 6.5-7.5),
as being used for human consumption in with moderate fertility. It hardly tolerates wa-
Ethiopia and the Caucasus. However, the seeds terlogging or drought.
and hay can have toxic effects (HCN-poisoning Propagation and planting Faba bean is
due to the cyanogenic glycoside vicianine; propagated by seed. The 1000-seed weight is
antinutritional effects of ß-cyanoalanine). The 400-2000 g. Faba bean does not require a fine
seeds of Vicia villosa Roth, cultivated for fod- seedbed, but the land should be ploughed to a
der in East Africa, and Vicia paucifolia Baker loose seedbed. The crop is broadcast or planted
are also said to be collected and eaten. Vicia in rows; in mechanized agriculture drilling is
monantha Retz. (bard vetch) has at least in common. The planting depth is 2—5 cm. Seed
former times been grown in oases in the Sa- rates vary widely; higher rates are required in
hara. However, the seeds of many Vicia spe- the cool high-altitude areas of the tropics,
cies, including Vicia villosa and Vicia monan- where crop growth is slower than in warm mid-
tha, are known to contain canavanine, a toxic altitude areas. Seed rates up to 150-200 kg/ha
arginine analogue. are recommended in Ethiopia, 70-120 kg/ha in
Growth and development Five principal Sudan and 95-100 kg/ha in Egypt. Spacings
stages have been distinguished in a key for vary from place to place. In Ethiopia 40 cm
faba bean development: germination and between rows and 5 cm between plants is rec-
emergence, vegetative development, reproduc- ommended. In Sudan a distance between rows
tive development, pod senescence and stem of 20-40 cm is recommended, with 5-20 cm
senescence. Vegetative development continues within rows and 1-3 plants per pocket. How-
after reproductive development has started, ever, small-scale farmers in Sudan, Ethiopia
thus both stages run concurrently. The onset of and Eritrea practice broadcasting. In Egypt
flowering strongly depends on environmental planting on ridges is the usual practice. In case
conditions (temperature, photoperiod), and of sowing on both sides of the ridges, a spacing
may range from 1 month to 7-8 months. The of 60 cm between ridges and 15-20 cm between
longer durations occur in winter-sown crops in pockets with 2 seeds/pocket is optimal. In
temperate regions. Flowering starts, on aver- Ethiopia common sowing dates are mid-June in
age, at node 7 and continues over as many as mid-altitude areas and late June to early July
20 nodes. Faba bean pollination habit is inter- in high-altitude areas. Planting in Egypt and
198 CEREALS AND PULSES

Sudan may start in mid-October and proceed Important virus diseases of faba bean are bean
until late November. Faba bean is grown as a yellow mosaic virus (BYMV), bean leaf roll
sole crop or in intercropping, e.g. with pea in virus (BLRV) and broad bean stain virus
Ethiopia, sugar cane in Egypt and various (BBSV). Root-knot nematodes (Meloidogyne
crops (wheat, rape, cotton and barley) in China. spp.), stem nematodes (Ditylenchus dipsaci)
Management Faba bean is sensitive to and root-lesion nematodes (Pratylenchus spp.)
weed competition and rigorous control of weeds also affect faba bean.
is needed from 3-8 weeks after seedling emer- Aphids (Aphis craccivora, Aphis fabae and
gence. Weeds are controlled manually or with Acyrthosiphon pisum) are major insect pests of
herbicides. One or 2 manual weedings may be faba bean, e.g. in Sudan and Egypt. Other in-
required, the first one at 3—4 weeks after sect pests are the leaf weevil (Sitona lineatus),
emergence, the second one at 6—8weeks. Faba the pod borer (Helicoverpa armigera), the root
bean is grown under irrigation in Egypt and nodule weevil (Sitona amurensis), cutworms
Sudan, whereas in Ethiopia and Eritrea it is (Agrotis spp.), the leaf miner (Liriomyza con-
grown entirely under rainfed conditions. Nitro- gesta) and the lesser armyworm (Spodoptera
gen application may not be necessary where exigua). Bruchids (Bruchus and Callosobruchus
Rhizobium leguminosarum is present, but in spp.) are major storage pests, e.g. in Ethiopia.
some countries 30—40 kg N/ha is applied as a In Europe, the Middle East and northern Af-
starter. Atmospheric nitrogen fixation rates of rica the parasitic herb Orobanche crenata
45-550 kg N per ha per year (on average Forssk. (bean broomrape) is a critical problem.
around 200 kg) have been recorded for faba No practical control measure is available.
bean. In areas where the bacteria are absent, Harvesting Harvesting of faba bean is done
inoculation of the seed with bacteria is an op- before full physiological maturity, because late
tion. Most small-scale farmers in Ethiopia do harvesting may result in pod shattering and
not apply chemical fertilizers. Experiments in rotting, particularly when rain is encountered.
Ethiopia have shown little or no response to N- The appropriate stage is when the leaves and
fertilization, but P-application often leads to the pods dry out and the seed moisture content
significant yield increases. In Sudan faba bean is reduced to 16—18%. Faba bean can be com-
is not normally responsive to application of N bine harvested, but in tropical Africa manual
and K, due to the presence of N-fixing bacteria harvesting is the common practice. Plants are
and high inherent K in the soil. However, P is hand-pulled or cut using a small knife or
limiting, as the soils are alkaline (pH>8) and sickle. Harvesting is usually done in the early
only a little P is available for the crop. Hence, morning or late afternoon to reduce losses from
placement of P close to the root system is rec- shattering. The harvested plants are gathered
ommended. In Egypt, 36 kg N and 30 kg P per into small heaps and left in the field for a few
ha is applied for traditional cultivars (yielding days to dry. Then they are transported to a
about 2.5 t/ha), whereas for improved cultivars threshing ground.
(yielding up to 5 t/ha) additional top dressings Yield The average seed yield of faba bean in
(at 40 and 70 days after sowing) of 50 kg K per Africa (1.3 t ha/ha) is below world average (1.5
ha are recommended as well as a micronutrient t/ha), while the average yields obtained in Asia
spray of 60 g Zn, 40 g Mn and 20 g Fe per ha. (1.7 t/ha) and Europe (2.2 t/ha) are higher. Ex-
Faba bean plays an important role in soil fertil- ceptionally high yields are obtained in Egypt
ity management as a rotation crop; it is often and Sudan where the crop is irrigated (3.0 and
grown in rotation with cereals, especially with 2.3 t/ha, respectively).
wheat or barley. Handling after harvest Threshing of faba
Diseases and pests The most important bean is traditionally done by beating the plants
fungal diseases of faba bean are chocolate spot with sticks or by trampling animals. Seeds
{Botrytis fabae and Botrytis cinerea), ascochyta should be stored under dry and cool conditions,
blight (Didymella fabae; synonym Ascochyta free of pests and prevented from absorbing
fabae), rust (Uromyces viciae-fabae), and black moisture. Cleaning seeds and storage struc-
root rot (Fusarium spp.). Chocolate spot and tures before storing is important. Seeds with a
rust have been recorded as causing up to 50% moisture content of 11-14% can be stored for
yield loss in Egypt. Suggested control measures 2-7 years at temperatures of 5-10°C and for1—
include use of resistant cultivars, cultural 4 years at 10-20°C.
practices (crop rotation, drainage, disease-free Genetic resources Over 25,000 faba bean
seed, burning of crop residues) and fungicides. accessions are currently conserved in different
 VICIA 199

countries. The International Center for Agri- tance to rust has been tagged, and quantitative
cultural Research in the Dry Areas (ICARDA) trait loci associated with seed weight, resis-
in Aleppo, Syria, holds about 10,700 faba bean tance to ascochyta blight and resistance to
accessions and 5900 accessions of wild Vicia bean broomrape have been located. The pres-
species. Other important collections are kept in ence of vicine and convicine in faba bean seeds
China (Institute of Crop Germplasm Resources is controlled by a single recessive gene that
(CAAS), Beijing; 3800 accessions) and Austra- reduces their content 20-fold. However, the
lia (Australian Temperate Field Crops Collec- same gene increases susceptibility to patho-
tion, Horsham; 2200 accessions). The largest gens and parasites. Two recessive genes elimi-
collection of faba bean germplasm in Africa nate tannin production in faba bean.
(2000 accessions) is kept at the Institute of Prospects Faba bean productivity is far
Biodiversity Conservation (IBC), Addis Ababa, below the potential in many countries of tropi-
Ethiopia. The collections include sources for cal Africa because of the biological limitations
multiple disease resistance, wild and primitive of the traditional cultivars and poor manage-
forms, lines carrying structural mutations, ment practices. However, faba bean will re-
breeding lines and cultivars of special interest. main an important crop in parts of tropical
The worldwide diversity available in faba bean Africa. Export demand is strong and regional
has not yet been adequately sampled, and the markets are emerging, e.g. between Ethiopia
available collections have not sufficiently been (exporter) and Sudan and Egypt (importers). In
characterized. Faba bean shows orthodox seed addition to the needs emanating from the
storage behaviour. physical environment, farming systems and
Breeding High yield and resistance/toler- local consumers' preferences, export qualities
ance to both biotic and abiotic stresses are the and standards also deserve priority in re-
prime objectives across faba bean breeding search. Efforts are being undertaken in some
programmes. Some breeding efforts to improve countries, e.g. China, to develop new higher-
the yield potential of conventional indetermi- value types superior in colour, smell and taste,
nate types have been promising. Sources of and these efforts, coupled with the wealth of
resistance to chocolate spot, ascochyta blight genetic diversity available, might result in new
and rust identified at ICARDA have been used opportunities.
in many national faba bean breeding pro- Major references Enneking, 1995; Hawtin
grammes. Australia has released cultivars re- & Webb (Editors), 1982; Hebblethwaite (Edi-
sistant to chocolate spot and ascochyta blight. tor), 1983; Jansen, 1989e; Jellis, Bond & Boul-
In Ethiopia the cultivars 'Wayu' and 'Selale' ton, 1998; Knight (Editor), 2000; Knott, 1990;
with resistance to black root rot disease on Lang et al., 1993; Muehlbauer & Kaiser (Edi-
waterlogged Vertisols have recently been re- tors), 1994;Thulin, 1989a.
leased. More than 10 cultivars have been re- Other references Bond, 1995; Bond et al.,
leased for different agro-ecological conditions 1985; Böttinger et a l , 2001; Ghizaw et al.,
in Ethiopia and Egypt, and 7in Sudan. Recent- 1999; Hanelt &Institute of Plant Genetics and
ly, export-quality seed (large seed size) has at- Crop Plant Research (Editors), 2001; ILDIS,
tracted attention in breeding programmes in 2002; Kay, 1979; Madar & Stark, 2002; Max-
China and Ethiopia. Attempts to develop hybrid ted, 1995; McPhee & Muehlbauer, 2002; Pol-
faba bean cultivars have not yet been success- hill, 1990; Purseglove, 1968; Roman et al.,
ful because of lack of an effective male sterility 2004; Singh & Saxena (Editors), 1993; Smartt,
system. Efforts to change the indeterminate 1976; Summerfield (Editor), 1988;Tindall, 1983;
growth habit into determinate types with in- USDA, 2004; Westphal, 1974; Zemede Asfaw &
creased yield through mutation breeding have Mesfin Tadesse, 2001.
also not been successful so far. Sources of illustration Jansen, 1989e.
In-vitro callus formation and plant regenera- Authors M.Jarso &G. Keneni
tion have been achieved with hypocotyl, cotyle-
don and embryo expiants. Stably transformed
faba bean lines have been produced using an
Agrobacterium-mediated gene transfer system.
Genetic linkage maps of the faba bean genome
have been constructed based on morphological
markers, isozymes, RAPDs, seed protein genes
and microsatellites. A gene controlling resis-
200 CEREALS AND PULSES

V I C I A H I E S U T A (L.) G r a y sphere and South America, with a few species

in Africa.
Protologue Nat. arr. Brit. pi. 2: 614 (1821). Vicia hirsuta is effectively nodulated by Rhizo-
Family Papilionaceae (Leguminosae - Papil- bium leguminosarum.
ionoideae, Fabaceae) Ecology In East Africa Vicia hirsuta is
Chromosome number 2n - 14 found in grassland, scrub, forest margins and
Vernacular names Hairy tare, tiny vetch, lava plains at 2000—3500 m altitude. Vicia hir-
hairy vetch (En). Ers velu, vesceron, vesce suta is a long-day plant. In many countries it is
hérissée (Fr). Cigerâo (Po). considered a weed.
Origin and geographic distribution Vicia Genetic resources and breeding The larg-
hirsuta is widely distributed in Europe, Asia est germplasm collections of Vicia hirsuta are
and Africa. In Africa it is native from northern maintained at the International Centre for
Africa through DR Congo and East Africa to Agricultural Research in Dry Areas (ICARDA),
Angola and South Africa. It is often introduced Aleppo, Syria (39 accessions) and the Interna-
and naturalized elsewhere, e.g. in the Indian tional Centre for Underutilised Crops, Univer-
Ocean islands. Vicia hirsuta is sometimes cul- sity of Southampton, Southampton, United
tivated as a pulse or as a fodder crop in India Kingdom (32 accessions). In tropical Africa
and was formerly grown in eastern Europe. some accessions are held in Kenya (National
Uses The seeds of Vicia hirsuta are collected Genebank of Kenya, Crop Plant Genetic Re-
from the wild and eaten cooked or roasted in sources Centre, KARI, Kikuyu, 9 accessions)
Ethiopia. They were eaten as a famine food in and Ethiopia (International Livestock Re-
Europe and Asia. The leaves and shoots are search Institute (ILRI), Addis Ababa, 5 acces-
used as a vegetable in Ethiopia. Vicia hirsuta sions). In view of its wide distribution and un-
is also a forage. specific habitat requirements Vicia hirsuta is
Properties The seeds of Vicia hirsuta con- not threatened with genetic erosion.
tain trypsin inhibitors, but heating for 20 min- Prospects Vicia hirsuta is only occasionally
utes at 100°C at pH 2.0 reduces the trypsin used as a pulse. It is unlikely that its impor-
inhibiting activity by 50%. The seeds also con- tance as a food crop will increase in the future.
tain the non-protein amino acid canavanine, a Still, more information would be useful on the
toxic arginine analogue. nutritional quality of the seed and appropriate
Botany Trailing or climbing annual herb up processing methods to eliminate its toxic com-
to 90 cm tall; stem glabrous or thinly hairy. pounds.
Leaves alternate, paripinnate, with 6-20 leaf- Major references Enneking, 1995; Gillett
lets; stipules semisagittate, 2-15 mm x 1.5—2.5 et al., 1971;Hanelt & Institute of Plant Genet-
mm, the upper part entire, the lower deeply ics and Crop Plant Research (Editors), 2001;
divided into 2-3 filiform segments; petiole 0— Thulin, 1983;Zemede Asfaw &Mesfin Tadesse,
5(-10) mm long, rachis usually terminating in 2001.
a branched tendril; petiolules c. 0.5 mm long; Other references Bohra & Sharma, 1981;
leaflets linear or narrowly oblong, 4-20 mm x Holm, Pancho &Herberger, 1979; ILDIS, 2005;
1-3 mm, almost glabrous. Inflorescence an Mutch &Young, 2004; Polhill, 1990; Sharma &
axillary raceme 2—6 cm long, 2—7-flowered; Lavania, 1977; Southon et al., 1994; Thulin,
peduncle 0.5-4 cm long. Flowers bisexual, 1989a.
papilionaceous; pedicel 0.5-2 mm long; calyx 5- Authors M. Brink
lobed, pubescent, with tube 1(—2.5) mm long
and lobes 1.5—2.5 mm long; corolla white, rose
or pale blue, standard obovate, 3-5 mm x 2 VIGNA ACONITIFOLIA (Jacq.) Maréchal
mm, wings and keel slightly shorter; stamens
10, 9 fused and 1free; ovary superior, hairy, 1- Protologue Bull. Jard. Bot. Belg. 39(2): 160
celled, style short, curved, stigma small. Fruit (1969).
an oblong pod 6-10 mm x 3-4 mm, com- Family Papilionaceae (Leguminosae - Papil-
pressed, pilose, dehiscent, (1—)2(—3)-seeded. ionoideae, Fabaceae)
Seeds globose, 2—3 mm in diameter, dark Chromosome number In = 22
brown or mottled pale and dark brown. Seed- Synonyms PhaseolusaconitifoliusJacq.(1768).
ling with hypogeal germination. Vernacular names Moth bean, moth gram,
Vicia comprises about 120 species, mainly in mat bean, dew bean, dew gram (En). Haricot
the temperate regions of the northern hemi- mat, mat, haricot papillon (Fr).
 VlGNA 201

Origin and geographic distribution Moth

bean is native to India, Pakistan and Myanmar
where it grows both wild and cultivated. It is
also grown in other parts of Asia, Africa, the
United States and Cuba. As a pulse it is mostly
grown in India and Thailand; elsewhere it is
mostly a forage, green manure or cover crop. In
tropical Africa it has been recorded from Su-
dan, Eritrea, Somalia, Kenya and Botswana.
Uses The ripe whole or split seeds of moth
bean are eaten cooked or fried. Sprouted and
cooked seeds are preferred as breakfast items
in India whereas fried splits are consumed in
the form of a ready to eat product. The seeds
are sometimes ground into flour, which is
mixed with other flours to make unleavened
bread. The immature pods are sometimes
eaten boiled as a vegetable. In India the pod
walls and residues left after the preparation of
dhal are fed to animals. Moth bean is also
grown for green manure, forage and hay and as
a cover crop. Seeds are used medicinally in
diets to treat fevers; roots are said to be nar-
cotic.
Production and international trade In In-
dia moth bean is grown on 1.5 million ha pro-
ducing annually about 0.4 million t of seed Vigna aconitifolia - 1, flowering branch; 2,
which is traded and consumed within the coun- infructescence; 3, seeds.
try. Worldwide moth bean is grown on about 2 Source: PROSEA
million ha.
Properties Mature, raw moth bean seeds late, c. 12 mm long, peltate; petiole 5-10 cm
contain per 100 g edible portion: water 9.7 g, long, grooved; stipels small; leaflets 5-12 cm
energy 1435 kJ (343 kcal), protein 22.9 g, fat long, deeply divided into 3—5 narrow lobes.
1.6 g, carbohydrate 61.5 g, Ca 150 mg, Mg 381 Inflorescence an axillary, head-like, dense false
mg, P 489 mg, Fe 10.9 mg, Zn 1.9 mg, vitamin raceme; peduncle 5-10 cm long. Flowers bisex-
A 32 IU, thiamin 0.56 mg, riboflavin 0.09 mg, ual, papilionaceous; pedicel 5-8 mm long; calyx
niacin 2.8 mg, vitamin Be 0.37 mg, folate 649 campanulate, c. 2.5 mm long; corolla yellow,
Hg and ascorbic acid 4.0 mg. The essential standard orbicular, up to 8 mm long, wings c. 6
amino-acid composition per 100 g edible por- mm long, keel sickle-shaped, c. 7 mm long;
tion is: tryptophan 147 mg, lysine 1248 mg, stamens 10, 9 united and 1 free; ovary supe-
methionine 220 mg, phenylalanine 1028 mg, rior, sessile, c. 4 mm long, style incurved. Fruit
valine 734 mg, leucine 1541 mg and isoleucine a cylindrical pod 2.5-5 cm x 0.5 cm, brown,
1138 mg. The principal fatty acids are per 100 covered with short stiff hairs, 4—9-seeded.
g edible portion: linoleic acid 485 mg, palmitic Seeds rectangular to cylindrical, 3—5 mm x
acid 313 mg, linolenic acid 265 mg, oleic acid 1.5-2.5 mm, whitish green, yellow to brown,
129 mg and stearic acid 51 mg (USDA, 2005). often mottled with black; hilum white, linear.
The protein has a lower digestibility than that Seedling with epigeal germination.
of mung bean (Vigna radiata (L.) R.Wilczek). Vigna comprises about 80 species and occurs
The digestibility of the starch and protein is throughout the tropics. Vigna aconitifolia be-
considerably improved by processing treat- longs to subgenus Ceratotropis, which also in-
ments such as soaking, removal of the seed cludes Vigna radiata (L.) R.Wilczek (mung
coat, germination and pressure cooking. bean), Vigna umbellata (Thunb.) Ohwi &
Botany Annual, slender, hairy herb with H.Ohashi (rice bean), Vigna mungo (L.) Hepper
short, angular, erect stem up to 40 cm tall and (black gram) and Vigna angularis (Willd.)
many prostrate branches up to 150 cm long. Ohwi & H.Ohashi (adzuki bean). In India nu-
Leaves alternate, 3-foliolate; stipules lanceo- merous landraces and cultivars of moth bean
202 CEREALS AND PULSES

exist. by nematodes, especially Meloidoigyne incog-

For germination of moth bean a temperature of nita. It is parasitized by several Striga species.
25-27 c C is optimal. Vegetative development Bruchids (Callosobruchus spp.) feed on the
starts slowly. Moth bean is predominantly self- seed during storage. Plants are difficult to har-
pollinated and takes 75-90 days after sowing vest with a mower because of the prostrate
to mature. It effectively nodulates with Bra- branches. They are usually cut with a sickle,
dyrhizobium strains from the cowpea cross- left to dry for one week, then threshed and
inoculation group. winnowed. Average seed yields of moth bean
Ecology In India moth bean is the most are only 70-270 kg/ha, although in the United
drought-resistant pulse crop and particularly States and Australia experimental seed yields
cultivated in hot, arid to semi-arid regions. For of up to 2600 kg/ha have been obtained. Yield
optimum production it requires an average of green matter for forage is 37-50 t/ha and of
temperature of 24-32°C, but it withstands day- hay 7.5-10 t/ha.
time temperatures up to 45°C. In India moth Genetic resources and breeding The larg-
bean is grown from sea-level up to an altitude est germplasm collection of moth bean is at the
of 1300 m. Moth bean thrives with a well- National Bureau of Plant Genetic Resources
distributed annual rainfall of 500-750 mm, but (NBPGR), New Delhi, India, where more than
it is also grown successfully in areas with as 1000 accessions are held. Smaller collections
low as 200-300 mm annual rainfall. Even with are available in the United States (USDA
as little as 50-60 mm in 3-4 showers during Southern Regional Plant Introduction Station,
the growing period, some yield can be obtained. Griffin, Georgia, 56 accessions), Kenya (Na-
Moth bean is a quantitative short-day plant, tional Genebank of Kenya, Crop Plant Genetic
but day-neutral types are also known. It grows Resources Centre, Kikuyu, 47 accessions) and
on many soil types but is particularly suitable the Russian Federation (N.I. Vavilov All-
for dry light sandy soils. It does not tolerate Russian Scientific Research Institute of Plant
waterlogging. Some degree of salinity and a Industry, St. Petersburg, 56 accessions). In-
wide pH range (3.5-10) are tolerated. creased efforts in germplasm collection, charac-
Management Moth bean is propagated by terization, evaluation and preservation are
seed; the 1000-seed weight is 10-35 g. It should recommended. Improved moth bean cultivars
be sown on a well-prepared seedbed. Moth have been developed and released in India, e.g.
bean is usually broadcast, at a seed rate of10— 'RMO-40', 'RMO-225', 'RMO-257', 'RMO-435'
20 kg/ha when grown for seed as a sole crop and 'Jwala'. Genetic transformation of moth
and 7-34 kg/ha when grown for forage. When bean has been achieved using particle bom-
sown in rows the seed rate is 2-5 kg/ha for bardment orAgrobacterium-mediated transfer.
pure stands; it is sown in rows 30-90 cm apart Prospects Moth bean is considered to be one
at a depth of 2.5—4 cm. When grown as a rain- of the most drought-tolerant pulse crops, but
fed crop in arid regions best results were ob- its spreading habit, which makes harvesting
tained in India by planting equal amounts of difficult, and the lack of information on its po-
early and late types in alternate rows. Moth tential and on appropriate management prac-
bean is frequently sown towards the end of the tices limits its spread and use. Although re-
rainy season and grown on residual soil mois- corded from various countries, it has not be-
ture. Weed control is important until a full come important in tropical Africa. It could,
canopy has developed. Irrigation and fertilizer however, increase production of food and forage
applications are rare. In India moth bean is in arid and semi-arid regions, and protect the
grown as a sole crop or intercropped with pearl soil against erosion. The ecological limits, op-
millet, sorghum or other cereals, occasionally timal cultivation practices and most appropri-
with pulses. It is grown as a green manure in ate cultivars should be investigated. Priorities
rotation with cotton. The most important dis- for breeding include the development of erect,
eases of moth bean are mung bean yellow mo- early maturing types, resistance to diseases
saic virus (MYMV) transmitted by white fly and high nutritional quality ofthe seed.
(Bemisia tabaci), and root rot and seedling Major references Kay, 1979; Narain, Singh
blight caused by Macrophomina phaseolina, & Kumar, 2000; Negi, Boora & Khetarpaul,
which is soil- and seed-borne. Cultivars resis- 2001; Thulin, 1983;van Oers, 1989a.
tant to yellow mosaic are available; some culti- Other references Bogdan, 1977; Hanelt &
vars are moderately resistant to Macro- Institute of Plant Genetics and Crop Plant Re-
phomina phaseolina. Moth bean is also affected search (Editors), 2001; Kamble et al., 2003;
 VlGNA 203

Kathju et al., 2003; Khatri, 2004; National pressed-hairy on both sides, venation reticu-
Academy of Sciences, 1979; Nimkar, Mandwe late. Inflorescence an axillary false raceme 5 -
& Dudhe, 2005; Rathore, 2001; Thulin, 1993; 30 cm long, 6-12-flowered; peduncle up to 25
USDA, 2005. cm long, rachis 2-7 cm long. Flowers bisexual,
Sources of illustration van Oers, 1989a. papilionaceous; pedicel 2-3 mm long, with
Authors M. Brink &P.C.M. Jansen ovate-oblong bracteoles 3—4mm x 2 mm; calyx
Based on PROSEA 1:Pulses. with tube 3-4 mm long, the 3 lower lobes fal-
cate or narrowly oblong, 3-5 mm long, the up-
per pair fused into a short, bifid lip, sparsely
VlGNAADENANTHA(G.Mey.) Maréchal, pubescent; corolla with almost circular stan-
Mascherpa & Stainier dard, 1—2.5 cm x 2-2.5 cm, rose or white with
green veins and a green basal eye surrounded
Protologue Taxon 27: 202 (1978). by violet-purple inside, wings c. 3 cm long,
Family Papilionaceae (Leguminosae - Papil- white-tinged violet, green and yellow at the
ionoideae, Fabaceae) base, keel c. 5 cm long, with a long beak, spi-
Chromosome number 2n = 22 rally incurved for about 3 turns, white to vio-
Synonyms Phaseolus adenanthus G.Mey. let-blue; stamens 10, 9 fused but upper one
(1818). free; ovary superior, appressed-hairy, style
Vernacular names Wild bean (En). Pois slender, strongly curved. Fruit an oblong pod
marron (Fr). Fava caranguejo (Po). 7-15 cm x 0.5-1.5 cm, slightly curved, flat-
Origin and geographic distribution Vigna tened, glabrous or slightly hairy, 9-15-seeded.
adenantha most probably originated from the Seeds reniform, 5.5-7.5 mm x 4.5-6 mm x2.5—
Neotropics, where it has its greatest variabil- 5 mm, dark reddish brown; hilum central,
ity. It is distributed pantropically and is occa- small, white.
sionally cultivated. In tropical Africa it occurs Vigna comprises about 80 species and occurs
in most countries, but it has not been recorded throughout the tropics. However, studies of the
from Ethiopia, Namibia, Botswana, Zambia, embryological characters indicate that Vigna
Zimbabwe or Mozambique. In the Indian Ocean adenantha is possibly better classified in the
islands it is found in Madagascar, the Sey- genus Phaseolus.
chelles and Réunion. The seed has a large cavity between the cotyle-
Uses The green pods and ripe seeds of Vigna dons which enables it to float, and the distribu-
adenantha are eaten as emergency food. In tion pattern of the species indicates that seeds
Liberia the plant is or has been cultivated for are sometimes dispersed by sea water.
its edible tuberous roots, which are cooked and Ecology Vigna adenantha is found in humid
eaten. The tuberous roots are also eaten in or swampy locations, along the sea shore and
times of food scarcity in India. Cattle in Sudan rivers, and in cultivated and disturbed areas at
browse the plant. In Nigeria a decoction of the low altitudes. Vigna adenantha is a short-day
whole plant is used as a medicine for gonor- plant.
rhoea, and mixed with rice water to treat dia- Management For uniform and faster ger-
betes. With its large pink and white flowers mination, seeds need scarification.
which turn yellow with age, Vigna adenantha Genetic resources and breeding The Centro
may be grown as an ornamental climber. Internacional de Agricultura Tropical (CIAT),
Properties In tropical America Vigna ade- Cali, Colombia holds 143 accessions of Vigna
nantha provides a good forage containing adenantha. In tropical Africa the International
17.4% crude protein and 0.18% P. Institute of Tropical Agriculture (IITA), Ibadan,
Botany Perennial climbing herb up to 4 m Nigeria, holds 18 accessions. Vigna adenantha
long, with tuberous roots; stem twining, gla- is widespread pantropically and is not in dan-
brous or sparsely hairy, rooting at the lower ger of genetic erosion.
nodes. Leaves alternate, 3-foliolate; stipules Prospects Vigna adenantha will remain of
oblong-ovate, 3-6 mm long, base slightly cor- minor importance as an emergency food. More
date, apex acute, conspicuously veined; petiole research is needed to evaluate its potential as
1-14 cm long, rachis 0.5-2 cm long; petiolules food, forage, medicinal and ornamental crop.
3-4 mm long, hairy; leaflets ovate to rhombic, Major references Burkill, 1995; du Puy et
lateral ones slightly asymmetric, (2.5-)5-10(- al., 2002; Faigón Soverna, Galati & Hoc, 2003;
14) cm x (1.5-)2.5-6.5(-8) cm, base rounded or Gillett et al., 1971;Hanelt &Institute of Plant
truncate, apex obtuse to acute, sparsely ap- Genetics and Crop Plant Research (Editors),
204 CEREALS AND PULSES

2001. are available. Major producers of the crop are

Other references CSIR, 1969; Dalziel, China (670,000 ha), Japan (60,000 ha), South
1937; Friedmann, 1994; Hepper, 1958; ILDIS, Korea (25,000 ha) and Taiwan (15,000 ha).
2005; Lai & Pitman, 1987; Maréchal, Japan produces about 100,000 t/year and con-
Mascherpa & Stainier, 1978; Pitman & Singer, sumes about 140,000 t/year; it imports from
1985;Tateishi, 1988;Thulin, 1993. China, Taiwan, the United States, Thailand
Authors M. Brink &P.C.M. Jansen and Canada. Average export from China in the
1990s was 25,000-40,000 t/year. Both the seed
and the seed flour are important trade items in
VIGNA ANGULARIS (Willd.) Ohwi & H.Ohashi oriental markets.
Properties Mature, raw adzuki bean seeds
Protologue Journ. Jap. Bot. 44(1): 29 contain per 100 g edible portion: water 13.4 g,
(1969). energy 1377 kJ (329 kcal), protein 19.9 g, fat
Family Papilionaceae (Leguminosae - Papil- 0.5 g, carbohydrate 62.9 g, dietary fibre 12.7 g,
ionoideae, Fabaceae) Ca 66 mg, Mg 127 mg, P 381 mg, Fe 5.0 mg, Zn
Chromosome number 2n - 22 5.0 mg, vitamin A 17 IU, thiamin 0.46 mg, ri-
Synonyms Phaseolus angularis (Willd.) boflavin 0.22 mg, niacin 2.6 mg, vitamin B6
W.Wight (1909). 0.35 mg, folate 622 |ig and ascorbic acid 0 mg.
Vernacular names Adzuki bean, azuki bean The essential amino-acid composition per 100 g
(En). Haricot adzuki (Fr). Feijâo adzuki (Po). edible portion is: tryptophan 191 mg, lysine
Origin and geographic distribution The 1497 mg, methionine 210 mg, phenylalanine
exact origin of adzuki bean is not known; wild 1052 mg, threonine 674 mg, valine 1023 mg,
types occur in Nepal, south-eastern China, leucine 1668 mg and isoleucine 791 mg. The
Taiwan, Korea and Japan. Cultivation is principal fatty acids are per 100 g edible por-
known since ancient times from northern Ko- tion: linoleic acid 113 mg and oleic acid 50 mg
rea, China and Japan. Adzuki bean has been (USDA, 2005). Adzuki bean seeds have a
introduced to many countries in the world. In sweet, nutty taste.
Africa experimental plantings have been car- Enzyme-resistant fractions of adzuki bean
ried out in DR Congo, Kenya and Angola, but seeds have shown hypocholesterolaemic effects
up-to-date information is lacking. Adzuki bean in rats. Hot water extracts have shown in-vivo
has also been recorded for Madagascar and the hypoglycaemic and antitumour properties. Wa-
Seychelles. ter extracts of the seed coat have shown hepa-
Uses The dried seeds of adzuki bean are toprotective activity.
eaten, either cooked whole or made into flour Botany Annual, usually bushy and erect
for use in soups, cakes, confectionery and ice herb up to 90 cm tall, sometimes climbing or
cream. Adzuki bean is particularly popular in prostrate and rooting at the nodes; taproot40—
China, Taiwan, Korea and Japan ('azuki an'), 50 cm long. Leaves alternate, 3-foliolate; stip-
where the red seeds have a cultural value re- ules small, peltate, often bifid with basal ap-
lated to birth, wedding and death. Immature pendages; stipels lanceolate; leaflets lanceolate
seeds and sprouted seeds are eaten as a vege- to ovate, 5—10 cm x 5-8 cm, acuminate, entire
table. The seeds may be popped like maize to 3-lobed. Inflorescence an axillary false ra-
grain, used as coffee substitute or eaten can- ceme, 2-20-flowered; peduncle long in lower
died. nodes to very short in upper nodes. Flowers
Adzuki bean is also grown for forage, as green papilionaceous, bisexual; pedicel short, bearing
manure and for soil conservation. Flour is also an extrafloral nectary at base; bracteoles
used for shampoos, to make facial creams and longer than calyx; calyx campanulate, with
as ingredient in culture media. In China the short teeth; corolla 15-18 mm long, bright yel-
seeds are used to treat kidney problems, con- low, standard orbicular, wings oblong, keel
stipation, abscesses, certain tumours, threat- turned towards the right, with a horn-shaped
ened miscarriage, retained placenta, non- spur on the left side; stamens 10, 9 fused and 1
secretion of milk and for improvement of blood free; ovary superior, shortly hairy, style
circulation and urination. The leaves are said abruptly bent in upper part, hairy on one side
to lower fever and the sprouts are used to avert near top, stigma lateral, discoid. Fruit a cylin-
threatened abortion caused by injury. drical pod 5-13 cm x 0.5 cm, pendulous,
Production and international trade No sta- slightly constricted between the seeds, nearly
tistics on the world production of adzuki bean glabrous, pale yellow, blackish or brown, 2-14-
 VIGNA 205

for at least 5 years when stored with about

13% moisture content, at 15% relative humid-
ity. Germination requires a soil temperature
above 6-10°C, with 30-34°C being optimal.
Emergence takes 7-20 days. Growth is slow
compared to other pulses. Flowering lasts 30-
40 days and can occur up to 3 times when
planted early in the growing season. Self-
pollination is predominant, but cross-polli-
nation also occurs. The growth duration is (60-)
80-120(-190) days. Nitrogen fixation levels up
to 100 kg N/ha have been observed, the amount
depending on soil moisture content and pH.
Adzuki bean effectively nodulates with Brady-
rhizobium bacteria.
Ecology Adzuki bean performs best in sub-
tropical and warm temperate climates. It re-
quires average temperatures of 15-30°C for
optimal growth. It tolerates high temperatures
but is sensitive to frost. In the tropics it is more
suitable for higher altitudes. Adzuki bean
grows in areas with average annual rainfall of
500-1750 mm. It is a quantitative short-day
plant but day-neutral cultivars exist. Adzuki
bean can be grown on a wide range of soils (pH
5-7.5), provided they are well drained.
Vigna angularis - 1,fruiting branch; 2, flower; M a n a g e m e n t Propagation of adzuki bean is
3, seed. by seed. The 1000-seed weight is 50-200 g.
Source: PROSEA Sowing practices differ greatly but usually seed
is sown 2-3 cm deep, in rows 30-90 cm apart
seeded. Seeds cylindrical with rounded ends, and 10—45 cm within the row; sometimes it is
flattened, 5-7.5 mm x 4-5.5 mm, smooth, wine broadcast. Seed rates vary widely (8-70 kg/ha).
red, occasionally buff, creamish, black or mot- Because of the relatively slow growth of adzuki
tled. Seedling with hypogeal germination; pri- bean, weed control is very important, particu-
mary leaves simple, opposite, cordate. larly between germination and flowering. Fer-
Vigna comprises about 80 species and occurs tilizer application differs widely. An adzuki
throughout the tropics. Vigna angularis be- bean crop yielding 2160 kg/ha was recorded to
longs to subgenus Ceratotropis, which also in- have an uptake per ha of 74 kg N, 18 kg P and
cludes Vigna radiata (L.) R.Wilczek (mung 50 kg K. Irrigation of adzuki bean is not nor-
bean), Vigna umbellata (Thunb.) Ohwi & mally done. In China adzuki bean is often
H.Ohashi (rice bean), Vigna mungo (L.) Hepper intercropped with maize, sorghum and millet.
(black gram) and Vigna aconitifolia (Jacq.) In Japan adzuki bean is grown in rotation with
Maréchal (moth bean). Cultivated plants of many crops (e.g. rice, wheat, sweet potato,
Vigna angularis have been classified as var. yam). The seed may be sown directly in rice
angularis, wild plants as var. nipponensis stubble at a high rate to reduce weed problems.
(Ohwi) Ohwi & H.Ohashi. Wild adzuki bean Numerous fungi and bacteria are known to
has an indeterminate growth habit with thin cause diseases in adzuki bean, including pow-
twining stems, small leaves, short and strongly dery mildew (Erysiphe polygoni, synonym: Ery-
dehiscent black to grey pods and black-mottled siphe betae), brown stem rot (Cephalosporium
seeds. Numerous cultivars have been recorded gregatum, synonym: Phialophora gregata) and
within Vigna angularis, differing in time to bacterial blight (Xanthomonas campestris).
maturity, seed colour and plant habit. Inter- Several insect pests, such as the adzuki pod
mediate types between wild and cultivated worm (Matsumuraeses phaseoli), the Japanese
plants, called weedy types, have been found in butterbur borer (Ostrinia scapulalis pacified)
Japan. and cutworm (Spodoptera litura) attack the
The seeds of adzuki bean retain their viability crop. Bean weevil (Callosobruchus chinensis)
206 CEREALS AND PULSES

attacks the stored seed. In general the pods of VlGNAMUNGO (L.) Hepper
adzuki bean do not shatter readily and the crop
can be harvested with a mower or bean har- Protologue Kew Bull. 11(1): 128 (1956).
vester. Traditionally, plants are cut by hand Family Papilionaceae (Leguminosae - Papil-
and allowed to cure on the ground for several ionoideae, Fabaceae)
days before being stacked into drying piles. Chromosome number In =22
Drying occurs until moisture content of the Synonyms Phaseolus mungo L. (1767).
seed is about 16% and threshing can start. Vernacular names Black gram, urd bean,
Some pods are very thin and in wet conditions urad bean (En). Haricot urd, urd (Fr). Feijào
seed may germinate in the pods. For hay, ad- urida (Po).Mchooko mweusi (Sw).
zuki bean should be cut when the pods are Origin and geographic distribution Black
about half mature. For seed, cutting is done gram was most probably domesticated in India
when all pods are mature. Seed yields up to from its wild ancestral type, which is also
3500 kg/ha are obtained. In an experimental found in Bangladesh, Pakistan and Myanmar.
planting in Kenya seed yields were 500-600 At present black gram cultivation is of major
kg/ha. importance in India only, but it is also grown to
Genetic resources and breeding Large some extent throughout tropical Asia. In Africa
germplasm collections of adzuki bean are held it is grown in Gabon, DR Congo, Kenya,
in China (Institute of Crop Germplasm Re- Uganda, Tanzania, Malawi, Mozambique,
sources (CAAS), Beijing, more than 3700 acces- South Africa, Madagascar and Mauritius. It is
sions) and Japan (Tokachi Agricultural Ex- grown in the United States and Australia
periment Station, Hokkaido-ken, about 2500 mainly as a fodder crop.
accessions). Uses Black gram seeds are eaten as a pulse,
In China, Japan, Korea and Taiwan breeding direct or in various preparations (whole or
has resulted in locally adapted better yielding split, boiled or roasted, ground into flour for
cultivars, e.g. 'Baihong No 1' (China), 'Erimo' cake, bread or porridge). It is with the flour of
(Japan), 'Chungwonpat' (Korea) and 'Kaohsi- black gram that in India the flat biscuits 'pa-
ung No 3' (Taiwan). In Japan alone more than padum' are made. Seed sprouts are also con-
300 cultivars, landraces and breeding lines sumed. Green pods are eaten as a cooked vege-
have been registered. In-vitro adzuki bean table. Small quantities of the pods and foliage
plants are routinely obtained using epicotyls as are used to supplement cattle feed or as forage.
explants. A genetic transformation system for Sometimes black gram is sown as a cover crop
adzuki bean has been established using Agro- and for green manure. The pod walls are fed to
bacterium-mediated transfer. A genetic linkage cattle. Flour from the seed is used as a substi-
map has been constructed using molecular tute for soap; it makes the skin soft and
(RAPD, RFLP) and morphological markers. smooth. In traditional medicine, the seed is
Prospects Adzuki bean is a suitable crop for used for its suppurative, cooling and astringent
the subtropics and the high-altitude tropics. properties, e.g. pounded and applied as a poul-
The potential of adzuki bean as an anti-erosion tice on abscesses.
crop should not be overlooked either. Further Production and international trade In In-
research on its potential in the high-altitude dia, the major producer and consumer, average
regions oftropical Africa is recommended. annual production of black gram seed is about
Major references Kay, 1979; Lumpkin & 1.3 million t from 3 million ha. Thailand pro-
McClary, 1994; Schuster et al., 1998;van Oers, duces annually about 90,000 t which is mainly
1989b; Zong et al., 2003. exported to Japan, where seed sprouts from
Other references Duke, 1981; Han et al., black gram are preferred to those from green
2003; Han et al., 2004; Hanelt & Institute of gram (Vigna radiata (L.) R.Wilczek) because of
Plant Genetics and Crop Plant Research (Edi- their longer shelf life. Annual production in
tors), 2001;Itoh et al., 2004; Itoh, Umekawa & Pakistan is about 28,000 t from 57,000 ha, and
Furuichi, 2005; Kaga et al., 1996a; USDA, in Sri Lanka 6000 t from 8000 ha. Sri Lanka
2005; Yamaguchi, 1992;Yamada et al.,2001. additionally imports 6000 t/year.
Sources of illustration van Oers, 1989b. Properties Black gram seeds contain per
Authors P.CM. Jansen 100 g edible portion: water 8.6 g, energy 1470
Based on PROSEA 1: Pulses. k j (351 kcal), protein 25.1 g, fat 1.8 g, carbohy-
drate 61.0 g, crude fibre 4.4 g, Ca 196 mg, Mg
260 mg, P 575 mg, Fe 6.8 mg, Zn 3.1 mg, vita-
 VlGNA 207

min A 114 IU, thiamin 0.36 mg, riboflavin 0.28 lipsoid, up to 5 mm long, with square ends, and
mg, niacin 1.8 mg, vitamin Ik 0.28 mg, folate raised and concave hilum, usually black or
628 |Xgand ascorbic acid 4.8 mg. The essential mottled, sometimes green. Seedling with
amino-acid composition ofblack gram seeds per epigeal germination.
g nitrogen is: tryptophan 65 mg, lysine 415 mg, Vigna comprises about 80 species and occurs
methionine 91 mg, phenylalanine 365 mg, throughout the tropics. Vigna mungo belongs
threonine 217 mg, valine 351 mg, leucine 518 to subgenus Ceratotropis, which also includes
mg and isoleucine 319 mg (Haytowitz & Mat- Vigna radiata (L.) R.Wilczek (mung bean),
thews, 1986). Black gram seeds have shown Vigna umbellata (Thunb.) Ohwi & H.Ohashi
anti-atherogenic activity in guinea pigs. (rice bean), Vigna angularis (Willd.) Ohwi &
Botany Erect, hairy annual herb up to 100 H.Ohashi (adzuki bean) and Vigna aconitifolia
cm tall, sometimes twining, with a well- (Jacq.) Maréchal (moth bean). There has been
developed taproot; stem diffusely branched confusion on the taxonomie status of Vigna
from the base, furrowed. Leaves alternate, 3- mungo and Vigna radiata; because they are
foliolate; stipules peltate, ovate; petiole 6-20 closely related it was proposed that they be
cm long; stipels falcate; leaflets ovate or rhom- grouped into a single species. However, at pre-
bic-ovate, 4—10cm x 2-7 cm, entire, acuminate. sent they are considered as 2 separate species
Inflorescence an axillary false raceme; pedun- with as major differences: flower colour (bright
cle up to 18 cm long. Flowers bisexual, papil- yellow in Vigna mungo, pale yellow in Vigna
ionaceous, small; bracteoles linear to lanceo- radiata), pocket on the keel (longer in Vigna
late, exceeding the calyx; calyx campanulate; mungo than in Vigna radiata), fruit shape
corolla yellow, standard 12—16mm wide, wings (pods of Vigna mungo are shorter and erect on
about as long as standard, keel spirally coiled the peduncle, in Vigna radiata the pods are
with a terminal horn-like appendage; stamens longer and spreading or pendulous).
10, 9 united and 1 free; ovary superior, style Three taxa are distinguished within Vigna
spirally curved. Fruit a cylindrical pod 4-7 cm mungo:
x 0.5 cm, erect or almost so, with long hairs - var. mungo, with large, black-seeded and
and short hooked beak, 4-10-seeded. Seed el- early-maturing cultivars;
- var. viridis Bose, with greenish dull or glossy
seeds and late-maturing cultivars;
- var. silvestris Lukoki, Maréchal & Otoul, the
wild type; compared to cultivated types it is
smaller, more climbing, more hairy, with
denser inflorescences and small seeds with
prominent raised aril; it is considered the
ancestor ofthe cultivated black gram.
For cultivated types a classification into culti-
vars and cultivar groups would be more appro-
priate.
Germination of black gram normally takes 7—
10 days. Flowering starts 30—60 days after
sowing. Flowers are normally self-pollinating,
with the pollen shedding before the flower
opens. Maturity is reached in 60-140 days af-
ter sowing. Black gram effectively nodulates
with Bradyrhizobium bacteria.
Ecology Black gram is basically a warm
season crop, but in India it is grown in both
summer and winter, up to 1800 m altitude. It
is quite drought resistant but intolerant of
frost and prolonged cloudiness. It is normally
grown in areas with an average temperature of
25-35°C and an annual rainfall of 600-1000
Vigna mungo - 1, part of fruiting branch; 2, mm. In higher rainfall areas it may be grown
flower; 3, seed. in the dry season on residual moisture. Heav-
Source: PROSEA ier, well-drained soils such as black-cotton soils
208 CEREALS AND PULSES

with pH 6-7 are preferred, but black gram is fer.

also grown on lighter soils. P r o s p e c t s It would be worthwhile trying
Management Black gram is propagated by black gram on a much larger scale in tropical
seed. The 1000-seed weight is 15-60 g. It is Africa because of its highly nutritious seeds
sown broadcast or in rows at a depth of 1—1.5 and its wide ecological applicability. Germ-
cm. The seed rate is 10—30 kg/ha, space be- plasm diversity needs to be exploited to obtain
tween rows 25—40 cm, space within the row suitable cultivars for Africa.
10-20 cm. Thorough field preparation is not Major references Arora & Shri S. Mauria,
required; rough tillage suffices. Weeding is 1989; Dikshit et al., 2004; Kay, 1979; Lawn &
done only once or twice until the canopy is es- Ahn, 1985; Souframanien & Gopalakrishna,
tablished. The crop is mainly rainfed and fertil- 2004.
izer application is not common. In the wet sea- Other references CSIR, 1976; Ghafoor et
son in India it is mainly intercropped, with al., 2001;Gillett et a l , 1971;Haytowitz & Mat-
sugar cane, cotton, groundnut, sorghum or thews, 1986;ILDIS, 2005;Maréchal, Mascherpa
pigeon pea as the main crops. In the dry season & Stainier, 1978; Midya et a l , 2005; Purse-
it is often sole cropped on rice fallow. Impor- glove, 1968; Saini &Jaiwal, 2005;Srivastava &
tant diseases of black gram are mung bean Joshi, 1990.
yellow mosaic virus (MYMV), Cercospora leaf Sources of illustration Arora & Shri S.
spot (caused by Cercospora sp.), web blight Mauria, 1989.
{Rhizoctonia solani, synonym: Thanatephorus Authors P.C.M. Jansen
cucumeris) and powdery mildew (Erysiphe po- Based on PROSEA 1: Pulses.
lygoni, synonym: Erysiphe betae). The most
serious pests are white fly and thrips. In stor-
age the seeds are attacked by bruchids (Cal- VlGNARADIATA(L.) R.Wilczek
losobruchus spp.). Black gram must be har-
vested before the pods are fully ripe to prevent Protologue Fl. Congo Beige 6: 386 (1954).
shattering. The plants are cut or uprooted, Family Papilionaceae (Leguminosae - Papil-
stacked to dry for up to 7 days, and threshed by ionoideae, Fabaceae)
beating with sticks or animal trampling. Alter- Chromosome number In = 22
natively, the pods may be handpicked. Yield of Synonyms Phaseolus radiatus L. (1753),
dry seed averages 350-800 kg/ha but it can Phaseolus aureus Roxb. (1832).
reach 1500-2500 kg/ha. In India black gram Vernacular names Mung bean, green gram,
seeds are usually processed into split seeds golden gram (En). Haricot mungo, mungo, am-
(dhal). bérique, haricot doré (Fr). Feijâo mungo verde
Genetic resources and breeding About (Po). Mchooko, mchoroko (Sw).
2100 accessions of black gram are maintained Origin and geographic distribution Mung
by the National Bureau of Plant Genetic Re- bean originated in India or the Indo-Burmese
sources (NBPGR), New Delhi, India, at its region where it has been cultivated for millen-
various research stations. The USDA Southern
Regional Plant Introduction Station, Griffin,
Georgia, United States, holds 300 accessions,
and the Asian Vegetable Research and Devel-
opment Centre (AVRDC) in Taiwan maintains
a collection of 200 accessions. Breeding pro-
grammes for improvement of this pulse aim for
a plant type combining determinate growth
habit with a plant height of 30 cm, early ma-
turity (60-90 days), and suitability for many
different agroclimatic regions. Sources of resis-
tance against most current diseases are avail-
able and several resistant cultivars have been
released. Genetic variability of black gram is
great, allowing development of suitable culti-
vars for most tropical and subtropical climates.
Genetic transformation of black gram has been
achieved using Agrobacterium-mediated trans- Vigna radiata -planted
 VlGNA 209

nia. T h e ancient cultivation of m u n g b e a n in duced in India is for domestic consumption. In

India is supported by fossilized r e m a i n s discov- most p a r t s of Africa w h e r e t h e r e a r e Asian
ered in c e n t r a l India a n d dated 1500-1000 BC. communities, m u n g bean food products a r e sold
M u n g b e a n cultivation spread in early t i m e s to in t h e cities.
most other Asian countries a n d later to Africa, P r o p e r t i e s The composition of m a t u r e m u n g
Australia, t h e Americas a n d t h e West Indies. It b e a n seeds per 100 g edible portion is: w a t e r
h a s not become a major crop outside Asia, al- 9.1 g, energy 1453 k J (347 kcal), protein 23.9 g,
t h o u g h m u n g bean is cultivated in m a n y tropi- fat 1.2 g, c a r b o h y d r a t e 62.6 g, dietary fibre 16.3
cal African countries. In certain a r e a s of g, Ca 132 mg, Mg 189 mg, P 367 mg, Fe 6.7 mg,
Kenya, especially t h e E a s t e r n Province, m u n g Zn 2.7 mg, v i t a m i n A 114 IU, t h i a m i n 0.62 mg,
b e a n is t h e principal cash crop. riboflavin 0.23 mg, niacin 2.3 mg, v i t a m i n B6
U s e s M a t u r e m u n g b e a n seeds or flour e n t e r 0.38 mg, folate 625 (ig a n d ascorbic acid 4.8 mg.
a v a r i e t y of dishes such as soups, porridge, T h e essential amino-acid composition per 100 g
snacks, bread, noodles a n d even ice-cream. In edible portion is: t r y p t o p h a n 260 mg, lysine
K e n y a m u n g b e a n is most commonly consumed 1664 mg, m e t h i o n i n e 286 mg, p h e n y l a l a n i n e
as whole seeds boiled w i t h cereals such as 1443 mg, t h r e o n i n e 782 mg, valine 1237 mg,
maize or sorghum. Boiled whole seeds a r e also leucine 1847 m g a n d isoleucine 1008 m g
fried w i t h m e a t or vegetables a n d e a t e n as a (USDA, 2004). T h e s t a r c h consists of 28.8%
relish w i t h thick maize porridge ('ugali') a n d amylose a n d 71.2% amylopectin. M u n g b e a n
p a n c a k e s ('chapatti'), w h e r e a s consumption of seed is highly digestible a n d low in a n t i n u t r i -
split seeds (dhal) is common a m o n g people of tional factors. It causes less flatulence t h a n t h e
A s i a n descent. In Ethiopia t h e seeds a r e used seed of most other pulses, m a k i n g it suitable
in sauces. In M a l a w i t h e seeds a r e cooked as a for children a n d older people. M u n g b e a n
side dish, mostly after removing t h e seed coat s t a r c h is considered to h a v e a low glycaemic
by grinding. In India a n d P a k i s t a n t h e dried index, i.e. to raise t h e blood s u g a r level slowly
seeds a r e consumed whole or after splitting a n d steadily.
into dhal. Split seeds a r e e a t e n fried a n d salted T h e composition of s p r o u t e d m u n g b e a n seeds
as a snack. T h e seeds m a y also be p a r c h e d a n d p e r 100 g edible portion is: w a t e r 90.4 g, energy
ground into flour after removing t h e seed coat; 126 k J (30 kcal), protein 3.0 g, fat 0.2 g, carbo-
t h e flour is used in various I n d i a n a n d Chinese h y d r a t e 5.9 g, d i e t a r y fibre 1.8 g, Ca 13 mg, Mg
dishes. T h e flour m a y be further processed into 21 mg, P 54 mg, Fe 0.9 mg, Zn 0.4 mg, v i t a m i n
highly valued s t a r c h noodles, bread, biscuits, A 21 IU, t h i a m i n 0.08 mg, riboflavin 0.12 mg,
vegetable cheese a n d extract for t h e soap in- niacin 0.75 mg, v i t a m i n B6 0.09 mg, folate 61
dustry. Sprouted m u n g b e a n seeds a r e e a t e n ug a n d ascorbic acid 13.2 mg. The essential
r a w or cooked as a vegetable; in F r e n c h t h e y amino-acid composition p e r 100 g edible por-
a r e erroneously called 'germes de soja', in Eng- tion is: t r y p t o p h a n 37 mg, lysine 166 mg, me-
lish 'bean sprouts'. thionine 34 mg, p h e n y l a l a n i n e 117 mg,
I m m a t u r e pods a n d young leaves a r e e a t e n as t h r e o n i n e 78 mg, valine 130 mg, leucine 175
a vegetable. P l a n t residues a n d cracked or m g a n d isoleucine 132 m g (USDA, 2004).
w e a t h e r e d seeds a r e fed to livestock. M u n g S p r o u t i n g especially leads to an increased
b e a n is sometimes grown for fodder, green ma- ascorbic acid concentration.
n u r e or as a cover crop. T h e seeds a r e said to M u n g b e a n h a y contains: m o i s t u r e 9.7%, crude
be a t r a d i t i o n a l source of cures for paralysis, protein 9.8%, fat 2.2%, crude fibre 24.0%, ash
r h e u m a t i s m , coughs, fevers a n d liver a i l m e n t s . 7.7%, N-free extract 46.6%, digestible crude
P r o d u c t i o n and international t r a d e Reli- protein 7.4%, total digestible n u t r i e n t s 49.3%.
able production statistics for m u n g b e a n a r e Aqueous e x t r a c t s of m u n g b e a n seed h a v e
difficult to obtain, as its production is often shown in-vivo hypotensive a n d hepatoprotec-
l u m p e d together with t h a t of other Vigna a n d tive effects in r a t s . E x t r a c t s from m u n g b e a n
Phaseolus spp. India is t h e m a i n producer, seeds a n d h u s k s h a v e shown antioxidative ef-
with a n e s t i m a t e d production in t h e l a t e 1990s fects.
of about 1.1 million t. C h i n a produced 891,000 D e s c r i p t i o n A n n u a l , erect to semi-erect,
t (19% of total pulse production in China) from slightly pubescent h e r b up to 1.3 m tall; root
772,000 h a in 2000. No m u n g b e a n production s y s t e m consisting of a well-developed taproot
statistics a r e available for Africa. w i t h deeply placed l a t e r a l roots; stem m u c h
China exported 110,000 t in 1998, 290,000 t in branched, with a tendency to t w i n e a t t h e tips,
1999 a n d 88,000 t in 2000. All m u n g b e a n pro- a n g u l a r , covered with long s p r e a d i n g h a i r s .
210 CEREALS AND PULSES

(7-)10—15(—20)-seeded, somewhat constricted

between the seeds. Seeds 2.5-4 mm x 2.5-3
mm x 2.5-3 mm, globose to ellipsoid or cube-
like, commonly green but sometimes yellow,
olive, brown, purplish brown or black, marbled
or mottled with black patches, glossy or dull;
hilum white, conspicuously flat, c. 1.5 mm x 0.5
mm; seed coat often with ridges, making the
seed rough to the touch. Seedling with epigeal
germination.
Other botanical information Vigna com-
prises about 80 species and occurs throughout
the tropics. Vigna radiata belongs to the sub-
genus Ceratotropis, a relatively homogenous
and morphologically and taxonomically distinct
group, primarily of Asian distribution. Other
cultivated Asiatic Vigna species in this subge-
nus include Vigna aconitifolia (Jacq.) Maréchal
(moth bean), Vigna angularis (Willd.) Ohwi &
Ohashi (adzuki bean), Vigna mungo (L.) Hep-
per (black gram or urd bean), Vigna trilobata
(L.) Verde, (pillipesara) and Vigna umbellata
(Thunb.) Ohwi & Ohashi (rice bean). Hybrids
have been obtained between many of these
species. The species have often been con-
founded, especially Vigna radiata and Vigna
Vigna radiata - 1,part of flowering branch; 2, mungo.
part offruiting branch; 3, seeds. The wild types of mung bean, which are usu-
Source: PROSEA ally smaller in all parts than cultivated types,
are usually classified into 2botanical varieties:
Leaves alternate, 3-foliolate (sometimes 5- - var. sublobata (Roxb.) Verde, occurring in
foliolate), dark green; stipules 5-18 mm x 3-10 India, Sri Lanka, South-East Asia, northern
mm, peltate, ovate, rhomboid or obovate- Australia (Queensland), in tropical Africa
oblong; petiole 5-21 cm long, rachis 1.5-4.5 cm from Ghana to East Africa, southern Africa
long; stipels conspicuous, 5-10 mm long; petio- and Madagascar;
lules 3-6 mm long; leaflets entire or 2—3-lobed, — var. setulosa (Dalzell) Ohwi & Ohashi, with
5-18 cm x 3-15 cm, elliptical, rhomboid or large, almost orbicular stipules and dense
ovate, base broadly cuneate or rounded, apex long hairs on the stem, and occurring in In-
acuminate, glabrous or hairy on both surfaces, dia, China, Japan and Indonesia.
distinctly 3-veined from the base, the lateral The cultivated types of mung bean are grouped
leaflets unequal-sided. Inflorescence an axil- as Vigna radiata var. radiata, although a clas-
lary false raceme up to 20 cm long, 4—15(—30)- sification into cultivar groups would be more
flowered. Flowers bisexual, papilionaceous; appropriate. Two types of mung bean cultivars
pedicel 2-3 mm long; calyx campanulate, tube are usually distinguished, based mainly on
3-4 mm long and glabrous, lobes narrowly tri- seed colour:
angular, 1.5-4 mm long, ciliate, upper pair — golden gram, with yellow seeds, low seed
united into a bifid lobe; corolla yellow or green- yield and pods shattering at maturity; often
ish, standard 11 mm x 16 mm, glabrous, wings grown for forage or green manure;
c. 11 mm x 7 mm, keel c. 10 mm long, often - green gram, with bright green seeds, more
tinged grey or reddish, with a long beak in- prolific, ripening more uniformly, less ten-
curved almost 360°, and with a distinct pocket dency for pods to shatter.
on one side; stamens 10, 9 united and 1 free; Two additional types are recognized in India,
ovary superior, sessile, c. 7 mm long, hairy. one with black seeds and one with brown seeds.
Fruit a linear-cylindrical pod (2.5—)4-9(-15) cm Growth and development The minimum
x 4-9 mm, usually straight, black or tawny temperature for seed germination of mung
brown, with brown short spreading pubescence, bean is about 12°C, the optimum temperature
 VlGNA 211

around 25°C. Seedlings emerge in 3-7 days. more modern cultivars ripening in 60-75 days,
Mung bean is a short-duration crop, flowering maximum yields are obtained at plant densi-
within 30-70 days and maturing within 50- ties of 300,000-400,000 plants/ha. The later-
120 days after sowing. Self-pollination is the maturing traditional cultivars generally need
rule, but up to 5% outcrossing may occur. wider spacing. Recommended spacings for sole-
Flowers are usually pollinated during the cropped mung bean in Kenya are 45 cm be-
night, before they open early in the morning. It tween rows and 15 cm within the row, with a
takes 3-4 weeks from flower opening to mature seed rate of 6-10 kg/ha and a sowing depth of
pod. Flower abscission is prevalent and may 4-5 cm.
reach 90%. Mung bean mostly has a determi- Mung bean can be grown mixed with other
nate growth habit, but because the inflorescen- crops such as sugar cane, maize, sorghum or
ces remain meristematic and may redevelop tree crops in agroforestry systems. Short-
flowers after a period of adverse conditions, it duration mung bean is often relay-cropped to
flowers and fruits over a period of several make use of a short cropping period. In Kenya
weeks. Green leaves, open flowers, green pods mung bean is usually intercropped with maize,
and ripe pods occur simultaneously on the sorghum or millet; it is occasionally grown in
same plant. A large part of the dry matter ac- pure stands or intercropped with other pulses.
cumulated during seed filling may still be par- The usual practice here is to place 1-2 rows of
titioned to vegetative parts and thus rapid se- mung bean between rows of a cereal, or to
nescence does not occur. Mung bean nodulates plant mung bean in the cereal row.
readily with Bradyrhizobium strains from the Management In pure stands, 1—2 weedings
cowpea cross-inoculation group. Because those are necessary during the early stages of
strains are rather common, mung bean shows growth. In Kenya weeding is done using hoes
little response to inoculation. and machetes. Farmers do not normally apply
Ecology Mung bean is a warm-season crop any inorganic fertilizer to a mung bean crop.
and grows mainly within a mean temperature Mung bean uses residues from fertilizer appli-
range of 20-40°C, the optimum being 28-30°C. cations to the main crops in the system, though
It can therefore be grown in summer and au- it responds well to phosphorus. Nutrient re-
tumn in warm temperate and subtropical re- moval per t of seed harvested (dry weight) is
gions and at altitudes below 2000 m in the 40-42 kg N, 3-5 kg P, 12-14 kg K, 1-1.5 kg Ca,
tropics. It is sensitive to frost. Mung bean is 1.5-2 kg S and 1.5-2 kg Mg. The nutrient re-
mostly grown in regions with an average an- moval is much higher when crop residues are
nual rainfall of 600-1000 mm, but it can do removed to be used for fodder.
with less. It withstands drought well, by cur- In its major area of cultivation, the monsoon
tailing the period of flowering and maturation, tropics, mung bean is mainly grown as a rainy
but it is susceptible to waterlogging. High hu- season crop on dryland or as a dry-season crop
midity at maturity causes damage to seeds after the monsoon in rice-based systems on
leading to seed discoloration or sprouting while wetland, making use of residual moisture or
still in the field. Mung bean cultivars differ supplementary irrigation. In some areas where
markedly in photoperiod sensitivity, but most adequate early rains occur, an early-season
genotypes show quantitative short-day re- crop can be grown before the monsoon. In semi-
sponses, flower initiation being delayed by arid regions of Kenya with 600-800 mm rain-
photoperiods longer than 12-13 hours. fall evenly distributed over 2 rainy seasons, 2
Mung bean grows in a wide range of soil types, mung bean crops are grown per year. In the
but prefers well-drained loams or sandy loams Wei Wei Integrated Development Project in
with pH (5-)5.5-7(-8). Some cultivars are tol- Sigor, Kenya, mung bean is grown under irri-
erant to moderate alkaline and saline soils. gation. In India mung bean is often sown as a
Propagation and planting Mung bean is fallow crop on rice land as a green manure.
propagated by seed. The 1000-seed weight is Diseases and p e s t s The most important
15-40 g. There is no seed dormancy, but ger- and widespread fungal diseases of mung bean
mination can be affected by a hard seedcoat. are Cercospora leaf spot (Cercospora canescens)
Mung bean is broadcast or dibbled in hills or in and powdery mildew (Erysiphe polygoni). Less
rows. Recommended sowing rates are 5—30 serious are scab {Elsinoë iwatae), anthracnose
kg/ha for sole cropped mung bean, and 3-4 (Colletotrichum lindemuthianum) and rust
kg/ha under intercropping. Recommended (Uromyces spp.). Important bacterial diseases
spacings are 25-100 cm x 5-30 cm. For the are blights caused by Xanthomonas and Pseu-
212 CEREALS AND PULSES

domonas spp. Mung bean suffers from several tions of mung bean are held in the Philippines
virus diseases but they are not well described, (National Plant Genetic Resources Laboratory,
except for mung bean yellow mosaic virus University of the Philippines Los Banos
(MYMV),which is widespread in South Asia. (UPLB), Los Banos, about 6900 accessions),
The main insect pests are aphids (Aphis fabae, Taiwan (Asian Vegetable Research and Devel-
Aphis craccivora), bean fly (Ophiomyia phase- opment Centre (AVRDC), Shanhua, about 5600
oli), thrips (Megalurothrips sjostedii), pod bor- accessions), United States (Southern Regional
ers (Heliothis spp., Etiella zinckenella, Maruca Plant Introduction Station, Griffin, Georgia,
testulalis) and pod-suckers such as the green about 3900 accessions), India (National Bureau
stink bug (Nezara uiridula). In the drier areas of Plant Genetic Resources, New Delhi, about
of Kenya the apion weevil (Apion soleatum) 3000 accessions) and China (CAAS, Beijing,
may cause heavy losses. Stored mung bean about 3000 accessions).
seed is attacked by bruchids (Callosobruchus In tropical Africa germplasm collections of
spp.). In Africa it is common to use ash made mung bean are held in Kenya (National Gene-
from neem (Azadirachta indica A.Juss.) leaves bank of Kenya, Crop Plant Genetic Resources
or cow dung to protect seeds against storage Centre, KARI, Kikuyu, 330 accessions), Nigeria
pests. Insecticides are seldom used on mung (International Institute ofTropical Agriculture,
bean in tropical Africa. Ibadan, 125 accessions) and Ethiopia (Interna-
Harvesting When grown for the mature tional Livestock Research Institute, Addis
seed, mung bean is usually harvested when the Ababa, 40 accessions).
pods begin to darken. Harvesting is highly la- Breeding In many traditional mung bean
bour intensive as the pods of most local culti- growing regions farmers still grow old land-
vars of mung bean are highly susceptible to races. Many cultivars have been developed
shattering and mature at different times. from those landraces by pure-line selection.
Mung bean is generally harvested in 2-5 hand- The traditional late robust types may be re-
pickings at weekly intervals. In Kenya individ- placed by new types useful for short seasons
ual pods are picked as they mature. Where the and multiple cropping systems with mung bean
crop matures uniformly, the entire plant is occupying the land for short periods between
harvested and sun-dried before threshing. major crops. These new types are short plants
Short-duration cultivars, which ripen more with high harvest index, reduced photoperiodic
uniformly, may be processed as whole plants sensitivity and a relatively uniform maturity.
with small rice threshers. Cultivars differ Many modern cultivars with improved resis-
markedly in harvesting efficiency, depending tance to major diseases and pests have been
on position (above or within canopy) and size of released in the major producing countries.
pods. Sources of resistance have been identified in
Yield Average mung bean yields are low: germplasm of mung bean and related species.
300-700 kg/ha. Under irrigation in Kenya Among the Asiatic Vigna species, black gram
yields are obtained of 1.25 t/ha. Yields over 3 (Vigna mungo) shows most promise for inter-
t/ha have been obtained in trials. specific hybridization with mung bean. AVRDC
Handling after harvest Handpicked pods in Taiwan is working on the development of
are dried in the sun. Shattering can be speeded mung bean tolerant to diseases (leaf spot, pow-
up by beating with a stick or by trampling. dery mildew) and pests (bean fly, bruchids).
Seed is cleaned by screening and winnowing, Information on mung bean breeding in Africa
and dried to a moisture content of 10-12% be- is scanty. In Kenya mung bean improvement
fore storage. Properly dried mung bean seeds work has been carried out by the National Dry-
maintain high viability over a long period. Seed land Farming Research Station, Machakos
stored by small farmers for sowing is often of since the late 1970s. Germplasm was collected
poor quality because ofbruchid damage. locally and was introduced from elsewhere,
To prepare mung bean sprouts, the seeds are mainly from India and AVRDC. Promising
soaked overnight, drained, placed in containers lines were selected and 2 cultivars ('KVR22'
in the dark, sprinkled with warm water every and 'KVR26') have been released. 'KVR22' has
few hours, and kept for 4-5 days at a tempera- a determinate growth habit and golden-yellow
ture of 24°C and a relative air humidity of 6 0 - seed colour; it flowers in 55—60 days and ma-
70%. One kg of mung bean seed produces 6-10 tures uniformly in 80-90 days after germina-
kg sprouts. tion. It has shown high resistance to MYMV,
Genetic resources Large germplasm collec- moderate resistance to powdery mildew and
 VlGNA 213

tolerance to aphids, but it is susceptible to VlGNASUBTERRANEA (L.) Verde.

thrips and apion weevil. 'KVR26' has a deter-
minate growth habit and a green seed colour; it Protologue Kew Bull. 35(3): 474 (1980).
flowers in 40-45 days, and matures fairly uni- Family Papilionaceae (Leguminosae - Papil-
formly in 60-65 days. It is much appreciated ionoideae, Fabaceae)
for its high yield, earliness and large seed size. Chromosome number In = 22
Direct in-vitro plant regeneration in mung Synonyms Glycine subterranea L. (1763),
bean is possible using cultured shoot tips, coty- Voandzeia subterranea (L.) DC. (1825).
ledons and cotyledonary node expiants. Regen- Vernacular names Bambara groundnut,
eration through organogenesis from callus has bambarra groundnut, earth pea, jugo bean
also been reported. Somatic embryogenesis has (En). Voandzou, pois de terre, pois bambara
been induced from mature cotyledons, hypo- (Fr). Mancara de Bijagó, jinguba de Cabambe
cotyl, nodal segments and leaf expiants. (Po). Njugu mawe (Sw).
AVRDC uses molecular markers to select for Origin and geographic distribution The
resistance to diseases and pests and has centre of origin of bambara groundnut is
worked on gene mapping using RFLP and iso- probably north-eastern Nigeria and northern
zyme electrophoresis. Agrobacterium-mediated Cameroon. It is found in the wild from central
genetic transformation of mung bean has been Nigeria eastwards to southern Sudan, and is
achieved. now cultivated throughout tropical Africa, and
Prospects Mung bean is a suitable crop for to a lesser extent in tropical parts of the
tropical Africa, especially the semi-arid re- Americas, Asia and Australia. Its use as a
gions, because of its short crop cycle and nutri- pulse in West Africa was recorded by Arabic
tional quality. Furthermore, it has a niche on travellers in the 14th Century. Its importance
the international market for the production of declined after the introduction of groundnut
bean sprouts. However, it has not yet become from the New World tropics.
important in tropical Africa, which may be due Uses Bambara groundnut is grown primarily
to its low yields, susceptibility to diseases and for its seeds, which are used in many types of
pests, high labour requirement (weeding, har- foods, some of which are an important part of
vesting), lack of suitable cultivars and good the diet and play a role in traditional ceremo-
quality planting material, and lack of informa- nies (e.g. funeral rites) and gift exchanges.
tion on its potential benefits. It therefore de- Mature, dry seeds are boiled and eaten as a
serves increased attention from research and pulse. Dried seeds, either whole or split, are
extension. also mixed with maize or plantains and then
Major references Dana &Karmakar, 1990; boiled. The seeds may be ground into flour,
Kay, 1979; Lawn, 1995; Lawn & Ahn, 1985; sometimes after roasting, to prepare a por-
Mayeux, 1990; Ministry of Agriculture and ridge. They are also added to maize flour to
Rural Development, 2002; Muthoka & Sha- enrich traditional preparations. Sometimes
koor, 1988; Poehlman, 1991; Siemonsma & seeds are soaked in water and ground into a
Arwooth Na Lampang, 1989;Westphal, 1974.
Other references Avenido, Motoda & Hat-
tori, 2001; Burkill, 1995; Chiu & Fung, 1997;
Devi et al., 2004; Dookun, 2001;Duke, 1981;du
Puy et al., 2002; Gillett et al., 1971; Hafeez,
Asad & Malik, 1991; Hanelt & Institute of
Plant Genetics and Crop Plant Research (Edi-
tors), 2001; Huijie et al., 2003; ILDIS, 2005;
Jaiwal et al., 2001; Joshi & Saxena, 2002;
Madar & Stark, 2002; Mugova & Mavunga,
2000; Thulin, 1989a; Tindall, 1983; USDA,
2004; Wu et al.,2001.
Sources of illustration Siemonsma & Ar-
wooth Na Lampang, 1989.
Authors K.K. Mogotsi
Based on PROSEA 1:Pulses.

Vigna subterranea -planted

214 CEREALS AND PULSES

paste which is used to prepare fried or steamed lenic acid and no oleic acid, however, has also
dishes. Immature seeds are often boiled with been recorded. The ratio of saturated to un-
salt and eaten as a snack. They are eaten dur- saturated fatty acids is approximately 1:2. The
ing the 'hungry gap' during the growing sea- oil content is too low for the seed to be used as
son, when stores are empty and crops are not a source of oil. Trypsin inhibition occurs. The
yet ready for harvest. Vegetable milk and fer- seeds contain tannins, mainly in the seed coat.
mented products similar to tempeh (from Gly- In comparative studies in Botswana and
cine max L.) and dawadawa (Parkia biglobosa Ghana, tannin levels were found to be lowest in
(Jacq.) R.Br, ex G.Don) can be made from the cream-coloured seeds, intermediate in red
seeds. seeds and highest in black seeds. Cooking and
The seeds are fed to pigs and poultry, and the other forms of processing (e.g. soaking, milling,
leafy shoots are used as fodder. In Senegal leaf hulling, germination, fermentation) reduce the
preparations are applied to abscesses and in- concentration of antinutritional factors. Ripe
fected wounds, leaf sap is applied to the eyes to seeds are very hard and usually have to be
treat epilepsy, and the roots are sometimes cooked longer than those of other legumes.
taken as an aphrodisiac. Pounded seeds mixed Cream-coloured seeds are often preferred to
with water are administered to treat cataracts. red and black seeds, because they are less bit-
The Igboin Nigeria use the plant to treat vene- ter ('sweeter') and take less time to cook. Large
real diseases. seeds are preferred over smaller ones, e.g. for
Production and international trade Reliable use as snack; smaller seeds are ground into
production figures for bambara groundnut are flour for use in various recipes. Dried leaves for
difficult to obtain, because the crop is mainly fodder contain crude protein 15.9%, crude fibre
grown for home consumption and sale at local 31.7%, ash 7.5%and fat 1.8%.
markets. In the early 1980s the estimated an- Description Annual herb with creeping
nual world production was 330,000 t, 45-50% stems branching just above ground level; root-
of which was produced in West Africa. The
major producers are Burkina Faso, Chad, Côte
d'Ivoire, Ghana, Mali, Niger and Nigeria, but
the crop is also widely grown in eastern and
southern Africa and in Madagascar. The main
exporting countries are Burkina Faso, Chad,
Mali, Niger and Senegal; they supply markets
in Benin, Ghana, Nigeria and Togo.
Properties Raw immature bambara ground-
nut seeds contain per 100 g edible portion: wa-
ter 57.3 g, energy 636 k j (152 kcal), protein 7.8
g, fat 3.1 g, carbohydrate 30.0 g, fibre 3.0 g, ash
1.8 g, Ca 14 mg, P 258 mg and Fe 1.2 mg. The
composition of mature dry seeds per 100 g edi-
ble portion is: water 10.3 g, energy 1537 kJ
(367 kcal), protein 18.8 g, fat 6.2 g, carbohy-
drate 61.3 g, fibre 4.8 g, ash 3.4 g, Ca 62 mg, P
276 mg, Fe 12.2 mg, ß-carotene 10 |lg, thiamin
0.47 mg, riboflavin 0.14 mg, niacin 1.8 mg and
ascorbic acid traces (Leung, Busson & Jardin,
1968). The content of essential amino acids per
100 g food is: tryptophan 192 mg, lysine 1141
mg, methionine 312 mg, phenylalanine 991 mg,
threonine 617 mg, valine 937 mg, leucine 1385
mg and isoleucine 776 mg (FAO, 1970). As in
other pulses, the sulphur-containing amino
acids cystine and methionine are limiting. The
main fatty acids in the seed oil are palmitic
acid 18-24%, stearic acid 5-12%, oleic acid 18- Vigna subterranea - 1,habit offlowering plant;
24%, linoleic acid 34-40%, linolenic acid 2-3% 2, flower; 3,fruits; 4,seed.
and behenic acid 3-7%. A content of 21% lino- Source: PROSEA
 VIGNA 215

system consisting of a tap root with lateral continue after reproductive development has
roots lower down, with rounded and sometimes started. Flowering starts 30-55 days after sow-
lobed nodules. Leaves alternate, 3-foliolate, ing and may continue until the plant dies. Self-
glabrous; stipules c. 3 mm long, spurred, stri- pollination is the rule. After fertilization, the
ate; petiole erect, grooved, up to 30 cm long, peduncle grows and pods form on or below the
thickened at base, rachis (0.1-)1—2.5 cm long; ground. Pods reach their maximum size in
stipels ovate-oblong, up to 3 mm long; petio- about 30 days. The seeds expand and reach
lules 1-3 mm long; leaflets elliptical to maturity during the following 10 days, when
oblanceolate, 3-10 cm x 1-5 cm. Inflorescence the parenchymatous layer surrounding the
an axillary false raceme, close to the ground, embryo has disappeared and brown patches
(l-)2(-3)-flowered; peduncle 0.5-2 cm long. appear on the outside of the pod. Seeds are
Flowers bisexual, papilionaceous, shortly pedi- mature 3-6 months after germination. Bam-
celled; calyx with tube c. 1mm long and 5 lobes bara groundnut is able to fix atmospheric ni-
c. 1 mm long; corolla whitish-yellow, standard trogen by nodulating with bacteria of the Bra-
obovate, 4-7 mm long, wings and keel slightly dyrhizobium group.
shorter; stamens 10, 9 with filaments fused for Ecology Bambara groundnut is cultivated in
more than half their length and 1 free; ovary the tropics at altitudes up to 2000 m. A frost-
superior, 1-celled, style bent. Fruit an almost free period of at least 3 months is necessary.
globose indéhiscent pod c. 2.5 cm in diameter, Average day temperatures of 20—28°C and full
usually 1-seeded. Seed 8.5-15 mm x 6.5-10 sun are preferred. The crop tolerates drought
mm x 5.5-9 mm, variously coloured from white and is cultivated successfully in areas with an
to cream, red, black or brown, sometimes mot- average annual rainfall of 600—750 mm,
tled, blotched or striped; eye around the hilum though optimum yields are obtained when
sometimes present, colour and shape variable. rainfall is higher (900—1200 mm/year). It is
Seedling with hypogeal germination. also grown in humid conditions, e.g. in north-
Other botanical information Vigna com- ern Sierra Leone, where the annual rainfall
prises about 80 species and occurs throughout exceeds 2000 mm. There are considerable dif-
the tropics. However, it is likely that the ferences between genotypes in their response
American species will be placed in a separate to temperature and photoperiod. In many geno-
genus in the near future. There are consider- types, flowering is photoperiod-insensitive,
able morphological difference between wild and while the onset of podding is retarded by long
domesticated types of bambara groundnut. photoperiods. In some genotypes both flowering
Wild bambara groundnut produces long run- and the onset of podding are delayed by long
ners, the pods are thin and do not wrinkle upon photoperiods. Podding may also be delayed by
drying, and the seeds are small (9—11mm long) drought.
and uniform in size. Domesticated types are The plant grows on any well-drained soil, but
more compact, with longer, less slender and light sandy loams with a pH of 5.0—6.5 are
more erect petioles, fleshy pods which wrinkle most suitable. Soils rich in phosphorus and
on drying, and larger seeds (11-15 mm long). potassium are suitable, but calcareous soils are
Morphological and isozyme data indicate a not. Nitrogen-rich soils promote vegetative
gradation from wild to domesticated bambara growth at the expense of seed yield. Sandy soils
groundnut through weedy populations. Wild enhance pod penetration into the soil, but
and domesticated types are sometimes distin- nematode incidence is generally higher on
guished as var. spontanea (Harms) Hepper sandy than on loamy soils.
(wild) and var. subterranea (cultivated). No Propagation and planting Bambara ground-
cultivars of bambara groundnut have been nut is propagated by seed. The seeds are or-
named, but genotypes are distinguished on the thodox and can be stored below 0°C. The 1000-
basis of seed attributes (colour, size, hardness) seed weight is 500-750 g; sowing rates range
and plant form (bushy or spreading). Some- from 25-160 kg/ha, depending on cropping
times names are based on the location where system and climate. Seed to be sown is usually
the seed was collected. retained from the previous harvest or bought
Growth and development The optimum at local markets. Planting material is usually
temperature for germination of bambara selected after harvesting on the basis of seed
groundnut is 30—35°C; below 15°C and above characteristics and not on plant characteris-
40°C, germination is very poor. Emergence tics. Often large seeds are selected for planting.
takes 5-21 days. Vegetative development may Seeds are stored in bags, bottles, gourds or
216 CEREALS AND PULSES

calabashes sometimes sealed with mud. They nut takes place 1—3 times, often with a hoe.
should be shelled just before sowing to retain Earthing up to cover the young pods is com-
maximum viability, but otherwise are rarely mon, and may be done by hand, with a hoe or
pretreated. with ox-drawn equipment. Earthing up im-
Bambara groundnut is not usually sown im- proves yields, but is labour intensive; it is often
mediately after the first rains, because staple combined with weeding. Nitrogen needs may
food and cash crops tend to receive priority. be met by symbiotic nitrogen fixation. Nitro-
Sowing dates vary considerably within loca- gen-fixation rates of up to 100 kg/ha have been
tions. In Zambia and Botswana, for example, reported, but sufficient phosphorus availability
sowing takes place from November to Febru- is essential for nodulation. The use of animal
ary. Late sowing, however, may result in large manure or chemical fertilizers is not common.
yield reductions. Sometimes phased planting Research in Botswana has shown that under
occurs, e.g. in Sukumaland, Tanzania. Land is the prevailing conditions nitrogen fertilization
cleared, and may be ploughed and ridged be- is not advisable, whereas phosphorus applica-
fore sowing. In Botswana, fields are sometimes tion is only beneficial when it is done close to
ploughed after the seed has been broadcast. the seedlings within 2 weeks of sowing and
The crop performs best on deeply ploughed when the soil during this period is moist.
fields with a fine seedbed, eventually allowing Bambara groundnut is used in rotations, e.g.
the plant to bury its developing fruits. Ridging with maize, sorghum, pearl millet, cassava and
is advisable if the soil is shallow or prone to yam. Farmers in Swaziland and in parts of
waterlogging. Bambara groundnut may be South Africa prefer to sow bambara groundnut
sown on mounds, e.g. in Ghana. When sowing a immediately after fallow, to maximize yields.
new field, inoculation with soil from an old Diseases and pests Although bambara
bambara groundnut field is recommended to groundnut is considered to be generally less
promote nodulation with rhizobial bacteria. affected by diseases and pests than groundnut
Bambara groundnut is sown in rows or broad- or cowpea, several diseases and pests can cause
cast; densities range from 2,500 plants/ha serious damage to the crop. The most impor-
(intercropping in Botswana) to 250,000 tant fungal diseases are Cercospora leaf spot
plants/ha (sole cropping in Nigeria). Rows can (Cercospora spp.), powdery mildew (Erysiphe
be 20-40 cm apart (Nigeria) to 50-100(-400) polygoni) and Fusarium wilt {Fusarium oxy-
cm apart (Botswana). Spacings can be 20-40 sporum). Symptoms of Cercospora leaf spot are
cm within rows (Nigeria) to 10—100 cm (Bot- reddish-brown circular spots on the leaves, as
swana). Dry matter production of bambara well as lesions on the stems, petioles, pedun-
groundnut is low, so high plant densities are cles and pods. The lesions may coalesce to give
recommended. However, high densities are the appearance of blight. In cases of severe
only possible where rainfall and soil fertility attack, defoliation occurs and plants may die
are adequate. Furthermore, close spacing prematurely. Crop rotation and burning of crop
makes earthing up difficult. Seed is often dib- debris of the previous season are recommended
bled, dropping 1-4 seeds in the hole and cover- to reduce damage, but the best solution is to
ing with soil. Sometimes a planter is used, or use more resistant types. Symptoms of pow-
the seed is sown immediately behind a plough. dery mildew are a whitish powder on both
Under rainfed conditions in sandy soils a sow- sides of the leaves, especially on the upper sur-
ing depth of at least 6 cm is advisable, but face. Infected leaves dry out and die. Treat-
farmers often sow less deep. Thinning may be ment with a chlorothalonil-based fungicide has
practised, often in combination with weeding. sometimes been effective. Fusarium wilt causes
When establishment problems occur, gaps are vascular discolouration, yellowing, necrosis
sometimes filled in with seeds or plants and wilting and plants become stunted and
thinned out elsewhere. Bambara groundnut eventually die. Crop rotation may help, but
may be grown in intercropping with cereals, planting more resistant types is the best con-
other pulses, root and tuber crops, or vegeta- trol. Other fungal pathogens affecting bambara
bles. It is often grown together with maize, groundnut include Macrophomina phaseolina
sorghum, pearl millet, groundnut and cowpea. (charcoal rot), Phomopsis sp.(blight), Phyllo-
Bambara groundnut is mainly grown by small- sticta spp. (Phyllosticta leaf spot) and Scle-
holders, often women, usually on small fields rotium rolfsii (southern blight and pod rot).
(less than 0.5 ha). Virus diseases include cowpea mottle virus
Management Weeding of bambara ground- (CPMoV), cowpea aphid-borne mosaic virus
 VIGNA 217

(CABMV) and peanut mottle virus (PeMoV). ginal for groundnut.

Genotypes resistant to cowpea mottle virus Handling after harvest The pods of bam-
have been identified. Root-knot nematodes bara groundnut are sun-dried to a moisture
(Meloidogyne incognita, Meloidogyne javanica) content of 12% and stored in bags or drums in
can seriously affect yields. granaries or in the house. They may be shelled
Pests of germinating seeds include rodents, first with mortar and pestle, flails or modified
termites, ants and cutworms (Agrotis). The groundnut shellers. The shelling percentage
standing crop may be attacked by insect pests ranges from 70-77% by pod weight. Bambara
such as aphids, groundnut jassid (Empoasca groundnut is a typical dual-purpose crop: usu-
facialis), groundnut hopper (Hilda patruelis), ally part of the harvest is sold and the rest is
brown leaf beetle (Ootheca mutabilis), and kept for own consumption. Canning of bambara
bean leaf webber (Hedylepta indicata, syno- groundnut seeds has been done in Ghana and
nym: Lamprosema indicata). A serious pest in Zimbabwe.
Swaziland is the American bollworm (Heli- Genetic resources The largest germplasm
coverpa armigera). Control measures of insect collection of bambara groundnut (2000 acces-
pests include the use of insecticides, e.g. sions from sub-Saharan Africa) is held by UTA
malathion against aphids. Leaves may also be (International Institute of Tropical Agricul-
eaten by mammals, such as duikers. Maturing ture), Ibadan, Nigeria. Most of the accessions
seeds may be attacked by rodents, ants, wild (1400) in this collection have been character-
pigs, monkeys and bush babies (Galago spp.). ized, evaluated and documented. Other large
Important storage pests are the bruchid beetles collections are found at the IRD (Institut de
Callosobruchus maculatus and Callosobruchus Recherche pour le Développement), Montpel-
subinnotatus, and the maize weevil Sitophilus lier, France (about 1200 cultivated and 60 wild
zeamais. Infestation often begins in seeds rip- accessions from Cameroon, of which 50 were
ening in the field and is later carried into the morphologically characterized), the University
stores. Seeds stored in the pod shell suffer less of Zambia, Lusaka, Zambia (460 accessions),
from deterioration and infestation by insects the Grain Crops Institute, Potchefstroom,
than shelled seeds. Stored seeds are sometimes South Africa (200 accessions) and the Plant
protected by applying ash, chemical products Genetic Resources Centre, Accra, Ghana (170
(malathion, carbamyl) or plant products, such accessions). In many African countries smaller
as ground tobacco leaves, ground peppers or collections are maintained.
the leaves of basil (Ocimum basilicum L.). The In studies of genetic diversity in cultivated
parasitic plants Alectra vogelii Benth. and bambara groundnut with RAPD and AFLP
Striga gesnerioides (Willd.) Vatke may reduce markers, considerable genetic variation was
yields considerably. found, with accessions clustering mainly ac-
Harvesting Bambara groundnut is har- cording to their geographical origin. Some-
vested 90-180 days after sowing, depending on times, e.g. in Swaziland, farmers sow a mixture
genotype, ecological conditions and farmers' of landraces as a buffer to biotic and abiotic
objectives. As the seeds may be consumed ei- stresses, thus helping to maintain the diversity
ther unripe or ripe, different harvesting meth- of the crop.
ods exist. Unripe seeds may be harvested in Breeding Bambara groundnut breeding has
several rounds from the same plants. Mature mainly been confined to selection between and
seeds are harvested when the leaves turn yel- within populations for yield, disease resistance
low and fall, and when the pods have become (Fusarium wilt and Cercospora leaf spot) and
hard. In the latter case, harvesting is usually drought tolerance. From the UTA germplasm
done by uprooting the plants by hand or with a collection genotypes have been identified with
hoe. The leaves are left in the field or fed to a longer and denser root system, which may be
animals. useful in breeding for drought tolerance.
Yield Yield fluctuations between years are Breeding of genotypes with a shorter growth
large in bambara groundnut and mainly de- period also seems useful for drier regions. Se-
pend on rainfall. The highest recorded seed lection of the most effective combinations of
yield under field conditions is 4 t/ha. Average genotypes and rhizobial strains seems promis-
yields are 300-800 kg/ha, but yields of less ing to improve nitrogen fixation and increase
than 100 kg/ha are not uncommon. Bambara crop yields.
groundnut still gives some yield under condi- Artificial hybrids between cultivated genotypes
tions (poor soils, drought) which are submar- and between cultivated and wild accessions
218 CEREALS AND PULSES

have been made in the United Kingdom and VlGNAUMBELLATA (Thunb.) Ohwi & H.Ohashi
Swaziland, but success rates are generally low.
A genetic linkage map of bambara groundnut Protologue Journ. Jap. Bot. 44(1): 31
using AFLP markers is being developed in the (1969).
United Kingdom as well. Micropropagation of Family Papilionaceae (Leguminosae - Papil-
bambara groundnut is possible using stem ionoideae, Fabaceae)
nodal cuttings or embryo axes. Chromosome number 2n = 22
Prospects Bambara groundnut is a suitable Synonyms Phaseolus calcaratus Roxb.
crop for semi-arid regions, because it tolerates (1832), Vigna calcarata (Roxb.) Kurz (1876).
drought and poor soil conditions and appears to Vernacular names Rice bean, red bean,
be less affected by diseases and pests than climbing mountain bean (En). Haricot riz (Fr).
cowpea or groundnut. Farmers also value its Feijào arroz (Po).
multiple uses and good taste. Although bam- Origin and geographic distribution Rice
bara groundnut will remain an important sec- bean originates from Asia, where it is found
ondary food crop in Africa, the area under cul- wild from India and central China through
tivation will probably decline, because of high Indo-China to Malaysia. It was introduced by
labour requirements, especially for earthing up the Arabs into Egypt, along the eastern coast
and harvesting, the absence of an export mar- of Africa and to the Indian Ocean islands.
ket outside Africa, and the competition from Nowadays, rice bean is widely cultivated in
groundnut and cowpea. The prospects of bam- tropical Asia, and to a more limited extent
bara groundnut as a food crop can be improved in Fiji, the United States, Australia, south-
by developing high-yielding cultivars with im- western Asia and tropical Africa and America.
proved disease resistance and lower anti- In tropical Africa it is grown in West Africa,
nutritional factors. The development of new East Africa and the Indian Ocean islands, and
food product composites with cereals may also less frequently in Central and southern Af-
lead to increased use ofthe crop. rica.
Major references Anchirinah, Yiridoe & Uses Mature, dry rice bean seeds are eaten
Bennett-Lartey, 2001; Brink, 1998; Brink, Col- as a pulse. They are usually boiled and eaten
linson & Wigglesworth, 1997; Heller, Bege- with rice or instead of rice, often in soups or
mann & Mushonga (Editors), 1997; Linne- stews, e.g. in Ghana. In Madagascar flour of
mann, 1989; Linnemann, 1994; Linnemann & dried germinated rice bean seeds is included in
Azam-Ali, 1993;Massawe et al., 2003; Pasquet, complementary foods for children. Rice bean is
Schwedes & Gepts, 1999; Sesay, Saboleh & not very popular in India, because it cannot
Yarmah, 1997. easily be processed into dhal as it contains a
Other references Allen & Lenné, 1998; fibrous mucilage, which prevents easy hulling
Amarteifio, Karikari & Moichubedi, 1998; and separation ofthe cotyledons.
Azam-Ali (Editor), 2003; Azam-Ali et a l , 2001; The leaves, young pods and sprouted seeds of
Burkill, 1995;Collinson et a l , 1997;Dijkstra et rice bean are eaten boiled as a vegetable. In
al., 1995;Doku &Karikari, 1971;du Puy et al.,
2002; FAO, 1970; Gillett et al., 1971; Goli,
1997; Kannaiyan & Haciwa, 1993; Lacroix,
Assoumou & Sangwan, 2003; Leung, Busson &
Jardin, 1968; Linnemann, 1988; Linnemann,
1990; Massawe, Azam Ali & Roberts, 2003;
Ofori, Kumaga & Bimi, 2001; Ramolemana,
1999.
Sources of illustration Linnemann, 1989.
Authors M. Brink, G.M. Ramolemana &
K.P. Sibuga

Vigna umbellata - planted

 VlGNA 219

India the young pods are sometimes eaten raw.

The whole plant is used as fodder and made
into hay and silage. The seeds are sometimes
used as feed for livestock. Rice bean is also
sown as a cover crop, green manure and living
hedge.
Production and international trade Rice
bean production statistics are not available,
but the crop is mainly produced in tropical
Asia. Little of the rice bean production enters
international trade. Japan is the main im-
porter; the main exporters are Thailand,
Myanmar and China. Madagascar also exports
some rice bean; its average annual export in
1998-2000 has been estimated at 1100 t.
Properties The composition of dried rice
bean seeds per 100 g edible portion is: water
13.3 g, energy 1369 kJ (327 kcal), protein 20.9
g, fat 0.9 g, carbohydrate 60.7 g, fibre 4.8 g, Ca
200 mg, P 390 mg, Fe 10.9 mg, thiamin 0.49
mg, riboflavin 0.21 mg and niacin 2.4 mg
(Leung, Busson & Jardin, 1968). The Ca con-
tent is high compared with other pulse crops,
and rice bean is considered good for lactating
mothers. Antinutritional factors in rice bean
include trypsin-inhibiting factors, phytates, Vigna umbellata 1, flowering and fruiting
tannins and oligosaccharides. Soaking, sprout- branch; 2, seeds.
ing, hulling and cooking can reduce these Source: PROSEA
antinutritional factors considerably. Rice bean
seeds do not contain cyanogenic glycosides. c. 0.5 cm long; leaflets broadly ovate to ovate-
Protein concentrates prepared from rice bean lanceolate, 5-10(-13) cm x 1.5-6(-7) cm, entire
seeds have shown in vivo anti-hyper- or 2-3-lobed, the lateral leaflets unequal-sided,
cholesterolaemic effects in hamsters. A peptide membranous, almost glabrous. Inflorescence
isolated from rice bean seeds has shown strong an erect axillary false raceme, 3-10 cm long, 5 -
antifungal activity against Botrytis cinerea, 20-flowered, with flowers usually in pairs; pe-
Fusarium oxysporum, Rhizoctonia solani and duncle up to 20 cm long. Flowers bisexual,
Mycosphaerella arachidicola. Furthermore, it papilionaceous; pedicel c. 5 mm long; calyx
has shown mitogenic and anti-HrV-1 reverse campanulate, c. 4 mm long, 5-toothed; corolla
transcriptase activity. bright yellow, standard 1.5-2 cm in diameter;
In the vegetative stage the rice bean plant wings large and broad, enclosing the keel; keel
(moisture content 84%) contains on a dry mat- with a curved beak and a conical pocket on one
ter basis: 18.0% crude protein, 1.1% fat, 31.5% side; stamens 10, 9 connate into a tube, upper
crude fibre, 39.9% N-free extract, 9.5% ash, one free; ovary superior, 1-celled, style broad-
1.4% Ca and 0.35% P. In the flowering stage ened and curved. Fruit a linear-cylindrical pod,
(moisture content 76%) it contains on a dry 6-13 cm x 0.3-0.6 cm, deflexed, glabrous,
matter base: 14.5% crude protein, 1.0% fat, green when young, black-brown at maturity,6—
32.1% crude fibre, 41.6% N-free extract, 10.8% 8(-16)-seeded. Seeds oblong, 5-10 mm x 2-5
ash, 1.2% Ca and 0.4% P. Its vigorous vegeta- mm x 3-4 mm, smooth, yellow, green, dark
tive growth makes rice bean suitable for use as red, brown, black, speckled or mottled; hilum
a cover crop and green manure. excentric, elongate, hidden by a cream-coloured
Description Annual climbing herb, with rim. Seedling with hypogeal germination.
stems up to 3 m long; stem grooved, usually Other botanical information Vigna com-
clothed with fine, deciduous, deflexed hairs; prises about 80 species and occurs throughout
taproot up to 1.5 m long. Leaves alternate, the tropics. However, the tropical American
trifoliolate; stipules lanceolate, c. 1.5 cm long; species are likely to be placed in a separate
petiole 5-10 cm long; stipels linear-lanceolate, genus in the near future, which would reduce
220 CEREALS AND PULSES

the genus to 50-60 species. Vigna umbellata ies widely, from 30-120(-230) g. Rice bean is
forms part of the subgenus Ceratotropis, which usually broadcast, after 2-3 ploughings, at a
also includes Vigna radiata (L.) R.Wilczek seed rate of 70-90 kg/ha. It is also sown in
(mung bean), Vigna mungo (L.) Hepper (black rows 30-90 cm apart, at a seed rate of 20—70
gram), Vigna angularis (Willd.) Ohwi & kg/ha. A normal seed rate in India is 40-50
H.Ohashi (adzuki bean) and Vigna aconitifolia kg/ha if grown for seed or 60-70 kg/ha if grown
(Jacq.) Maréchal (moth bean). Vigna umbellata as a catch crop for fodder. In India rice bean is
is closely related to Vigna angularis, with normally a 'kharif crop, sown in June—July
which it can be crossed using embryo culture and harvested in October—November. In Asia
and with rice bean as the female parent. Vigna rice bean is mostly grown as an intercrop, es-
minima (Roxb.) Ohwi & H.Ohashi, a wild spe- pecially of maize.
cies from tropical Asia, is even more closely Management Rice bean usually receives
related to Vigna umbellata. little care. Seedlings are able to smother weeds
Within Vigna umbellata 2 types have been due to their vigorous growth. Fertilizers are
distinguished, commonly designated as varie- seldom applied to a rice bean crop, although in
ties: India superphosphate at a rate of 17-26 kg P
- var. gracilis (Prain) Maréchal, Mascherpa & per ha is recommended. In Asia rice bean was
Stainier, the wild type with slender branch- formerly widely planted after the harvest of the
lets, narrow leaflets and long peduncles, traditional long-season rice crop (hence the
found from India to Malaysia, the Philip- name 'rice bean'), but multiple cropping of
pines and central China. This taxon closely short-duration rice cultivars has led to a de-
resembles Vigna minima, or should even be cline of this practice.
united with it. Diseases and pests Rice bean is seldom
— var. umbellata, the cultivated types. seriously affected by diseases and pests, but it
Cultivars have mainly been identified on the is susceptible to root-knot nematodes (Meloi-
basis of maturity period and seed colour. In dogyne spp.). Rice bean seeds are considered
Madagascar 2 types are distinguished: yellow resistant to storage pests such as the bruchids
rice bean ('tsiasisa mavo') and red rice bean Callosobruchus analis, Callosobruchus chinen-
('tsiasisa mena'). sis and Callosobruchus maculatus, due to the
Growth and development Rice bean seed- presence of compounds in the cotyledons with
lings grow vigorously and establish themselves growth-inhibiting effects on them.
quickly. In Madagascar time to flowering is70— Harvesting The viny habit and the shatter-
75 days, time to seed maturity 100-135 days. ing of pods make rice bean difficult to harvest.
In Angola the time from sowing to maturity Harvesting of mature seeds, green pods and
can be as short as 60 days. In the Philippines leaves is usually by hand. If grown for fodder,
time to flowering averages 64 days, to maturity rice bean should be harvested when the pods
92 days; in India early-maturing types behave are immature, since the leaves drop easily
likewise, whereas late types ripen in 130-150 when the plant reaches maturity. As a green
days. Flowers are self-compatible, but cross- manure, rice bean can already be ploughed at
pollination also occurs. about 30 days after sowing.
Ecology Rice bean is typically suited to hu- Yield The average seed yield of rice bean is
mid tropical lowlands, but some cultivars are only 200-300 kg/ha, but the low yields are re-
adapted to subtropical or temperate conditions. lated to the often very short crop cycle. Ex-
It is found in areas with average temperatures perimental yields up to 2500 kg/ha have been
of 18-30°C. It is susceptible to frost. In the obtained in India. Fresh fodder yields of 35 t/ha
tropics the crop can be grown up to 2000 m have been obtained.
altitude. In Madagascar it is grown up to 1000 Handling after harvest Rice bean seeds
m altitude. Rice bean prefers a rainfall of are normally dried in the sun and threshed by
1000-1500 mm/year; it tolerates moderate hand. The seed stores well.
drought. It is a quantitative short-day plant. Genetic resources Large germplasm collec-
Rice bean can be grown on a wide range of soil tions of rice bean are kept in China (Institute
types, but grows best on fertile loams. The op- of Crop Germplasm Resources (CAAS), Beijing;
timum pH is 6.8—7.5. Wild rice bean types are 1363 accessions) and India (National Bureau of
found in open locations and on roadsides. Plant Genetic Resources (NBPGR), New Delhi;
Propagation and planting Rice bean is 902 accessions). No germplasm collections of
propagated by seed. The 1000-seed weight var- rice bean are known in Africa. Rice bean is not
 VlGNA 221

threatened with genetic erosion, but more VlGNAUNGUICULATA(L.) Walp.

germplasm collection and characterization
work are required. Protologue Repert. bot. syst. 1: 779 (1843).
Breeding Breeding programmes of rice bean Family Papilionaceae (Leguminosae - Papil-
have been established mainly in India, where ionoideae, Fabaceae)
several improved cultivars have been devel- Chromosome number In =22
oped and released. In tropical Africa no rice Synonyms Vigna sinensis (L.) Hassk.
bean breeding programmes are known to exist. (1844).
Prospects Rice bean is a valuable crop that Vernacular names
deserves more testing throughout the tropics, - Cowpea, black-eye bean, black-eye pea,
because of its tolerance of high temperature China pea, marble pea (En). Niébé, haricot à
and humidity, short crop cycle, resistance to l'œil noir, pois yeux noirs, cornille, voème,
diseases and pests, nutritious seeds and multi- haricot dolique, dolique mongette (Fr).
ple uses. Limitations of rice bean production Caupi, feijâo frade, feijâo da China, feijâo
are low yields and easy shattering of the pods, miûdo, feijâo macundi, makunde (Po).
which make harvesting difficult. Limited Mkunde (Sw).
availability of germplasm and the lack of tech- - Yard-long bean, asparagus bean (En). Hari-
nical information on its cultivation are serious cot-kilomètre, dolique asperge (Fr). Feijâo de
bottlenecks. The possibilities of industrial metro, feijâo chicote, feijâo espargo, feijâo
processing of the seeds into derived products, frade alfange (Po).
such as flour, are still poorly known. Research - Catjang cowpea, Bombay cowpea (En). Cat-
priorities for rice bean include the development jang (Fr).
of quick-maturing, day-neutral, high-yielding Origin and geographic distribution Vigna
and non-shattering, erect cultivars that are unguiculata originated in Africa, where a large
nematode-resistant. Moreover, investigations genetic diversity of wild types occurs through-
are needed on agronomical aspects (e.g. time of out the continent, southern Africa being rich-
sowing, plant density and fertilizer require- est. It has been introduced in Madagascar and
ments) and post-harvest technology. other Indian Ocean islands, where it is some-
Major references Arora et al., 1980; times found as an escape from cultivation. The
Burkill, 1995; du Puy et al., 2002; Goei, Raina greatest genetic diversity of cultivated cowpea
& Ogihara, 2002; Kashiwaba et al., 2003; Kay, is found in West Africa, in the savanna region
1979; Lawn, 1995; Maréchal, Mascherpa & of Burkina Faso, Ghana, Togo, Benin, Niger,
Stainier, 1978; National Academy of Sciences, Nigeria and Cameroon. Cowpea was probably
1979; van Oers, 1989c. brought to Europe around 300 BC and to India
Other references Chau, Cheung & Wong, 200 BC. As a result of human selection in
1998; CSIR, 1976; Das & Dana, 1987; Ellis et China, India and South-East Asia, cowpea un-
al., 1994; FAO, 1989; FAO, 1998; Gopinathan, derwent further diversification to produce two
Babu & Shivanna, 1986; Hanelt & Institute of cultivar-groups, Sesquipedalis Group with long
Plant Genetics and Crop Plant Research (Edi-
tors), 2001; ILDIS, 2002; Kaga et a l , 1996b;
Khanda, Mohapatra & Misra, 2001; Leung,
Busson & Jardin, 1968; Polhill, 1990; Rabena-
rivo, 1992; Ralison et al., 2004; Saharan,
Khetarpaul & Bishnoi, 2002; Saikia, Sarkar &
Borua, 1999; Schuster et al., 1998; Tindall,
1983;Ye &Ng, 2002.
Sources of illustration van Oers, 1989c.
Authors R. Rajerison
Based on PROSEA 1:Pulses.

_ ,

Vigna unguiculata - wild and planted

222 CEREALS AND PULSES

pods used as a vegetable, and Biflora Group Cowpea is used as fodder in West Africa, Asia
grown for the pods, dry seeds and for fodder. (especially India) and Australia; it is used for
Cowpea was probably introduced to tropical grazing or cut and mixed with dry cereals for
America in the 17thcentury by the Spanish and animal feed. In the United States and else-
is widely grown in the United States, the Car- where cowpea is grown as a green manure and
ibbean region and Brazil. cover crop. In Nigeria special cultivars are
Cowpea is the most important pulse crop in the grown for the fibre extracted from the peduncle
savanna regions of West and Central Africa, after retting; the strong fibre is especially suit-
where it is also an important vegetable and a able for fishing gear, and produces a good-
valuable source of fodder. In East and southern quality paper. The dry seeds have been used as
Africa it is also important both as a vegetable coffee substitute.
and a pulse. Only in humid Central Africa is it Various medicinal uses of cowpea have been
less prominent. reported: leaves and seeds are applied as a
Uses Cowpea is the preferred pulse in large poultice to treat swellings and skin infections,
parts of Africa. The mature seeds are cooked leaves are chewed to treat tooth ailments,
and eaten alone or together with vegetables, powdered carbonized seeds are applied on in-
spices and often palm oil, to produce a thick sect stings, the root is used as an antidote for
bean soup, which accompanies the staple food snakebites and to treat epilepsy, chest pain,
(cassava, yam, plantain). In West Africa the constipation and dysmenorrhoea, and unspeci-
seeds are decorticated and ground into a flour fied plant parts are used as a sedative in
and mixed with chopped onions and spices and tachycardia and against various pains.
made into cakes which are either deep fried Production and international trade Accord-
('akara balls'), or steamed ('moin moin'). In ing to FAO statistics the total annual world
Malawi the seeds are boiled with their seed production of dry cowpea seeds in 1999-2003
coat, or the latter is removed by soaking and was about 3.6 million t from 9.5 million ha.
leaving the seeds in the soil for a few hours. Other estimates indicate a higher production:
Small quantities of cowpea flour are processed over 4.5 million t from about 14 million ha.
into crackers, composite flour and baby foods in According to FAO 3.3 million t was produced
Senegal, Ghana and Benin. annually in sub-Saharan Africa, from 9.3 mil-
The leaves and the immature seeds and pods of lion ha, mainly in West Africa (3 million t/year
cowpea are eaten as vegetables. Cowpea leaves from 8.8 million ha), the main producers being
are served boiled or fried and are usually eaten Nigeria (2.2 million t/year from 5.1 million ha)
with a porridge. The leaf may be preserved by and Niger (400,000 t/year from 3.3 million ha).
sun-drying or boiling and then sun-drying to be Brazil, which is not included in the FAO cow-
used during the dry season. Leaves to be pre- pea statistics, is estimated to produce about
served for later use are generally plucked to- 0.6-0.7 million t/year from 1.1-1.9 million ha.
wards the end of the season. It is believed that Cowpea seeds are produced for local consump-
leaves developed towards the end of the season tion and surpluses are sold in local markets.
are tastier as they tend to grow under condi- International trade is mainly within West Af-
tions of stress. In Botswana and Zimbabwe rica, with the exporting countries in the drier
boiled cowpea leaves are kneaded to a pulp and Sahelian zone, and the importing countries in
squeezed into small balls, which are dried and the more densely populated humid region
stored. Immature, green and still soft seeds are along the coast. It has been estimated that at
cooked to a thick soup and used as relish. The least 285,000 t was traded between West Afri-
tender seedless cowpea pods are sometimes can countries in 1998, mainly from Niger to
used as a cooked vegetable, as are young pods Nigeria, but the total trade is probably larger.
of yard-long bean. In Asia this is the most im- There are no statistical data on the quantity of
portant use of cowpea, in Africa it is uncom- leaves and pods harvested, but it is likely to be
mon. In Benue State, Nigeria, the stringless considerable. Fresh and dried leaves are much
coiled pods with little parchment of a landrace sold in urban markets and some are traded to
called 'Eje-O'Ha' are parboiled for a few min- neighbouring countries. Dried leaves in the
utes, opened and split in half. The seeds are form of black balls are exported from Zim-
eaten directly while the pod walls are dried babwe to Botswana and South Africa. Yard-
and preserved for later use. Pods are also eaten long bean is grown in Asia on hundreds of
locally in Benin. The roots are sometimes thousands of hectares, but is of minor impor-
eaten, e.g. in Ethiopia and Sudan. tance in Africa.
 VlGNA 223

Properties The nutritional composition of minous plants are used as substitutes for im-
leafy stem tips of cowpea per 100 g edible por- mature cowpea seeds, e.g. those of pea (Pisum
tion is: water 89.8 g, energy 121 kJ (29 kcal), sativum L.), common bean and lima bean
protein 4.1 g, fat 0.3 g, carbohydrate 4.8 g, Ca {Phaseolus lunatus L.).
63 mg, Mg 43 mg, P 9 mg, Fe 1.9 mg, Zn 0.3 Description Climbing, trailing or more or
mg, vitamin A 712 IU, thiamin 0.35 mg, ribo- less erect annual or perennial herb, cultivated
flavin 0.2 mg, niacin 1.1 mg, folate 101 fig, as an annual; taproot well developed, with
ascorbic acid 36 mg. Young cowpea pods with many lateral and adventitious roots; stem up to
seeds contain per 100 g edible portion: water 4 m long, angular or nearly cylindrical, slightly
86.0 g, energy 184 kJ (44 kcal), protein 3.3 g, ribbed. Leaves alternate, 3-foliolate; stipules
fat 0.3 g, carbohydrate 9.5 g, Ca 65 mg, Mg 58 ovate, 0.5-2 cm long, spurred at base; petiole
mg, P 65 mg, Fe 1.0 mg, Zn 0.3 mg, vitamin A up to 15(—25) cm long, grooved above, swollen
1600 IU, thiamin 0.15 mg, riboflavin 0.15 mg, at base, rachis (0.5-)2.5-4.5(-6.5) cm long;
niacin 1.2 mg, folate 53(xg,ascorbic acid 33 mg. stipels small; leaflets ovate or rhombic to
Yard-long bean pods contain per 100 g edible lanceolate, (1.5-)7-14(-20) cm x (l-)4-10(-17)
portion: water 87.9 g, energy 197 kJ (47 kcal), cm, basal ones asymmetrical, apical one sym-
protein 2.8 g, fat 0.4 g, carbohydrate 8.4 g, Ca metrical, entire, sometimes lobed, glabrous or
50 mg, Mg 44 mg, P 59 mg, Fe 0.5 mg, Zn 0.4 slightly pubescent, 3-veined from the base.
mg, vitamin A 865 IU, thiamin 0.1 mg, ribofla- Inflorescence an axillary or terminal false ra-
vin 0.1 mg, niacin 0.4 mg, folate 62 |ig, ascorbic ceme up to 35 cm long, with flowers clustered
acid 19 mg. Immature cowpea seeds contain near the top; rachis tuberculate. Flowers bi-
per 100 g edible portion: water 77.2 g, energy sexual, papilionaceous; pedicel 1-3 mm long,
377 kJ (90 kcal), protein 3.0 g, fat 0.4 g, carbo- with spatulate, deciduous bracteoles; calyx
hydrate 18.9 g, fibre 5.0 g, Ca 126 mg, Mg 51 campanulate, tube c. 5 mm long, lobes nar-
mg, P 53mg, Fe 1.1 mg, Zn 1.0 mg, vitamin A 0 rowly triangular, c. 5 mm long; corolla pink to
IU, thiamin 0.1 mg, riboflavin 0.15 mg, niacin purple, sometimes white or yellowish, standard
1.45 mg, folate 168 |xg, ascorbic acid 2.5 mg. very broadly obovate, hood-shaped, c. 2.5 cm
Mature cowpea seeds contain per 100 g edible
portion: water 12.0 g, energy 1407 kJ (336
kcal), protein 23.5 g, fat 1.3 g, carbohydrate
60.0 g, fibre 10.6 g, Ca 110 mg, Mg 184 mg, P
424 mg, Fe 8.3 mg, Zn 3.4 mg,vitamin A 50 IU,
thiamin 0.85 mg, riboflavin 0.23 mg, niacin 2.1
mg, vitamin Be 0.36 mg, folate 633 (ig, ascorbic
acid 1.5 mg. The essential amino-acid composi-
tion per 100 g mature, raw cowpea seeds is:
tryptophan 290 mg, lysine 1591 mg, methion-
ine 335 mg, phenylalanine 1373 mg, threonine
895 mg, valine 1121 mg, leucine 1802 mg and
isoleucine 956 mg. The principal fatty acids are
per 100 g edible portion: linoleic acid 343 mg,
palmitic acid 254 mg, linolenic acid 199 mg and
oleic acid 88 mg (USDA, 2004). The approxi-
mate fatty acid composition of fat from cowpea
seeds is: saturated fatty acids 25%, mono-
unsaturated fatty acids 8%, polyunsaturated
fatty acids 42%. Cowpea protein is relatively
rich in lysine, but poor in S-containing amino
acids. Cowpea seed is lower in antinutritional
components such as lectins and trypsin inhibi-
tors than common bean (Phaseolus vulgaris L.),
and is easier and quicker to cook.
Adulterations and substitutes The pods
of common bean are often used for the same Vigna unguiculata - 1, inflorescence; 2, fruiting
dishes as yard-long bean, although the taste is branch; 3,seed.
not the same. Immature seeds of several legu- Source: PROSEA
224 CEREALS AND PULSES

long, wings obovate, c. 2 cm long, keel boat- - Melanophthalmus Group: originating from
shaped, c. 2 cm long; stamens 10, 9 fused and 1 West Africa; plant able to flower quickly
free; ovary superior, c. 1.5 cm long, laterally from the first nodes under inductive condi-
compressed, style upturned, with fine hairs in tions, pods comparatively few-seeded, seed
upper part, stigma obliquely globular. Fruit a coat thin, often wrinkled, partly white.
linear-cylindrical pod 8-30(-120) cm long, - Textilis Group: a small group only grown in
straight or slightly curved, with a short beak, Nigeria for the fibre extracted from the long
glabrous or slightly pubescent, pale brown peduncles; at the beginning of the 20 th cen-
when ripe, 8—30-seeded. Seeds oblong to almost tury this group was distributed from the in-
globose, often laterally compressed, 0.5-1 cm terior delta ofthe Niger river eastward to the
long, black, brown, pink or white; hilum ob- Lake Chad basin, but it is gradually disap-
long, covered with a white tissue, with a black- pearing.
ish rim-like aril. Seedling with epigeal germi- In Africa there are numerous landraces and
nation; cotyledons oblong or sickle-shaped, improved cultivars within Unguiculata Group.
thick; first two leaves simple and opposite, Leaves are traditionally picked in cowpea fields
subsequent leaves alternate, 3-foliolate. grown primarily for the dry seed and belong to
Other botanical information Vigna com- the top ten most popular leafy vegetables in
prises about 80 species and occurs throughout many African countries. In addition, special
the tropics. However, the tropical American types with erect plant habit or prostrate stems
species are likely to be placed in a separate with long tender shoots are grown as a leafy
genus in the near future, which would reduce vegetable, sometimes also for the immature
the genus to 50—60species. seeds or young pods. The use of dual purpose
Vigna unguiculata is extremely variable, both types (seeds and leaves) is becoming very popu-
in wild and cultivated plants. Several subspe- lar in some countries as the leaves are the
cies (up to 10) have been distinguished, most of main vegetable during the early rainy season.
them comprising perennial wild types, but Various cultivars of yard-long bean are offered
subsp. unguiculata includes annual wild types by Asian seed companies, with a large varia-
and cultivated ones. tion in plant characters.
In cultivated Vigna unguiculata 5 cultivar- Growth and development Germination of
groups are generally recognized, although the cowpea takes 3-5 days at temperatures above
groups can be crossed readily and overlap: 22°C. The optimum temperature for germina-
- Unguiculata Group (common cowpea): pulse tion is about 35°C. Flowers open in the morn-
and vegetable types, grown for the dry or ing and close before noon; they fall the same
immature seeds, young pods or leaves; plant day. In dry climates cowpea is almost entirely
habit prostrate to erect, up to 80 cm tall, late self-pollinated, but in areas with high air hu-
flowering, pods 10-30 cm long, pendent, hard midity cross-pollination by insects may amount
and firm, not inflated when young, many- to 40%. Only fairly large insects are heavy
seeded and seeds not spaced; most African enough to open the keel. The length of the re-
cultivars belong to this group. productive period is very variable, with the
- Sesquipedalis Group (yard-long bean, syno- earliest cultivars taking 30 days from planting
nyms: Dolichos sesquipedalis L., Vigna ses- to flowering, and less than 60 days to mature
quipedalis (L.) Fruhw.): grown for the young seeds. When leaves are harvested during the
pods; plant climbing, stem up to 4 m long, early growth stages, senescence starts 1.5-2
pods 30-120 cm long, pendent, inflated when months after sowing and the plant dies after 3—
young, many-seeded and seed spaced; impor- 4 months, depending on crop health and inten-
tant vegetable in South-East Asia, but of mi- sity of harvesting. Late cultivars with indeter-
nor importance in tropical Africa, where only minate growth take 90-100 days to flower and
cultivars introduced from Asia are grown. up to 240 days for last pods to mature. Cowpea
- Biflora Group (catjang cowpea): grown for forms N-fixing nodules with Sinorhizobium
the seeds, tender green pods and for fodder; fredii and several Bradyrhizobium species.
plant habit prostrate to erect, up to 80 cm Ecology Wild types of Vigna unguiculata
tall, early flowering, pods 7.5—12 cm long, grow in savanna vegetation, often in disturbed
erect or ascending, hard and firm, not in- localities or as a weed, up to 1500 m altitude,
flated when young, few-seeded and seeds not but some can be found in grassland subject to
spaced; important in India and South-East regular burning, sandy localities close to the
Asia, locally also in Africa (e.g. Ethiopia). coast, woodland, forest edges or swampy areas,
 VlGNA 225

occasionally up to 2500 m altitude. plants spread over a 2-3 m radius. In the Su-
Cowpea grows best at day temperatures of 2 5 - dan savanna cowpea is intercropped with pearl
35CC; night temperatures should not be less millet, sorghum and/or groundnut, in diverse
than 15°C and consequently cultivation is re- and complex traditional intercropping patterns
stricted to low and medium altitudes. At alti- with varying interplant distances and planting
tudes above 700 m growth is retarded. Cowpea sequences of component crops. For instance, in
does not tolerate frost, and temperatures above some areas of Kano state in Nigeria (Minjibir
35°C cause flower and pod shedding. It per- and Gezawa areas) pearl millet is planted first
forms best under full sunlight but tolerates in rows 1.5-3 m apart at the onset of the rains
some shade. Cowpea is generally grown as a (May-June), with 1 m distance within the row,
rainfed crop in sub-Saharan Africa, but in Asia resulting in 4000-6000 hills/ha. When the
it is sometimes grown on residual moisture rains become more stable towards the end of
after an irrigated rice crop. Short-duration June, pulse-type early cowpea cultivars are
determinate types can be grown with less than planted between alternate pearl millet rows at
500 mm rainfall per year; in experiments in a distance of 1 m. Fodder-type, late-maturing
Senegal 'Ein al Ghazal' produced 2400 kg/ha of cowpea is planted later, in mid-July, in the
seeds with only 450 mm rain. Long-duration remaining rows. When grown as a sole crop,
types require 600—1500 mm. Yard-long bean cowpea is sown at densities ranging from
tolerates high rainfall; a fully-grown crop has a 22,000 plants/ha for prostrate types to 100,000
water requirement of 6—8 mm per day. Cultiva- plants/ha for erect types. Recommended plant-
tion in the dry season with ample irrigation is ing distances for sole-cropped cowpea in Kenya
practised, as well as cultivation during the are 60 cm between rows and 20 cm within the
rainy season, although sowing during the rainy row. In Swaziland spacings are 50 cm between
season can result in damage to the emerging or rows and 15 cm within the row for erect culti-
young plants. Most cowpea cultivars are quan- vars. For landraces the spacings are much
titative short-day plants, but day-neutral types wider, especially for the dual purpose types.
also exist. Cowpea can be grown on a wide Often 2-3 seeds are sown per pocket, with
range of soil types with pH 5.5-6.5(-7.5), pro- thinning afterwards, e.g. during weeding. The
vided they are well drained. It is moderately sowing depth is 4—5 cm. Cowpea requires soil
sensitive to salinity and exhibits greater salt with fine tilth for good root growth. Generally,
tolerance during later stages of growth. deep ploughing followed by harrowing provides
Propagation and planting Farmers nor- an adequate tilth. In intercropping systems,
mally use farm-saved seed for planting. The tillage normally follows the crop in which cow-
1000-seed weight of cowpea is 150-300 g. The pea is interplanted.
seed rate for pure stands is 15-30 kg/ha. Seed Peri-urban vegetable farmers use special culti-
dressing with an insecticide and a fungicide vars for ratoon cropping of the leaves. They
(e.g. thiram) prior to planting is recommended. broadcast the seed on raised beds, made on
In tropical Africa cowpea is mostly grown well-manured soil, aiming at a dense stand of
intercropped or in relay with other crops such about 25plants per m2.
as yam, maize, cassava, groundnut, sorghum or Farmers in Africa use yard-long bean seed
pearl millet. Pure stands are not common ex- harvested from a previous crop, in contrast to
cept in the coastal areas of East Africa, and South-East Asia, where many farmers procure
also in Asia and Western countries. In the for- healthy seed from improved cultivars. The
est and Guinea savanna zones of West Africa 1000-seed weight of yard-long bean is lower
cowpea is mainly intercropped with maize, than that of cowpea, 100-150 g. Seed is sown
cassava, yam or groundnut, at a very low den- in pockets of 2-4 seeds. Cultivation is usually
sity (1000-5000 hills/ha). In the northern on raised beds for good drainage and easy sur-
Guinea savanna zone cowpea is intercropped face irrigation and for easy staking and har-
with groundnut and/or sorghum. The compo- vesting. Earthing-up the young plants protects
nent crops are normally planted in rows with the shallow root system and gives support to
systematic intercropping patterns, which may the seedlings. Some farmers apply mulch of
vary from alternate row intercropping to rice straw, but this is not a common practice.
within-row intercropping with varying dis- Management Cowpea derives a significant
tance, giving a grid of groundnut or sorghum amount of its nitrogen requirements from the
rows crossed by the cowpea rows every 2-3 m. atmosphere and may leave 75—150kg/ha in the
The cowpea population is low, with individual soil for the benefit of the succeeding crop. If
226 CEREALS AND PULSES

cowpea is grown in localities where it has not nas campestris pv. vignaeunguiculatae) re-
been grown recently, inoculation with nitrogen- ported from Nigeria. These bacteria are seed-
fixing bacteria has been found to be beneficial. transmitted and secondary spread occurs by
Cowpea requires phosphorus for nodulation wind-driven rain. Control measures include the
and root growth. Incorporation of 25 kg/ha P is use of pathogen-free seeds, seed treatment
adequate for plant growth in phosphorus- with a mixture of antibiotics and fungicides
deficient soils. In soils known to be deficient in such as streptocycline plus captan, and strict
potassium, application of 25 kg/ha K is recom- crop rotation. Resistance genes are available
mended. Cowpea must be kept weed free dur- for bacterial blight and bacterial pustules.
ing the early stages of growth. Two to three Many viruses attack Vigna unguiculata. Some
weedings during the first 6 weeks after plant- viruses of economic importance are cowpea
ing are recommended; once the crop is estab- aphid-borne mosaic potyvirus (CABMV), cow-
lished it outcompetes weeds. Weeding is usu- pea mottle carmovirus (CPMoV), cowpea yellow
ally done by superficial hoeing. mosaic virus (CYMV), black eye cowpea mosaic
Cowpea grown as a vegetable and yard-long potyvirus or bean common mosaic potyvirus
bean have a high mineral uptake. In soils of (BCMV), cucumber mosaic cucumovirus (CMV-
average fertility an application is recom- CS) and cowpea golden mosaic virus (CPGMV).
mended of 5-10 t/ha of farmyard manure dur- Some of the viruses are seedborne, while
ing soil preparation, together with N 20 kg/ha, aphids, white flies and beetles perform field
K 25 kg/ha and P 40 kg/ha. Three weeks after transmission. Control measures include use of
emergence a top dressing of 50 kg/ha urea is healthy seed of resistant cultivars if available,
given. In yard-long bean, 2-2.5 m long stakes and weeding to remove alternative hosts. In
are inserted near the seed beds before sowing poor sandy soils, cowpea is attacked by root-
or during the first two weeks after emergence, knot nematodes (Meloidogyne spp.). It is also a
before the plants have reached a height of 30 host plant of, among others, reniform nema-
cm. Acheap method of staking is to relay-plant todes (Rotylenchus spp.), root-lesion nematodes
yard-long bean next to the stems of maize be- (Pratylenchus spp.) and lance nematodes (Hop-
fore orjust after the cobs are harvested. lolaimus spp.). Crop rotation and resistant
Diseases and pests Cowpea is susceptible cultivars are used to control nematodes.
to a wide range of diseases and pests. Yard- Insect pests are also a major factor limiting
long bean suffers from the same diseases and cowpea production and may even cause total
pests as cowpea but seems less susceptible seed loss. In tropical Africa much damage is
than cowpea under humid conditions. Fungal caused by cowpea aphids (Aphis craccivora),
diseases are more troublesome during the flower thrips (Megalurothrips sjostedti), legume
rainy season, whereas insect and mite pests pod borers (Maruca vitrata, Etiella zincke-
and virus diseases cause more damage during nella), pod bugs and seed suckers (e.g. Clavi-
the dry season. gralla tomentosicollis, synonym: Acanthomia
The major fungal diseases are anthracnose tomentosicollis). Lygus beetle (Lygus hesperus),
(Colletotrichum lindemuthianum), Ascochyta cowpea curculio (Chalcodermus aeneus) and
blight (Phoma exigua), brown blotch (Colleto- green leafhoppers (Empoasca spp.) are of less
trichum truncatum), leaf smut (Protomycopsis importance. Yard-long bean is especially at-
phaseoli), leaf spot (Cercospora canescens, Sep- tractive to aphids (Myzus persicae, Aphis gos-
toria vignae, Mycosphaerella cruenta synonym: sypii), green stink bug (Nezara uiridula) and
Pseudocercospora cruentà), brown rust (Uromy- red spider mite (Tetranychus spp.); greasy cut-
ces appendiculatus), scab (Elsinoë phaseoli), worms (Agrotis ipsilon) often cause damage
powdery mildew (Erysiphe polygoni), pythium just after emergence. The bean shoot fly
soft stem rot (Pythium aphanidermatum), stem (Ophiomyia phaseoli) is a common pest; the
canker (Macrophomina phaseolind) and web larvae tunnel in the leaves and stems, and
blight (Thanatephorus cucumeris, synonym severely attacked young plants will die,
Rhizoctonia solani). Crop rotation and the use whereas older plants will suffer from hampered
of chemicals and resistant cultivars are neces- growth and serious yield reduction. Lodging
sary for integrated disease control. Bacterial incidence is generally high in infested fields;
diseases include bacterial blight (Xanthomonas tolerant cultivars may produce aerial roots
campestris pv. vignicola), which occurs world- above the wound. Another common pest is the
wide, and bacterial pustules (Xanthomonas bean pod fly (Melanagromyza sojae). The lar-
axonopodis pv. glycines synonym: Xanthomo- vae damage the petioles and young pods. Con-
 VlGNA 227

trol of insect pests involves protecting the seed set of the dry season, between October and
with a systemic insecticide (e.g. carbofuran) at November, when the leaves show signs of wilt-
sowing or applied as a solution to the emerging ing. The fodder types are uprooted or cut from
seedlings in the planting holes. Plant debris the base and rolled into bundles with the
and affected plants must be burned. Cowpea leaves intact. These bundles are then kept on
seeds are extremely vulnerable to storage roof tops or in tree forks for drying, and are
pests, with the cosmopolitan cowpea weevil used or sold in the peak dry season.
(Callosobruchus maculatus) being the major The first picking of yard-long bean pods in the
storage pest. Measures to reduce pest damage desirable stage takes place 6—7 weeks after
include application of inoffensive vegetable oil, planting, depending on cultivar and market
neem (Azadirachta indica A.Juss.) oil or wood requirements. Normally the pods are picked
ash, roasting and bagging the seeds in airtight when the outline of the seeds is just visible.
plastic bags, and storing as whole pods. Picking must be meticulous, because pods
Use of chemicals, resistant cultivars, biological which are passed over until the next harvest
control and proper crop management such as will become tough and discoloured, with swol-
intercropping and weeding are necessary for len seed, and may exhaust the plant. Succes-
integrated pest management. Chemical control sive harvests take place at least once a week
of insects is common practice on yard-long (twice a week for a better tuned grading) dur-
bean, but not on cowpea. Because of the risks ing 4-8 weeks.
for farmer and consumer (especially when Yield Farmers may harvest up to 400 kg/ha
leaves are harvested), these sprayings must be of cowpea leaves in a few rounds with no no-
reduced to the strict minimum. ticeable reduction of seed yields. In Nigeria
Two parasitic weeds are a serious problem: climbing cultivars yielded 9-17 t/ha of fresh
Alectra vogelii Benth. prevalent in the south- pods, whereas decumbent cultivars yielded 6—
ern savanna regions ofWest Africa, East Africa 15 t/ha. The mean dry seed yield of the same
and southern Africa, and Striga gesnerioides cultivars was 1.4—1.7 t/ha. The world average
(Willd.) Vatke prevalent in the savanna regions yield of dry cowpea seed is low, 240 kg/ha, and
ofWest and Central Africa. Crop rotation, deep for fodder it is 500 kg/ha (air-dried leafy
cultivation, intercropping, early planting and stems). Average yield of dry cowpea seeds un-
use of resistant cultivars reduce infestation by der subsistence agriculture in tropical Africa is
these parasitic weeds. 100-500 kg/ha. The average seed yield in Niger
Harvesting Cowpea leaves are picked in a is 120 kg/ha, in Nigeria 400 kg/ha, and in the
period from 4 weeks after emergence of the United States 900 kg/ha. Apart from the effects
seedlings to the onset of flowering. In crops of diseases and pests, the low yields are partly
grown for the seed, farmers often harvest 10— explained by the fact that the crop is mostly
20% of the leaves before the start of flowering grown at low densities in intercropping sys-
with little detrimental effect on the seed yield. tems, shaded by taller cereals. Furthermore,
Stronger defoliation increasingly reduces flow- cowpea is often sown later in the rainy season,
ering, fruiting and seed yield. Growers of leafy which results in a shorter crop duration due to
cowpea types cut the plants at about 10 cm photoperiod-sensitivity. A yield potential of 3
above the ground for a succession of new shoots t/ha of seed and 4 t/ha of hay can be achieved
(ratooning). Green pods are harvested when in sole-cropping with good management. In the
the seed is still immature, 12-15 days after United States seed yields up to 7 t/ha have
flowering. Harvesting of dry seed is done when been obtained.
at least two-thirds of the pods are dry and yel- For yard-long bean, a total yield of 15 t/ha in a
low. In indeterminate types harvesting is com- harvest period of at least one month is consid-
plicated by prolonged and uneven ripening; for ered satisfactory, but yields as high as 30 t/ha
some landraces harvesting may require 5-7 have been reported.
rounds. Mature seeds are usually harvested by Handling after harvest Harvested leaves
hand. Sometimes plants are pulled out when cannot be kept for long; they have to be sold
most of the pods are mature. In the complex within 2 days. The shoots can be kept longer by
traditional intercrop patterns of Kano state putting them in a basin with water. Cowpea
(Nigeria), early cowpea and sorghum cultivars leaves are frequently dried in the sun for pres-
are harvested at the end of August or the be- ervation, either after boiling and squeezing to
ginning of September. The late cowpea and black balls, or directly as whole or broken
sorghum cultivars are harvested after the on- leaves, or as powder. Green yard-long bean
228 CEREALS AND PULSES

pods are tied in bundles of 20-40 and packed in and fodder for livestock in the dry season. Im-
baskets or crates for transport to the market. proved cultivars have also been developed for
Yard-long bean is less susceptible to loss of intercropping. National programmes in many
weight by transpiration and to transport dam- countries have released improved cowpea cul-
age than most other vegetables. In cool storage tivars with resistances to bacterial blight, cow-
(8°C) the pods will keep for 4 weeks. Immature pea aphid-borne mosaic potyvirus, cowpea
fresh cowpea seeds have a limited shelflife if aphids, cowpea curculio, root-knot nematodes,
stored at ambient temperatures, but at 8°C cowpea weevil and parasitic weeds. New early
they can stay fresh for 8 days. In Europe, the maturing cultivars were developed for hot and
United States and Japan, immature tender dry conditions, e.g. 'Ein al Ghazal' and
green pods are sometimes frozen or canned. As 'Mouride'. Improved cultivars are often short,
a pulse, the threshed seed should be dried erect, determinate types selected for optimal
thoroughly to a moisture content of 14%or less dry seed production and less suitable for the
for good storability. traditional leaf picking. Wild African Vigna
Genetic resources The International Insti- species have been successfully crossed with
tute of Tropical Agriculture (UTA), Ibadan, Vigna unguiculata.
Nigeria holds a collection of over 15,000 acces- Breeding work on African vegetable types is
sions of the cultivated cowpea and 1000 acces- scarce. Simlaw Seeds in Kenya has commer-
sions of related wild Vigna; the University of cialized 'Kenduke-1', a semi-trailing type se-
California, Riverside, United States holds 5000 lected for large leaves with an attractive green
accessions. UTA characterized 8500 accessions colour and good taste and that can be picked
for resistance to Maruca pod borer and sucking for a long time. In Senegal the leaf vegetable
bugs, and 4000 for resistance to flower thrips, 'Fuuta' with a vegetative period of up to 50
bruchids and viruses. The level of resistance to days was selected. The Crop Breeding Institute
insect pests is high in the wild species Vigna in Harare, Zimbabwe, selected dual-purpose
vexillata (L.) A.Rich., especially to pod sucking cultivars with high leaf and seed yield; the
bugs and Maruca pod borer. Many accessions Zimbabwean cultivar 'Chigwa' is specially
of wild Vigna species possess high levels of suited for use as a leaf vegetable because of
resistance to the storage weevil. late flowering. 'Melakh' is a dual-purpose culti-
Small collections of yard-long bean are present var bred for dry and fresh seed production in
at the Asian Vegetable Research and Develop- Senegal.
ment Center (AVRDC), Shanhua, Taiwan and Breeding of improved cultivars of yard-long
the Institute of Crop Germplasm Resources bean by backcrossing and pedigree selection
(CAAS), Beijing, China and in national insti- has been performed in South-East Asia. Yield
tutes in Asia. Only very small collections of is strongly correlated with pod length and the
catjang cowpea exist. In Asia landraces of vege- number of pods per plant. Resistance to bean
table and pulse types of Vigna unguiculata are flies would be welcome but seems difficult to
in danger of being lost since improved cultivars achieve. East-West Seed Company in Thailand
are widely grown. This process has also started selected cultivars adapted to a wide range of
in Africa. growing conditions, e.g. 'Aba', with early ma-
Breeding Much work has been performed turity (first harvest 45 days after sowing), high
on Vigna unguiculata breeding, mostly for cul- yield, greyish green pods 60-70 cm long, and
tivars grown as a pulse, and in South-East excellent market quality.
Asia for yard-long bean. In the United States Genetic linkage maps of cowpea have been
special cowpea cultivars for harvesting pods constructed using RAPD, AFLP and RFLP; the
and young seeds have been developed. Selec- linkage maps have been used to locate genes
tion criteria for cowpea concern resistances (to conferring resistance to Striga gesnerioides,
insect pests, diseases, nematodes, parasitic several viruses and root-knot nematodes, as
weeds, drought), plant type, seed type, yield well as to locate quantitative trait loci (QTLs)
and cropping system. UTA has a large breeding for time to flowering, time to maturity, pod
programme and distributes cowpea germplasm, length, pod and seed weight, and resistance to
breeding material and cultivars. In collabora- aphids. Direct organogenesis of cowpea has
tion with the International Livestock Research been achieved using hypocotyl, epicotyl or coty-
Institute (ILRI), UTA initiated a breeding pro- ledon tissue. Regeneration of cowpea via so-
gramme to develop improved cowpea cultivars matic embryogenesis has been attempted, but
that provide both seed for human consumption callus failed to regenerate plants at an accept-
 ZEA 229

able frequency. Genetic transformation has 1989.

been proposed, e.g. to achieve resistance to Authors R. Madamba, G.J.H. Grubben, I.K.
pests by incorporating Bacillus thuringiensis Asante &R. Akromah
(Bt) genes and a-amylase inhibitor genes, but a
robust system for stable genetic transformation
ofcowpea is not yet available. ZEAMAYSL.
Prospects Cowpea serves as a cheap source
of plant protein, especially in West Africa. It Protologue Sp. pi. 2:971 (1753).
plays an important role in multiple cropping Family Poaceae (Gramineae)
systems and is a major component of inte- Chromosome number 2re= 20
grated crop/livestock systems in West Africa. Vernacular names Maize, corn, Indian
Diseases and pests are the major constraints in corn (En). Maïs (Fr). Milho (Po). Mhindi, mu-
cowpea production. Resistance breeding could hindi (Sw).
be of utmost importance to overcome these Origin and geographic distribution Maize
constraints, with an increasingly important was domesticated in southern Mexico around
role for biotechnological tools. Future im- 4000 BC. Early civilizations of the Americas
provement also relies on the collection of land- depended on maize cultivation. When the
races and their wild relatives and their incor- Europeans arrived in the Americas, maize had
poration into breeding programmes. already spread from Chile to Canada. Maize
The prospects for vegetable cowpea in Africa was reported for the first time in West Africa
are bright. Apart from traditional dual-purpose in 1498, six years after Columbus discovered
cowpea cultivars (harvested as pulse and for the West Indies. The Portuguese brought
the leaves) there is a need for special vegetable floury grain types from Central and South
types. As a leaf vegetable: dwarf plants with America to Sâo Tomé, from where they spread
erect or prostrate habit, long vegetative period, to the West African coast. Portuguese and Arab
tender shoots and leaves. For immature seed: traders introduced Caribbean flint maize types
dwarf plants with erect or prostrate, determi- into East Africa in the mid 1500s, from where
nate habit. For fresh pods: pods about 15 cm they spread to southern Africa. Through the
long (replacing French bean in hot lowland trans-Saharan trade, the Arabs introduced the
regions). flinty types that had been brought to northern
As a fruit vegetable, it seems logical to replace Africa into sub-Saharan Africa. The flinty
cowpea by yard-long bean, because of its supe- types still predominate in northern parts of
rior yield and quality. Asian cultivars should West Africa while the floury types prevail in
be tested on suitability for tropical African the southern parts, with some variation from
conditions because, if combined with market this pattern. Maize had become a staple food in
development, yard-long bean has the potential East and southern Africa by the 1930s.
to become an excellent enrichment of the avail- Maize has an extremely wide distribution. It is
able vegetable assortment. grown from latitude 58°N in Canada and Rus-
Major references Ehlers, 1997; Grubben,
1993; Hall & Coyne (Editors), 2003; Langyin-
tuo et al., 2003; Ng & Singh, 1997; Pandey &
Westphal, 1989; Pasquet & Baudoin, 1997;
Singh & Rachie (Editors), 1985; Singh et al.
(Editors), 1997a; Vanderborght & Baudoin,
2001.
Other references Allen et a l , 1998; Bhat,
Etejere & Oladipo, 1990; Burkill, 1995; de
Vries & Toenniessen, 2001; Ezedinma, 1973;
Hall et a l , 2003; Kahn, 1993;Madamba, 1997;
Madamba, 2001; Magkoko, 2001; Messiaen,
1989; Ouédraogo et a l , 2002; Pasquet, 1998; .—^y /}
Popelka, Terryn & Higgins, 2004; Schippers,
2000; Singh et a l , 2003; Ubi, Mignouna & • J — , — —
Thottapilly, 2000; Uguru, 1996; Uguru, 1998;
USDA, 2004.
Sources of illustration Pandey &Westphal, Zea mays -planted
230 CEREALS AND PULSES

sia, throughout the tropics, to latitude 42°S in female inflorescences ('baby cobs') are a fancy
New Zealand and South America, and in areas vegetable in Western countries and in Asia.
helow sea level in the Caspian Plain up to ar- Mature maize plants are used for animal feed.
eas as high as 3800 m in Bolivia and Peru. It is Silage maize is one of the leading crops in in-
grown in all countries of Africa, from the coast dustrialized Western countries, where special
through savanna regions to the semi-arid re- cultivars and production technologies have
gions of West Africa, and from sea-level to the been developed. The stalks are used for fuel,
mid- and high-altitudes of East and Central fodder and thatching and as compost. The fibre
Africa. in the stems and the inner leaves surrounding
Uses Maize grain is used for three main the cob are made into paper. These cob leaves
purposes: as a staple food, as feed for livestock are often used to wrap foods, and may also be
and poultry, and as a raw material for many made into cloth or mats, and be used for mat-
industrial products. In tropical Africa nearly tress filling. Ash of the burnt stem is some-
all maize grain is used for human food, pre- times a substitute for salt. The cob is made into
pared and consumed in many ways. It may be pipe-bowls. In southern Africa the incinerated
eaten fresh on the cob and simply roasted, but cob is included in a snuff.
the grain is usually ground and the meal is Maize has a range of uses in traditional Afri-
boiled into porridge or fermented into beer. In can medicine. Urino-genital problems are
tropical Africa maize is mainly consumed as treated with prescriptions based on the whole
thick porridge ('ugali' in East Africa, 'sadza' in or parts of the maize plant, especially a decoc-
Zimbabwe). It is commonly eaten with cooked tion of the styles, which is also used to treat
vegetables and, when available, meat. A thin jaundice. A leaf maceration is drunk to treat
porridge ('uji' in East Africa, 'ogi' in Nigeria, fever. Charcoal made from the culms is in-
'koko' in Ghana) is also commonly eaten espe- cluded in medicines to treat gonorrhoea; an
cially as weaning food. In Ethiopia local beer infusion from the burnt cob is used to wash
('tella') and spiritual liquor ('arakie') are pre- wounds.
pared from maize grain malt. Popcorn is a Production and international trade Accord-
popular snack. ing to FAO estimates, the average world pro-
The main industrial products obtained from duction of maize in 1999-2003 amounted to
maize are breakfast products such as corn- 611 million t/year from 139 million ha. The
flakes, starch, sugar and oil. The main product main producing countries are the United
is starch that is used for human consumption States (243 million t/year in 1999-2003, from
or made into syrup, alcohol, but also among 28 million ha), China (117 million t/year from
others as laundry starch and as a source mate- 24 million ha), Brazil (38 million t/year from
rial for many chemical products. Most indus- 12 million ha), Mexico (19 million t/year from 7
trial products are obtained by the wet-milling million ha), France (15 million t/year from 2
process, in which the grain is first steeped in million ha), Argentina (15 million t/year from
water, after which the germ and bran are sepa- 3 million ha) and India (12 million t/year from
rated from the endosperm. The various prod- 7 million ha). South Africa produced 9.4 million
ucts are subsequently obtained by physical or t/year from 3.6 million ha. Maize production in
chemical processes, and e.g. sugars from maize tropical Africa in 1999-2003 was 26.6 million
now account for half of the sugars used in hu- t/year from 21.2 million ha. The main produc-
man nutrition. Dry milling produces grits, con- ing countries in tropical Africa are Nigeria (4.7
sisting of coarsely ground endosperm from million t/year from 4.2 million ha), Ethiopia
which most of the bran and the germ have been (2.9 million t/year from 1.6 million ha), Tan-
separated. The germ yields an oil that can be zania (2.6 million t/year from 1.6 million ha),
refined for human consumption, widely used as Kenya (2.5 million t/year from 1.6 million ha)
cooking or salad oil and in margarines. It is the and Malawi (2.0 million t/year from 1.5 million
second most widely consumed vegetable oil in ha). From 1961-1965 to 1999-2003 the annual
the United States and is also made into soap or maize production in tropical Africa increased
glycerine. The residues from the production of from 9.1 to 26.6 million t/year, and the har-
starch or oil, together with the bran, are used vested area from 10.2 to 21.2 million ha.
in animal feeds (corn gluten meal and corn Average world export of maize amounted to
gluten feed). 80.1 million t/year in 1998-2002, with the
Unripe cobs are consumed as vegetable or United States (47.5 million t/year), Argentina
green maize, boiled or roasted. Very young (10.3 million t/year), France (7.9 million t/year)
 ZEA 231

and China (7.4 million t/year) as main export- sumption, due to its flavour, colour and stabil-
ers. Export of maize from tropical Africa was ity and the presence of linoleic acid and vita-
only 307,000 t/year, with Zimbabwe (143,000 min E.
t/year), Tanzania (42,000 t/year) and Uganda Maize grain in tropical Africa often contains
(25,000 t/year) as main exporters. The main mycotoxins such as aflatoxins and fumosinins,
importers were Japan (16.3 million t/year) and which are harmful to humans and livestock.
South Korea (8.3 million t/year). Maize imports Aflatoxins are produced by Aspergillus spp.,
into tropical Africa were 1.8 million t/year. especially Aspergillus flavus; they are powerful
Properties The composition of mature white carcinogens, especially affecting the liver, and
maize grain per 100 g edible portion is: water have immunosuppressive properties. Fumos-
10.4 g, energy 1527 k j (365 kcal), protein 9.4 g, inins are produced by Fusarium spp., espe-
fat 4.7 g, carbohydrate 74.3 g, dietary fibre 7.3 cially Fusarium verticillioid.es; they have been
g, Ca 7 mg, Mg 127 mg, P 210 mg, Fe 2.7 mg, implicated in various animal diseases. Human
Zn 2.2 mg, thiamin 0.39 mg, riboflavin 0.20 mg, health risks due to fumosinins are possible, but
niacin 3.6 mg, vitamin B60.62 mg, folate 19 ug so far there is no conclusive evidence, although
and ascorbic acid 0 mg. The essential amino- correlation studies have suggested a link be-
acid composition per 100 g edible portion is: tween consumption of maize with fumosinins
tryptophan 67 mg, lysine 265 mg, methionine and high incidence of human oesophageal car-
197 mg, phenylalanine 463 mg, threonine 354 cinoma.
mg, valine 477 mg, leucine 1155 mg and isoleu- Description Robust annual grass up to 4(-6)
cine 337 mg. The principal fatty acids per 100 g m tall; root system consisting of adventitious
edible portion are: linoleic acid 2097 mg, oleic roots, developing from the lower nodes of the
acid 1247 mg and palmitic acid 569 mg (USDA, stem near the soil surface, usually limited to
2004). Maize is deficient in tryptophan and the upper 75 cm of the soil, but single roots
lysine, but cultivars with higher content of sometimes penetrating to a depth of over 2 m;
these amino acids have been bred using the stem (culm) usually single and simple, solid.
recessive gene Opaque-2 with modifiers. These
cultivars are referred to as Quality Protein
Maize (QPM). In general 100 kg of whole
maize, with 16%moisture content, yields about
64 kg starch and 3 kg oil. The endosperm,
which accounts for 80% of the weight of the
grain, is poor in phosphorus and calcium and
contains most of the starch and two-thirds of
the protein. More than 80%of the fat and most
minerals are in the embryo or germ, which
constitutes about 12% of the grain. The starch
of the endosperm usually consists of a mixture
of about 75% amylopectin and 25% amylose,
but waxy maize contains only amylopectin. The
most common grain colours are yellow and
white. Yellow maize predominates in the
United States, China and Brazil, whereas
white maize predominates in tropical Africa,
Central America and the northern part of
South America. White maize has harder grain
and gives sweeter, more flavourful products; it
is primarily grown for food, whereas yellow
maize is mainly used as animal feed. Yellow
maize contains the provitamin A cryptoxan-
thin. Most vitamins are found in the outer lay-
ers of the endosperm and in the aleurone layer.
Maize is deficient in gluten and therefore un- Zea mays - 1, basal plant part; 2, central plant
suitable for making leavened bread; it is toler- part with female inflorescences; 3, upper plant
ated by patients with coeliac disease. Maize oil part with male inflorescence; 4, infructescence.
is considered a premium oil for human con- Source: PROSEA
232 CEREALS AND PULSES

Leaves alternate, simple; leaf sheaths overlap- peared first in the United States at the end
ping, auricled at the top; ligule c. 5 mm long, of the 18 th century, and spread to Europe in
colourless; blade linear-lanceolate, 30-150 cm the 20 th century, where it is widely culti-
x 5-15 cm, acuminate, margins smooth, midrib vated;
pronounced. Male and female inflorescences - Pod maize: this is the most primitive type of
separate on the same plant; male inflorescence maize in which the grain is enclosed in
('tassel') a terminal panicle up to 40 cm long, bracts; it is not grown commercially;
lateral branches with paired spikelets 8-13 - Pop maize or popcorn: it has small grains
mm long, one sessile, the other on a short pedi- with a high proportion of very hard corneous
cel, each spikelet with 2 glumes and 2 florets, endosperm and a little soft starch in the cen-
each floret with an ovate lemma, a thin palea, tre; on heating the steam generated inside
2 fleshy lodicules and 3 stamens; female inflo- the grain causes it to pop and explode, the
rescence a modified spike, usually 1-3 per endosperm becoming everted about the em-
plant in leaf axils about half way up the stem, bryo and hull to produce a palatable white
composed of a thick spongy axis with paired fluffy mass ('popcorn'); in 'rice popcorn' culti-
sessile spikelets in 8-20 longitudinal rows and vars the grains are pointed and in 'pearl pop-
enclosed by 8-13 modified leaves (spathes), corn' cultivars rounded; popcorn is most im-
spikelet with 2 glumes and 2 florets, lower flo- portant in the United States and Mexico, but
ret sterile, consisting solely of a short lemma has also become a popular snack in tropical
and palea, upper floret with a short, broad Africa;
lemma and palea, a single superior ovary and a - Flour maize or soft maize: the grain can have
long threadlike style and stigma ('silk') up to all types of colours, it usually has no dent
45 cm in length and emerging from the top of and the endosperm consists of soft starch,
the inflorescence, receptive throughout most of when parched it can be chewed more easily
its length. Fruit a caryopsis (grain), usually than flint maize and it is also easier to grind,
obovate and wedge-shaped, variously coloured but it is susceptible to mould and breakage
from white, through yellow, red and purple to during handling; it is one of the oldest maize
almost black, up to 1000 together in an in- types and was widely grown in the drier
fructescence ('cob') enclosed by modified leaves parts of the United States, western South
up to 45 cm x 8 cm. America and South Africa, and it is still
Other botanical information Zea com- widely grown in the Andes and small
prises 5 species, including cultivated Zea mays amounts are grown in the United States;
and 4 wild relatives, all from tropical America people in the southern parts of West Africa
and called teosintes. Zea mays is a heterogene- relish flour maize;
ous species and cultivars can be divided into 8 - Sweet maize: the grain contains glossy en-
types (or cultivar groups) according to the dosperm with little starch, giving a wrinkled
structure and shape of the grain: appearance after drying, and it is usually
- Dent maize: the sides of the grain have cor- eaten in the immature state as a fresh vege-
neous endosperm, but the inside has soft table; it is mostly grown in the United
white starch, extending to the apex, shrink- States, but has become popular among the
ing on drying to produce the characteristic elites in African countries;
dent, the wedge-shaped grains are usually - Waxy maize: the starch is composed entirely
yellow or white; it is the principal maize in of amylopectin and is used for the manufac-
the United States and northern Mexico; ture of adhesives; it is mainly grown in east-
- Flint maize: the grain can be coloured vari- ern Asia for human consumption, but also in
ously and consists mainly of hard endosperm the West for industrial applications.
with a little soft starch in the centre, it has Within the various grain types, there are many
rounded ends and is generally smaller than cultivars grown in different parts ofthe world.
the grain of dent maize, it matures earlier, is Growth and development The first leaf of
harder, and when dry it is more resistant to maize emerges from the soil usually 4—6 days
insect attack; it is the predominant type after planting. The minimum temperature for
grown in Europe, Asia, Central and South germination is 10°C; the optimum around
America and parts oftropical Africa; 20CC. The plant sometimes has a few tillers
- Flint-dent maize: this group resulted from that are of value in low density stands. At a
hybridization between flint and dent maize, later stage some whorls of aerial roots ('brace
and has intermediate characteristics; it ap- roots') may develop from the lower nodes above
 ZEA 233

the ground which partly help to anchor the does not tolerate waterlogging and is sensitive
plant while also contributing to the uptake of to salinity. Since a young crop leaves much of
water and nutrients. Flower initiation is gen- the ground uncovered, soil erosion and water
erally 20-30 days after germination. Maize is losses can be severe and attention should be
protandrous: in cultivars that mature in 4 paid to adequate soil and water conservation
months the male inflorescence emerges 50-60 measures.
days after planting and the styles of the female Propagation and planting Maize is prop-
inflorescence appear about a week later. Maize agated by seed and direct sowing is common.
is mature 7-8 weeks after flowering. The pe- The 1000-grain weight is 150-300 g. Sowing
riod from planting to harvesting varies consid- should preferably be done early in the season,
erably. It may be as short as 70 days in some as soon as soil conditions and temperature are
extra early cultivars and as long as 200 days in favourable and the rainfall is well established.
some very late cultivars. Climatic conditions, Smallholders plant maize by hand while me-
latitude and altitude influence growth dura- chanical planting is practised on large com-
tion. In tropical highland areas it may take 9- mercial farms. Planting by hand requires 5-10
10 months to maturity. Maize is predominantly man-days/ha. Seed is dropped in the plough
cross-pollinating (90—95%), but is self-fertile. furrow or in holes made with a planting stick
Maize follows the C4-cycle photosynthetic or hoe. Planting may be done on hills or in
pathway. rows, on flat land or on ridges. Ridging or
Ecology Maize is adapted to a wide range of heaping is usually done on heavy soils, to im-
environments, but it is essentially a crop of prove drainage. The seed rate is up to 25 kg/ha
warm regions where moisture is adequate. The in sole cropping, and 10-15 kg/ha in intercrop-
bulk of the crop is grown in tropical and sub- ping. When maize is sown in rows, the spacing
tropical regions. In West and Central Africa is usually 75-90 cm between rows and 25-50
the Guinea savanna zone offers the best eco- cm within the row, with 1—3 seeds per pocket,
logical conditions for maize. The mid-altitude resulting in a plant density of 40,000-80,000
regions of East and southern Africa are also plants/ha. Wide spacing results in more weed
suitable for maize production. In Ethiopia, for growth and increases erosion. To obtain a high
instance, maize is mainly grown at 1000-2400 yield, a uniform crop stand is very important,
m altitude. Maize is generally less suited to as the tillering capacity of maize is limited. The
semi-arid or equatorial climates, although sowing depth is commonly 3-8 cm, depending
drought-tolerant cultivars adapted to semi-arid on soil conditions and temperature. Deep sow-
conditions are now available. The crop requires ing is recommended on light, dry soils. On
an average daily temperature of at least 20°C smallholdings the land is usually cultivated by
for adequate growth and development; the op- hand or by animal traction. The usual depth of
timum temperature for growth and develop- ploughing is 8-10 cm and ploughing is done
ment is 25—30°C; temperatures above 35° re- just before or at planting time. Sometimes
duce yields. Frost is not tolerated. Maize re- animal manure or fertilizers are applied at the
quires abundant sunlight for optimum yields. time of planting.
The time of flowering is influenced by photope- Maize may be grown as a sole crop or in inter-
riod and temperature; maize is considered a cropping with other food crops such as common
quantitative short-day plant. Maize is less bean, cowpea, pigeon pea, groundnut, yam,
drought-resistant than sorghum, pearl millet cassava, sweet potato, pumpkin, melon or wa-
and finger millet. In the tropics it does best termelon. In some parts of tropical Africa two
with 600-900 mm well-distributed rainfall crops of maize are planted per year. In areas
during the growing season. It is especially sen- where the rainy season is shorter, the crop is
sitive to drought and high temperatures planted only once, although a second planting
around the time of flowering. is possible under irrigation, on residual mois-
Maize can be grown on a wide range of soils, ture on heavy soils or on hydromorphic soils.
but performs best on well-drained, well- Management Maize is very sensitive to
aerated, deep soils containing adequate organic weed competition during the first 4-6 weeks
matter and well supplied with nutrients. The after emergence, and weed control is very im-
high yield of maize is a heavy drain on soil portant. The crop should be planted as soon as
nutrients and maize is therefore often grown as possible after the preparation of the seed-bed.
a first crop in the rotation. It can be grown on Interrow cultivation to control weeds and to
soils with a pH of 5-8, but 5.5-7 is optimal. It break up a crusted soil surface may be done
234 CEREALS AND PULSES

until the plants reach a height of about 1 m. diseases are now available in tropical Africa.
Weeding is mostly done by hand, requiring at Measures to reduce mycotoxin contamination
least 25 man-days/ha. Chemical weed control is of the cob include early harvesting, rapid dry-
gaining importance in tropical Africa, because ing, sorting out of damaged and infected
hand weeding is time-consuming and expen- grains, sanitation (removal of crop residues,
sive as a result of the increasing scarcity of cleaning of stores, removal of heavily damaged
labour. Ridging or earthing-up is sometimes cobs), improved storage, and the use of fungi-
practised. Most maize production in tropical cides. The most important virus disease of
Africa is rainfed. Occasionally it is grown on maize is maize streak virus (MSV), which is
bunds in irrigation schemes. Maize usually restricted to Africa and may cause 100% yield
responds well to fertilizers. A maize crop yield- loss. It is transmitted by leafhoppers (Cica-
ing 2 t grain and 5 t stover per ha removes dulina spp.) and is most serious in late-planted
about 60 kg N, 10 kg P and 70 kg K per ha crops. Cultivars resistant to maize streak virus
from the soil. Nitrogen uptake is slow during are available. Of lesser importance in tropical
the first month after planting, but increases to Africa are maize dwarf mosaic virus (MDMV),
a maximum during formation of the inflores- sugar cane mosaic virus (SCMV) and maize
cences. Maize has a high demand for nitrogen, chlorotic mottle virus (MCMV). Maize is rela-
which is often the limiting nutrient. High ni- tively tolerant to nematodes occurring in tropi-
trogen levels should be applied in 2 doses; the cal soils.
first dose at planting or 2-3 weeks after emer- The most serious insect pests of maize in tropi-
gence and the second one about 2 weeks before cal Africa are cutworms (Agrotis spp.), stem
flowering. Phosphate is not taken up easily by borers (especially Busseola fusca, Eldana sac-
maize and, moreover, many tropical soils are charina, Sesamia calamistis and Chilo partel-
deficient in available phosphate. It is advisable lus), cob borer (Mussidia nigrivenella), cotton
to apply organic manures before ploughing to bollworm (Helicoverpa armigera), armyworm
improve soil structure and supply nutrients. (Spodoptera exempta), leafhoppers (Cicadulina
Smallholder farmers in tropical Africa apply spp.) and less commonly variegated grasshop-
little or no fertilizer to the maize crop. When per (Zonocerus variegatus). Occasionally ter-
they do, it is usually only once, about 4 weeks mites and locusts also infest maize fields. Ap-
after planting when the crop is knee high. plication of insecticides may be necessary to
Maize is grown in rotation with groundnut, control these pests. Cultural methods for insect
common bean, cowpea, cotton and tobacco. Ro- control include early planting and burying or
tation with soya bean is gaining popularity in burning of crop residues. Although biological
northern Nigeria; it increases maize yields by control of stem borers using natural enemies
providing nitrogen and by reducing parasitism. has not been very successful, it is still consid-
In the United States maize is often grown in ered a potentially viable control option. Maize
rotation with soya bean. is not prone tobird damage.
Diseases and pests The most important Common storage pests of maize are grain
fungal diseases of maize in tropical Africa are moths (Sitotroga cerealella and Ephestia cau-
rots affecting female inflorescences (Fusarium tella), grain weevils (Sitophilus spp.) and the
spp. and other fungi), the stalk-rot complex larger grain borer (Prostephanus truncatus).
(Diplodia maydis, Fusarium moniliforme, Grains may be mixed with small amounts of an
Macrophomina phaseoli and Pythium apha- insecticide (e.g. malathion) to control these
nidermatum) and leaf blights (Exserohilum pests. Rodents are also major storage pests in
turcicum and Bipolaris maydis). Of more local tropical Africa.
importance are downy mildew (Peronosclero- The parasitic witchweed (Striga spp.) is a seri-
spora sorghi), smut (Ustilago maydis) and ous constraint to maize production in many
rusts (Puccinia sorghi and Puccinia polysora). parts of tropical Africa, especially Striga her-
Grey leaf spot (Cercospora zeae-maydis) is im- monthica (Delile) Benth. in West and Central
portant in East and southern Africa, but in Africa and Striga asiatica (L.) Kuntze in
West and Central Africa it occurs only in mid- southern Africa. No single control measure is
altitude regions. Host-plant resistance is the effective for this weed, and an integrated ap-
most effective disease control measure. Culti- proach is recommended, involving planting
vars resistant to Exserohilum turcicum leaf maize seed that is free from Striga seeds,
blight and downy mildew are available. Maize planting resistant cultivars, adequate fertilizer
cultivars that possess resistance to multiple application (especially N), crop rotation (e.g.
 ZEA 235

with cotton, soya bean or cowpea), and removal Genetic r e s o u r c e s The largest germplasm
of Striga plants before they flower. collections of maize are held in India (Indian
Harvesting Maize is usually harvested by Agricultural Research Institute, New Delhi,
hand. Mechanical harvesting is practised on 25,000 accessions), Mexico (International
large farms. Indicators of maturity are yellow- Maize and Wheat Improvement Center
ing of the leaves, yellow dry papery leaves (CIMMYT), Mexico City, 22,140 accessions),
around the cobs, and hard grains with a glossy the United States (USDA-ARS North Central
surface. In the dry season, maize is often left in Regional Plant Introduction Station, Iowa
the field until the moisture content of the grain State University, Ames, Iowa, 17,910 acces-
has fallen to 15-20%. In case of harvesting by sions) and China (Institute of Crop Germplasm
hand, the cobs should be broken off with as Resources (CAAS), Beijing, 15,840 accessions).
little attached stalk as possible. Cobs may be In tropical Africa substantial germplasm col-
harvested with the surrounding leaves still lections are held in Kenya (Kenya Agricultural
attached. These may be turned backward and Research Institute (KARI), National Agricul-
used to tie several cobs together and hung up tural Research Centre, Kitale, 1780 acces-
to dry. Alternatively, the leaves are completely sions), Malawi (Malawi Plant Genetic Re-
removed from the cobs, which are then stored sources Centre, Chitedze Agricultural Re-
in cribs to dry. search Station, Lilongwe, 970 accessions),
Yield Maize has the highest yield potential Rwanda (Institut des Sciences Agronomiques
among the cereal crops. The current average du Rwanda (ISAR), Butare, 580 accessions).
world yield of maize is 4.4 t/ha, but grain yields Breeding Maize breeding in tropical Africa
over 20 t/ha are possible. Average grain yields started with introduction of improved materi-
of maize in tropical Africa are about 1.25 t/ha, als from Central and South America. Some of
varying greatly from less than 1 t/ha for small- the cultivars were multiplied and distributed
holders to about 6 t/ha in commercial farms. to farmers directly, while others were subjected
Yields higher than 10 t/ha have been recorded, to genetic improvement. Breeding for resis-
but these are exceptional. In 2001 the average tance to various diseases, such as rust, blight,
yields of maize in the different sub-regions of smut and leaf spots, was a major objective.
tropical Africa were: West Africa 1.3 t/ha, Cen- Many cultivars resistant to the prevalent dis-
tral Africa 1.0 t/ha, East Africa 1.6 t/ha and eases were released. At the initial stages of
southern Africa 1.4 t/ha. maize improvement in the region, composites
Handling after harvest The major post- (mixtures of genotypes from various sources
harvest problems of maize in most production that are maintained by normal pollination) and
areas are reducing the moisture content of the synthetic cultivars (cultivars produced by
grain to 12-15%, protection from insects and crossing a number of genotypes in all possible
rodents, and proper storage. Ahigh grain mois- combinations, with subsequent maintenance by
ture content combined with high ambient tem- open pollination) were developed for the farm-
peratures can cause considerable damage, ers. Fi hybrids (first generation progeny of
making the product unsuitable for consump- crosses between genetically distinct parents)
tion by humans and livestock. Maize grain for with greatly increased grain yield were pro-
home consumption is either sun-dried for sev- duced in the United States and in some tropi-
eral days by hanging up whole cobs tied to- cal African countries during the early part of
gether by their leaves, or these are put in a the 20 th century. Zimbabwe, for example,
well-ventilated store or crib. Shelling (the re- adopted hybrids at that time; most other tropi-
moval of grains from the cob) is usually carried cal African countries could not produce hybrids
out by hand, although mechanical shellers are because there were no seed companies that
available. The average shelling percentage is could produce and distribute hybrid seed in
about 75%.The shelled grain is dried for a few commercial quantities. Two international re-
days and then stored in bags, tins or baskets. search institutes, IITA (International Institute
The optimum moisture content for storage is of Tropical Agriculture) and CIMMYT, estab-
12-13%, but often it is not below 18%. Small- lished breeding programmes in the region and
holder farmers generally select seed for the greatly boosted the development of improved
next crop from the last harvest. The selected cultivars. With time, many African countries
cobs are stored at home in the surrounding also established their own breeding pro-
leaves above the fireplace to prevent insect grammes and developed maize cultivars for
damage. their special needs. Germplasm from CIMMYT
236 CEREALS AND PULSES

and UTA has been widely used in these pro- Bt maize for tropical Africa, especially to con-
grammes. The demand for maize continued to trol stem borers. Industrial and academic re-
increase, thus necessitating attempts to im- search is testing transgenes capable of improv-
prove the yield of the cultivars grown by the ing grain quality, e.g. by increasing the lysine
farmers. Hybrid seed is commonly used in content. Maize was one of the earliest crops to
high-input farming with high fertilizer use and be subjected to molecular mapping; the first
adequate facilities for seed production. In molecular map was reported in 1986. Many
tropical Africa breeding methods such as re- genetic linkage maps have been constructed
current selection, inbreeding and hybridization since then, using mainly RFLP, SSR and SNP
have been used in maize breeding. Seed com- markers; maps have been integrated into a
panies are now being established in many of high-density linkage map. Quantitative trait
the countries, thereby making it possible to loci (QTLs) for a wide range of traits have been
produce hybrid seed in commercial quantities. localized, including grain yield, resistance to
In low-input farming, composite or synthetic diseases and pests, drought tolerance, and oil
cultivars may be preferable, as they permit and protein contents of the grain. Genome se-
farmers to save seed from one crop to the next, quencing of maize is difficult because of its
while their wider genetic base provides a better large size (2500 Mbp), complexity and highly
adaptation to variable growth conditions. Em- repetitive character.
phasis in maize breeding is on incorporation of Prospects Maize will continue to play a
resistance to biotic and abiotic stresses. Sev- large and important role in Africa's food pro-
eral open-pollinated cultivars and Fi hybrids duction. It is the principal staple food in large
that are resistant to one or more stress factors, parts of East and southern Africa. Although
including Striga, diseases, insect pests, less important in West and Central Africa, it is
drought and low soil N, are now available for a major source of energy in these regions, espe-
farmers in tropical Africa. 'Obatanpa', a Qual- cially in parts of Côte d'Ivoire, Ghana, Benin
ity Protein Maize (QPM) with higher contents and Nigeria. Of the cereals, maize gives the
of tryptophan and lysine, developed by maize highest yield per man-hour invested; it is usu-
breeders in Ghana, is widely grown in West ally the first crop to be harvested for food dur-
and Central Africa, and also in some East and ing the hunger period of the year; it is easy to
southern African countries. Compared to other grow as sole crop or intercropped with other
crops, the adoption level of improved maize crops; it is easy to harvest, it does not shatter
cultivars is relatively high in tropical Africa. It and is not liable to bird damage. Many maize
has been estimated that 35-50% of the maize technologies have been developed in national
area in tropical Africa is planted with improved and international research stations in Africa
open-pollinated cultivars and Fi hybrids, but but most of these are yet to be adopted by the
large differences exist between countries. farmers. This has led to a large yield gap be-
A range of techniques is available for in-vitro tween the researchers' and the farmers' fields.
regeneration of maize, using callus tissue, cell High-quality seed is in short supply because
suspensions, excised plant parts and immature most countries, especially in West and Central
embryos. Genetic transformation of maize is Africa, do not have adequately organized seed
possible using Agrobacterium-mediated and sectors. Farmers also need improved access to
biolistic methods, but the efficiency of the lat- fertilizers, crop protection chemicals and other
ter is relatively low. Genetic transformation of inputs. Cultivars and cropping techniques that
maize is now routinely and commercially em- fit well into the prevailing cropping systems
ployed, although genotype-independent tech- are now being developed in collaboration with
niques are not yet available. In 2001 the world farmers in what is called participatory plant
area under transgenic maize was estimated at breeding.
9.8 million ha, and maize was only second to Major references Abalu, 2001; Badu-Apraku
soya bean in area planted to transgenic crops. et al. (Editors), 2003b; Byerlee & Eicher (Edi-
The main transgenic maize types planted are tors), 1997; UTA (International Institute of
Bt maize (maize that possesses genes from Tropical Agriculture), 1992; Kling & Ed-
Bacillus thuringiensis conferring resistance to meades, 1997;Koopmans, ten Have & Subandi,
the European maize borer, Ostrinia nubilalis), 1996; Kulp, K. & Ponte, J.G. (Editors), 2000;
herbicide-tolerant maize or types with both Ristanovic, 2001;Smith, Betrân &Runge (Edi-
traits. Bt maize has been commercially re- tors), 2004;White &Johnson (Editors), 2003.
leased in South Africa. CIMMYT is working on Other references Aljanabi, 2001; Badu-
 ZEA 237

Apraku et al., 2003a; Bankole &Adebanjo, 2003;

Blackie, 1994; Buddenhagen & Bosque-Pérez,
1999; Burkill, 1994; Cope, 1995; de Vries &
Toenniessen, 2001; Dowswell, Paliwal &
Cantrell, 1996; Evenson & Gollin (Editors),
2003;James, 2002;Marchand et a l , 1997;Neu-
winger, 2000; Phillips, 1995; Polaszek (Editor),
1998; Rybicki & Pietersen, 1999; Sprague &
Dudley, 1988; Taba, 1997; USDA, 2004; van
Wyk &Gericke, 2000.
Sources of illustration Koopmans, ten Have
& Subandi, 1996.
Authors B.Badu-Apraku &M.A.B. Fakorede
Based on PROSEA 10: Cereals.
238 CEREALS AND PULSES
 239

Cereals and pulses with other primary use

List of species in other commodity groups (parenthesis), which are used also as ce-
real or pulse. Synonyms are given in the indented lines (9 November 2005).
The names listed here have not been repeated in the Index of scientific plant names
(p. 289).

Abrus precatorius (medicinal plants) Brachystegia eurycoma (timbers)

Acacia macrostachya (medicinal plants) Brachystegia nigerica (timbers)
Acacia polyacantha (essential oils and exu- Bussea massaiensis (timbers)
dates) Bussea occidentalis (timbers)
Acacia campylacantha Cajanus scarabaeoides (auxiliary plants)
Acacia Senegal (essential oilsand exudates) Atylosa scarabaeoides
Mimosa Senegal Calpocalyx aubrevillei (timbers)
Senegalia Senegal Calpocalyx breuibracteatus (timbers)
Acacia tortilis (forages) Canavalia africana (vegetables)
Acacia spirocarpa Canavalia virosa
Acacia raddiana Canavalia ensiformis (vegetables)
Adenanthera pavonina (timbers) Canavalia gladiata (vegetables)
Afzelia pachyloba (timbers) Cathormion altissimum (medicinal plants)
Afzelia parviflora (timbers) Albizia altissima
Afzelia bracteata Pithecellobium altissimum
Amaranthus cruentus (vegetables) Cenchrus ciliaris (forages)
Amaranthus paniculatus Cenchrus setigerus (forages)
Amaranthus sanguineus Chenopodium album (vegetables)
Amaranthus hybridus Chenopodium murale (vegetables)
Amaranthus graecizans (vegetables) Chloris lamproparia (forages)
Amaranthus angustifolius Combretum aculeatum (fibres)
Amaranthus silvestris Cordyla pinnata (auxiliary plants)
Amaranthus hypochondriacus (vegetables) Cyamopsis tetragonoloba (forages)
Amblygonocarpus andongensis (timbers) Dactyloctenium aegyptium (forages)
Amblygonocarpus schweinfurthii Dactyloctenium giganteum (forages)
Anthephora nigritana (forages) Daniellia oliveri (essential oils and exudates)
Anthephora pubescens (forages) Delonix elata (ornamentals)
Anthephora hochstetteri Detarium microcarpum (medicinalplants)
Anthonotha macrophylla (timbers) Detarium senegalense (timbers)
Macrolobium macrophyllum Dichrostachys cinerea (medicinal plants)
Baikiaea insignis (ornamentals) Dichrostachys glomerata
Baphia nitida (dyes and tannins) Dichrostachys nyassana
Boerhavia repens (medicinal plants) Digitaria ciliaris (forages)
Bombax rhodognaphalon (timbers) Digitaria debilis (forages)
Bombax stolzii Digitaria delicatula (forages)
Brachiaria comata (forages) Digitaria leptorhachis (forages)
Brachiaria kotschyana Digitaria longiflora (forages)
Brachiaria jubata (forages) Digitaria nuda (forages)
Brachiaria fulva Dioclea reflexa (medicinal plants)
Brachiaria lata (forages) Dolichos trilobus (auxiliary plants)
Urochloa lata Echinochloa colona (forages)
Brachiaria ramosa (forages) Panicum colonum
Brachiaria serrifolia (forages) Echinochloa crus-galli (forages)
Brachiaria stigmatisata (forages) Echinochloa crus-pavonis (forages)
Brachiaria villosa (forages) Echinochloa pyramidalis (forages)
Brachiaria distichophylla Eleusine indica (forages)
240 CEREALS AND PULSES

Entada gigas (fibres) Mucuna poggei (dyes and tannins)

Entada scandens Mucuna rubro-aurantiaca
Entada rheedei (fibres) Mucuna pesa
Entada pursaetha Mucuna pruriens (auxiliary plants)
Enteropogon prieurii (forages) Mucuna cochinchiniensis
Chloris prieurii Mucuna aterrima
Eragrostis cilianensis (forages) Mucuna nivea
Eragrostis megastachya Mucuna sloanei (dyes and tannins)
Eragrostis ciliaris (forages) Nymphaea lotus (carbohydrates)
Eragrostis curvula (forages) Oxytenanthera abyssinica (timbers)
Eragrostis minor (forages) Oxytenanthera borzii
Eragrostis pooides Panicum fluviicola (forages)
Eragrostis pilosa (forages) Panicum aphanoneurum
Eragrostis tenella (forages) Panicum humile (forages)
Eragrostis tremula (forages) Panicum walense
Eragrostis turgida (forages) Panicum maximum (forages)
Eriochloa fatmensis (forages) Panicum jumentorum
Eriochloa nubica Panicum pansum (forages)
Eriochloa acrotricha Panicum kerstingii
Eriosema macrostipulum (carbohydrates) Panicum subalbidum (forages)
Eriosema erectum Panicum longijubatum
Erythrina variegata (auxiliary plants) Parkia bicolor (timbers)
Erythrina indica Parkia biglobosa (spices and condiments)
Faidherbia albida (auxiliary plants) Mimosa biglobosa
Acacia albida Parkia africana
Gilbertiodendron dewevrei (timbers) Parkia clappertoniana
Macrolobium dewevrei Parkia filicoidea
Gilletiodendron glandulosum (auxiliary Parkia filicoidea (timbers)
plants) Parkinsonia aculeata (auxiliary plants)
Glinus lotoides (medicinal plants) Paspalum scrobiculatum (forages)
Guibourtia coleosperma (timbers) Paspalum orbiculare
Guibourtia ehie (timbers) Paspalum polystachyum
Hyparrhenia nyassae (fibres) Paspalum lamprocaryon
Indigofera cordifolia (forages) Pennisetum unisetum (forages)
Intsia bijuga (timbers) Beckeropsis uniseta
Afzelia bijuga Piliostigma reticulatum (fibres)
Ipomoea eriocarpa (vegetables) Bauhinia reticulata
Ipomoea hispida Piliostigma thonningii (fibres)
Ipomoea sessiliflora Bauhinia thonningii
Ischaemum afrum (forages) Polygala butyracea (fibres)
Ischaemum rugosum (forages) Prosopis africana (timbers)
Kyllinga alba (essential oils and exudates) Prosopis glandulosa (auxiliary plants)
Lablab purpureus (vegetables) Psophocarpus scandens (vegetables)
Lablab niger Psophocarpus longepedunculatus
Lablab vulgaris Psophocarpus tetragonolobus (vegetables)
Dolichos lablab Pterocarpus santalinoides (timbers)
Lantana camara (medicinal plants) Pueraria phaseoloides (auxiliary plants)
Lasiurus scindicus (forages) Requienia obcordata (forages)
Lasiurus hirsutus Tephrosia obcordata
Leptothrium senegalense (forages) Rottboellia cochinchinensis (forages)
Leucaena leucocephala (auxiliary plants) Rottboellia exaltata
Leucaena glauca Saccharum spontaneum (auxiliary plants)
Limeum viscosum (forages) Sacciolepis africana (forages)
Lotus arabicus (forages) Schotia brachypetala (timbers)
Macroptilium lathyroides (forages) Senna obtusifolia (vegetables)
Monopetalanthus pteridophyllus (timbers) Cassia obtusifolia
 CEREALS AND PULSES WITH OTHER PRIMARY USE 241

Sesamum indicum (vegetable oils) Sporobolus festivus (forages)

Sesamum orientale Sporobolus pyramidalis (forages)
Sesbania rostrata (auxiliary plants) Sporobolus indicus
Sesbania sesban (auxiliary plants) Sporobolus spicatus (auxiliary plants)
Sesbania aegyptiaca Sporobolus virginicus (auxiliary plants)
Setaria geminata (forages) Stenotaphrum secundatum (auxiliary plants)
Paspalidium geminatum Sterculia africana (fibres)
Setaria palmifolia (forages) Sterculia mhosya (medicinal plants)
Setaria pumila (forages) Sterculia quinqueloba (timbers)
Setaria pallide-fusca Sterculia rhynchocarpa (medicinal plants)
Setaria sphacelata (forages) Stipagrostis pungens (forages)
Setaria anceps Stipagrostis uniplumis (forages)
Setaria aurea Tamarindus indica (fruits)
Setaria verticillata (forages) Tetrapleura tetraptera (medicinal plants)
Sorghum arundinaceum (forages) Themeda triandra (forages)
Sorghum aethiopicum Tribulus terrestris (forages)
Sorghum verticilliflorum Trigonella foenum-graecum (spices and con-
Sorghum virgatum diments)
Sorghum halepense (forages) Triplisomeris explicans (timbers)
Sorghum purpureo-sericeum (forages) Anthonotha explicans
Sphenostylis marginata (carbohydrates) Vigna angivensis (carbohydrates)
Sphenostylis erecta Vigna trilobata (auxiliary plants)
Sphenostylis schweinfurthii (carbohydrates) Xeroderris stuhlmannii (timbers)
Sphenostylis stenocarpa (carbohydrates) Ostryoderris stuhlmannii
Sphenostylis congensis Xylia evansii (timbers)
Sporobolus africanus (forages)
242 CEREALS AND PULSES

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 289

Index of scientific plant names

Page numbers printed in bold refer to main treatment; page numbers in italics refer to line drawings.
The species names listed in the section on 'Cereals and pulses with other primary use' (p.239) are not
repeated here.

Acacia nilotica, 173 Ctcerarietinum, 42, 43

Aegilops tauschii, 176 Cicer bijugum, AA
Amaranthus caudatus, 17, 18 Cicer chorassanicum, 44
Amaranthus caudatus subsp. caudatus, 18 Cicer cuneatum, 44
Amaranthus caudatus subsp. mantegazzianus, 19 Cicer echinospermum, 44
Amaranthus cruentus, 17, 18, 19 CicerJudaicum, A4
Amaranthus hybridus, 18 Cicerpinnatifidum, 44
Amaranthus hybridus L. subsp. quitensis, 17 Cicer reticulatum, 44
Amaranthus hypochondriacus, 18, 19 Ciceryamashitae, AA
Amaranthus mantegazzianus, 19 Citrullus lanatus, 11
Anogeissus leiocarpa, 173 Coix lacryma-jobi, 46, 47
Arachis duranensis, 23 Coix lacryma-jobi var. lacryma-jobi, 48
Arachis hypogaea, 20, 22 Coix lacryma-jobi var. ma-yuen, 48
Arachis hypogaea subsp. fastigiata, 23 Cordeauxia edulis, 49, 50
Arachis hypogaea subsp. hypogaea, 23 Craibia brownii, 52
Arachis ipaensis, 23 Craibia elliottii, 52
Arachis monticola, 23 Crotalaria karagwensis, 52
Arachis stenosperma, 23 Crotalaria lachnophora, 53
Arachis villosulicarpa, 23 Crotalaria lugardiorum, 52
Avena abyssinica, 28, 31 Digitaria barbinodis, 55
Avena barbata, 28 Digitaria ciliaris, 55
Avena byzantina, 30, 31 Digitaria cruciata, 55
Avena fatua, 30, 31 Digitaria exilis, 34, 54, 55, 58
Avena sativa, 28, 30 Digitaria exilis var. densa, 56
Avena sterilis, 29, 30, 31,32 Digitaria exilis var. gracilis, 56
Avena vaviloviana, 28 Digitaria exilis var. mixta, 56
Azadirachta indica, 38, 132, 212, 227 Digitaria exilis var. rustica, 56
Bauhinia esculenta, 187 Digitaria exilis var. stricta, 56
Bauhinia fassoglensis, 190 Digitaria fuscescens, 55
Bauhinia kirkii, 190 Digitaria iburua, 55, 57
Bauhinia macrantha, 33 Digitaria leptorhachis, 55
Bauhinia petersiana, 33 Digitaria longiflora, 54, 55
Bauhinia petersiana subsp. macrantha, 33 Digitaria nuda, 55
Bauhinia petersiana subsp.petersiana, 33 Digitaria sanguinalis, 55
Brachiaria deflexa, 34, 55, 63 Digitaria ternata, 55, 57, 58
Brachiaria deflexa var. sativa, 34 Dolichos biflorus, 102
Brachiaria ramosa, 34 Dolichos sesquipedalis, 224
Cajanus acutifolius, 39 Dolichos uniflorus, 102
Cajanus albicans, 39 Echinochloa colona, 60
Cajanus cajan, 35, 3ff, 53 Echinochloa obtusiflora, 58
Cajanus cajanifolius, 39 Echinochloa pyramidalis, 60
Cajanus indicus, 35 Echinochloa scabra, 59
Cajanus kerstingii, 37 Echinochloa stagnina, 59
Cajanus platycarpus, 39 Eleusine africana, 62
Cajanus scarabaeoides, 37, 39 Eleusine coracana, 60, ff2, 123, 163
Cenchrus biflorus, 40, 42 Eleusine coracana subsp. africana, 62
Cenchrus prieurii, 41 Eleusine indica, 63
Cicer anatolicum, 44 Eleusine indica subsp. coracana, 60
290 CEREALS AND PULSES

Eragrostis abyssinica, 68 Lupinus angustifolius, 97, 99

Eragrostis aethiopica, 65 Lupinus graecus, 98
Eragrostis annulata, 65 Lupinus termis, 98
Eragrostis cilianensis, 66, 69 Macrotyloma geocarpum, 99, 700
Eragrostis ciliaris, 69 Macrotyloma geocarpum var. geocarpum, 101
Eragrostis curvula, 68, 69, 72 Macrotyloma geocarpum var. tisserantii, 101
Eragrostis cylindriflora, 69 Macrotyloma uniflorum, 102, 205
Eragrostis denudata, 66 Macrotyloma uniflorum var. benadirianum,
Eragrostis gangetica, 69 103
Eragrostis nindensis, 66 Macrotyloma uniflorum var. stenocarpum, 103
Eragrostis pilosa, 65, 68, 69, 72 Macrotyloma uniflorum var. uniflorum, 103
Eragrostis pilosa subsp. abyssinica, 68 Macrotyloma uniflorum var. verrucosum, 103
Eragrostis plana, 67 Mucuna gigantea, 104
Eragrostis tef, 45, 68, £3, 86 Mucuna gigantea subsp. quadrialata, 105
Eragrostis tremula, 69 Mucuna longipedicellata, 104
Eragrostis turgida, 69 Mucuna quadrialata, 104
Fagopyrum esculentum, 72, 74 Ocimum basilicum, 217
Fagopyrum homotropicum, 76 Oryza oarireü, 105, 106, 108, 109, 110, 111,
Fagopyrum tataricum, 73, 76 112, 115, 119
Glycine hispida, 76 Oryza breviligulata, 105, 106
Glycine max, 76, 75,214 Oryzaglaberrima, 106, 207, 111, 112, 113,115,
Glycine max subsp. max, 79 119, 120
Glycine max subsp. so/a, 79 Oryza longistaminata, 60, 106, 108, 109, 110,
Glycine soja, 79 112, 115, 119, 120
Glycine subterranea, 213 Oryza nivara, 115
Guizotia abyssinica, 71 Oryza punctata, 111, 119
Harpagocarpus snowdenii, 74 Oryza rufipogon, 115
Hordeum sativum, 82 Oryza sativa, 106, 107, 108, 109, 110, 111, 112,
Hordeum spontaneum, 84
Hordeum vulgare, 82, 54 Oryza stapfii, 105
Hordeum vulgare subsp. spontaneum, 84 Panicum kalaharense, 120
Hordeum vulgare subsp. vulgare, 84 Panicum laetum, 121
Kerstingiella geocarpa, 99 Panicum miliaceum, 61,122, 223
Kerstingiella tisserantii, 101 Panicum miliaceum subsp. miliaceum, 124
Lathyrus amphicarpos, 89 Panicum miliaceum subsp. ruderale, 124
Lathyrus aphaca, 89 Panicum turgidum, 126
Lathyrus cicera, 87, 89 Parkia biglobosa, 214
Lathyrus odoratus, 89 Pennisetum americanum, 128
Lathyrus pseudocicera, 89 Pennisetum americanum subsp. monodii, 129
Lathyrus sativus, 43,87, 55 Pennisetum americanum subsp.
Leresculinaris, 91, 52 stenostachyum, 129
Lewsculinaris subsp. culinaris, 93 Pennisetum dalzielii, 129
Leresculinaris subsp. odemensis, 93 Pennisetum fallax, 129
Leresculinaris subsp. orientalis, 93 Pennisetum glaucum, 61, 123, 128, 225, 163
Leres culinaris subsp. tomentosus, 93 Pennisetum purpureum, 129
Lereservoides, 93, 95 Pennisetum sieberianum, 129
Leresesculenta, 91 Pennisetum spicatum, 128
Leres lamottei, 93 Pennisetum stenostachyum, 129
Leresnigricans, 93, 95 Pennisetum typhoides, 128
Leresorientalis, 93 Pennisetum violaceum, 128, 129
Leres tomentosus, 93 Phaseolusaconitifolius, 200
Limeum indicum, 96 Phaseolus acutifolius, 133, 235, 139, 148, 150,
Limeum obovatum, 96 151
Lupinus albus, 96 Phaseolus acutifolius var. acutifolius, 135
Lupinus albus subsp. albus, 98 Phaseolus acutifolius var. latifolius, 135
Lupinus albus subsp.graecus, 98 Phaseolus acutifolius var. tenuifolius, 135
 INDEX OF SCIENTIFIC PLANT NAMES 291

Phaseolus adenanthus, 203 Sorghum vulgare, 169

Phaseolus angularis, 204 Sporobolus fimbriatus, 175
Phaseolus aureus, 208 Sporobolus panicoides, 176
Phaseolus calcaratus, 218 Tephrosia vogelii, 53
Phaseolus coccineus, 137, 138, 148, 150 Trigonella foenum-graecum, 68
Phaseolus costaricensis, 139 Triticum aestivum, 176, 275, 184
Phaseolus dumosus, 139 Triticum aethiopicum, 184
Phaseolus jaliscanus, 145 Triticum dicoccon,183, 185
Phaseolus lunatus, 136, 139, 141, 142, 223 Triticum durum, 183, 185
Phaseolus lunatus var. lunatus, 143 Triticum monococcum, 179, 184
Phaseolus lunatus var. Silvester, 143 Triticum polonicum, 185
Phaseolus maculatus, 145 Triticum spelta, 176, 179
Phaseolus mungo, 206 Triticum turgidum, 176, 177, 179, 183, 254
Phaseolus polyanthus, 139 Triticum urartu, 183
Phaseolus polystachyus, 145 Triticum vulgare, 176
Phaseolus radiatus, 208 Tylosema esculentum, 187,2&S
Phaseolus salicifolius, 145 Tylosema fassoglense, 189, 190
Phaseolus vulgaris, 134, 137, 139, 140, 146, Urochloa brachyura, 192
247, 157, 223 Urochloa mosambicensis, 191, 193
Pisum fulvum, 155 Urochloa trichopus, 192, 193
Pisum sativum, 31,90, 151, 253, 223 Vatovaea pseudolablab, 194
Plumbago zeylanica, 61 Vicia faba, 155, 195, 23S
Pseudobrachiaria deflexa, 34 Vicia galilaea, 197
Secale africanum, 160 Vicia hirsuta, 200
Secale cereale, 158, 255 Vicia hyaeniscyamus, 197
Secale cereale subsp. ancestrale, 160 Vicia Zens, 91
Secale cereale subsp. cereale, 160 Vicia monantha, 197
Secale strictum, 161 Vicia narbonensis, 197
Secale strictum subsp. africanum, 160 Viciapaucifolia, 197
Sesbania rostrata, 117 Vicia sativa, 197
Setaria finita, 163 Vicia villosa, 197
Setaria italica, 61,122, 162, 2£3 Vi£7iaaconitifolia, 200, 202, 205, 207, 210, 220
Setaria italica subsp. viridis, 164 Vig/ia adenantha, 203
Setaria pumila, 163 Vigna angularis, 201,204, 205, 207, 210, 220
Setaria sphacelata, 163 Vigna angularis var. angularis, 205
Setaria verticillata, 163 Vigna angularis var. nipponensis, 205
Setaria viridis, 162, 164 Vigna calcarata, 218
Sorghum xdrummondii, 170 Vigna minima, 220
Sorghum ankolib, 169
Vi£7ia mn/igo, 201,205, 206, 207, 210, 212, 220
Sorghum arundinaceum, 170
Vigna mungo var. mungo, 207
Sorghum aterrimum, 170
Vigna mungo var. silvestris, 207
Sorghum bicolor, 165, 2£5
Vigna mungo var. viridis, 207
Sorghum bicolor subsp. arundinaceum, 170
Vigna radiata, 201,205, 206, 207, 208, 220,
Sorghum bicolor subsp. bicolor, 169, 170
Sorghum bicolor subsp. drummondii, 170 220
Sorghum bicolor subsp. verticilliflorum, 170 Vigna radiata var. radiata, 210
Sorghum caudatum, 169 Vigna radiata var. setulosa, 210
Sorghum cernuum, 169 Vigna radiata var. sublobata, 210
Sorghum dochna, 169 Vigna sesquipedalis, 224
Sorghum durra, 169 Vigna sinensis, 221
Sorghum halepense, 169 Vigna subterranea, 213, 224
Sorghum membranaceum, 169 Vigna subterranea var. spontanea, 215
Sorghum nigricans, 169 Vigna subterranea var. subterranea, 215
Sorghum propinquum, 169 Vigna trilobata, 210
Sorghum subglabrescens, 169 Vigna umbellata, 201,205, 207, 210, 218, 225
Sorghum sudanense, 170 Vigna umbellata var. gracilis, 220
Vigna umbellata var. umbellata, 220
292 CEREALS AND PULSES

Vigna unguiculata, 221, 223 Voandzeia subterranea, 213

Vigna unguiculata subsp. unguiculata, 224 Zea mays, 229, 231
Vigna vexillata, 228
 293

Index ofvernacular plant names

Abyssinian oat, 28 Cacahuète, 20 Earth-nut, 20

Adlay, 46 Camel's foot, 33, 187 Egyptian lupin, 96
Adzuki bean, 204 Capim chorâo, 67 Eleusine, 60
African millet, 60 Capim colonial, 60 Eragrostis annelé, 65
African rice, 106 Capim teff, 67 Eragrostis d'Afrique du Sud, 67
African spinach, 17 Caranga, 20 Eragrostis vivace, 66
Amarante-grain, 17 Case knife bean, 137 Ers velu, 200
Amarante- de cauda, 17 Catjang, 221 Erva dos rosârios, 46
Ambérique, 208 Catjang cowpea, 221 Ervanço, 42
Ambrevade, 35 Cattail millet, 128 Ervilha, 151
Amendoim, 20 Cauda de raposa, 17 Ervilha de Angola, 35
Animal fonio, 34 Caupi, 221 Ervilha do Congo, 35
Arachide, 20 Centeio, 158 Ervilha torta, 151
Arroz, 112 Cevada, 82 Ethiopian oat, 28
Asian rice, 112 Chicharo, 87 Faba bean, 195
Asiatic rice, 112 Chicharo comun, 87 Fagópiro, 72
Asparagus bean, 221 Chickling pea, 87 False signal grass, 34
Aveia, 28 Chickling vetch, 87 Famine grass, 176
Aveia-amarela, 28 Chickpea, 42 Fava, 195
Avoine, 28 China pea, 221 Fava caranguejo, 203
Avoine cultivée, 28 Chingando, 33 Favalinha, 102
Avoine d'Abyssinie, 28 Cigeräo, 200 Faveira, 195
Azuki bean, 204 Climbing mountain bean, 218 Feijào, 146
Bambara groundnut, 213 Coffee neat's foot, 33 Feijào adzuki, 204
Bambarra groundnut, 213 Common bean, 146 Feijào arroz, 218
Barley, 82 Common lentil, 91 Feijào chicote, 221
Bauhinia rampant, 190 Common millet, 122 Feijào da China, 221
Beech wheat, 72 Common oat, 28 Feijào da Espanha, 137
Bengal gram, 42 Common urochloa, 191 Feijào de Lima, 141
Black acha, 57 Common wheat, 176 Feijào de metro, 221
Black fonio, 57 Congo pea, 35 Feijào escarlata, 137
Black gram, 206 Coracan, 60 Feijào espadinho, 141
Black-eye bean, 221 Corn, 229 Feijào espargo, 221
Black-eye pea, 221 Cornille, 221 Feijào favona, 141
Blé, 176 Cowpea, 221 Feijào frade, 221
Blé des Incas, 17 Cram-cram, 40 Feijào frade alfange, 221
Blé dur, 183 Creeping bauhinia, 190 Feijào guandu, 35
Blé noir, 72 Crotalaire à toison, 53 Feijào maeundi, 221
Blé tendre, 176 Culita, 102 Feijào miûdo, 221
Bombay cowpea, 221 Desert grass, 126 Feijào mungo verde, 208
Bourgou, 59 Desert panic, 121 Feijào tepari, 133
Bread wheat, 176 Dew bean, 200 Feijào trepador, 137
Brède malabar, 17 Dew gram, 200 Feijào urida, 206
Bredo, 17 Discipline des religieux, 17 Feijoeiro, 146
Broad bean, 195 Dohi, 99 Feijoeiro de lagartixa, 102
Buckwheat, 72 Doï, 99 Fève, 195
Bulrush millet, 128 Dolique asperge, 221 Fève de(s) marais, 195
Burgu grass, 59 Dolique mongette, 221 Fève de Kandale, 99
Burny bean, 104 Dropseed, 175 Fèverole, 195
Bushveld signal grass, 191 Durra, 165 Fèvette, 195
Butter bean, 141 Durum wheat, 183 Field bean, 195
Cacahouète, 20 Earth pea, 213 Field pea, 151
294 CEREALS AND PULSES

Finger millet, 60 Horse bean, 195 Marble pea, 221

Fonio, 54 Horse grain, 102 Massambala, 165
Fonio à grosses graines, 34 Horse gram, 102 Massango liso, 128
Fonio blanc, 54 Hungry rice, 54 Mat, 200
Fonio noir, 57 Ibourou, 57 Mat bean, 200
Fonio sauvage, 121 Iburu, 57 Mbaazi, 35
Foxtail, 17 Inca wheat, 17 Mchetez, 111
Foxtail millet, 162 Indian corn, 229 Mchicha, 17
Fringed dropseed, 175 Indian sandbur, 40 Mchooko, 208
Froment, 176 Indian spinach, 17 Mchooko mweusi, 206
Garbanzo, 42 Italian millet, 162 Mchoroko, 208
Garden pea, 151 Jataco, 17 Mdengu, 42, 91
Gemsbok bean, 187 Jégé, 34 Mfiwi, 141
Geocarpa groundnut, 99 Jinguba de Cabambe, 213 Mharagwe, 146
German millet, 162 Job's tears, 46 Mhindi, 229
Gesse, 87 Jugo bean, 213 Mil, 128
Gesse blanche, 87 Kalahari white bauhinia, 33 Mil à chandelle, 128
Gesse commune, 87 Karanga, 20 Mil pénicillaire, 128
Golden gram, 208 Kersting's groundnut, 99 Mil rouge, 60
Gourgane, 195 Kibi, 122 Milheto, 128
Grain amaranth, 17 Kidney bean, 146 Milho, 229
Grain de cheval, 102 Kimanga, 162 Milho de canario, 122
Gram, 42 Kolo rassé, 34 Milho miûdo, 122, 165
Grào de bico, 42 Koracan, 60 Milho painço, 162
Grass pea, 87 Kulthi, 102 Milho painço de Itâlia, 162
Gravanço, 42 Lâgrimas de Job, 46 Milho preto, 128
Great millet, 165 Lâgrimas deNossa Senhora, 46 Milho zaburro, 128
Green gram, 208 Larmes de Job, 46 Millet commun, 122
Gros fonio, 34 Larmilles, 46 Millet d'Italie, 162
Gros mil, 165 Lentil, 91 Millet de Guinée, 34
Ground bean, 99 Lentilha, 91 Millet des oiseaux, 162
Groundnut, 20 Lentille, 91 Mjugu nyasa, 20
Guinea corn, 165 Lentille d'Espagne, 87 Mkunde, 221
Guinea millet, 34 Lentille de terre, 99 Mnjugu nyasa, 20
Hairy tare, 200 Lentillon, 91 Moncos de peru, 17
Hairy vetch, 200 Liane cadoque, 104 Monkey nut, 20
Haricot, 146 Liane caiman, 104 Morama bean, 187
Haricot à l'œil noir, 221 Lima bean, 141 Mort aux rats, 104
Haricot adzuki, 204 Long-awned water grass, 59 Moth bean, 200
Haricot bean, 146 Love-lies-bleeding, 17 Moth gram, 200
Haricot commun, 146 Luco, 60 Mpunga, 112
Haricot d'Espagne, 137 Lupin, 96 Mtama, 165
Haricot de Lima, 141 Lupin blanc, 96 Mtasbihi, 46
Haricot dolique, 221 Macaroni wheat, 183 Mtasubihu, 46
Haricot doré, 208 Madagascar bean, 141 Mtera, 104
Haricot mat, 200 Madras gram, 102 Muhindi, 229
Haricot mungo, 208 Maïs, 229 Multiflora bean, 137
Haricot papillon, 200 Maize, 229 Mung bean, 208
Haricot riz, 218 Makunde, 221 Mungo, 208
Haricot tépari, 133 Mancara de Bijagó, 213 Mwele, 128
Haricot urd, 206 Mandinka rice, 105 Mwimbi, 60
Haricot-kilomètre, 221 Mandobi, 20 Nachenim, 60
Haze, 121 Mange-tout, 151 Navy bean, 146
Herbe à chapelets, 46 Manne noire, 57 Ngano, 176
Hippo grass, 59 Marama, 187 Niébé, 221
Hog millet, 122 Marama bean, 187 Njegere, 151
 INDEX OF VERNACULAR PLANT NAMES 295

Njengere, 151 Rice bean, 218 Tépari, 133

Njugu mawe, 213 Ring windgrass, 65 Tepary bean, 133
Oat, 28 Ringed lovegrass, 65 Texas bean, 133
Oats, 28 Riz, 112 Tick bean, 195
Orge, 82 Riz africain, 106 Tiny vetch, 200
Paddy, 112 Riz asiatique, 112 Tough lovegrass, 67
Painço, 162 Riz de Casamance, 106 Tremoceiro, 96
Panis, 162 Riz de marais, 105 Tremoceiro branco, 96
Pea, 151 Riz sauvage, 105 Tremoceiro da Beira, 96
Pea pod, 151 Riz sauvage annuel, 105 Tremoço, 96
Peanut, 20 Riz sauvage vivace, 110 Trigo, 176
Pearl millet, 128 Riz vivace, 110 Trigo duro, 183
Perennial dropseed grass, 175 Roseau à miel du Niger, 59 Trigo mole, 176
Perennial lovegrass, 66 Roseau sucré, 59 Trigo rijo, 183
Petit mil, 54, 128 Runner bean, 137 Trigo sarraceno, 72
Petit pois, 151 Rye, 158 Trigo-mourisco, 72
Petit-pois, 151 Sabi grass, 191 Turgid panic grass, 126
Pigeon pea, 35 Sarrasin, 72 Ulezi, 60
Pistache de terre, 20 Scarlet runner bean, 137 Urad bean, 206
Pois, 151 Sea bean, 104 Urd, 206
Pois bambara, 213 Seigle, 158 Urd bean, 206
Pois cajan, 35 Self-sown rice, 105 Vesce hérissée, 200
Pois carré, 87 Shayiri, 82 Vesceron, 200
Pois chiche, 42 Sincho, 87 Voandzou, 213
Pois d'Angole, 35 Snap pea, 151 Voème, 221
Pois de terre, 213 Snow pea, 151 Wadi rice, 111
Pois du Cap, 141 Soja, 76 Wether lovegrass, 66
Pois gourmand, 151 Sorgho, 165 Wheat, 176
Pois mangetout, 151 Sorgho des teinturiers, 165 White fonio, 54
Pois marron, 203 Sorgho rouge, 165 White lupin, 96
Pois savon, 141 Sorghum, 165 White pea, 87
Pois sec, 151 Sorgo, 165 Wild bean, 203
Pois souche, 141 South-African lovegrass, 67 Wild coffee bean, 33
Pois yeux noirs, 221 Soya, 76 Wild fonio, 121
Proso millet, 122 Soya bean, 76 Wild rice, 105, 110
Queue de renard, 17 Soybean, 76 Yard-long bean, 221
Red bean, 218 Sprawling bauhinia, 190 Yeheb, 49
Red gram, 35 Sugar pea, 151 Yeheb bush, 49
Red rice, 106, 110, 111 Tef, 68 Yeheb nut, 49
Red-hot cattail, 17 Teff, 68
Rice, 112 Teff grass, 68
296 CEREALS AND PULSES

pROTA
Plant Resources ofTropical Africa
•*? '"7 Ressources végétales del'Afrique tropicale

•O

PROTA in short

The Plant Resources of Tropical Africa (PROTA) programme was initiated in 2000
and developed into an international partnership of 11institutions in 11 countries
during the Preparatory Phase 2000-2003. Since 19 February 2003, PROTA operates
as an international foundation domiciled in Wageningen, Netherlands.

PROTA is a major 'information brokerage and knowledge repatriation' programme.

The objectives are to bring the 'world literature' on the useful plants of Tropical Af-
rica, now accessible only to the resourceful happy few, into the (African) public do-
main, and contribute to greater awareness and sustained use of the plants, with due
respect for traditional knowledge and intellectual property rights. PROTA will de-
scribe the estimated 7,000 useful plants during the Implementation Phase 2003-
2012. The information carriers will be freely accessible Web databases
(www.prota.org), a low-price Handbook and CD-Rom series featuring 16 Commodity
groups, and Special Products per commodity group for rural development, educa-
tion, research and policy actors (all in English and French).

PROTA 1: Cereals and pulses (2006) PROTA 9: Auxiliary plants

PROTA 2: Vegetables (2004) PROTA 10 Fuel plants
PROTA 3: Dyes and tannins (2005) PROTA 11 Medicinal plants
PROTA 4: Ornamentals PROTA 12 Spices and condiments
PROTA 5: Forages PROTA 13 Essential oils and exudates
PROTA 6: Fruits PROTA 14 Vegetable oils
PROTA 7: Timbers PROTA 15 Stimulants
PROTA 8: Carbohydrates PROTA 16 Fibres

PROTA, P.O.Box 341,6700 AH Wageningen, Netherlands (www.prota.org)

 297

CTA
partageons lesconnaissances au profit des communautés rurales

sharing knowledge, improving rural livelihoods

CTA in short

The Technical Centre for Agricultural and Rural Cooperation (CTA) was established
in 1983 under the Lomé Convention between the ACP (African, Caribbean and Pa-
cific) Group of States and the European Union Member States. Since 2000, it has
operated within the framework of the ACP-EC Cotonou Agreement.

CTA's tasks are to develop and provide services that improve access to information
for agricultural and rural development, and to strengthen the capacity of ACP coun-
tries to produce, acquire, exchange and utilise information in this area. CTA's pro-
grammes are designed to: provide a wide range of information products and services
and enhance awareness of relevant information sources; promote the integrated use
of appropriate communication channels and intensify contacts and information ex-
change (particularly intra-ACP); and develop ACP capacity to generate and manage
agricultural information and to formulate ICM strategies, including those relevant
to science and technology. CTA's work incorporates new developments in method-
ologies and cross-cutting issues such as gender and social capital.

CTA, P.O.Box 380, 6700 AJ Wageningen, Netherlands (www.cta.int)

WEST AFRICA CENTRAL AFR ICA SOUTHERN AFI

1. Cape Verde 17. S ä o Tomé et Principe 35. Malawi

2. Mauritania 18. Cameroon 36. Zambia
3. S e n e g a l 19. Chad 37. Angola
4. Gambia 20. Central African Republic 38. Namibia
5. Guinea B i s s a u 21. Equatorial Guinea 39. B o t s w a n a
6. Guinea 22. Gabon 40. Zimbabwe
7. Sierra Leone 23. Congo 41. Mozambique
8. Liberia 24. D e m o c r a t i c Republic of Congo
9. Côte d'Ivoire 25. R w a n d a INDIAN OCEAN
10. Mali 26. B u r u n d i ISLANDS
11. B u r k i n a F a s o
12. Ghana EAST AFRICA 42. Comoros
13. Togo 43. Mayotte (Fr)
14. B e n i n 27. S u d a n 31. Somalia 44. Madagascar
15. N i g e r 28. Eritrea 32. Kenya 45. S e y c h e l l e s
16. Nigeria 29. Ethiopia 33. U g a n d a 46. R é u n i o n (Fr)
30. Djibouti 34. Tanzania 47. Mauritius