Annona Species Monograph.pdf - Crops for the Future

Annona species 

Authors : 

A. C. de Q. Pinto 

M. C. R. Cordeiro 

S. R. M. de Andrade 

F. R. Ferreira 

H. A. de C. Filgueiras 

R. E. Alves 

D. I. Kinpara 

Editors: 

A. Hughes 

C. R. Clement 

N. Haq 

R.W. Smith 

J.T. Williams (Chief editor)

First published in 2005 by: 

International Centre for Underutilised Crops, University of Southampton, 

Southampton, SO17 1BJ, UK 

© 2005 International Centre for Underutilised Crops 

Printed at RPM Print and Design, West Sussex, UK 

The text in this document may be reproduced free of charge in any format 

or media without requiring specific permission. This is subject to the materials 

not being used in a derogatory manner or in a misleading context. The 

source of the material must be acknowledged as [ICUC] copyright and the 

title of the document must be included when being reproduced as part of 

another publication or service. 

Copies of this handbook, as well as an accompanying manual and factsheet, 

in English, can be obtained by writing to the address below: 

ICUC@IWMI 

127 Sunil Mawatha, Pelawatte, Battaramulla, Sri Lanka 

British Library Catalogue in Publication Data 

Annona 

1. tropical fruit trees 

i Hughes ii Clement iii Haq iv Smith v Williams 

ISBN 0854327851 

Citation: A. C. de Q. Pinto, M. C. R. Cordeiro, S. R. M. de Andrade, F. R. 

Ferreira, H. A. de C. Filgueiras, R. E. Alves and D. I. Kinpara (2005) 

Annona species, International Centre for Underutilised Crops, University 

of Southampton, Southampton, UK. 

Cover photographs: Annona fruit tree, transport and marketing, supplied 

by A. C. de Q. Pinto

DFID/FRP and DISCLAIMERS 

This publication is an output from a research project funded by the United 

Kingdom Department for International Development (DFID) for the benefit 

of developing countries. The views expressed are not necessarily those 

of DFID [R7187 Forestry Research Programme]. 

The opinions expressed in this book are those of the authors alone and do 

not imply acceptance or obligation whatsoever on the part of ICUC, 

ICRAF or IPGRI. 

MEMBERS OF THE ADVISORY COMMITTEE 

R. K. Arora 

International Plant Genetic Resources Institute (IPGRI), Office for South 

Asia, New Delhi, India 

S. Azam-Ali 

Private Consultant, UK 

C. R. Clement 

National Research Institute of Amazonia (INPA), Manaus, Brazil 

N. Haq 

International Centre for Underutilised Crops (ICUC), University of 

Southampton, Southampton, UK 

A. Hughes 

International Centre for Underutilised Crops (ICUC), University of 

Southampton, Southampton, UK 

A. J. Simons 

World Agroforestry Centre (ICRAF), Nairobi, Kenya 

R. W. Smith 

International Centre for Underutilised Crops (ICUC), UK 

P. Vantomme 

Food and Agriculture Organization of the United Nations (FAO), Rome, 

Italy 

J. T. Williams 

Board of Trustees, International Centre for Underutilised Crops (ICUC), 

UK

ICUC 

The International Centre for Underutilised Crops (ICUC) is an autonomous, nonprofit, 

scientific research and training centre. It was established in 1992 at the University 

of Southampton in the UK and has now moved to the International Water 

Management Institute (IWMI) in Sri Lanka. The Centre was established to address 

ways to increase the use of under-utilised crops for food, nutrition, medicinal and 

industrial products. The enhancement of currently under-utilised crops is a key to 

food security, to the conservation of biological diversity and to the preservation 

and restoration of fragile and degraded environments throughout the world. 

World Agroforestry Centre 

The World Agroforestry Centre (ICRAF), established in Nairobi in 1977, is an 

autonomous, non-profit research body supported by the Consultative Group on 

International Agricultural Research (CGIAR). ICRAF aims to improve human welfare 

by alleviating poverty, improving food and nutrition security and enhancing 

environmental resistance in the tropics. 

IPGRI 

The International Plant Genetic Resources Institute (IPGRI) is an international 

research institute with a mandate to advance the conservation and use of genetic 

diversity for the well-being of present and future generations. It is also a centre of 

the CGIAR. 

Also available in this series: 

Tamarind - Tamarindus indica by H.P.M. Gunasena and A. Hughes 

(ISBN 0854327274) 

Ber - Ziziphus mauritiana by O. P. Pareek (ISBN 0854327525) 

Safou - Dacryodes edulis by J. Kengue (ISBN 0854327649) 

Baobab - Adansonia digitata by M. Sidibe and J.T. Williams (ISBN 0854327762) 

Forthcoming in this series: 

Jackfruit - Artocarpus heterophyllus by N. Haq 

Mangosteen - Garcinia mangostana by M. bin Osman and A. R. Milan 

Ndjanssang - Ricinodendron heudelotii by Z. Tchoundjeu 

Monkey orange - Strychnos cocculoides by C. K. Mwamba 

Sapota - Pouteria sapota by C. Azurdia

Table of Contents 

Abbreviations ................................................................................................. i 

Acknowledgements ...................................................................................... iv 

Preface .......................................................................................................... iv 

1. Introduction............................................................................................... 1 

2. Taxonomy and Botany.............................................................................. 3 

2.1 General ............................................................................................ 3 

2.2 Specific and common names ........................................................... 3 

2.3 Botanical description....................................................................... 6 

2.3.1 Description of the genus ..................................................... 6 

2.3.2 Description of the species ................................................... 7 

3. Origin and Distribution.......................................................................... 19 

4. Major and Minor Production Areas ..................................................... 23 

4.1 Major Production Areas ................................................................ 23 

4.2 Minor Production Areas ................................................................ 26 

4.3. Demand ........................................................................................ 27 

5. Ecological Factors ................................................................................... 28 

5.1 Physiography and climate.............................................................. 28 

5.2 Soil ................................................................................................ 31 

5.3 Phenology...................................................................................... 32 

6. Properties................................................................................................. 35 

6.1 Chemical properties....................................................................... 35 

6.2 Pulp properties............................................................................... 37 

7. Uses........................................................................................................... 41 

7.1 Food products................................................................................ 41 

7.2 Industrial food uses ....................................................................... 42 

7.3 Medicinal uses............................................................................... 44 

7.4 Other uses...................................................................................... 46 

8. Genetic Resources ................................................................................... 48 

8.1 The annona genepool..................................................................... 48 

8.2 In situ conservation ....................................................................... 50 

8.3 Ex situ conservation....................................................................... 51 

9. Genetic Improvement ............................................................................. 55 

9.1 Introduction ................................................................................... 55

9.2 Cytogenetics and genetic aspects .................................................. 55 

9.3 Characteristics of annona ideotypes .............................................. 56 

9.4 Breeding programme..................................................................... 56 

9.4.1 Limiting factors and major constraints ............................. 57 

9.4.2 Breeding objectives........................................................... 59 

9.4.3 Methods and strategies...................................................... 61 

9.4.4 Selection and cultivar development .................................. 64 

9.5 Role of modern biotechnology ...................................................... 68 

9.5.1 Tissue culture.................................................................... 68 

9.5.2 Genetic transformation...................................................... 69 

9.5.3 Molecular markers ............................................................ 69 

10. Agronomy .............................................................................................. 71 

10.1 Propagation.................................................................................. 71 

10.1.1 Seed propagation............................................................. 71 

10.1.2 Vegetative propagation ................................................... 75 

10.2 Field establishment...................................................................... 85 

10.2.1 Orchard location.............................................................. 85 

10.2.2 Land preparation ............................................................. 85 

10.2.3 Time of planting.............................................................. 91 

10.2.4 Direct seeding ................................................................. 91 

10.2.5 Transplanting and spacing .............................................. 92 

10.3 Orchard management................................................................... 93 

10.3.1 Windbreaks ..................................................................... 93 

10.3.2 Pruning............................................................................ 93 

10.3.3 Orchard maintenance, intercropping and cover-cropping99 

10.3.4 Flowering, pollination and fruit set............................... 100 

10.3.5 Nutrition and fertilization.............................................. 103 

10.3.6 Irrigation ....................................................................... 111 

10.3.7 Pest and disease management ....................................... 116 

10.3.7.1 Pests.................................................................. 117 

10.3.7.2 Diseases............................................................ 122 

10.3.8 Physiological disorders ................................................. 126 

11. Harvest, Postharvest and Processing................................................. 127 

11.1 Introduction ............................................................................... 127 

11.2 Harvest ...................................................................................... 128 

11.3 Postharvest handling.................................................................. 132 

11.3.1 Physiological changes................................................... 132 

11.3.2 Handling........................................................................ 133 

11.4 Storage....................................................................................... 134 

vi

11.5 Processing.................................................................................. 136 

12. Economic Information........................................................................ 139 

12.1 Economics of production........................................................... 140 

12.1.1 Production cost, price and income ................................ 140 

12.1.2 Production, productivity and production value............. 143 

12.1.3 Social improvement ...................................................... 145 

12.2 Marketing and commercialization.................................... 146 

13. Conclusions and Research Needs....................................................... 150 

13.1 Research requirements and technology transfer ........................ 152 

13.1.1 Genetic resources and genetic improvement................. 152 

13.1.2 Propagation studies ....................................................... 153 

13.1.3 Studies on crop management ........................................ 154 

13.1.4 Postharvest and processing ........................................... 154 

13.1.5 Industrialisation and marketing..................................... 154 

References.................................................................................................. 156 

A. Common chemical compounds found in annonas............................. 192 

B. Uses of Annona species in medicine .................................................... 198 

C. Institutions and Individuals Engaged in Annona Research and 

Development .................................................................................... 203 

D. Countries and Institutions with Collections of Germplasm ............. 229 

Glossary ..................................................................................................... 247 

vii

List of Tables 

2-1. Botanical or specific, common and vernacular names and their 

synonyms of the five Annona species studied ....................................... 4 

5-1. A quick reference guide to monthly rainfall (mm) in some important 

Annona production areas ..................................................................... 34 

6-1. Chemical composition of 100g of edible pulp of cherimoya, custard 

apple, soursop and sugar apple fruits................................................... 39 

8-1. Centres of origin and diversity of some Annona species....................... 49 

8-2. Number of Annona accessions in germplasm collections around the 

world. .................................................................................................. 51 

9-1. Main characterisitics of cherimoya, custard apple and sugar apple 

ideotypes (adult plants) ....................................................................... 59 

9-2. Some selections and cultivars of cherimoya, atemoya, soursop, sugar 

apple and custard apple that are currently planted in various countries 

...................................................................................................................... 66 

10-1. Time of storage to assure 90% seed viability, time for germination, 

germination percentage, seedling age for transplanting and age if used 

for grafting........................................................................................... 71 

10-2. Seed and vegetative propagation methods, commercial 

recommendations and success for different Annona species............... 77 

10-3. Rootstock x scion compatibility and the recommended vegetative 

propagation methods for nine annona species..................................... 78 

10-4. Recommendation of NPK fertilization for cherimoya in Spain based on 

tree age after planting during the first three years of orchard 

establishment..................................................................................... 104 

10-5. Recommendation of P and K for soursop trees of different ages 

according to the amount of N fertilization applied and levels of P 2 

0 5 

and K 2 

0 in the soil analysis in the semi-arid region of Brazil............ 104 

10-6. Recommendation of P and K for sugar apple trees of different ages 

according to the amount of N fertilization applied and levels of P 2 

0 2 

and K 2 

0 in the soil analysis in the semi-arid region of Brazil............ 105 

10-7. The average normal and deficient levels of macro and micronutrients in 

leaves of cherimoya, soursop and sugar apple................................... 106 

10-8. A guide for nitrogen fertilization for adult soursop trees in different 

regions of Colombia, according to age and nutrient content in the soil 

.................................................................................................................... 107 

viii

10-9. A guide for phosphorus fertilisation for adult soursop trees in different 

regions of Colombia, according to age and nutrient content in the soil 

……………………………………………………………………………..107 

10-10. A guide for potassium fertilization of adult soursop trees in Colombia, 

according to tree age and potassium content in the soil .................... 107 

10-11. Removal of macronutrients per tonne of soursop and sugar apple fruits 

produced............................................................................................ 110 

10-12. Removal of micronutrients per tonne of soursop and sugar apple fruits 

produced............................................................................................ 110 

10-13. Major and selected minor insect pests of Annona species. .............. 117 

10-14. Major and selected minor diseases of Annona species. ................... 122 

11-1. Harvesting season of the four major Annona species in different 

countries and regions......................................................................... 128 

12-1. Mean costs to establish and maintain one hectare of soursop cv 

Morada, based on 204 plants per hectare, and estimated gross and net 

incomes.............................................................................................. 142 

12-2. Total area, production, productivity and value of three important 

Annona species in some of the major producing countries ............... 144 

12-3. Carton types for classification and packing of sugar apple fruit in the 

São Paulo market, Brazil ................................................................... 148 

A-1. Some of the Most Common Chemical Compounds Found in Different 

Parts of Cherimoya (Annona cherimolia), Custard apple (A. reticulata), 

Soursop (A. muricata), Wild soursop (A. senegalensis) and Sugar 

Apple (A. squamosa). ........................................................................ 192 

B-1. Some Uses of Annona Species in Medicine, Cherimoya (Annona 

cherimolia), Custard apple (A. reticulata), Soursop (A. muricata), Wild 

soursop (A. senegalensis) and Sugar apple (A. squamosa)................ 198

List of Figures 

2-1. Botanical characteristics of some plant parts of cherimoya (Annona 

cherimola Mill.) .................................................................................... 9 

2-2. Botanical characteristics of some plant parts of soursop (Annona 

muricata L.)......................................................................................... 12 

2-3. Botanical characteristics of some plant parts of custard apple (Annona 

reticulata L.)........................................................................................ 14 

2-4. Botanical characteristics of some plant parts of wild soursop (Annona 

senegalensis L.)................................................................................... 16 

2-5. Botanical characteristics of some plant parts of sugar apple (Annona 

squamosa L.) ....................................................................................... 18 

3-1. Global distribution and occurence of 5 annona species......................... 18 

10-1. Type and sequence of seed germination of annona species................. 74 

10-2. A sketch (plan view) of an irrigated nursery for production of grafted 

soursop and sugar apple trees showing the cement block supports, 

wires, micro-sprinklers and plastic bags, their distances and 

characteristics ...................................................................................... 76 

10-3. A sketch (side view) of an irrigated nursery for production of grafted 

soursop and sugar apple trees showing the cement block supports, 

wires, micro-sprinklers and organization of plastic bags, their distances 

and characteristics ............................................................................... 77 

10-4. Splice grafting in Annona .................................................................... 81 

10-5. Steps of the inverted T - budding technique in Annona....................... 82 

10-6. Steps of the topworking technique used to regenerate an unproductive 

canopy of an annona tree..................................................................... 84 

10-7. Soursop planting systems .................................................................... 86 

10-8. Planting systems according to the slope of the land ............................ 90 

10-9. Stylized diagram of cherimoya tree formation with length of the 

growing branches of subsequent years ................................................ 95 

10-10. Two types of pruning for soursop tree formation .............................. 96 

10-11. Pruning for rejuvenation of a soursop tree......................................... 97 

10-12. A typical scheme for establishment of a drip irrigation system in the 

field ................................................................................................... 113 

10-13. Water distribution (degree of opening) of different micro-sprinklers 

that can be used in annona orchards .................................................. 115 

11-1. Sketch of a mechanized system to process fruit pulp ........................ 136 

x

12-1. Commercialization channels for distribution of cherimoya in 

California, USA................................................................................. 147 

List of Plates 

1. Purple skinned sugar apple is commercialized as an exotic fruit, since its 

colour makes it look somewhat like a rotten fruit thus limiting its 

acceptance at consumer market………………….………………….. 87 

2. A solarization system is used as a pre-planting treatment for germination 

and seedling growth media to control fungi and nematode attacks ..... 87 

3. An intercropping system using sugar apple and papaya can help growers to 

earn additional income ........................................................................ 88 

4. Small plastic containers (e.g. empty film-roll holders) can be used to carry 

pollen................................................................................................... 88 

5. Small plastic containers to carry pollen should be kept in the operator’s 

pocket, to facilitate hand pollination………………………………….89 

6. Annona fruit borer, seed borer and trunk borer are the most important 

annona pests ........................................................................................ 89 

7. Soursop damaged by brown rot disease caused by the fungus Rhizopus 

stolonifer.............................................................................................. 90 

8. Mature sugar apple fruit at its “harvest point”, and fully ripened fruit at 

its “consumption point” ……………………..………………………. 90

Abbreviations 

BAP - Benzylamino purine 

EDTA - Ethylene Diamino Tetra Acetic Acid 

EMBRAPA - Brazilian Corporation for Agricultural Research 

FAO - Food and Agriculture Organization of the United Nations 

FOB price - Free on Board price 

GA - Gibberellic Acid 

GXE – Genotype by Environment Interaction 

HIV - Human Immunodeficiency Viral Disease 

IBA - Indole Butyric Acid 

IAA - Indole Acetic Acid 

IPGRI - International Plant Genetic Resources Institute 

MS - Murashige and Skoog growth medium 

MTH - Monotetrahydrofuran 

NAA - Napthalene Acetic Acid 

NAS - National Academy of Sciences 

NPK - Nitrogen, Phosphorus, Potassium (fertilizer) 

NRC - National Research Centre 

ODEPA - Oficina de Estudios y Politicas Agrarias del Chile 

PROCIANDINO - Programa Cooperativo de Investigación y Transferencia 

de Tecnología Agropecuaria para la Region Andina 

RAPD - Random Amplified Polymorphic DNA 

RH - Relative Humidity 

THF - tetrahydrofuran 

UFAL - Federal University of Alagoas 

USDA - United States Department of Agriculture 

i

Acknowledgement 

Acknowledgements 

Any major project involves the contribution of many people. This Annona 

monograph is no exception. 

First of all, my thanks to Embrapa Cerrados directorate, especially the 

Research and Development Director, Dr. Eduardo Assad, for the challenge, 

support and encouragement. To the other members of Embrapa Cerrados 

staff that contributed through their support and help, my thanks. To my 

colleagues Francisco Ricardo Ferreira of Embrapa Genetic Resources and 

Biotechnology Research Center, Maria Cristina Rocha Cordeiro, Solange 

Rocha Monteiro de Andrade and Daniel Ioshiteru Kinpara of Embrapa 

Cerrados, Ricardo Elesbão Alves and Heloisa da Cunha Filgueiras of 

Embrapa Tropical Agroindustry for agreeing to co-author this monograph. 

A special thanks to my friends Dr. Charles Roland Clement, the monograph´s 

local editor, and Ms. Angela Hughes, for reviewing the English text, for the 

excellent support in searching for and selecting bibliographic references, and 

to both for their encouragement to complete this study. 

Thanks to Miss Alessandra Alves Rodrigues for the help in collecting and 

selecting necessary literature important for this monograph. Thanks to my 

colleague Victor Hugo Vargas Ramos, Embrapa Cerrados, and his Peruvian 

brother, Antonio Isaias Vargas, for the information on area and production of 

cherimoya and soursop in Peru. Thanks also go to my sister-in-law Fátima 

Maria Eugênio de Sousa Oliveira for help in searching Annona 

bibliographies at the library of Ministry of Agriculture, in Brasilia. Also, 

thanks to Mr. Chaile Cherne for his help with the botanical designs. 

I would also like to extend my thanks to Mr. Gonzalo B. Gómez from the 

Embassy of Spain, and to colleagues Wouter Conradie (South Africa), 

Victoria Morales (Venezuela), José Maria Hermoso, Victor Galan Sáuco and 

José Farré (Spain), Zora Singh (Australia), D.K. Sharma and C.P. A. Iyer 

(India), X. Scheldeman (Colombia), E. Lemos and R. Kavati (Brazil), for 

providing information on economics, production data and management of 

Annona species in their respective countries, provinces or states. 

Finally, to my family, especially to my wife Rita and my children Liane and 

Fabiano, whose willingness to tolerate some intolerable demands on our time 

ii

Acknowledgement 

together combined with their loving support and encouragement has made 

my research on tropical fruit crops a reality. 

Alberto Carlos de Queiroz Pinto, Brasília, DF, Brazil 

iii

Preface 

Increasing demand for exotic fruits for human consumption has become 

evident in many countries, both tropical and temperate. The indigenous and 

often "under-utilised" fruits in many developing countries can provide the 

vitamins, mineral salts and fibre in their diets. 

The majority of the Annona species are considered to be under-utilised, 

hence information on them is scarce and widely scattered. However, the 

areas under production have increased more rapidly than the contributions 

from science and technology. There is a need for better information for 

academics, researchers, extension workers and growers. 

The information available varies from species to species. Far less is known 

about A. senegalensis and A. reticulata than the other three species. It is 

hoped that the information provided will make this account useful to all those 

interested in annonas and encourage their wider cultivation and use. 

This is a reference text bringing together available information on five 

Annona species, with the aim of identifying gaps in knowledge and thus 

research needs. With an accompanying extension manual, this will provide 

opportunities for both resource-poor as well small-scale commercial farmers 

to develop this species for income generation. The monograph is written for 

researchers and extension agency offices. The extension manual is designed 

to meet the needs of farmers and small-scale producers. We hope that this 

work may encourage further production, processing and marketing of 

annonas particularly at the village level, and researchers and scientists to 

further explore the benefits of indigenous tropical fruit trees such as annona. 

For further information visit: www.civil.soton.ac.uk/icuc. 

Preparation and publication of this book has been funded by the Department 

for International Development (DFID), UK as part of a project called "Fruits 

for the Future". In addition to ICUC, other partner organisations involved are 

the World Agroforestry Centre (ICRAF) and the International Plant Genetic 

Resources Institute (IPGRI). 

Mention of specific products or commercial organisations in the text is 

intended solely for purposes of illustration and does not imply any 

endorsement by the sponsoring organisations. 

We would like to express our sincere thanks to Dr. A. C. de Q. Pinto and his 

colleagues Francisco Ricardo Ferreira, Maria Cristina Rocha Cordeiro, 

iv

Preface 

Solange Rocha Monteriro de Andrade, Ricardo Elesbão Alves, Heloisa 

Almeida da Cunha Filgueiras and Daniel Ioshiteru Kinpara who have 

prepared a very detailed and thorough manuscript for this publication. Also 

to Dr. Charles Clement and Prof. Trevor Williams for their help in reviewing 

and editing the manuscript, Ms. Barbara Richie, CABI for providing her 

advice on the pests and disease sections, the Editorial Board who have 

contributed their comments and help on the manuscript and to all other 

collaborators who provided information and research papers for analysis and 

citation. 

Lastly, we would like to thank DFID-FRP for funding the project without 

which this publication was not possible. 

Editors, 2005 

v

Chapter 1. Introduction 


Four of the five species covered in this monograph originated in South or 

Meso-America and one seems to have originated in eastern Africa. The first 

group includes Annona cherimola Mill, cherimoya, A. muricata L., soursop, 

A. reticulata L., custard apple, and A. squamosa L., sugar apple. The African 

species is A. senegalensis Pers., wild soursop. Cherimoya is the only species 

adapted to subtropical or tropical highland conditions; the others are mostly 

adapted to the tropical lowlands but can be grown in the subtropics. Two, 

cherimoya and soursop, received international attention in publications of the 

US National Academy of Sciences (NAS, 1975, NRC, 1989). These five 

species have been selected for attention because available evidence suggests 

the possibility of expanding their use and importance. This does not mean 

that other Annona species have no potential. Three others, at least, have been 

recommended for attention. A. purpurea Moc. and Sassé, soncoya, is a small 

tree of Mexico and Central America that is occasionally cultivated for its 

agreeably flavoured fruit (see Gauthier and Poole, 2003, Enhancing the role 

of forest fruits in sustaining livelihoods of forest margin communities (R7349 

- FRP)). A. scleroderma Saff., posh té, is from the same area and has a fruit 

that is reputed to taste better than soursop (Uphof, 1959). In the Brazilian 

Cerrados (the savannahs of central Brazil), A. crassiflora Mart., araticum, is a 

popular minor fruit, also used in traditional medicine (Almeida et al., 1998), 

and is now receiving research attention. Other species, supposedly with 

promise, were introduced to certain areas but did not become important. The 

case of A. glabra L., pond apple, taken to S. E. Asia is an example. 

The strongest consumer demand, and hence production, is for cherimoya, 

soursop and sugar apple. The fruits of these species are delicately flavoured 

and are marketed mainly in local, regional or national trade, only rarely in 

international trade. Pulp of these fruits is sold fresh or frozen, and is usually 

used for desserts or made into sherbets and ice cream. Although custard 

apple and wild soursop are less important economically, their fresh fruits are 

sold in the markets of some developing countries, and their pulp and seeds 

are also used for medicinal purposes more commonly than those of the major 

species. 

1

Chapter 1. Introduction 

In many countries, the species are grown in small areas by small scale 

farmers, who generally have difficulty accessing internal and external 

markets. Major limiting factors are (1) climatic limitations and other growing 

conditions, (2) ineffective or poor agronomic techniques, (3) limited postharvest 

knowledge for harvesting, handling and transporting, (4) lack of 

agribusiness initiatives to stimulate production, marketing and research, and 

(5) lack of knowledge about the fruits' nutritional values, and techniques for 

the elaboration of processed products. 

The major species, such as cherimoya in countries with subtropical climates, 

and soursop and sugar apple in tropical regions, are becoming better known 

and are finding their way into commercial marketing channels. Nonetheless, 

better technical and scientific knowledge is needed to sustain and enhance 

the development of these species in appropriate regions. 

This monograph systematizes information from different areas of study, such 

as taxonomy and botany, origin and distribution, ecology, properties and 

uses, agronomy, harvesting and post-harvest processing and the economics of 

production. This monograph is expected to be useful to students, research 

and development specialists, annona farmers and others interested in these 

fruits. The companion extension manual is expected to be useful to the same 

set of people, but is prepared especially for annona farmers and fruit 

extension agents. 

2

Chapter 2. Taxonomy and Botany 

2.1 General 


The number of genera and species in the family Annonaceae is still debated. 

Bailey (1949) affirmed that Annonaceae has 46 genera and between 500 and 

600 species, while Fries (1959), cited by Geurts (1981), affirmed that it 

contains 119 genera and over 2,000 species. Popenoe (1974 a) described the 

family as having 40 to 50 genera and more than 500 species, most of which 

are shrubs and small trees. A limited number of species produce edible fruits, 

including many gathered from the wild, and some that have been 

domesticated (Ochse et al., 1974). Most of the species are found in the 

tropics, with only a few genera present in the temperate zone. 

According to Geurts (1981), of the 119 species described in the genus 

Annona, 109 are native to tropical America and 10 to tropical Africa. All of 

the domesticated species are American, while one African species (A. 

senegalensis) is probably in the process of domestication. 

2.2 Specific and common names 

The names of the five annona species dealt with in this monograph are 

presented in Table 2.1. The number of common names is large for some of 

the species, and they need to be used with caution because some names may 

be applied to two or more species in different countries, or even in different 

regions of the same country. Clearly therefore, the botanical descriptions are 

essential for extensionists, students and growers to distinguish one species 

from another. 

The relevant botanical details of the species are outlined in section 2.3 below. 

Mistaken identities among botanically similar annonas are relatively 

frequent. For instance, atemoya (a hybrid between cherimoya and sugar 

apple) was mistakenly called custard apple for many years (Morton, 1987), 

when this name more properly relates to A. reticulata. Custard apple is 

sometimes confused with A. glabra, and A. montana has been confused by 

some Brazilian growers with A. muricata. Key botanical literature includes 

3


Bailey (1949), Geurts (1981), Léon (1987), Ochse et al. (1974), Pinto and 

Silva (1996) and Popenoe (1974 a, b). 

Table 2-1. Botanical or specific, common and vernacular names and 

their synonyms of the five Annona species studied 

Botanical Synonyms Common Other common names 

A. cherimola 

Mill. 

A. tripetala Aiton, 

A. pubescens Salisb. 

Cherimolia, 

Cherimoya 

Cherimoya, anona del Peru, 

chirimoyo del Peru, catuche, 

momora (Spanish), 

chérimolier (French), 

cherimoya, cherimoyer, 

annona, custard apple 

(English), honumanaphala 

(Kannada), lakshamanphal 

(India), noina ostrelia (Thai), 

anon (Spanish, Guatemala), 

cherimólia, anona do Chile, 

cabeça de negro 

(Portuguese), cerimolia 

(Italian), chirimoyabaum, 

peruanischer flaschenbaum, 

flachsbaum (German) 

A. muricata L. A. muricata L. 

A. bonplandiana 

Kunth.; A. cearensis 

Barb. Rodr.; 

A. macrocarpa 

Werckle; 

A. muricata 

var.borinquensis 

Morales; 

Guanabanus 

muricatus Gómez 

A. reticulata L. A. excelsa Kunth; 

A. laevis Kunth; 

A. longifolia Moc. 

and Sessé; A. riparia 

Soursop 

Custard apple 

Guanábana (Spanish), 

corossolier (French), zuurzak 

(German), munolla (India), 

mulluseeta, pullupala 

(Tamil), mullu ramaphala 

(Kannada), mullanchakka, 

vilathinura (Malayalan), 

graviola (Portuguese), durian 

belanda (Malaysian), mamon 

(Spanish, Philippines) 

Bullock's heart, corazon 

(English), condessa e 

coração-de-boi (Portuguese), 

buah nona (Indonesian), 

4



Kunth. 

ramphal (India), ramaseeta 

(Tamil), ramasitapalam 

(Tegelu), vilathi 

(Malayalan), ramaphala 

(Kannada) 

A. senegalensis 

Pers. 

A. arenaria Thonn.; 

A. chrysophylla Boj.; 

A. chrysophylla var. 

porpetac Bail.; 

A. porpetac Bail.; 

A. senegalensis var. 

porpetac Bail. Wild 

Wild soursop 

Mchekwa, (kishwahili), 

mtomoko (kichaga), 

mtopetope (kirufiji), gishta 

gaba (Arabic), annone 

africain, pomme cannelle du 

Sénégal (French), nhonokono 

nwitu, ntokwe, mtokwe 

(Kenya), nchakwa 

(Tanzania), mposa, muroro, 

mponjela, mulembe 

(Malian), ntantanyerere, 

mtopa (Zambia), gishit'a 

(Ethiopia), dau-ha, dyangara 

(Bambara), moupa (Dierma), 

bu bualansambu, goritsaa 

tibu, iuboualansahu 

(Gourmancho), barkudugo, 

bakikudiga, barduki, 

barkudugo, barkoudouga 

(Moore), barkoutahe, 

dokumi, doukouhi (Peulh), 

digor, dugor, jorqut (Wolof) 

A. squamosa L. A. asiatica L.; 

A. cinerea Dunal; 

Guanabanus 

squamosus Gomez 

Sugar apple 

Sweetsop, sugar apple, 

custard apple (English), ata, 

pinha or fruta do conde 

(Portuguese, Brazil), attier 

(French), saramuya and 

Aztec (Mexico), sitaphal, 

(Tamil), seethapalam, 

athichakku (Malayalan), 

nona sri kaya (Malaysian) 

seethapandu (Tegelu), 

5



amritaphala, seethaphala 

(Kannada), aatoa, shariffa, 

sitaphal (Hindi), ata, luna 

(Bengali), sita pandu 

(Tebgu), noina (Thai) 

2.3 Botanical description 

6 

2.3.1 Description of the genus 

The name annona derives from the Latin "annual harvest" (Lizana and 

Reginato, 1990). The genus presents numerous unifying characteristics, 

especially relating to plant height, root system, bark, stem, floral biology, 

pollination, fruit set and fruit type (Ochse et al., 1974; Geurts, 1981; León, 

1987). There are important variations among annona seedlings in the same 

species, affecting not only the mature foliage and productivity of the plants, 

but also the fruit size, form, colour, quality and number of seeds in the fruit. 

These variations are often pronounced enough to have resulted in several 

botanical names for the same species. 

In general, the annonas are shrubs or small trees, whose height varies from 5 

to 11 m depending on several factors, such as species, climate, soil and crop 

management. They are erect or somewhat spreading in habit, with greybrown 

bark, often rough and corrugated (León, 1987). Generally, annona 

stems are ferruginous to greyish, and tomentose when young but later 

becoming glabrous. With few exceptions, annonas are deciduous, even 

tropical species, especially when cultivated in areas with dry or cool seasons 

and without irrigation. 

The root system has abundant thin lateral roots and a taproot that is not as 

strong as in other tropical fruit trees, such as mango (Mangifera indica L.). 

Although the taproot is not generally pronounced, the lighter the soil texture 

the longer the taproot will grow. The taproot of an adult soursop tree can 

reach approximately 1.5 to 1.8 m in depth in oxisols of the Cerrado 

ecosystem (Brazilian savannah) in Central Brazil (Pinto and Silva, 1996). 

Annona flowers are hermaphrodite, usually somewhat fragrant, solitary or in 

fascicles with 2 to 4 flowers, with three green sepals and six petals arranged 

into two verticils. The external verticil has three yellow-greenish petals and


the internal one has three yellowish petals. The flowers have several 

conglomerated and spirally arranged stamens below and around an upper 

globose (conical) shaped dome of numerous united carpels, which have one 

ovule each. After fertilization, the united carpels will form a syncarp or 

composite fruit. 

Flowering starts when the plant is three to four years old, although it may 

occasionally occur earlier depending upon environmental conditions. 

Anthesis (flower opening) starts slowly, with the separation of the apex of 

the external petals and takes from 6 to 8 h to complete. 

Pollination is mainly carried out by insects or sometimes by wind. The fact 

that flowers are protogynous (pistils are mature before pollen is liberated 

from anthers) suggests that self-pollination is not the rule for annonas. Due to 

the protogynous flowers, fruit set may be poor when the pollinating insect 

populations are small, and small, asymmetric fruits may be produced, since 

fruit size and form depends on the number and position of fertilized ovules. 

As a result, the number of fruits per plant, and fruit size and shape are highly 

dependent on abundant insect pollination and amenable to control via hand 

pollination where the pollinating insects are rare. 

2.3.2 Description of the species 

1. Annona cherimola (cherimola) 

The name cherimoya derives from the Quechua name "chirimuya", which 

means "cold seeds" (Lizana and Reginato, 1990). It is a small, erect and/or 

somewhat spreading, deciduous tree, rarely reaching a height of more than 

7.5 m. Its stem frequently divides at ground level into several stems (NRC, 

1989). It was domesticated in the mid-elevation Andes of South America. 

It has simple, alternate, 2 to 4 ranked leaves, which are ovate-lanceolate to 

elliptical in shape, 10 to 25 cm long, glabrous on the ventral surface and 

pubescent dorsally, with leaf shedding in the spring. 

The single, protogynous, fragrant flower emerges from the leaf axils, and 

possesses a short peduncle, ca 2.5 cm in length (Fig. 2.1). Flowering occurs 

once a year, the season depending on the environment and it starts when the 

tree is 3 to 4 years old. Flower anthesis starts in the early morning and it 

takes 8 h to attain complete opening. 

The fruit is normally heart-shaped, conical, oval or somewhat irregular in 

form due to irregular pollination. Fruits measure 7.5 to 12.5 cm in length and 

7


weigh from 200 to 700 g (Fig. 2-1). The fruit surface is smooth in some 

varieties; in others, it is covered with small conical protuberances over the 

carpels. The fruit rind is delicate and thin, and is greenish-yellow when ripe 

(Popenoe, 1974 a). The white, subacid flesh has a fragrant, delicate flavour, 

like that of pineapple and banana. The fruit has numerous seeds (21 to 41 

seeds/fruit), which are 1.5 to 2.0 cm in length and approximately 1.0 cm in 

width (Manica, 1997). 

8


Figure 2-1. Botanical characteristics of some plant parts of cherimoya 

(Annona cherimola Mill.) 

Popenoe (1974 a) cites five botanical forms differentiated on fruit shape and 

skin design (due to the variable structure of the carpel protuberances and 

outlines over the carpel segments called areoles). 

9


Forma impressa: Finger-printed - This form is called anona de dedos 

pintados (finger printed annona) in Costa Rica. The fruit is conoid or 

subglobose in shape, and has a smooth surface covered with U-shaped 

areoles resembling finger-prints in wax. Fruits tend to be of good quality, 

although the fruit contains many seeds. 

Forma laevis: Smooth - This form is called cherimoya lisa (smooth 

cherimoya) in South America and anon in Mexico City. It is often mistaken 

for A. glabra or A. reticulata because of the general appearance of the fruit 

and on account of the name anon, which is also applied to A. reticulata. This 

is one of the finest botanical forms. 

Forma tuberculata: Tuberculate - This is the commonest form. The fruit is 

heart-shaped and has wart-like tubercles near the apex of each areole. The 

cultivar Golden Russet belongs to this botanical form. 

Forma mamillada: Mammillate - This form is called "cherimoya de tetillas" 

(nippled cherimoya) in some South American countries. It is said to be 

common in the Nilgiri Hills in southern India, and to be one of the best types 

grown on Madeira Island. 

Forma umbonada: Umbonate - This form is called "cherimoya de púas" 

(barbed cherimoya; spiny cherimoya) and "anona picuda" (pointy anona) in 

Latin America. It has a thick skin, with more acidic flesh than other types, 

and the seeds are numerous. The fruit is oblong-conical, with the base 

somewhat umbilicate and the surface studded with protuberances, each of 

which corresponds to a carpel component. 

Numerous cultivars have been described, both in the area of origin, where 

every valley has its particular type, and in the numerous areas where 

cherimoya was introduced. The germplasm introduced to Spain alone appears 

to have included about 200 traditional cultivars. Modern commercial 

cultivars have been developed since these introductions. NRC (1989) 

mentioned such well known North American cultivars as ‘Booth’, ‘White’, 

‘Pierce’, ‘Knight’, ‘Bonito’, ‘Chaffey’, ‘Ott’, ‘Waley’ and ‘Orchard’, and 

stated that cultivars exhibit a wide variation in climatic and soil requirements 

(see also Table 9.2). 

2. Annona muricata (soursop) 

The soursop has an erect growth habit with a high canopy height-to-diameter 

ratio (Pinto and Silva, 1996), although it tends to be low-branching and 

bushy, with upturned limbs (NAS, 1975). It is a small, slender, evergreen 

10


tree, 4 to 8 m tall when fully mature. It was domesticated in lowland South 

America as a garden plant. 

The stems are rounded, rough and not pubescent, with a dark-brown colour. 

The leaves have short petioles, and are oblong-ovate to cylindrical, 14 to 16 

cm in length and 5 to 7 cm in width. Because of similarities of plant canopy 

and leaf form, soursop and mountain soursop (A. montana Macf.) are often 

confused. The flowers of soursop are much larger than those of the other four 

species listed in this monograph, being 3.2 to 3.8 cm in length (Fig 2-2). 

11


Figure 2-2. Botanical characteristics of some plant parts of soursop 

(Annona muricata L.) 

The flowers start to open in the early morning and complete anthesis takes 

approximately 6 h, depending on the climate. Flowering is more or less 

12


continuous. This species also experiences inefficient natural pollination 

(normally done by beetles) and frequently poor fruit set; hence hand 

pollination is an important orchard management practice. 

Soursop produces an ovate, conical or heart-shaped fruit, that is dark green 

when unripe and a slightly lighter green when ripe. The rind has many short, 

fleshy, pointed carpel protuberances and is popularly regarded as 'spiny'. The 

soursop has the largest fruit in the genus, weighing from 0.9 to 10 kg, and 

averaging 4 kg. Its white, cottony-fibrous, juicy flesh resembles that of 

cherimoya in colour. The flavour is more acid and less sweet than cherimoya, 

and calls to mind a mixture of pineapple and mango. 

The fruit has 127 to 170 seeds, scattered throughout the pulp. They are toxic. 

Seed size varies from 1 to 2 cm in length and from 0.33 to 0.59 g in weight, 

with a black colour soon after harvest, but becoming dark-brown later (Pinto 

and Silva, 1996). 

Few cultivars of A. muricata exist, and comparisons among them have not 

been made to assess their validity. In particular, those with good-sized, lowfibre 

fruits need to be identified (NAS, 1975). All known selections are Latin 

American. 

3. Annona reticulata (custard apple) 

The tree reaches 6.0 to 7.5 m in height, with many lateral branches; stems are 

cylindrical, with lenticels and very short coffee-coloured hairs. It is 

considered the most vigorous of the annonas described in this publication. 

Custard apple is thought to have been domesticated, even though the fruit is 

considered to be of inferior quality. It can be distinguished from cherimoya 

by its long, narrow, glabrous leaves, from sugar apple by its solid, compact 

fruit, as well as its larger leaves, and from A. glabra by its small, dark brown 

seeds (León, 1987). 

The leaves are oblong-lanceolate and dark-green, measuring 25 to 30 cm in 

length and 7 cm wide, with 10 to 20 vein pairs and a pubescent petiole. 

Flowers are similar in form to those of sugar apple, except that they are 

grouped in a short inflorescence with 2 to 10 flowers, with pedicels 

measuring 1.5 to 3.0 cm in length. This species also presents inefficient 

natural pollination and poor fruit set. 

Fruits weigh from 0.1 to 1.0 kg and are commonly heart-shaped, but may be 

conical, ovate or irregular in form, and 10 to 12 cm in length (Fig. 2-3). They 

are coriaceous and have a reddish-yellow surface colour, with impressed 

lines (around 5 to 6 angled areoles) above the carpels. The flesh is milk-white 

13


and sweet, although insipid in flavour, being considered the least tasty of the 

cultivated annonas. There are commonly more than 40 oblong, dark coffeecoloured 

seeds per fruit (León, 1987). 

Figure 2-3. Botanical characteristics of some plant parts of custard apple 

(Annona reticulata L.) 

14


4. Annona senegalensis (wild soursop) 

Wild soursop is a spreading shrub or small, semi-deciduous tree, 1.5 to 11.0 

m (averaging usually about 3.5 m) in height, with a stem diameter up to 28 

cm at breast height (FAO, 1983, 1988). It has a greyish-black bark, often 

rough and corrugated, branching near the ground, with young stems mostly 

ferruginous, velvety to greyish or red-brown tomentose, later becoming 

glabrous. It is not strictly domesticated, but some trees are 'protected' due to 

preferred qualities. 

15


Figure 2-4. Botanical characteristics of some plant parts of wild soursop 

(Annona senegalensis L.) 

Source: FAO (1983) 

The leaves are ovate, oblong-elliptical or oblong-ovate in form, 8 to 17 cm 

by 4 to 10 cm, with an acute, obtuse, rounded or slightly emarginate apex, 

and upper surface smooth, lower surface pale brown and hairy. Like other 

annonas, the leaves are simple, alternate, with 0.5 to 2 cm long petioles. 

The flowers are inconspicuous, green, single or grouped on long smooth 

stalks (in fascicles with 2 to 4 flowers). They are fleshy, up to 3 cm diameter 

and usually fragrant. Although there is no available information on flower 

16


opening, pollination and fruit set of this species, it seems obvious from its 

flower biology (FAO, 1983) that it has problems similar to the other annonas 

mentioned here. 

The fruit has an ovate, globose or subglobose form, measuring 2.5 to 5.0 cm 

in length and 2.5 to 4.0 cm in width (Fig. 2-4). The unripe fruit is green with 

white specks turning yellow or orange when ripe. The white to yellow edible 

flesh, which has many seeds, has a pleasant aroma, resembling pineapple, but 

tasting of apricot. One hundred seeds weigh ca. 40 g. There is a recognized 

botanical variety, var. posteide (Bail.) Diels., and no known cultivars. There 

is a dwarf form in Malawi where the plant is so small that the fruits grow 'on 

the ground' and this form is thought to taste the best by the locals 

(Williamson, 1974). 

5. Annona squamosa (sugar apple) 

The sugar apple tree is deciduous and much smaller than the soursop, 

reaching a maximum of 6.0 m in height, with many lateral branches. The 

stems present lenticels, while the young shoots are pubescent and the oldest 

are smooth. It was domesticated in the circum-Caribbean or northern South 

American lowlands. 

Like other annona species, it has deciduous leaves that are brilliant green 

above and bluish green below, with petioles 0.7 to 1.5 cm in length. The 

leaves are oblong-elliptical in form, measuring 5 to 17 cm in length and 2 to 

7 cm in width, with an obtuse or acuminate apex. The blade has 15 to 17 

pairs of veins (Ochse et al., 1974). 

The flowers measure 2.0 to 2.5 cm in length and are much smaller than 

soursop flowers, being similar in size and form to those of cherimoya. 

Pollination and fruit set problems are similar to those of other annonas. 

Pollen germination is low and may influence final fruit set, which varies 

from 5.4% to 5.6% (Thakur and Singh, 1965). 

The fruit is rounded, heart-shaped, ovate or conical, 5 to 7.5 cm in diameter, 

6 to 10 cm in length and weighing 120 to 330 g (Fig. 2-5). Fruit size depends 

on cultivar, pollination, nutrition and other factors, but its form resembles a 

hand-grenade, with a tuberculate surface covered with a whitish bloom. The 

white, custard-like pulp has a pleasant sweet-sour flavour. The fruit contains 

35 to 45 black seeds, each 1.5 to 2.0 cm in length and 0.6 to 0.8 cm in width. 

There are a few recognized cultivars of sugar apple, with the majority of 

these in India, and their names give some idea of their origin as 

introductions: ‘Mammoth’, ‘Barbados’, ‘British Guinea’, ‘Balondegar’, ‘Red 

17


Sitaphal’, and ‘Sindhan’, the last being local to Gujarat (Singh, 1992). A 

dwarf cultivar is ‘Lal Sitiphal’. 

Figure 2-5. Botanical characteristics of some plant parts of sugar apple 

(Annona squamosa L.) 

18

Chapter 3. Origin and Distribution 


The origin of most of the species treated in this book is South America and 

the Antilles, however wild soursop is thought to have originated in Africa. 

The current distribution of these five species covers almost all continents, 

with soursop and sugar apple showing the widest distribution, mainly in 

tropical regions (Fig. 3-1). 

Although there is controversy about the origin of cherimoya, the majority of 

the literature (Fouqué, 1972; Ochse et al., 1974; Popenoe, 1974 a) attributes 

the area of origin to the Andean Valleys of Ecuador, Peru and Chile, at 

altitudes of 1,600 to 2,000 m. The primary centre of diversity probably 

occurs there, corresponding roughly to Vavilov's South American centre. 

Hermoso et al. (1999) suggested a secondary centre of diversity in Central 

America, based on work by Perfectti (1995) using molecular markers. 

Cherimoya is an ancient domesticated crop: seeds have been identified in 

archaeological sites in Peru and fruits are depicted on pre-Inca pottery (NRC, 

1989). Wild populations can be found in Ecuador, Peru and Bolivia (Smith et 

al., 1992) and the Loja area of SW Ecuador appears to be a centre of 

diversity of wild material. 

The early Spanish explorers introduced cherimoya to Mediterranean 

countries, as well as to Asia, via Africa (Ochse et al., 1974). The cherimoya 

was introduced to the USA in 1871 by Judge Ord, of Santa Barbara, 

California, from Mexico (Popenoe, 1974 a). Currently, Spain and Chile are 

the main producing countries and also distribute cherimoya germplasm 

around the world. 

Soursop most likely originated in Central America, the Antilles or Northern 

South America, and is found in the Andean valleys in Peru, presumably as an 

ancient introduction. Spanish colonizers distributed it to other tropical 

regions of the world (Popenoe, 1939; Purseglove, 1968). The existence of 

several wild types of soursop in the Amazon region (Cavalcante, 1976) 

suggests that this may be a primary centre of diversity, but the types could be 

remnants from cultivated introductions. Wild populations of soursop are well 

known in the West Indies and on Barro Colorado Island, Panama (Croat, 

1978; Smith et al., 1992). 

19


Figure 3-1. Global distribution and occurence of 5 annona species 

1. Cherimoya (Annona cherimola); 2. Soursop (Annona muricata); 3. Custard apple (Annona 

reticulata); 4. Wild soursop (Annona senegalensis); 5. Sugar apple (Annona squamosa) 

20


In south-eastern Brazil, cultivated soursop was introduced during the 

sixteenth century. Nowadays it is found in almost all Brazilian states, except 

in the southernmost states, where low temperatures and occasional snowfall 

do not allow the tree to grow and produce (Pinto and Silva, 1996). Soursop is 

now a popular fruit in Cuba, Mexico, Central America and throughout South 

America. It is also found in Sri Lanka up to elevations of 460 m, in China 

and many parts of Polynesia. In the USA it is grown in southern Florida. 

Custard apple or bullock's heart is believed to have originated in the Antilles 

(Fouqué, 1972) and other parts of the circum-Caribbean region. From the 

Antilles, Spanish explorers may have distributed this species to Mexico, and 

certainly did to Asia and Africa. Primitive germplasm was transported by 

Native Americans and wild populations in Costa Rica probably represent 

this. Although widely distributed in the tropical world today, it is a fruit of 

little commercial value. 

Custard apple is very commonly found in home gardens of coastal towns 

throughout tropical America. It is also cultivated in India, Sri Lanka, the 

Malay Archipelago, Polynesia, the Philippines, Australia and most of the 

countries of Africa. The vernacular name "custard apple" is usually applied 

to sugar apple in India. 

There is no precise information on the origin and diversity of wild soursop. 

FAO (1983) states that this species is most widely distributed in Tanzania, 

Kenya and Mozambique, and in the Zanzibar and Pemba Islands, suggesting 

that eastern Africa may be the region of origin and diversity. In Sudan, this 

species is found where rainfall is greater than 500 mm, typically in tall-grass 

savannah areas. It is distributed across the Sahel in semi-arid to sub-humid 

areas. It also grows in Angola, Senegal and Mauritania (Vogt, 1995). 

The sugar apple originated in lowland Central America, where it is 

indigenous. From there, it was distributed to Mexico and throughout tropical 

America. In the lowlands of Mexico it is found in a naturalized or wild state. 

It is grown from Central America southwards to northern South America, 

extending to north-eastern Brazil, where it is one of the most popular fruits. It 

is believed that it was first introduced into Brazil via Bahia state, in 1626, by 

Conde de Miranda, which explains the vernacular name in Brazil - "fruta do 

Conde". It was later taken to the Philippines and Asia via the West Indies 

(Antilles) and the Cape of Good Hope (Popenoe, 1974 a; León, 1987). 

In India, there is a very large, diverse population of sugar apple, and its 

commercial importance is so great that some botanists have considered it to 

21


be a native fruit of that country (León, 1987). However, this is a secondary 

centre of diversity, created during the last 500 years. Some of the arguments 

used by those who favour an Asiatic origin for this species include: the 

occurrence of common names for it in Sanskrit; the existence of large, 

apparently wild populations in several parts of India; and the presence of 

carvings and wall-paintings, maybe representing the fruit, in the ruins of 

ancient Muttra and Ajanta temples (Popenoe, 1974 a). In Asia, it grows not 

only in India, but also in south China, where it is known as fan-li-chi, or 

foreign lichi. 

Safford, cited by Popenoe (1974 a), suspected that the name "ata" is not of 

American origin. He said that it may be from the Malayan name "atis", 

meaning heart, and that it was carried to Mexico from the Philippines in early 

colonial days. Coronel (1994) cites the vernacular name "atis" in the 

Philippines. 

In Cuba, the sugar apple ranks with mango as one of the favourite fruits and 

it is common in other islands of the West Indies. In the USA, it grows 

successfully in southern Florida but has never been grown to fruiting size in 

California (Popenoe, 1974 a). 

Sugar apple and cherimoya have been hybridized and produced a new fruit 

called atemoya. The crosses were made by P.J. Webster in 1907 in Florida, 

for the USDA. However, crossing occurred naturally in the field in Australia 

in 1850 and again in Palestine in 1930 (NRC, 1989). The hybrid is in 

commercial production in Australia (where it is confusingly called custard 

apple), as well as the USA, Israel, South Africa, the Philippines and 

numerous parts of Central and South America. This hybrid is preferred 

because there appear to be no pollination difficulties. Nonetheless, major 

selection programmes from diverse seedling progenies have not been 

vigorously pursued in any major production area. 

22

Chapter 4. Major and Minor 

Production Areas 


Statistics on minor fruits, such as Annona species, are unavailable in many 

countries, and where reported they often lack reliability, uniformity and 

continuity. Generally, production data relates only to plantation or orchard 

crops grown for sale for international markets, e.g. banana, grape and mango. 

Production of minor fruits from scattered trees used mainly for home 

consumption is not collected. 

Some developed countries, such as Spain and Australia, have produced a 

body of technical knowledge on cherimoya production, which has 

contributed to better international marketing by these countries. 

Consequently, cherimoya is well known commercially, and has good 

production and export performance, so that it is more important in the 

external market and world consumption than soursop and sugar apple. These 

latter species have their major production areas in developing tropical 

countries, and they are produced mainly for internal markets, principally for 

consumption as fresh fruit or for processing. The custard apple and wild 

soursop, the less important annonas, have the smallest areas of production. 

4.1 Major Production Areas 

The estimated production area of cherimoya in the world in 1994 was 13,500 

ha and, considering an average yield of 6 MT/ha, the total production was 

estimated as 81,000 MT. In Chile, the average production of cherimoya has 

been estimated at 25 MT/ha, which is 4 times higher than the world average 

(PROCIANDINO, 1997). 

Commercial cherimoya production occurs mainly in Spain, Peru and Chile. 

Smaller production areas occur in some countries of Central America, 

Mexico, Israel and the USA (California). 

Spain is considered the most important cherimoya producer in the world, 

with a cultivated area of 3,266 hectares in 1999 (Guirardo et al., 2001, cited 

by Scheldeman, 2002). Granada is the major producing province, 

23

Chapter 4. Major and Minor Production Areas 

24 

representing ca. 90% of the total area of cherimoya in Spain (Farré and 

Hermoso, 1997). However, Agustin (1997) commented that the production 

area of cherimoya in Spain was 1800 ha, which is approximately 55% of that 

reported by Guirardo et al. (2001) in the year of 1999. Up-to-date data show 

a total cultivated area of 3,090 ha of cherimoya in Spain, with 99% under 

irrigation, which suggests a total production of approximately 29,000 MT 

(Gómez, G.B., Embassy of Spain in Brazil, July 2000, personal 

communication). 

Peru had an area of 1975 ha in 1998 producing 14,606 MT and a yield of 7.4 

MT/ha. The Nor Oriental del Marañon is the most important producing 

province with 665 ha of cultivated cherimoya (Vargas, A.I., Oficina de 

Información Agraria del Peru, July 2000, personal communication). 

Chile had 785 ha in production in 1996 (Agustín, 1997). In 1998, 1,152 ha 

were reputed to be in production (Furche, C., Director of the Oficina de 

Estudios y Politicas Agrarias del Chile - ODEPA, July 2000, personal 

communication ), which represents a 68% increase in two years. 

Carlos Furche indicated that in the same year (1998), Peru had an area of 

1,800 ha, Bolivia 1,000 ha, Ecuador 700 ha and Australia 500 ha Crane and 

Campbell (1990) and Grossberger (1999) commented that California had 

100-120 ha of cherimoya, with an estimated production of 453 MT in the 

1989-90 season. Crane and Campbell (1990) also noted that Thailand, the 

Dominican Republic and Costa Rica were important exporters to the USA. 

Soursop is cultivated in many tropical areas in countries such as Angola, 

Brazil, Colombia, Costa Rica, Cuba, Jamaica, India, Mexico, Panama, Peru, 

USA (Porto Rico), Venezuela and S.E. Asia (Pinto and Silva, 1996). There is 

a dearth of production data for most of the South, Central and North 

American countries, except Mexico, Venezuela and Brazil, which seem to be 

the major producing countries of this species. 

Mexico is the most important soursop producing country in the Americas and 

in 1990 had an area of 598 ha, with production of 4,087 MT. Rebollar- 

Alviter et al. (1997) estimated the cultivated area in Mexico at 4,890 ha in 

1996, which means that in six years the cultivated area had increased nine 

fold. On the other hand, Hernández and Angel (1997) stated that the Mexican 

area planted to soursop in the same year was equivalent to 5,915 ha with a 

production of 34,900 MT, easily the largest in the world. However, the 

yield/ha had decreased from 6.8 MT/ha in 1990 to 5.9 MT/ha in 1996. 

Nayarit, with approximately 380 ha, is the most important province for 

soursop production in Mexico.


Venezuela had a cultivated area of 3,496 ha in 1987, with a total production 

of 10,096 MT. Zulia is the most important producing state (Diego, 1989). 

Brazil, with approximately 2,000 ha, has an estimated production of 8,000 

MT of fruits per year (average of 4 MT/ha), almost totally devoted to the 

internal market. Because of its climatic conditions, the Northeast is the major 

production region, representing around 90% of the total production of 

soursop. Recent government support for the development of agroindustry on 

small farms (1 to 5 ha), through processing fruits by freezing pulp, and 

making jellies, syrups and ice creams, has promoted the expansion of soursop 

production in Brazil, especially in the Northeast. Ceará state, in the 

Northeast, with an estimated area greater than 500 ha (Bandeira and 

Sobrinho, 1997), is the most important producer of soursop in Brazil, largely 

because many juice industries operate in that region. 

The cultivated area of soursop in Peru was estimated at 443 ha in 1998, with 

a total production of 3,262 MT and a yield of 7.4 MT/ha (Dr. Antonio Isaias 

Vargas, Oficina de Información Agraria del Peru, July 2000, personal 

communication ). Although Venezuela and Brazil have larger production 

areas than Peru, this country has a larger yield/ha. 

Although sugar apple production data are scarce, the information collected 

shows that the potential for expanding the sugar apple market is high in many 

countries. This species is grown commercially in the West Indies and 

Dominican Republic, the USA (Florida), the Middle East, India, Malaysia 

and Thailand (Crane and Campbell, 1990). Although it is still considered a 

backyard fruit used mainly for domestic consumption in the Philippines, this 

country's production is considered one of the largest in the world. The 

Bureau of Agricultural Economics of the Ministry of Agriculture reported 

that in 1978 there were 2,059 ha of sugar apple in the Philippines, with a 

production of 6,262 MT of fruits. Western Visayas (975 ha with 1,844 MT) 

and Southern Tagalog (390 ha with 1,302 MT) were the largest producing 

regions (Coronel, 1994). 

In Brazil, sugar apple production is concentrated in Alagoas and São Paulo 

States. The area of production of sugar apple in Alagoas State appears to 

have increased greatly, since it was estimated at 500 ha in 1995 

(Albuquerque, 1997), and 814 ha in 1996 (Dr. Eurico Lemos, Federal 

University of Alagoas-UFAL, July 2000, personal communication). This 

increase was due to increasing demand in the north-eastern market. São 

Paulo State had, in this same year, approximately 240,000 sugar apple and 

atemoya trees (Piza Jr. and Kavati,1997), which is an estimated production 

25


26 

area of 480 ha and has been expanding. Like soursop, the development of 

agroindustry and the reasonable price of fresh fruits have encouraged sugar 

apple growers to expand cultivated areas. Currently, the sugar apple and 

atemoya areas are moving into north-eastern and northern Minas Gerais 

State, mainly to the major irrigation projects, where small fruit growers 

produce excellent fruits and sell them to retailers in the Brasilia and Belo 

Horizonte markets. 

4.2 Minor Production Areas 

Several factors impede the production and marketing of the lesser known 

annonas, and scattered cultivation and harvesting from the wild continues, 

e.g., wild soursop in Africa and custard apple in Brazil. Other important 

factors are management of pollination, pests and diseases, financial support 

for growers, highly seasonal harvesting period, organoleptic quality, short 

shelf-life, other commercial opportunities and, finally, marketing. Each 

minor production area is limited by one or more of these factors. 

Carlos Furche (Diretor of the Oficina de Estudios y Politicas Agrarias del 

Chile - ODEPA, July 2000, personal communication ) indicated that Israel, 

with 50 ha, represents one of the important countries with minor production 

of cherimoya. Palacios Rangel and Cano Garcia (1997) state that there was a 

small area of cherimoya in Mexico (31 ha in 1990). 

Portugal (Madeira) had an area of 85 ha of cherimoya in 1996 (Nunes, 1997). 

This area was very important in supporting Portuguese demand for this fruit. 

In Italy, an area of 30 ha under cherimoya has been reported, located in the 

coastal part of Calabria (Monastra, 1997). 

There are no official statistics on the production areas of cherimoya in Brazil, 

although Paraná, São Paulo and Minas Gerais States have small areas with 

appropriate microclimates and altitudes above 1,400 m. Generally, these 

cherimoya areas are cultivated by fruit growers of European origin settled in 

Brazil. Bonaventure (1999) stated that cherimoya and atemoya occupy an 

area of 80 ha in Brazil. However, the cultivated area of atemoya in Northeast 

of Brazil is expanding very quickly. The reason for increasing the area 

planted to cherimoya and atemoya is their excellent organoleptic qualities, 

which make these fruits ideal for export. 

In Mexico, sugar apple was produced in an area of 12 ha, with a total 

production of 73 MT, in 1990, and an estimated yield of 6 MT/ha. Egypt is 

also a representative of the minor areas of production, since the total acreage


of cherimoya and sugar apple in that country in 1991 was 50 ha, with yield of 

about 170 MT of fruit (Mansour, 1997). 

There are no data on cultivated areas and production of custard apple and 

wild soursop. Custard apple is grown in small backyard orchards or harvested 

from the wild in most of its North, Central and South American distribution, 

while the wild soursop is found scattered as wild or dooryard plants in many 

African countries. Custard apple has been widely spread around the tropics 

and has become a prized backyard plant in many parts of Africa. 

In India, sugar apple is cultivated in rain-fed orchards mainly in Maharashtra, 

Gujarat, Andhra Pradesh, Karnataka, Madhya Pradesh, Uttar Pradesh, Bihar, 

Assam and Orissa (Singh, 1992) and in the 1980s the area was estimated to 

be 44,100 ha (Pareek, 1985). Few of the plantings are commercial except for 

areas of Gujarat. Most fruits come to market from semi-wild forests of the 

Deccan Plateau where sugar apple has gone wild. 

4.3. Demand 

(see also section 12.2) 

There has been little market research on international fruit markets for 

annonas. The experience with cherimoya and the sale of improved types 

(swollen skin, round shape, good flavour, juiciness, low seed content, 

resistant to bruising and adequate packing: NRC, 1989) shows small but 

steady increase in demand in Chile, Argentina, Portugal, Spain, the USA, the 

UK, France and Japan. 

27

Chapter 5. Ecological Factors 

M. C. R. Cordeiro and A. C. de Q. Pinto 

5.1 Physiography and climate 

The Annonaceae contains species which are mostly tropical and subtropical, 

although some species can be grown in temperate climates (Donadio, 1997; 

Silva and Silva, 1997). In general, the annonas grow at a range of altitudes, 

and those with the widest adaptation to altitude are also those with the widest 

adaptation to latitude. No photo-period responses have been reported 

(Nakasone and Paull, 1998). 

Most annonas do not adapt to low temperatures. However, highland species, 

such as cherimoya, wild soursop and, to some extent, custard apple, are better 

adapted to cold weather than the lowland soursop and sugar apple. 

Heavy shading reduces fruit set in annonas. Consequently, appropriate 

pruning and spacing are very important and should be adjusted to each 

species (see Chapter 10). Rainfall influences the efficiency of pollination 

(Nakasone and Paull, 1998), generally reducing it significantly when rains 

occur during peak flowering periods. 

Wind is a factor that effects annona cultivation, often severely, as it can 

reduce humidity around the stigma and reduce pollination. Wind can also 

break branches, especially if laden with fruit, and fruits are sensitive to dry 

winds (Nakasone and Paull, 1998). Cherimoya, for example, is reported to be 

especially sensitive to dry winds, which can cause fruit loss (Belotto and 

Manica, 1994). 

Cherimoya is reported to grow at altitudes between 900 and 2500 m in its 

natural range (Popenoe, 1939; Zayas, 1966; Fouqué, 1972; Belloto and 

Manica, 1994) on plateaus and in mountain valleys in subtropical areas with 

a dry, cool climate (the Andes in Peru), and is cultivated mostly in dry, cool 

regions (Fouqué, 1972). Because it is adapted to high altitudes, it can grow 

and yield well in the subtropics - cherimoya is cultivated around the 

Mediterranean (Spain, Italy, Egypt, Israel), and southern coastal California 

and Portugal, as well as in South Africa, Argentina and Chile. In Spain, it is 

cultivated along the southern 'Sun coast', especially in Malaga and Granada. 

28


Cherimoya is now cultivated between latitudes 37° North and 37° South 

(Bonaventure, 1999). 

Cherimoya is not adapted to high humidity regimes and it is reported that the 

dry season favours fruiting (Popenoe, 1939). In Mexico, the cherimoya is 

cultivated in three types of climate: (A)C(m)(w), (A)C(w2) and (A)C(w), in 

the Köppen climate classification system. The first is considered subtropical, 

with a high rainfall regime in the summer (mean of 1692 mm/year in the 

summer and greater than 5% of this in winter); the other two types are 

considered subtropical, but with less abundant rainfall (1,047 to 1,182 

mm/year in summer and less than 5% of this in winter) (Agustín and Angel, 

1997). Water stress just before flowering can increase flower production 

(NRC, 1989). 

The best temperatures for cherimoya cultivation are 18 to 22°C in the 

summer and 5 to 18°C in the winter (Belotto and Manica, 1994; Nakasone 

and Paull, 1998). Consequently, it is considered to be tolerant of relatively 

low temperatures (but is less hardy than avocado or orange: NRC, 1989), and 

needs chilling periods (Nakasone and Paull, 1998). It is sensitive to high 

temperatures. 

Soursop is cultivated from sea level to 1,200 m altitude (Zayas, 1966; Pinto 

and Silva, 1994) and between latitudes 27° North and 22.5° South. Its 

northern extremes include southern Florida (USA), Culiacan, Chiapas, 

Veracruz and Yucatán (Mexico), Cuba, and the south of China, while it's 

southern extreme is in central Brazil. 

Soursop is the most tropical annona (Popenoe, 1939; Nakasone and Paull, 

1998) and is cultivated mainly in tropical moist regions, classified as A in the 

Köppen system (Pinto and Silva, 1994). All months have average 

temperatures greater than 18°C and annual precipitation exceeds 1500 mm. 

Tropical wet (Af) climates have year round precipitation with minor monthly 

temperature variations (less than 3°C). Tropical monsoon (Am) climates have 

annual rainfall equal to or greater than Af, but concentrated in the 7 to 9 

hottest months, with water deficits in the dry season. Soursop probably 

originated somewhere in these climate types. Tropical savanna (Aw) climates 

have an extended dry season during the winter, with less than 1,000 mm 

precipitation during the wet season (Ayoade, 1991). 

In Brazil, soursop is cultivated in warm and humid to semi-arid climates (the 

latter with rainfall near 1,000 mm/year), but only fruits if irrigated in the 

semi-arid regions (Pinto and Silva, 1994). The mean temperature in the semiarid 

winter is greater than 18°C. Nakasone and Paull (1998) reported that 15 

29


to 25°C is the minimum temperature range for good growth, while Belotto 

and Manica (1994) reported that the temperature range for its establishment 

is 18 to 29°C. Soursop is reported to be cultivated between 21 and 30°C and 

is susceptible to abrupt changes in temperature, especially if they go below 

12°C (Pinto and Silva, 1994). Consequently, even though the literature is 

variable, it is clear that soursop is sensitive to colder temperatures. 

Additionally, it does not tolerate dry, cold winds. It is the least hardy of the 

annonas (NAS, 1975). 

Soursop is reported to require high light intensity to grow (Villachica, 1996), 

although the wild populations reported by Cavalcante (1976) in Amazonia 

are apparently shade tolerant. This contrast suggests that it is a completely 

domesticated species, as proposed by Clement (1999). 

Custard apple is the most widely cultivated annona at low to medium 

elevations (0 to 1,500 m) (Popenoe, 1952). It grows between latitudes 25° 

North and South, and is reported to be found in almost all tropical areas of 

the world (Zayas, 1966; Nakasone and Paull, 1998). 

Custard apple cultivation is possible in both humid and semi-arid climates 

(Popenoe, 1952), although it is reported to prefer humid climates (Fouqué, 

1972). The average temperature recommended for custard apple cultivation is 

not reported anywhere, but Fouqué (1972) affirms that it is sensitive to long 

periods of cold. 

Wild soursop is adapted to various altitudes, being cultivated from 0 to 1,800 

m in Kenya and from 0 to 2,400 m in other parts of East Africa (FAO, 1983). 

Wild soursop is still essentially restricted to Africa, between latitudes 22.5° 

North and 22.5° South. It appears to have adaptation to very low to 

moderately high rainfall regimes, occurring generally in areas with 600 to 

1,200 mm (but 716 to 2,029 mm in Tanzania; FAO, 1989), while across 

Africa requirements are for more than 600 mm annual rainfall. It can 

withstand a relative humidity as low as 44% at midday. The best 

temperatures for wild soursop growth are between 16°C and 30 o C (FAO, 

1983). 

Sugar apple is usually cultivated in the lowlands, although in Cuba it is 

reported in cultivation up to 900 m (Zayas, 1966). Sugar apple is a lowland 

tropical or marginally subtropical species, growing between latitudes 22.5° 

North and South. 

Sugar apple is native to the warmest and driest places in Central America but 

is also reported yielding well in humid regions (Popenoe, 1952). It is also 

30


frequently reported in cultivation in semi-arid climates, such as north-eastern 

Brazil (Belotto and Manica, 1994). It is relatively drought-tolerant and does 

not fruit well in high rainfall regimes (Nakasone and Paull, 1998). 

Sugar apple is more adaptable to low temperatures than soursop and more 

tolerant of high temperatures than cherimoya (Belloto and Manica, 1994). 

Fouqué (1972) reported that this species is also sensitive to long periods of 

cold. 

5.2 Soil 

Soil characters are extremely important for annona cultivation, the most 

important factor being drainage. No annona grows well in soils with drainage 

problems. High water content in the soil causes root diseases (Nakasone and 

Paull, 1998). In general, annonas are not too demanding of soil type 

(Nakasone and Paull, 1998), but produce better in fertile, well aerated, well 

drained, deep soils rich in organic matter (Zayas, 1966). 

The best soil pH for cherimoya growth is around 6.0 to 6.5 (Villachica, 

1996). In Mexico, the best physical/chemical composition for cherimoya 

cultivation was pH 6.5, with an organic matter content of 6.2%, nitrogen of 

0.25%, phosphorus of 2.8 ppm, and potassium of 0.79 meq/100 g (Agustín 

and Angel, 1997). Cherimoya grown in soils which are poor in calcium, 

phosphorus or rich in aluminium does not produce well. 

Soursop prefers deep soils with good aeration (Melo et al., 1983; Ledo, 

1992) and can be grown on a wide variety of soil types. Pinto and Silva 

(1994) reported the best soil pH to be 6.0 to 6.5, while Zayas (1966) reported 

that it is between 6.0 and 7.5, and Belotto and Manica (1994) reported that it 

is between 5.5 and 6.5. 

Custard apple is well adapted to unfavourable soil conditions. It can grow in 

soils with pH 5.0 to 8.0. Because of this high tolerance to variable soil types, 

it is reported to be a good rootstock for cherimoya and soursop (Popenoe, 

1952; Zayas, 1966). 

Wild soursop occurs in a variety of soil types (FAO, 1983), but no precise 

information on limiting conditions is available. FAO (1989) mentions its 

occurrence on coral rocks dominated by sandy loam soils (Tanzanian coast), 

stony and ferruginous soils, and gravel banks. It is noted for regenerating on 

areas that have been burnt, taking advantage of the nutrient flush. As a 

31


component of natural or semi-natural vegetation, it occurs in grasslands, 

thickets and open woodlands. 

Sugar apple grows on a wide range of soils from sandy to heavy clays. It is 

relatively shallow-rooted and can tolerate salinity to a certain degree. It is 

typical of stony soils along rivers, along the coast and on fallow land, as well 

as on hills and slopes (Von Maydell, 1986). 

5.3 Phenology 

Flowering and fruiting seasons differ among annonas, depending upon the 

geographic location and climate where they are cultivated. Phenology is 

important for planning management, harvesting and commercialisation. In 

general, the period from pollination to fruit maturity averages 5 to 6 months. 

Annonas adapted to the highest latitudes or altitudes (like cherimoya) are 

described as responding to typical seasonal regimes (autumn, winter, spring 

and summer). In general, winter is the colder and drier season, and summer 

the warmer and wetter one. Most annonas, however, are cultivated in tropical 

areas, where temperatures do not vary very much and the seasons are divided 

into rainy and dry seasons (see Table 5.1). 

The fruiting season of cherimoya in Spain (37° North) occurs at the end of 

the dry season (September to October). It is less frequent, but possible, to 

harvest it in the wet season (November to December) (Farré and Hermoso, 

1997). On the other hand, in Mexico (22.5° North), cherimoya flowering and 

fruiting occur in dry and wet seasons (flowering: February to May and 

fruiting: March to October), respectively. Fruit development takes around 6 

months (Agustín and Alviter, 1996). 

Although soursop tends to flower and fruit continuously, there are fruiting 

seasons. The fruiting season in Florida (25° North) occurs in the wet season 

(June to September) (Mowry et al., 1941). In Mexico, flowering occurs in the 

dry season (December to January) and fruiting continues into the wet season 

(May to June). In Mexico, a second flowering can also occur in June to July 

(wet season) with fruiting from November to January (dry season). In Brazil 

(Brasília at 15° South), flowering occurs in the wet season (November to 

February) and fruiting during the beginning of the dry season (April to July). 

In Puerto Rico and the Caribbean region (15-20° North), the soursop fruiting 

season extends from February and March (dry season) to September (wet 

season), with a peak in the wet season (June to August) (Bueso, 1980). This 

32


annona requires around 6 months for fruit development (Pinto, A.C.Q., 

Embrapa Cerrados, July 2003, personal communication). 

Custard apple matures in Florida during late winter and early spring (dry to 

wet season) (Mowry et al., 1941). In Mexico, its flowering occurs in the 

transition from the wet to the dry season (August to November) and fruiting 

is in the dry season (March to April). 

In Tanzania (5° South), wild soursop flowers during the beginning of the wet 

season (October to December), and along the coast in the wet season 

(December to February), while fruit maturity occurs during the rainy season 

(the peak of rain is in April) (FAO, 1983). In western and eastern Tanzania, 

fruiting takes place in the wet season, in western Tanzania from December to 

March and in eastern Tanzania from March to May. 

In Florida, the sugar apple fruiting season begins in mid summer (wet season); 

its ripening is irregular, lasting 3 months (Mowry et al., 1941). In the 

Philippines (15-20° North), fruiting occurs during the beginning of the rainy 

season (summer). In India (10-22.5° North), fruiting also occurs in the wet 

season (August to mid September) and can occur from October to November 

(the end of the wet season) (Coronel, 1994). In Mexico, flowering occurs at 

the end of the dry season (March to May) and fruiting at the end of the wet 

season (September to November). In Brazil (Brasília), flowering occurs at the 

end of the dry season (March to May) and fruiting in the wet season (December 

to January). In Brasília, flowering can be induced in the wet season (December) 

and to fruiting in the dry season (May) (Pinto, A.C. de Q., Embrapa 

Cerrados, July 2003, personal communication). 

33


Table 5-1. A quick reference guide to monthly rainfall (mm) in some important Annona production areas 

Location Species grown Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec 

Spain (Málaga) Cherimoya 203 203 178 229 152 76 25 25 76 203 229 229 

Florida (Miami) Soursop, sugar apple, custard 51 53 61 76 150 224 152 198 216 178 79 46 

apple 

Caribbean 

Cherimoya, soursop, sugar 76 56 58 94 155 112 114 135 135 140 147 119 

(Puerto Rico) 

apple, custard apple 

Mexico (Mexico City) Cherimoya, soursop, sugar 8 5 13 20 48 107 130 122 109 43 15 8 

apple, custard apple 

India (Mumbai) Sugar apple, cuatard apple 0 0 0 0 13 566 650 488 356 89 5 0 

Ecuador (Quito) Cherimoya 114 130 152 175 124 48 20 25 79 130 109 104 

Peru (Lima) Cherimoya 0 0 0 0 0 3 5 3 3 3 0 0 

Brazil (Brasilia) Soursop, sugar apple 553 457 457 305 152 50 50 76 203 406 533 610 

Tanzania 

(Dar-es-Salaam) 

Wild soursop 71 64 130 269 183 33 28 25 28 48 84 94 

During the wet season the average precipitation is 203 mm in Málaga, 159 mm in Miami, 114 mm in Puerto Rico, 4 mm in Lima, 

83 mm in Mexico City, 471 mm in Brasilia, 130 mm in Quito, 310 mm in Mumbai and 128 mm in Dar-es-Salaam. 

34

Chapter 6. Properties 

M. C. R. Cordeiro and A. C. de Q. Pinto 

6.1 Chemical properties 

Leaves, roots, bark, fruits and seeds of annonas contain numerous bioactive 

chemical substances, such as acetogenins, alkaloids, terpenes, flavonoids and 

oils. At least some acetogenins have insecticidal, cytotoxic, antitumoral, 

antifeedant, antibacterial, immuno-suppressant, pesticidal or antihelminthic 

properties (Rupprecht et al., 1990). Alkaloids, terpenes and flavonoids are 

potentially useful in medicine. A list of some of the chemical compounds 

present in annonas is given in Appendix A. 

Acetogenins isolated and characterized from different annonas have 

monotetrahydrofuran (MTH) or bis-tetrahydrofuran (bis-THF), with adjacent 

and nonadjacent bis-THF systems, in their structures (Cortés et al., 1993 a, b; 

Duret et al., 1994). These substances can be extracted from seeds using 

ethanol, methanol or petroleum ether (Rupprecht et al., 1990). 

Cherimoya: At least 6 types of acetogenins have been identified in cherimoya 

roots (Cortés et al., 1993 b; Duret et al., 1994) and some of them exhibit 

cytotoxic and antiparasitic activities. Three alkaloids have been identified 

from the leaves and stem (Fresno and Cañavate, 1983). The stems also 

contain acetogenins, amides, kauranes, purine and steroids (Chen et al., 

1998). Ethanol extracts of cherimoya seeds also have bioactive acetogenins 

(Cortés et al., 1993 a, b; Sahpaz et al., 1996; Chen et al., 1999) and alkaloids 

(Fresno and Cañavate, 1983). Moreover, cherimoya seeds have oils 

containing oleic (43%), linoleic (35%), palmitic (12%), stearic (8%), 

linolenic (1%) and traces of arachidic acids (Lizana and Reginato, 1990). 

Soursop: Roots, stems and leaves of soursop have different kinds of 

acetogenins. Some of them have antitumoral activities and act preferentially 

against human cancer cell lines (Wu et al., 1995 a, b, c; Zeng et al., 1996; 

Kim et al., 1998 a, b). Acetogenins found in soursop leaves and stems are 

used to prepare extracts that have insecticidal activities. These compounds 

are similar to anonins and muricins (Pinto and Silva, 1994). Additionally, 

biogenetic intermediaries of acetogenins are found (Gleye et al., 1997). In 

soursop seeds there are amyloids (Kooiman, 1967), acetogenins (Myint et al., 

35


1991; Roblot et al., 1993; Philipov et al., 1994; Pinto and Silva, 1994; Wu et 

al., 1995 b; Rieser et al., 1996; Yu et al., 1998), and unsaturated and 

saturated fatty acids (Bueso, 1980; Castro et al., 1984; Pinto and Silva, 

1994). The main types of unsaturated fatty acids found in soursop seeds are 

oleic (41%), linoleic (33%) and palmitoleic (2%) acids, together making up 

76% of total fats. The saturated fatty acids are palmitic (19%) and stearic 

(5%), together making up 24% (Castro et al., 1984; Pinto and Silva, 1994). 

Custard apple: In custard apple leaves and stem bark there are acetogenins 

that have cytotoxic activity and potential use in cancer treatments (Hisham et 

al., 1994). Custard apple seeds have bioactive acetogenins (Chang et al., 

1998), diterpenoids, alkaloids and n-fatty acyl tryptamines as structural 

components (Maeda et al., 1993). Diterpenoids are represented by kaurane 

and kaurene types (Maeda et al., 1993). Dopamin is also present in seeds 

(Maeda et al., 1993). In fruits, there are essential oils which account for their 

characteristic perfume and flavour. In stem and root barks there are amino 

acids and ent-kaurenoids (Fatope et al., 1996). 

Wild soursop: The most important chemical constituents found in leaves of 

wild soursop are aliphatic ketone, alkanes, alkanols, fatty acids, flavonoids, 

sterols, monoterpenoids and sesquiterpenoids (Langason et al., 1994; You et 

al., 1995). Unidentified bioactive substances found in wild soursop leaves 

reduce the feeding activity of insects (Abubakar and Abdurahman, 1998). 

There are also alkaloids, such as aporphine and (-) roemerine, with cytotoxic 

activity (Cassady, 1990). These enhance the cytotoxic response mediated by 

vinblastine in multidrug resistance to KB V1 cells and interact with P 

glycoproteins (You et al., 1995). Seeds also contain cytotoxic acetogenins 

(Sahpaz et al., 1996). 

Sugar apple: Sugar apple leaves are rich in aporphines (Salluja and Santani, 

1990) and fruits contain diterpenoids. Bark contains acetogenins (Chao-Ming 

et al., 1997; Hopp et al., 1997; 1998). Squamotacin (similar to bullatacin) 

and molvizarin acetogenins have cytotoxic activity against prostate tumour 

cell lines (Hopp et al., 1996). Fatty acid composition of seeds is: stearic acid 

(9.3%), oleic acid (37%), linoleic acid (10.9%), arachidic acid (3.3%) and 

isoricinoleic acid (9.8%) (Leal, 1990). The seeds also contain terpene 

hydrocarbon essential oils, such as alpha pirene, beta pirene, limorene, beta 

farnesene and trans orimene (Leal, 1990). 

Sugar apple seeds are also rich in acetogenins, diterpenes and saponin 

(Salluja and Santani, 1990; Li et al., 1990; Nonfon et al., 1990; 

Mukhopadhhyay et al., 1993; Chao-Ming et al., 1997; Hernández and Angel, 

36


1997). The most important acetogenins are anonins or anonacins: asimicin, 

annonastatin, bullatacin, bullatacinone and squamocin. These substances 

have toxic effects when eaten by insects and can inhibit insect growth, 

development and reproduction. The cytotoxic anonins cause 70% mortality 

of Aedes aegypti with a concentration of only 10 ppm. They act by inhibiting 

respiration (Londershausen et al., 1991 a, b). Asimicin is effective against 

insect pests, such as A. aegypti, A. vittatum, A. gossypii, Colliphora vicina, 

Epilachna varivertis, Tetranychus urticae, and the nematode Caenoharbiditis 

elegans. This compound has 256 known isomers, of which bullatacin is the 

most toxic (Li et al., 1990). Bullatacin causes 80% mortality of A. aegypti, A. 

gossypii and Diabrotica undecimpunctata when in concentrations of 1, 10 or 

24 ppm, respectively. Another powerful isomer is bullatacinone (Hernández 

and Angel, 1997). Some of these acetogenins could be used as insect 

repellents (Hernández and Angel, 1997). 

6.2 Pulp properties 

Annona pulps are useful foods because they contain proteins, fatty acids, 

fibre, carbohydrates, minerals and vitamins (Bueso, 1980; Leal, 1990; Lizana 

and Reginato, 1990). However, annona fruits do not contribute many calories 

to the diet (Kalil et al., 1979). 

Nutrients in the diet are important because they have many biological 

functions, such as providing energy and matter for growth, and regulating 

biological reactions. These functions are modulated by the quality and 

quantity of the carbohydrates, lipids, proteins, minerals and vitamins in food. 

Nutrients are divided into protectors and non-protectors. Protector nutrients 

are essential to protect organisms against pathogens, while non-protector 

nutrients provide only calories. Milk, meat and egg (principally because they 

are rich in proteins), and vegetables and fruits (mainly because they are rich 

in minerals and vitamins) are examples of foods that provide protector 

nutrients (Evangelista, 1992). 

The most important factors for a healthy diet are: (1) nutrients must be 

ingested in sufficient quantity and quality to provide nutritional and caloric 

balance. Sex, age and physical activity effect requirements for nutrients. For 

example, a 65 kg person, with strong physical activity, needs 4,000 kcal/day 

in the diet, while a person with strong intellectual activity needs only 3,000 

kcal/day. (2) The balanced diet must provide a harmonious combination of 

proteins, carbohydrates, lipids, minerals and vitamins. For example, a normal 

37


person (with normal activity) needs 10-15% proteins, 25-35% lipids and 50- 

60% carbohydrates in the diet (all percentages are related to total caloric 

intake per day). So, if we consider that 1 g of protein, lipids or carbohydrates 

gives 4, 9 and 4 kcal, respectively (in a 2,800 kcal diet), the total diet must 

have approximately 70-105 g protein, 77-103 g lipids and 350-420 g 

carbohydrates to be well balanced. Animal or plant protein, calcium and 

vitamin D are also very important, but plant proteins do not have all the 

essential amino acids for human diets (Kalil et al., 1979). 

The nutritional value of annonas is not very high because their nutrient 

content is not high, contrary to some assertions in the literature. The 

carbohydrate content is reasonable and explains why some authors have 

referred to annonas as being fruits with high caloric value. Although the pulp 

is not nutritionally important, it is flavourful and is reasonably rich in 

minerals and vitamins, making it an agreeable input to a healthy diet. 

Cherimoya, soursop and sugar apple are the most widely consumed species 

and, consequently, more is known about their nutritional composition. The 

chemical-nutritional content of custard apple is only reported by Wu Leung 

and Flores (1961) and Zayas (1966). Even less is known about the chemicalnutritional 

content of wild soursop, except for its high content of vitamin C 

and moderate levels of minerals (FAO, 1988). Known chemical composition 

of 4 species is shown in Table 6.1 and should be referred to as an adjunct to 

the text below. 

Cherimoya is commonly eaten fresh as a dessert fruit. It can also be pureed 

and used as a sauce. In Chile, it is commonly used for ice cream. The flesh is 

white, melting in texture, and moderately juicy. The flavour is sweet and 

delicate, suggestive of pineapple and banana. The edible portion corresponds 

to 60% of the fruit weight. The physical-chemical analysis of the pulp varies 

among varieties and according to the horticultural practices and the climate 

where it is cultivated. In general, the pulp carbohydrate content is high, while 

acidity is low (Table 6.1). 

The sugar content represents a mix of fructose, glucose (11.75%) and sucrose 

(9.4%). The fibre combines cellulose, hemicellulose, lignin and pectic 

substances. The degree of ripening does not interfere in this proportion, 

suggesting that fibre content is determined early in ontogeny. The protein 

percentage is reported to be the highest among commercially important 

annonas (Popenoe, 1974 a), but this is not very important nutritionally. 

Various volatile hydrocarbons, such as esters, alcohols, carbonyls and other 

compounds, are responsible for pulp flavour and aroma (Idstein et al., 1984). 

38


These compounds could be used to flavour processed foods based on 

annonas. The vitamin A content is low, but it is a good source of thiamine, 

riboflavin and niacin (NRC, 1989). 

Soursop pulp is considered to be aromatic and exotic, and is consumed 

mostly after processing into cold beverages or sometimes fresh. The edible 

portion constitutes 67.5% of total fruit weight (Bueso, 1980). The 

characteristic flavour of this fruit is produced by amyl and geranyl caproic 

acids (Bueso, 1980; Pinto and Silva, 1994). The processed pulp is used to 

prepare juices and ice creams (Pinto and Silva, 1994). In Cuba, the pulp is 

processed to prepare an alcoholic drink called champola (Popenoe, 1974 b). 

The most important sugars are fructose (1.8%), glucose (2.3%) and sucrose 

(6.6%). The most common acid in its pulp is citric, with some malic and, less 

commonly, isocitric acid. Soursop fruit contains vitamins A and B 5 

. Also, it 

is the only annona with tannins in its pulp (Castro et al., 1984). It was 

suggested that pectin in the fruits could become an important by-product 

(NAS, 1975), but this has not been developed. 

Custard apple flesh is creamy yellow, rich and sweet, with low acidity 

(Wester, 1913). However, its flavour is not considered comparable to that of 

cherimoya or sugar apple. Wu Leung and Flores (1961) reported that the 

edible part of custard apple is 45%, of which 78.6 g/100 g is water, which is 

similar to Zayas' (1966) report. As in other annonas, other components are of 

low to moderate nutritional importance. 

Wild soursop flesh is scant, but sweet and aromatic (Wester, 1913). It has a 

white pulp and a pleasant pineapple-like odour (FAO, 1983). No physicalchemical 

composition data were reported in the literature available to us. 

Sugar apple pulp is slightly granular, creamy yellow or white, sweet, with a 

good flavour and low acidity (Mowry et al., 1941). It is considered the 

sweetest of the annona fruits (FAO, 1990) and is generally consumed fresh as 

a dessert fruit. The edible portion is 28-37% of the total fruit weight; seeds 

correspond to 31-41% and rind to 23-40% (Leal, 1990). The carbohydrates 

present in the pulp are fructose (3.5%), sucrose (3.4%), glucose (5.1%) and 

oligosaccarides (1.2-2.5%). 

39


Table 6-1. Chemical composition of 100 g of edible pulp of cherimoya, 

custard apple, soursop and sugar apple fruits 

Components Cherimoya Custard 

apple 

Soursop 

Water (g) 77.3±3 

75.8±2.8 81±2.5 

(74.6-83.3) 

(77.9-81.7) 

Proteins (g) 1.6±0.6 1.85±0.05 1±0.55 

(1.0-2.9) 

(0.69-1.7) 

Lipids (g) 0.3±0.2 0.35±0.15 0.6±0.3 

(0.1-0.5) 

(0.3-0.8) 

Carbohydrates (g) 18.42±4 18.7 17.25±0.1 

(11.7-22.0) 

(16.3-18.2) 

Fibre (g) 1.64±0.5 2.55±0.35 0.86±0.1 

(1.0-2.0) 

(0.78-0.95) 

Total acidity (g) 0.58±0.19 - 1.0±0.3 

(0.39-0.77) 

(0.7-1.3) 

Ash (g) 0.7±0.1 0.95±0.15 0.61±0.2 

(0.6-1.0) 

(0.4-0.86) 

Energy (calories) 68.6±13.4 75 65±5 

(56-101) 

(64-71) 

Calcium (mg) 27.14±5 24 15±7 

(21.7-34.0) 

(8.8-0.22) 

Phosphorous (mg) 35.2±18 26 28±1 

(30.2-47.0) 

(27.1-29) 

Iron (mg) 0.6±0.2 1.0 0.7±0.1 

(0.4-0.8) 

(0.6-0.82) 

Vitamin A (mg) - Traces 14.45±5.45 

(8.9-20) 

Vitamin B 1 

(mg) 0.09±0.03 0.07 0.07±0.01 

(0.06-0.12) 

(0.06-0.077) 

Vitamin B 12 

(mg) 0.12±0.2 0.12 0.08±0.035 

(0.11-0.14) 

(0.05-0.12) 

Vitamin B 5 

(mg) 0.8±0.2 0.7 1.2±0.3 

(0.6-1.02) 

(0.89-1.52) 

Ascorbic acid 11.5±5.5 30 19.4±3 

(mg) 

(4.3-17) 

(16.4-22) 

Sugar apple 

72.6±2.4 

(68.6-75.9) 

1.6±0.8 

(1.2-2.4) 

0.4±0.3 

(0.1-1.1) 

19.6±1 

(18.2-26.2) 

1.4±0.6 

(1.1-2.5) 

0.1 

0.7±0.1 

(0.6-1.3) 

96±10 

(86-114) 

26.2±6 

(17-44.7) 

42±14 

(23.6-55.3) 

0.8±0.5 

(0.3-1.8) 

0.005±0.001 

(0.004-0.007) 

0.1±0.01 

(0.10-0.11) 

0.13±0.05 

(0.057-0.167) 

0.9±0.3 

(0.65-1.28) 

37.38±4.62 

(34-42.2) 

Tannins (mg) ±0 ±0 85.30 ±0 

Sources: (Wu Leung and Flores, 1961; Zayas, 1966; Bueso, 1980; Castro et 

al., 1984; Leal, 1990; Lizana and Reginato, 1990). Means ± standard 

deviations. 

40

Chapter 7. Uses 

M. C. R. Cordeiro, A. C. de Q. Pinto and S. R. M. de Andrade 

7.1 Food products 

In general, annonas are consumed as fresh fruits, but they are also widely 

used in semi-processed and processed products, especially desserts. As world 

demand for exotic flavours and healthy foods expands, the use of annona 

fruits is also likely to expand. 

The cherimoya fruit is consumed mostly fresh, generally chilled and often 

with salt and lemon. Fruit pulp is often mixed with wine, milk (to make milk 

shakes) and yoghurt, processed into ice cream and sherbet, and baked into 

cookies and pastries (Bueso, 1980; Lizana and Reginato, 1990; Leal, 1990; 

Bonaventure, 1999). Most of these preparations, and others, can be made at 

home (Ibar, 1986). 

Soursop fruits are occasionally consumed fresh or more commonly made into 

juices, ice creams (Pinto and Silva, 1994) or sherbets (Popenoe, 1974 b). 

Most people consider it to be too acid for eating fresh, but it is esteemed for 

making refreshing drinks (Mowry et al., 1941), nectars, ice creams and 

similar foods. Nectar (sweetened pulp) can be prepared and used after 

dilution with 3 parts of water. In Java, Indonesia, fruits of soursop are added 

to soup (sajoer). 

The flavour of custard apple pulp is considered to be poor and hence of little 

commercial value (Popenoe, 1974 a), although it is a popular backyard fruit 

and attracts children. Chilling of this fruit, as well as other annonas, improves 

the flavour (Mowry et al., 1941). 

Wild soursop fruits are sold in local markets in Africa. The fruit has a 

pineapple-like odour and sweet taste (FAO, 1983). It keeps for only a few 

days. It is used in sherbets, ice creams and for making drinks (FAO, 1988). 

The sugar apple is consumed as a fresh dessert fruit, or used for preparing 

juice and ice cream. In the latter case, it should not be pasteurised or cooked, 

but simply blended into the semi-solid cream just before freezing (Sturrock, 

1959). Leal (1990) reports that it can also be used to make wine, as can 

cherimoya. 

41


7.2 Industrial food uses 

42 

Cherimoya is widely consumed in a processed form. Industrial processing 

depends on development of freezing techniques for pulp preservation. In a 

simple freezer, frozen cherimoya can be successfully preserved for 120 days. 

For freezing, the fruits should be peeled, preferably with stainless steel 

knives or by chemical peeling with caustic soda (20%). The pulp or fruit 

slices should be bagged in polyethylene prior to freezing and sugar can be 

added if desired (Lizana and Reginato, 1990). Additives, such as EDTA, 

ascorbic acid and citric acid, preserve against oxidation. 

Soursop is the other annona of which the pulp or nectar can be frozen, 

processed and used industrially (Beneto et al., 1971). It is perhaps the best 

annona for industrial processing and commercialisation because of its exotic 

taste and agreeable aroma. The processed pulp can be preserved by 

pasteurisation or freezing (Zayas, 1966) and conditions for these processes 

and storage have been developed. The quality of the processed product 

depends on total sugars and ascorbic acid retention, low acidity, viscosity and 

presence of pectinesterase activity. The final product should have an 

agreeable flavour and a good consistency. Temperatures of 93°C and 107°C 

decrease the quality in unsweetened and sweetened frozen soursop puree, 

respectively (Bueso, 1980). The high temperatures and exposure also 

influence pectinesterase activity and ascorbic acid retention. Moreover, 

soursop fruit composition differs among varieties (Pinto and Silva, 1994) and 

cultural treatments. Hence, it is necessary to work out the best conditions for 

each variety in each production environment. 

Several authors have reported on soursop pulp or nectar processing, freezing 

and canning (e.g., Sánchez-Nieva, 1953; Payumo et al., 1965). Bueso (1980) 

reported that fruits should be picked by hand, washed with chlorinated water 

and peeled by hand. The edible pulp should be extracted from the fruit using 

a blender, a pulper or dispersed in a sugar syrup. At this stage, care should be 

taken not to mix peel with pulp or to break the seeds, as these are dangerous 

because of the presence of bioactive compounds (see Chapter 6). After 

extraction, the pulp is strained through a screen. Sánchez-Nieva (1953, in 

Bueso, 1980) commented that extraction of the pulp should be done quickly 

so as to avoid aeration and oxidation. 

Holanda et al. (1980) reported that the fruit should be processed after 

selection and further maturation (3-5 days depending on temperature) in an 

acetylene acid maturation chamber at 12.5-16°C and 80% relative humidity


(RH), before finally being weighed and washed with 0.5% potassium sorbic 

acid. Fusagri (1982) demonstrated that 12.5°C is the best temperature to 

mature and store soursop fruits; at this temperature the fruits are preserved 

for up to 5 days. After maturation, the fruits are selected again, peeled and 

the pulp separated mechanically. 

Pulp can be pasteurised at 85°C and nectar at 90.6°C. Both can then be stored 

in cans for a year at 29.4°C (Bueso, 1980). More recently, Umme et al. 

(1997) established that the best conditions for pasteurisation of soursop pulp 

are a pulp : water mixture of 2:1, 78.8°C for 69 s at pH 3.7. Under these 

conditions, the inactivation of pectinesterase enzymatic activity is maximized 

and ascorbic acid is preserved, which helps to maintain quality. 

Frozen soursop puree can be stored for 400 days at -23°C (Bueso, 1980). To 

prepare this puree, the sugar content should be adjusted to 45-59°Brix; with 

45°Brix, ascorbic acid retention is higher in the pasteurised puree. Ascorbic 

acid should be added to the pasteurised puree at a rate of 0.5-1.5 g/0.45 kg; 

this improves the retention of the nectar's flavour and serves as an antioxidant 

to control polyphenol oxidase-mediated pulp darkening of the fruit 

juices (De Oliveira et al., 1994). The sweetened or unsweetened frozen 

nectar can be preserved by pasteurisation. Unsweetened nectar can be 

pasteurised at a temperature of 90.6°C, while higher temperatures give 

inferior final products. The same is observed for sweetened nectars. While 

higher temperatures reduce pectinesterase activity they also increase ascorbic 

acid retention. 

Another industrial application involves the extraction of essential oils present 

in soursop pulp. These oils, such as esters of aliphatic acids, have potential to 

improve the flavour of processed fruit products (Jirovetz et al., 1998). 

Sugar apple pulp can also be processed and frozen. Its industrial processing 

is less important than that of cherimoya and soursop, but it is used to prepare 

drinks, fermented liquors and ice creams (Prasada and Rao, 1984). For this 

purpose, the fruits should be peeled and cut by hand, and the seeds extracted 

from the pulp. The pulp is heated for 3 min at 70°C, stored in jars and then 

double boiled for 15 min at 95°C. Sealed jars can be stored at 27°C for 150 

days, during which time the acidity and total reducing sugar concentrations 

increase and ascorbic acid content decreases. A similar study was done on 

the nectar prepared with acclimatised (in an ethylene chamber at 16°C and 

RH 80%) and non-climatized (at room temperature for 72 h) fruits. Before 

the pasteurisation process, the pulp was adjusted with water, sugar and citric 

acid (pulp 1 kg, water 2.51 kg, sugar 0.37 kg and citric acid 1 g) (Leal, 1990). 

43


The same pasteurisation process was used for the pulp (for 3 min at 70°C, 

stored in jars and then double boiled for 15 min at 95°C). After 150 days of 

storage the only observed change was an increase in total sugars by the end 

of the first month and the complete loss of vitamin C content (Leal, 1990). 

7.3 Medicinal uses 

Various plant parts are also widely used in folk medicine, because of the 

bioactive compounds (mainly acetogenins, alkaloids and flavonoids) found in 

the roots, leaves, bark, fruits and seeds (listed in Chapter 6 and Appendix A). 

Acetogenins are potential anti-cancer treatments, as they have cytotoxic 

effects (Chang et al., 1993; Cortés et al., 1993 b). Flavonoids present in the 

seeds, roots, bark, stems and fruits are potential chemo-preventive agents, 

given evidence that they decrease tumour incidence (for a review, see 

Cassady, 1990). Appendix B provides a summary of the known uses, the 

most important of which are discussed below. 

When a herbal product finds widespread use as a medicine, particularly for 

primary health care of people with little access to modern health services, it 

is important that natural sources are not over collected and depleted. 

Cultivation becomes an imperative, as does the standardization of herbal 

preparations (Bajaj and Williams, 1995), and there is some evidence that this 

is occurring for A. squamosa as an anti-bacterial herb (Anjaria, 1989). In 

Brazil, the National Sanitary Vigilance Agency (ANViSa) has recently 

required both evidence of bioactivity and lack of toxicity for medicinal plants 

used as phytopharmaceuticals. The latter requirement is especially important 

in annonas, given the toxicity of many of the bioactive compounds. 

Cherimoya roots have aporphine alkaloids, such as roemerine, anonaine and 

dehydroroemerine. These have relaxant effects, provided by the blockage of 

calcium movement across the cell membrane through voltage-operated 

channels and disruption of the alpha-1 adreno-receptors connected to the 

receptor-operated channels (Chuliá et al., 1995). Ethanol extracts of 

cherimoya seeds are used in folk medicine for their insecticidal and antiparasitic 

activity (Bories et al., 1991). The dark-yellow resin extracted from 

the seeds contains substances that dilate pupils, intensify photophobia, cause 

dryness of the mouth, burning of the throat, nausea, vomiting and other 

symptoms resembling the effects of atropine remedies (Lizana and Reginato, 

1990). In Mexico, the powder of two seeds from a fruit, mixed with water or 

milk, is a potent emetic and cathartic remedy (Lizana and Reginato, 1990). 

44


Seeds contain a reddish oil and caffeine. Flowers of cherimoya are used to 

flavour snuff in Jamaica but whether this is used medicinally is not clear. 

Some soursop root acetogenins are known to have cytotoxic effects (Gleye et 

al., 1998): panatellin, uvariamicin IV, uvariamicin I, reticulatacin, 

reticulatacin 10-one and solamin. The bark contains alkaloids. The leaves 

have essential oils with parasiticide, anti-diarrhoea, rheumatological and antineuralgic 

properties (Moura, 1988). Boiled water infusions of leaves have 

anti-spasmodic, astringent, gastric properties (Calzavara et al., 1987; Khan et 

al., 1997), help treat diabetes and gastric upsets (Calzavara et al., 1987), and 

are used in kidney ailments (Duke, 1970). The cooked flowers and petals are 

used for healing eye inflammations; the treatment requires 2-3 washes a day 

(Calzavara et al., 1987). 

Immature soursop fruits have medicinal properties against dysentery, 

cankers, diuretic, scorbutic, anti-thermical processes, skin diseases, rashes, 

fever, malaria, peptic ulcers, colic and oedema (Khan et al., 1997). The peel 

from immature fruits has constituents that act against atonic dyspepsia, 

diarrhoea and chronic dysentery; it is astringent and provokes vomiting 

(Calzavara et al., 1987). The acid pulp is used to heal foot parasites and 

icteric liver diseases (Calzavara et al., 1987). The fruit also has properties 

that act on the biliary vesicle (Calzavara et al., 1987). The seeds have antispasmodic 

and anti-parasitic properties (Moura, 1988; Bories et al., 1991; 

Philipov et al., 1994). They contain amyloids, oleic acid and steroids 

(Kerharo and Adam, 1974; Asolkar et al., 1992). 

Wild soursop roots, leaves and bark are also used in folk medicine (FAO, 

1983). The roots are used to treat cancer, convulsions, venereal disease, 

diarrhoea, dysentery, fever, filariosis and male impotency, and have antineoplasic 

and anti-protozoal activities (Fatope et al., 1996). The leaves are 

used for diseases of the eye, stomach and intestines (Philipov et al., 1995; 

You et al., 1995). Alcoholic leaf extracts have anti-spasmodic and relaxant 

activity on the smooth muscles, anti-ulcer activity against indomethacin 

induced ulcers and reduce the effect of stress on ulcer induction. These 

effects are produced by various compounds, including flavonoids, alkaloids, 

tannins and saponins (Langason et al., 1994). Moreover, the leaves contain 

compounds that have insecticidal effects and are used to control insect pests 

(Abubakar and Abdurahman, 1998). The bark is utilised as a vermifuge and 

snakebite treatment (Philipov et al., 1995). The stem bark contains 4-entkaurenoids 

that have cytotoxic activity against tumour cell lines (Fatope et 

al., 1996). Other wild soursop uses in folk medicine include treatments for 

45


pain of the chest, swelling and trypanosomiasis (You et al., 1995; Fatope et 

al., 1996), and treatment of convulsions in children and against cancer. 

Sugar apple has many alkaloids, such as aporphine, roemerine, norcorydine, 

corydine, norisocorydine, glaucine and anonaine in different parts of the 

plant (Kowalska and Puett, 1990). The roots are used to treat acute dysentery, 

depression and spinal marrow diseases, while leaves have been used in cases 

of prolapse of the anus, sores and swelling (Chao-Ming et al., 1997). Tea 

made from the roots is highly purgative, while when it is made from the 

leaves is mildly laxative (Leal, 1990). It has a tonic effect on the digestive 

tract (Leal, 1990). Ethanol extracts of the bark appear to have anti-tumour 

activity (Hopp et al., 1996, 1997, 1998). The leaves have an alkaloid, 

higenamine, and this is a cardiotonic active principle (Wagner et al., 1980). 

Sugar apple fruits contain 16-b, 17-dihydroxykauran-19-oic acid, which has 

demonstrated anti-HIV activity (Wu et al., 1996). Seed extracts are very 

poisonous and have insecticidal properties (Pandey and Varma, 1977; Qadri 

and Rao, 1977; Hernández and Angel, 1997); saponin, extracted from the 

seeds, haemolyses red blood cells and is toxic to fish (Salluja and Santani, 

1990). In India, the extract of the seeds is used to provoke abortion by tribes 

in Madhya Pradesh State (Salluja and Santani, 1990), often combined with 

leaves of Plumbago zeylanica. Constituents of the leaves and tender stems 

are itemized in Asolkar et al. (1992). 

The folk and modern medicinal uses of the annonas are clear, but this chapter 

should not be used for self-medication, as the toxic properties of most of 

these compounds can have undesirable side effects. Caparros-Lefebvre et al. 

(1999) showed that the alkaloids present in the leaves, bark and seeds of 

annonas, when consumed for their sedative and hypnotic effects in the 

French West Indies, are responsible for inducing neurotoxic effects with 

symptoms of Parkinsonism. Hence, any medicinal use of the annonas should 

only be carried out with medical guidance. 

7.4 Other uses 

The annonas have a number of other non-medicinal important uses for their 

chemical constituents. The acetogenins with insecticidal properties, present 

in roots, stems, leaves and seeds, can be prepared domestically as powders or 

by extracting them with water, acetone, ethanol, petrol ether, ethylic ether or 

hexane solvents. These extracts can be very potent insecticides, even in 

diluted form, and proper protection should be used when handling. An oil can 

46


also be extracted from seeds and used as an insecticide (Hernández and 

Angel, 1997). The seeds of sugar apple yield an oil suitable for soap making, 

and the cake can be used as a manure (Mishra et al., 1979, Salunkhe and 

Desai, 1984). 

Wild soursop bark can be used to produce a yellow or brown dye (in Uganda) 

and its wood is used for making tool handles (FAO, 1983). As is the case 

with many woody species in areas of subsistence agriculture, the plant is 

multi-purpose. The leaves and young shoots are used as vegetables, the 

flower buds are used to flavour foods and the bark is used for rope making. 

Sugar apple prunings are valuable for thatch in India because they are not 

attacked by white ants (Singh, 1992). 

In general, the annonas offer potential for agroforestry, although this 

potential is seldom exploited. The presence of annonine in the leaves, stems 

and other parts make the plants bitter to goats or cattle. Aiyelaagbe (1994) 

reported on a system that improved the productivity in a cashew-coconut 

system in Kenya and which could also be adopted for annona production. 

However, care should be taken for annona production as the plants do not 

perform well under low light intensity conditions, which may be created with 

combined planting. 

Lastly, various species can be used as rootstocks to which other desirable 

species can be grafted. Since A. reticulata can withstand diverse ecological 

conditions and survive long dry periods, it is very useful as a vigorous 

rootstock. A. diversifolia also has a similar ecological amplitude, but has 

been less widely tested. 

47

Chapter 8. Genetic Resources 

F. R. Ferreira and A. C. de Q. Pinto 

8.1 The annona genepool 

Annona species are widely distributed and their genetic resource 

conservation has achieved a degree of world-wide attention. Until recently, 

the centres of diversity of Annona seemed to contain inexhaustible supplies 

of genetic materials for plant breeding. However, these genetic materials 

have been changing rapidly as a result of genetic erosion in both cultivated 

and wild annonas. Human pressure on natural ecological systems, leading to 

the destruction of wild species, and the introduction of improved new clonal 

varieties, which have replaced many landraces, has promoted the loss of the 

genetic variability that had accumulated over a period of thousands of years 

of natural evolution and human directed domestication (Ng, 1991). 

Chromosome numbers among Annona species do not vary significantly. 

Kessler (1993, cited by Scheldeman, 2002), reported that most of the Annona 

species present a chromosome number of 2n = 2x = 14 or 16, except for A. 

glabra which is a tetraploid species. 

The most important cultivated Annona species not treated in this book 

include: ilama or annona blanca (A. diversifolia Saff.) from Central America 

and Mexico; pond apple or alligator apple (A. glabra L.) from Tropical 

America and West Africa, cultivated as a medicinal plant rather than a fruit 

(Scheldeman, 2002). This author also mentioned mountain soursop or 

cimarrona (A. montana Macfad), soncoya or negro head (A. purpurea Moc. 

et Sessé), and posh té or cawesh (A. scleroderma Saff.), all three of which are 

from Central America. A. diversifolia, A. montana and A. muricata are quite 

similar morphologically, and they can be cross-grafted with reasonable 

compatibility. Along with the cross-fertility of A. cherimola and A. squamosa 

there are clearly many aspects of species relationships that are by no means 

well studied yet (George et al., 1999). 

There are very extensive areas in which diversity of numerous species has 

been observed (Table 8.1), which suggests that certain specific regions need 

targeted exploration. These include the mid-elevation valleys of the Andes, 

many parts of Brazil, Mexico, Guatemala, Honduras and the Antilles. 

48


Table 8-1. Centres of origin and diversity of some Annona species 

Species 

Centres of Origin 

A. aurantiaca Brazil (Mato Grosso, Goias and Minas Gerais) 

A. cacans Brazil (Savannah regions) 

A. cherimola Andean valleys of Ecuador, Peru and Chile 

A. coriacea Brazil (Mato Grosso do Sul) and Paraguay 

A. crassifolia Brazil (São Paulo, Goias and Bahia) 

A. diversifolia Southwestern Mexico, Guatemala and El Salvador 

A. furfuracea Brazil (Mato Grosso, São Paulo, Goias and Minas Gerais 

A. glabra Central America, Antilles, Ecuador, Brazil 

A. longifolia Mexico (Jalisco) 

A. longipes Mexico (Veracruz) 

A. montana West Indies, Antilles, tropical South America 

A. muricata Antilles, tropical America 

A. mutans Southern Brazil, Paraguay, Northern Argentina 

A. paludosa Guyana (Savannah regions) 

A. purpurea Southern Mexico and Central America 

A. reticulata Antilles, tropical America 

A. salzmannii Brazil (Pernambuco) 

A. scleroderma Southern Mexico, Guatemala 

A. senegalensis East Africa 

A. spinescens Brazil (Piauí, Bahia, Goias) 

A. spraguei Panama 

A. squamosa Antilles, tropical America 

A. testudinea Guatemala, Honduras 

A. xespertonium Brazil (Bahia) 

A. senegalensis is widespread in sub-Sahalian tropical Africa but nothing is 

known about patterns of variation. A related smaller species, A. stenophylla 

Engl. & Diels, occurs in Botswana, Namibia, Zimbabwe and Malawi, and is 

a seasonal staple for bushmen (FAO, 1983). 

Diversity is still to be found in most of the areas where annonas are backyard 

crops. In these agroecosystems, diverse seedlings are raised and fruit quality 

varies considerably. Commercial production using propagation by budding or 

grafting onto local rootstocks is rare. 

The conservation of genetic resources requires both in situ and ex situ 

conservation. In situ refers to the preservation and protection of genetic 

resources in their natural habitats (Lloyd and Jackson, 1986), while ex situ 

conservation is the preservation of genetic resources outside of natural 

habitats. 

49


8.2 In situ conservation 

The establishment of protected natural areas constitutes one of the principal 

strategies for in situ conservation of wild populations, allowing for their 

continued evolution. The major criteria to select areas by genetic reserves are 

diversity, intrinsic fragility, vulnerability, and high degree of endemism with 

current and potential use. Generally, in situ conservation is practised in preestablished 

protected areas, where inclusion of significant Annona genetic 

diversity is a random event, rather than planned. Nonetheless, conservation 

areas throughout the Americas and in central-eastern Africa should be 

surveyed for the presence of Annona populations, both wild populations of 

the species discussed in this book, and wild species and populations of other 

annonas. However, in situ conservation is not always possible or acceptable 

(Ndambuki, 1991). 

In situ conservation also includes on-farm (including backyard) conservation. 

Incentives could be given to large and small farmers, and to indigenous 

peoples, to continue cultivation using traditional agricultural methods. 

Ideally, agro-ecosystems should be preserved in their totality and should be 

evaluated comprehensively. 

The economic environment of the farm household determines the degree of 

genetic diversity used in its agricultural system (Goeschl, 1998) and, 

consequently, the amount available for on-farm conservation. The causal link 

between market conditions and conservation efforts on-farm offers scope for 

policy interventions, such as deliberate changes in economic parameters. 

Goeschl (1998) suggests both market and non-market incentives, the latter 

directed at the individual farm situation. 

In Tocantins State, northern Brazil, the Brazilian Corporation for 

Agricultural Research (EMBRAPA) has started a project with the Kraô, an 

indigenous group, to encourage the conservation of their genetic resources, 

especially on-farm genetic resources. The first step of this project is to carry 

out a survey of species occurring in this area. Ferreira and Bustamante (2000) 

commented that ethnobotany can help to establish new alternatives for 

conservation and use of genetic resources via on-farm conservation, 

including for Annona species. No other references have been found on the in 

situ conservation of Annona species. 

50


8.3 Ex situ conservation 

Ex situ conservation includes various strategies, such as seed storage, in vitro 

culture and field genebanking. Annona seeds show an orthodox response to 

desiccation and exposure to sub-zero temperatures. Cherimoya seeds tolerate 

desiccation to 4.8% moisture content, while soursop seeds tolerate 

desiccation to 5% moisture content. The seeds of sugar apple tolerate 

desiccation to 1.5% moisture content, and no viability loss occurred during 6 

months of hermetic storage at -20°C (Hong et al., 1996). These authors 

suggest that Annona seeds can be conserved in conventional seed genebanks 

under conditions of 18°C or less, in airtight containers at a seed moisture 

content of 5 ± 1%. 

In vitro culture techniques can be used for collecting, exchange and ex situ 

conservation of species that produce seedless fruits, as well as for 

vegetatively propagated plants, including annonas. In vitro storage can be 

done by using slow growth techniques, when medium-term preservation is 

sufficient, or by cryopreservation in liquid nitrogen at -196°C, if the need is 

for long-term preservation. Both techniques present great advantages for 

germplasm conservation. In vitro culture also offers the possibility of 

eliminating pathogens, and thus conserving and exchanging germplasm 

under disease free conditions. Despite their potential, in vitro conservation 

techniques are currently used to a limited extent only. This is due principally 

to the lack of research to develop protocols for each species (Ashmore, 

1997). Although several papers have appeared on annona tissue culture 

(Rasai et al., 1995; Lemos and Blake, 1996; Padilla, 1997; Castro et al., 

1999; Encina et al., 1999; Lemos, 2000), a great deal of work remains to be 

done on development of methods for in vitro propagation for germplasm 

conservation of Annona species. 

Emphasis on in vitro research should be placed on conserving specific clonal 

material which is well documented. Much of the range of variation can be 

conserved using seed storage, and this is more cost-effective than attempting 

large in vitro programmes. Despite the availability of seed and in vitro 

conservation techniques, in practice the majority of Annona genetic resources 

are stored in field genebanks, also called clonal repositories or collections of 

living plants, which face higher risks of disease, human error and 

environmental hazards than other conservation techniques (Ferreira, 2001). 

These collections seem to be mainly breeders' collections, and are rarely 

representative of the range of Annona variability that needs to be conserved. 

51


There is an urgent need to survey and collect wild materials, primitively 

cultivated forms and varieties of Annona species. However, primary 

emphasis needs to be on improving the agronomic and economic yields of 

each species in the range of habitats where they are grown. 

A total of 1,741 germplasm accessions of eleven identified species, one 

interspecific hybrid and various Annona spp. are documented (IPGRI, 2000), 

with a surprisingly low percentage of duplication across the 67 institutional 

collections in 34 countries (Table 8.2). Due to their commercial importance, 

the three species with the largest number of accessions are A. cherimola, A. 

muricata and A. squamosa. 

Considering that almost all conserved Annona germplasm is maintained in 

field collections, which are subject to abiotic and biotic stress conditions, 

such as flooding, drought, pathogen or insect infestations, the low percentage 

of duplication is a matter of considerable concern. The first step to remedy 

this has been taken by the Spanish government, in co-operation with IPGRI, 

through the establishment of a cherimoya genebank in Peru (Coppens 

d'Eeckembrugge et al., 1998). Besides this cherimoya genebank, Ecuador is 

establishing an Annona collection on the same basis as the Peru genebank (G. 

Coppens d'Eeckembrugge, Cali, 2001, personal communication). Other 

actions are needed, since these two actions only target cherimoya. 

A global strategy for collecting, evaluating and conserving germplasm needs 

to be thought out and implemented. This is particularly important since 

Annona collections are scarce in most of the major areas of diversity, such as 

Honduras, Mexico and the Antilles. In addition, the need of long-term 

financial commitment for germplasm banks, especially field genebanks of 

fruits in their centres of diversity, is important. It was observed in Amazonia, 

for example, that various collections were in advanced stages of 

deterioration. In part, this is due to a lack of breeders using the collections, 

although these collections are valuable as sources of genetic materials for 

testing in different areas or for exchanging among countries even if breeders 

are not locally available (Arkcoll and Clement, 1989). 

52


Table 8-2. Number of Annona accessions in germplasm collections around the world 

Species/Country che div ret squ mur mon pur gla pit cin scl A.che x 

A.squ 

spp No. 

Collections 

Total 

Accessions 

Australia 10 1 3 1 4 - - - - - - 22 - 3 41 

Brazil* 2 - 9 92 124 3 1 5 - - - 5 16 11 257 

Cameroon - - 4 1 4 - - - - - - - - 1 9 

Costa Rica 24 1 4 1 67 1 5 2 1 - - 2 4 5 112 

Cuba 2 - 6 5 9 1 1 1 - 2 - - 2 2 29 

Cyprus 4 - - - - - - - - - - - - 1 4 

Ecuador 218 - 1 1 5 - 20 - - - - - - 5 245 

El Salvador - 23 4 - 30 - 2 2 - - - - - 1 61 

France 4 - 2 4 13 - - - - - - 2 - 1 25 

Germany 1 - - 1 1 - - - - - - - - 1 3 

Ghana - - 1 1 1 - - - - - - 1 - 1 4 

Grenada - - - 2 - - - - - - - 2 - 1 4 

Guatemala - 3 6 - 4 - 5 8 - - 2 - 1 1 29 

Honduras - 1 - - 1 - 1 1 - - 1 - 1 2 6 

India - - - - - - - - - - - - 10 1 10 

Israel - - - - - - - - - - - - 20 1 20 

Jamaica - - - 1 1 - - - - - - - 9 2 1 

Malawi - - - - - - - - - - - - 3 1 3 

Mexico 3 - - 1 - - - - - - - - - 1 4 

Panama - - - - 13 - - - - 1 - - - 2 14 

Papua New Guinea - - - - - - - - - - - - 4 1 4 

53


Total 

Accessions 

spp No. 

Collections 

Species/Country che div ret squ mur mon pur gla pit cin scl A.che x 

A.squ 

54 

Peru 62 - - - 9 - - - - - - - - 5 71 

Philippines - - 11 43 7 - - - - - - 8 14 2 83 

Portugal 7 - - - - - - - - - - - - 1 7 

Saint Lucia - - - - 2 - - - - - - - - 1 2 

Seychelles - - - - 5 - - - - - - - - 1 5 

South Africa 11 - - - - - - - - - - - - 1 11 

Spain 291 - - 1 - 1 - 1 - - - 5 - 1 299 

Sudan - - - 7 - - - - - - - - - 1 7 

Suriname - - - - 3 - - - - - - - 3 2 6 

China 7 - 1 1 1 1 - 1 - - - 3 - 1 15 

Tanzania 2 - - - 2 - - - - - - - - 1 4 

USA - 3 17 13 263 - - 2 - - - - 31 4 329 

Venezuela - - - - 7 - - - - - - - - 1 7 

Total 648 32 69 176 576 7 15 43 1 3 3 50 118 67 1741 

Source: IPGRI, 2000; *Updated by authors; che = A. cherimola; squ = A. squamosa; div = A. diversifolia; ret = A. reticulata; mur = 

A. muricata; mon = A. montana; pur = A. purpurea; gla = A. glabra; pit = A. pittieri; cin - A. cinerea; scl = A. scleroderma; spp = 

Annona spp.

Chapter 9. Genetic Improvement 

9.1 Introduction 

A. C. de Q. Pinto and S.R.M. de Andrade 

Although restricted to only a few species, principally cherimoya, 

improvement programmes of annonas have made great contributions by 

producing important cultivars with good yields and fruit quality, and that 

more closely meet consumers' demands. The development of new cultivars is 

not a simple task. In many cases the major constraints are agronomic, as well 

as the lack of prolific cultivars to start improving. 

9.2 Cytogenetics and genetic aspects 

The chromosome numbers of cherimoya, custard apple, soursop and sugar 

apple are 2n = 14 to 16 (Nakasone and Paull, 1998; George and Nissen, 

1992; Koesriharti, 1992). Although there is some variation in chromosome 

number, they are all diploids, 2n = 2x. The chromosome number of wild 

soursop is not reported. This slight variation in chromosome number may 

explain the ease or difficulty of interspecific hybridisation and grafting, and 

warrants further work to determine if intra-specific variation also exists. 

Some related species, e.g., A. glabra, are known to be tetraploid (Kessler, 

1993, cited by Scheldeman, 2002). 

Generally, cross-pollination between annonas is conducted primarily to 

determine compatibility for increasing fruit set (Nakasone and Paull, 1998) 

and occasionally for new hybrid development. Samuel et al. (1991, cited by 

Nakasone and Paull, 1998) commented that crosses among soursop and other 

annonas, such as cherimoya, ilama, custard apple or sugar apple, have not 

been successful. This may reflect the genetic distance between soursop and 

the others. However, there is a dearth of information on species' 

relationships, so this is a topic that needs to be researched. 

The extensive morphological diversity, much of it genetically based, within 

all Annona species (Page, 1984) not only offers great potential for breeding, 

but also lowers the possibility of easily selecting a cultivar with all the 

possible desirable characters. Considerable variation exists among cultivars 

55


and seedlings of cherimoya, but sugar apple and soursop are reputed to be 

less variable (George and Nissen, 1992; Pinto and Silva, 1996). 

The influence of Annona rootstocks on scion behaviour is also quite marked, 

and genetic variability within seedling rootstock lines and between different 

rootstock species induces wide variability in scion performance (Page, 1984). 

This kind of genotype and environment interaction requires much more study 

than it has received to date. 

Seedling populations of some Annona species, such as sugar apple and 

soursop, are known to be rather uniform (George and Nissen, 1987). There 

are some growers using seedlings in commercial orchards in Brazil (Pinto 

and Ramos, 1999). Since annonas are considered out-crossing species 

(George and Nissen, 1987; Scheldeman, 2002) with high degrees of 

heterozygosity and do not generally produce true-to-type seedlings, 

commercial orchards should be clonally propagated to avoid possible 

influence of genetic variability. However, little has been done to identify and 

characterize the diversity in any of the Annona species. 

9.3 Characteristics of annona ideotypes 

There are several characters that are considered important in a superior 

commercial cultivar of an Annona species (Table 9-1). According to 

Mahdeem (1990), the most important characteristics of an ideotype, 

especially of cherimoya, are the following: a) vigorous and prolific plants, 

compatible with one or more rootstocks, regular-bearing, resistant to cold and 

dry conditions, as well as to pests and diseases; b) architecture of the canopy 

with acceptable form, which does not need pruning, and which is easy to 

harvest; c) abundant flowers with fertile pollen, and which attract insect 

pollinators; d) out-of-season fruit harvest for specific locations; e) fruit with 

symmetrical form, high natural fruit set, hard skin resistant to pests and 

diseases, as well as with long post-harvest life; f) excellent fruit quality with 

regards to flavour, with fine, fibreless and firm pulp texture, and a low 

number of free seeds in the pulp. Each species will have a slightly different 

ideal type, due to their inherent biological differences. 

9.4 Breeding programme 

Wester (1913) was the first scientist to realize the possibilities for genetic 

improvement of annonas and initiated breeding programmes in Florida and in 

56


the Philippines. However, he faced a lot of limiting factors and because he 

evaluated only a small number of progenies, no new cultivars were selected 

at that time. 

9.4.1 Limiting factors and major constraints 

Climate and soil are the factors with greatest influence on the variation in 

growth, fruit set, fruit size and quality of commercial annonas. They 

represent the main constraints in the establishment of an Annona breeding 

programme, since they directly influence response via the genotypeenvironment 

interaction. 

Rainfall and high humidity during the peak flowering season greatly enhance 

fruit production of most annonas by preventing desiccation of stigmas, 

prolonging their receptive period and increasing fruit set and early fruit 

growth (Nakasone and Paull, 1998). The sugar apple is the contrast, as it is 

probably the most drought-tolerant species, and it grows, but produces 

poorly, where rains are frequent. This is shown by the fact that sugar apple 

does much better in northern Malaysia, where dry periods occur, than in the 

southern part, which has year-round high moisture (Nakasone and Paull, 

1998). This climatic adaptation of sugar apple to semi-arid conditions is 

confirmed by Coronel (1994). Sugar apple's deciduous growth habit 

contributes to its drought resistance, as it does not have any leaves during 

most months of the dry season. In contrast, soursop grows and produces very 

well under high rainfall conditions in the Amazon region. However, both 

sugar apple and soursop grow and produce very well in the semi-arid 

conditions of north-eastern Brazil, with very low rainfall, but they both 

require irrigation. Given these good responses to environmental control, 

breeding of sugar apple and soursop have a greater likelihood of success 

under semi-arid conditions. 

Temperature is also a limiting factor, mainly for the tropical annonas, 

soursop, custard apple, sugar apple and wild soursop, since low temperatures 

(< 14°C) may damage or even kill young trees, although adult plants may 

show some tolerance. Poor pollination is frequent in all species when high 

temperatures (> 30°C) and low RH (< 30%) occur, even with handpollination 

(Nakasone and Paull, 1998). These authors also comment that 

cherimoya is more tolerant to low temperature (7-18°C) and soursop is the 

least tolerant (15-25°C). Therefore, improvement of cherimoya would be 

better in the northern hemisphere (temperate and subtropical regions) and 

57


soursop, like the other annonas, in the southern hemisphere under tropical 

conditions. 

Shading of the generally vigorous annona trees can greatly reduce fruit set. 

Therefore, pruning and spacing are cultural practices that need to be adjusted 

for enhancing fruit set in any breeding programme. No photoperiod 

responses have been reported in annonas, so this factor can safely be ignored. 

It is very common to observe trunk and stem breakage of soursop and sugar 

apple trees due to winds. Tree shaking may also be partially responsible for 

collar-rot by allowing penetration of pathogens, and the fruit skin is easily 

damaged by rubbing and exposure to drying winds (Marler et al., 1994). 

Control of fruit drop can be improved by windbreaks and under-tree 

sprinkling to raise RH above 60% (Nakasone and Paull, 1998). 

All Annona species can grow in a wide range of soil types, from sandy soils 

to clay loams. However, they prefer rich, well-drained soils, and breeders 

have the additional advantage of avoiding root-rot diseases in seedling 

populations. 

A small population of pollinator insects may limit fruit set of open pollinated 

annonas. The morphology and fragrance of flowers suggest that natural 

pollination is done by certain species of Coleoptera beetles (Coronel, 1994; 

Pinto and Silva, 1996). Low numbers of pollinator insects coupled with slow 

anthesis (flower opening) impede insect visits. Consequently, there is often 

very low fruit set from natural pollination, varying from 0% in some 

cherimoya orchards (Gardiazabal and Cano, 1999) to 26% in some soursop 

orchards (Pinto and Ramos, 1999). In contrast, the same authors obtained 

26% and 73% fruit set by using hand pollination on cherimoya and soursop, 

respectively. Due to protogyny, hand pollination is useful in breeding 

programmes, since breeders can select the parents to be crossed and may also 

improve a full-sib progeny population more quickly than a half-sib one. 

Apart from cherimoya, germplasm banks that contain Annona species are 

rare throughout the world (see Chapter 8), which is a limiting factor for 

selecting and crossing among elite cultivars. The length of the juvenile period 

is also a limiting factor for genetic improvement. Generally, the juvenile 

period lasts until the third year. Therefore, if one considers three years of 

seedling and fruit evaluations as a minimum requirement, the release of an 

F1 cultivar is only possible six years after the cross. 

Seedling rootstocks of annonas are generally derived from heterogeneous 

open-pollinated plants; hence, it is often difficult to fix specific characters in 

58


a short period. Early maturity, better fruit appearance and long post-harvest 

life for tropical annonas, and in the subtropics, greater cold tolerance, are 

objectives for cherimoya breeding (Nakasone and Paull, 1998). 

9.4.2 Breeding objectives 

The first objective that the breeder has in mind is the determination of the 

specific characteristics that are important for the new cultivar - the ideotype 

(Table 9-1). These characteristics may help a breeder to select parental 

groups to be used in an annona breeding programme, in order to obtain 

desirable progenies. These progenies may not have all desirable 

characteristics, but at least will have those most important to growers, 

retailers and consumers. For instance, a sweet sugar apple cultivar with long 

shelf life fruits (> 5 days), which is important from a consumer’s viewpoint, 

may not be selected, if it presents low yield (< 20 kg/tree/year) and fruit 

weight lower than 400 g (Table 9-1), since no grower will plant this cultivar. 

Indeed, yield is always an important trait in cultivar development for any 

crop species (Fehr, 1987 b) and annona is no exception. However, genetic 

improvement for yield is the most difficult and expensive of all breeding 

objectives, due to the complex nature of its inheritance and the numerous 

environmental factors influencing the trait. The different yield responses of 

the various Annona species and cultivars, such as soursop (Pinto and Silva, 

1996) and cherimoya (Gardiazabal and Cano, 1999), in the same region, 

prove the complexity of the genotype-environment interaction. 

Quality is another important characteristic for the improvement of annonas, 

as important or sometimes more so than yield, since market value is based on 

the fruit's appearance and its organoleptic characteristics. A complicating 

factor is that quality standards may not be the same for all markets. In 

cherimoya, a skin without protuberances above the carpel walls may be 

preferable, since this diminishes the susceptibility of the fruit to mechanical 

damage (Gardiazabal and Cano, 1999). Small, sweet soursop fruits are 

recommended for the fresh market, while large acid ones are more suitable 

for the processing industry (Pinto and Silva, 1996). However, ripe fruits 

become soft and perishable, with subsequent rapid fermentation. Besides the 

difficulty to determine a correct harvest point, handling and processing 

procedures have to be adapted to this postharvest problem. Therefore, it is 

difficult to produce fresh soursop fruits for export to distant markets without 

resolving this postharvest impediment. 

59


Table 9.1. Main characteristics of cherimoya, soursop and sugar apple 

ideotypes (adult plants). 

Characteristic Cherimoya Soursop Sugar Apple 

Plant 

- Vegetative vigour Medium to low Medium to low Medium to low 

- Reproductive vigour 

(yield) 

High ( > 30 kg/tree/year) High ( > 60 

kg/tree/year) 

High ( > 20 

kg/tree/year) 

- Bearing Regular Regular Regular 

- Rootstock/Scion Com- 

High High High 

patibility 

- Resistance 

Climate (Temperature) 

Pest and Disease 

High Temperature (> 22 

°C) 

High 

Abundant ( >150 flowers/tree) 

Low Temperature (< 

18 °C) 

High 

Abundant ( >100 

flowers/tree) 

Low Temperature (< 

18 °C) 

High 

Abundant ( >180 

flowers/tree) 

- Flower Number 

- Pollen Fertility (20 High (> 76%) High (> 76%) High (> 76%) 

°C) 

- Harvest Out-of-Season Throughout year Out-of-Season 

Fruit 

- Size/Weight 

For Industry 

For Fresh Consumption 

Large / > 650 g 

Medium / 300-600 g 

Large / > 2,5 kg 

Small / 0,8-2,5 kg 

Large / > 400 g 

Medium / 300 - 400 g 

- Shape Heart Conical Rounded to Heart 

- Fruit Set (naturally) High (> 27%) High (> 26%) High (> 25%) 

- Skin Smooth (Impressa) Short protuberances Tuberculate 

- Pulp Sweet, fibreless Sub-acid, low fibre Sweet, low fibre 

- Flavour Delicate Sub-acid flavour Pleasant acidulous 

- Seed number/100g of 

pulp 

- Shelf Life (15° to 

30°C) 

- Transport resistance High (pulp pressure > 1.5 

kg/cm2) 

Low or Absent / < 6 seeds Low / 10-30 seeds Low or Absent / < 10 

seeds 

Long ( > 10 days) Long ( > 5 days) Long ( > 5 days) 

High (not found) 

High (not found) 

Sources: Anderson and Richardson (1990); George and Nissen (1987); Guardiazabal 

and Cano (1999); Higuchi et al. (1998); Nakasone and Paull (1998); 

Pinto et al. (2001); Scheldeman and Van Damme (1999); Viteri et al. (1999). 

60


Breeding for pest and disease resistance is another important part of cultivar 

development. The soursop cultivar ‘Morada’ has shown less susceptibility to 

fruit borer than other cultivars under the environmental conditions of Brazil's 

Central Region (Pinto and Silva, 1996). Given the stage of domestication of 

the three principal annonas, it is not surprising that this type of resistance is 

rare, especially when most annona plantations are monoculture orchards. 

Quantitative characteristics, like yield, controlled by numerous major and 

minor genes, are more difficult to manipulate than qualitative ones. In 

addition, traits whose expression is influenced by environment are also more 

difficult to select for than those unaffected by environmental factors. As 

Annona breeding programmes have several objectives, it is the responsibility 

of the breeder to design cost-effective strategies to select genotypes with the 

maximum number of desired traits. 

9.4.3 Methods and strategies 

Most Annona species and cultivars differ in environmental adaptation, 

productivity and fruit quality. Therefore, different conventional methods can 

be used in their breeding. According to Fehr (1987a), there are three 

requirements for the development of an asexually propagated cultivar: a) a 

suitable source of genetic variability; b) evaluation of individuals from the 

population; c) asexual multiplication of a new cultivar for commercial use. 

Introduction of superior genotypes and/or cultivars to establish a germplasm 

collection is, basically, the first requirement of any breeding programme. 

This can be complemented by the introduction into the collection of some 

wild Annona species with useful genes, mainly for resistance to diseases. All 

accessions require comprehensive characterisation and documentation, 

followed by evaluation and selection. 

Several types of populations can be developed by hybridisation, from which 

superior clones are selected. Nonetheless, most of the existing commercial 

cherimoya cultivars in Chile and Spain were released after selection and 

asexual propagation of open-pollinated progenies (Hermoso and Farré, 1997; 

Gardiazabal and Cano, 1999), so hybridisation is not always necessary, 

although it can often accelerate a breeding programme. However, breeding 

(and selection) in cherimoya has been neglected and only a few new cultivars 

have been developed in the past 20 years, due mainly to the lack of breeding 

programmes and clear strategies. In contrast, other subtropical and tropical 

fruit species, such as mango, have been intensively selected and cloned from 

61


62 

more than several hundred thousand seedlings over more than 100 years 

(George et al., 1999), although some progress has recently been made in 

selecting new cherimoya cultivars. 

A very simple but interesting strategy, developed by members of the 

California Cherimoya Association, is to bring samples of new cultivars to 

show at their Annual Meeting. This helps to identify new materials that they 

might want to grow next year (Grossberger, 1999). This strategy has a long 

history in developed countries, such as the USA, and needs to be more 

widely adopted in developing countries. 

George et al. (1999) commented that a major Annona breeding programme, 

funded by the Australian Custard Apple Growers Association, started with 

the objective of developing high quality seedless cultivars of atemoya 

(Annona squamosa x A. cherimoya hybrids). The vernacular name 'custard 

apple' is usually and wrongly used to refer to Annona squamosa by Indian 

researchers (Pawshe et al., 1997) and to Annona hybrids or to Annona 

cherimola by UK and Commonwealth researchers (George and Nissen, 1987, 

Van Damme and Scheldeman, 1999). Therefore, the reader is advised to 

carefully identify the origin of articles in order to correctly identify the 

species which is referred to. 

In this breeding programme, the Australian breeders first produced 

tetraploids, either through gamma irradiation or colchicine, and then crossed 

these to diploids to produce seedless triploids. There are several other 

potentially useful methods of obtaining seedlessness in annonas, e.g., a) 

trying to identify progeny from diploid x diploid crosses with small seed 

sizes and numbers; b) producing triploids from diploid x tetraploid and 

tetraploid x diploid crossing, as done in Australia; c) producing triploids 

through protoplast fusion of diploids and haploids; d) irradiation of budwood 

to "knock out" genes for seed production; e) crossing of low-seeded parents 

(as the seed number is an heritable character); f) development of selfincompatible 

parthenocarpic hybrids; and g) using endosperm culture to get 

triploids. Not all of these strategies are currently being employed in annona 

breeding programmes (George et al., 1999). 

Progeny of a cherimoya x sugar apple cross produced fruits which were late 

maturing (spring in Queensland), and appeared to have inherited the 

flowering and fruiting characteristics of atemoya, with flowering in autumn 

and fruit maturity in late spring, under the subtropical conditions in that part 

of Australia. A similar study was carried out by Zill Nursery near Boyton 

Beach, Florida, and approximately 3,000 seedlings, mainly from interspecific


crosses, were planted. Since attractiveness is a key factor in selling fruits, the 

exciting possibility exists to develop new cultivars with external and internal 

pink-red colours. Crossing of newly introduced red and pink-skinned 

atemoya x cherimoya selections to red-skinned sugar apple types selected in 

Queensland, Australia, is currently in progress (George et al., 1999). 

Spain is more focused on germplasm collection and ex situ evaluation of 

numerous accessions (Farré Massip and Hermoso González, 1987, cited by 

Scheldeman, 2002). Pérez de Oteyza and Farré (1999) reported that the 

selection of a superior cultivar of cherimoya at the Experimental Station of 

La Mayora, Spain, is based on the following characters: a) regarding 

agronomic and commercial parameters - size, format and pilosity on leaves, 

length and colour of flowers, floral density (number of flowers per one metre 

of mature stem), susceptibility to fruit fly, season of maturation and harvest, 

defects on the skin and in the pulp, resistance to fungal attack and seed 

colour; and b) regarding fruit transport and consumption - type of skin, pulp 

firmness, taste (sugar content and acidity) and seed index (number of seeds 

per 100 g of fruit). 

In Chile, introduction and selection of cherimoya showed that the new 

cultivars from Spain were superior, with longer harvest periods and better 

fruit quality than Chilean cultivars (Gardiazabal and Cano, 1999). Ten 

cultivars from Spain were evaluated for fruit weight, shape and colour, skin 

type, number of days post-harvest to reach appropriate ripeness for eating, 

resistance of pulp to pressure, percentage by weight of fruit components, 

seed type, number of seeds per 100 g of pulp, sugar and acidity, and taste. 

The Spanish cultivar ‘Cholan’ showed the highest general rating. 

In Mexico, genetic improvement of cherimoya started in 1991, with 

evaluation of seedlings of local cultivars and evaluation of introduced 

cultivars from Spain, Chile and New Zealand (Román and Damián, 1999). 

Characterisation and selection of cherimoya fruits from trees collected in 

three regions of Michoacán State, Mexico, was done by Agustín (1999). 

While studying such characteristics as fruit weight, percentage of pulp and 

seed, fruit soundness, type of skin and earliness, he found great genetic 

variability among native fruits and proposed this as the basis for developing 

germplasm for commercial plantations. A very similar strategy has been used 

in Italy and Portugal (Madeira), with selection and evaluation of promising 

local types and introduced cultivars (Monastra, 1997; Nunes, 1997). 

The methods and strategies used in Madeira to develop superior cultivars of 

cherimoya were somewhat different. First, the agricultural service surveyed 

63


and mass selected local types. Plant behaviour was observed with respect to 

age, origin, farming practices, pest and disease resistance, organoleptic 

parameters and so on. At the same time, genetic material was introduced 

from other countries, especially the USA and Spain, increasing variability 

and allowing comparison of adaptability, productivity and quality with the 

local materials. Finally, clonal selection produced four improved cultivars: 

Madeira, Mateus I, Perry Vidal, and Funchal (Nunes, 1997). 

Soursop is still largely propagated by seed, and the progenies can be selected 

and separated into groups based on acidity and sweetness. Morada, an 

ecotype introduced into Brazil from Colombia, produces large fruits with an 

average weight of 3.8 kg, but its fruits have very acid pulp, which is a 

constraint for the fresh fruit market. A local soursop selection from Bahia 

State named "ecotype A" produces very small fruits (< 1.8 kg), but has sweet 

pulp, appropriate for the fresh market. Crossing between these two might 

produce a medium weight fruit with a commercially acceptable taste. Several 

genotypes and some related species are used in the soursop breeding work of 

Embrapa Cerrados, Brazil, from which a clone based on Morada has been 

released as a reliable option for tolerance to trunk borer (Pinto and Ramos, 

1997). At the same time, A. glabra, which has dwarf characteristics and good 

adaptation to damp areas, is being used in a breeding programme for 

rootstock improvement, since there is acceptable rootstock/scion 

compatibility between these two species. 

Wild soursop has been used for insect control in Africa, since it has 

significant amounts of secondary metabolites with insecticidal activity (see 

Chapter 7). However, there is no specific breeding programme to improve 

this characteristic, although Abubakar and Abdurahman (1998) mentioned a 

project to prospect, collect and chemically identify wild soursop variation 

with insect growth-regulating activity in Kaduma State, Nigeria (work at the 

Department of Biological Sciences, Ahmadu Bello University, Zaria). 

9.4.4 Selection and cultivar development 

Like any other fruit tree, the type of selection or cultivar development of 

annonas is strongly influenced by commercial use. Also, the feasibility of 

using a particular type of selection or cultivar may depend on the nature of 

the organisations that produce and distribute grafted plants for commercial 

use, these being much better established in more developed countries. This 

explains why annona cultivar development concentrates on cherimoya and 

soursop (and atemoya), and has been carried out only by public institutions. 

64


In short, the cost of annona cultivar development is too high compared with 

the demand for superior grafted material, therefore, only a few private 

companies or nurseries can afford a breeding program, e.g., Zill Nursery in 

Boyton Beach, Florida. 

Nakasone and Paull (1998) stated that only cherimoya and atemoya have 

important named clonal cultivars. For other annonas, such as soursop, sugar 

apple and custard apple, there are some selections and very few named 

cultivars (Table 9.2). 

In California, some old cultivars of cherimoya, such as ‘McPherson’, 

‘Deliciosa’ and ‘Bays’, were selected and cloned from seedling plantings 

(Nakasone and Paull, 1998). Grossberger (1999) stated that cv. ‘White’ is the 

most important cultivar grown in California because of its large size. 

However, ‘Booth’, as a very sweet cultivar, and ‘Pierce’, as a cultivar with 

the best shape for packing, have been cultivated for a long time. 

In Peru, considerable work has been done on the development of annona 

cultivars, but they are not widely known outside Peru. Chile, Spain and New 

Zealand grow cherimoya, as it is more tolerant of cold temperatures and has 

more successful self-pollination than atemoya. ‘Reretai’ and ‘Burton’s 

Wonder’ are the most important New Zealand cultivars while ‘Madeira’, 

‘Mateus I’, ‘Perry Vidal’ and ‘Funchal’ are important cherimoya cultivars 

grown on Madeira island (Nunes, 1997). Numerous cherimoya cultivars have 

been reported in Spain and Fino de Jete identified as the finest commercial 

cherimoya cultivar, mainly because of its good postharvest life (Hermoso and 

Farré, 1997). In Ecuador, there are no single-cultivar orchards of cherimoya; 

rather most orchards consist of plants propagated by seed. In some cases 

there are plants grafted onto the local ecotypes known as Jaramillo and 

Chumina, selected by farmers on the basis of yield and quality (Fuentes, 

1999). 

There is no well-established breeding programme for custard apple. 

Nonetheless, a few cultivars have been mentioned, such as ‘Camino Real’, in 

Guatemala, and ‘Fairchild Purple’ and ‘Young’, in Florida, USA (George et 

al., 1999). 

Soursop is largely planted by seed in most countries, including Mexico and 

Brazil. Soursop clones are separated into groups, such as acid (for the 

processing market) and low acid (for the fresh market), or juicy and nonjuicy 

types. Seedling populations of soursop have been established in 

Mexico, Malaysia and Brazil, permitting the selection of superior clones with 

better yield and improved processing qualities (Pinto and Silva, 1996; 

65


Nakasone and Paull, 1998; Lemos, 2000 b). Morada is a soursop type 

belonging to the acid group and producing large fruits (average of 3.8 kg), 

which was introduced into central Brazil in 1980, and has shown high yield 

and good tolerance to trunk and fruit borers. ‘Giant of Alagoas’ developed by 

Lemos (2000 b) is a clonal cultivar selected from seedling of Morada 

soursop, which shows the same performance as the mother plant. Several 

soursop selections in the low acid group have been evaluated in Pernambuco 

State, Brazil, and some of them have given high yields - up to 70 

kg/plant/year (Lederman and Bezerra, 1997). 

Table 9-2. Some selections and cultivars of cherimoya, atemoya, soursop, 

sugar apple and custard apple that are currently planted in various 

countries 

Cultivar/Selection Country of Origin Cultivar/Selection Country of Origin 

Cherimoya 

Atemoya 

Alvaro (1) Mexico African Pride (1) S. Africa/Israel 

Andrews (1) Australia Bradley (1) USA/California 

Bays (1) USA/California Jennifer (1) Israel 

Booth (1) USA/California Kabri (1) Israel 

Burton's Wonder (1) New Zealand Malalai (1) Israel 

Bronceada (1) Chile Nielsen (1) Australia 

Campas (1) Spain Island Gem (1) Australia 

Chaffey (8) USA/California Page (1) USA/Florida 

Cholan (1) Spain Pink's Mammoth (1) Australia 

Concha Lisa (1) Chile Soursop 

Cortes II-31 (1) Mexico Morada (3) Colombia/Brazil 

E-8 (1) Ecuador Lisa (3) Colombia/Brazil 

Fino de Jete (1) Spain Blanca (3) Colombia/Brazil 

Funchal (2) Portugal/Madeira Giant of Alagoas (4) Brazil/Alagoas 

Gangemi (1) Italy Ibimirim Selection (5) Brazil/Pernambuco 

Cherimoya 

Golden Russet (8) USA/California Sugar Apple 

Kempsey (1) Australia IPA Selections (5) Brazil/Pernambuco 

Leone (1) Italy Molate (6) Philippines 

Libby (7) USA/California Cuban Seedless (1) Cuba 

Lisa (1) USA/California Lobo (7) Philippines 

Madeira (2) Portugal/Madeira Noi (7) Thailand 

Mateus I (2) Portugal/Madeira Access 6333 (7) Philippines 

66


Cultivar/Selection Country of Origin Cultivar/Selection Country of Origin 

McPherson (8) USA/California Red Sugar Apple (7) USA/Florida 

Mossman (1) Australia Mammouth India 

Negrito (1) Spain Balangar India 

Ott (1) USA/California Sitaphal (red/lal) India 

Perry Vidal (2) Portugal/Madeira Borhodes India 

Pierce (1) USA/California Britishbaroa India 

Reretai (1) New Zealand Custard Apple 

Whaley (8) Australia Camino Real (7) Guatemala 

White (8) USA/California Fairchild Purple (7) USA/Florida 

Dr.León (7) 

Young (7) 

USA/Florida 

West Java 

Sources: (1) Nakasone and Paull (1998); (2) Nunes (1997); (3) Pinto and Silva 

(1996); (4) Lemos (2000 b). 

India, China and Taiwan have produced a few named cultivars of sugar apple 

that are propagated vegetatively. In Cuba, researchers developed ‘Cuban 

Seedless’, which is a seedless cultivar with medium-sized fruits, and another 

cultivar with low fibre content that is very important for the commercial 

market (Nakasone and Paull, 1998). In the Philippines, there are 3 forms of 

sugar apple fruits: (a) a green-fruited seedy form, which is grown in most 

parts of the country; (b) a purple-fruited seedy form, reportedly introduced 

from India; (c) and the green-fruited seedless form, which is a recent 

introduction and whose origin is unknown. The selection of superior strains 

is aimed in the direction of a green-fruited seedy form (Coronel, 1994). In 

Petrolina, Pernambuco state, Brazil, some sugar apple growers are producing 

and commercialising a purple sugar apple type (Plate 1), and north-eastern 

Brazilian consumers – mainly those with higher per capita income – are 

buying it much more as an exotic fruit, due to its colour, than because of any 

other characteristic, since taste and shape are similar to standard green sugar 

apples. Unfortunately, this sugar apple type has not been totally accepted in 

the market, because most of the consumers think that the purple fruit is 

already rotten. 

67


9.5 Role of modern biotechnology 

9.5.1 Tissue culture 

Tissue culture has many uses: a) micropropagation; b) maintenance of 

germplasm collections; c) embryo rescue; d) development of haploid plants; 

e) enhancement of variability by somaclonal variation; and f) to prepare 

explants for transformation methods (Encina et al., 1999; Herrera, 1999). 

However, for most annona work the main objective is micropropagation, as 

the conventional methods of propagation are slow and costly, and in some 

cases, such as with cherimoya, are also inefficient. This appears to be 

because the morphological potential for rooting of cherimoya is very low 

(Encina et al., 1999). 

Researchers have successfully micropropagated cherimoya, sugar apple, 

soursop and atemoya. Atemoya clonal propagation was described by Rasai et 

al. (1994). They obtained multiple shoot formation from hypocotyls and 

nodal cuttings of the cultivar ‘African Pride’. The explants were cultivated in 

MS medium supplemented with BAP, kinetin, biotin and calcium 

pantothenate. In spite of improved rooting by shoot pre-treatment in liquid 

MS medium containing IBA, the percentage of rooting was still low (40%) 

and remains a limiting factor for commercial micropropagation of atemoya. 

Benjoy and Hariharam (1992) described plantlet differentiation in soursop. 

They found a mean of 4.8 shoots per hypocotyl explant growing in an MS 

medium containing BAP and NAA. However, they reported only relative 

success in rooting and survival (35%). To improve the system, Lemos and 

Baker (1998) suggested the use of sorbitol to induce de novo shoot 

development and Lemos and Blake (1996 c) tried galactose and NAA to 

stimulate rooting. Nonetheless, no commercial protocol is ready for use. 

The first haploid plants induced by anther culture in fruit trees were reported 

by Nair et al. (1983) with sugar apple. The availability of haploids is very 

important for fruit-breeding, because of the long generation intervals, the 

highly heterozygous nature of most fruit species and the presence of 

parthenocarpy and self-incompatibility. These researchers obtained callus 

differentiation, and formation of triploid roots and shoots from sugar apple 

endosperm (Nair et al., 1986). Their aim was development of seedless fruits, 

but a complete plantlet was not obtained. 

68


Shoot proliferation of sugar apple was achieved with hypocotyls and nodal 

cuttings growing in Woody Plant Medium, supplemented with BAP and 

silver thiosulphate to control leaf abscission (Lemos and Blake, 1994; 1996 

a, b). Rooting was obtained when shoots were preconditioned in medium 

with activated charcoal, and then treated with NAA or IBA (Lemos and 

Blake, 1996 b). To improve rooting, they used galactose instead of sucrose in 

the rooting medium. Eighty percent of the plantlets were successfully 

acclimatized in the greenhouse (Lemos and Blake, 1996 b). This 

methodology now needs to be transformed into a commercial protocol. 

However, the greatest success in annona micropropagation was obtained in 

cherimoya. Encina et al. (1994) described in vitro morphogenesis of juvenile 

cherimoya and achieved a micropropagation system for adult cherimoya 

materials, obtaining 50% rooting and exceptionally good acclimatisation 

(Encina et al., 1999). To increase the success of acclimatisation, Azcòn- 

Aguilar et al. (1994) inoculated cherimoya with arbuscular mycorrhizal fungi 

to improve growth, survival and development of cherimoya produced in 

vitro. Currently they are working on several methodologies (Encina et al., 

1999): a) somatic embryogenesis; b) adventitious organogenesis and cellular 

cultures; c) ploidy manipulation; d) autotrophy induction; and e) genetic 

transformation. However, the authors did not mention that these 

methodologies have already been applied commercially. 

Although there are numerous experimental protocols for Annona tissue 

multiplication, the final price of the plantlets is still too high for commercial 

use. Given the potential of this technology, further research is needed to 

transform experimental protocols into commercial protocols. 

9.5.2 Genetic transformation 

Encina et al. (1999) have started studying genetic transformation of 

cherimoya, mainly to optimize the protocol for Agrobacterium 

transformation. The objectives are to control ripening, to change post-harvest 

characteristics, and to provide pest and disease resistances. For other Annona 

species there are no advanced studies. 

9.5.3 Molecular markers 

Samuel et al. (1991) suggested the use of allozymes to study diversity and 

systematics in Annonaceae. They considered these systems to be efficient for 

investigations of the origin of polyploids for breeding programmes. A 

69


preliminary study, using eleven isoenzyme loci, was developed with five 

Annona species. Strangely, soursop, mountain soursop (A. montana), and 

pond apple (A. glabra) presented no variation between or within the 

populations studied. 

Some isoenzymes studies with cherimoya have been carried out. Groups at 

the University of California (USA) and University of Granada (Spain) 

studied the variation in isoenzyme patterns of cherimoya cultivars from the 

USA and Spain. Both groups found sufficient variation to distinguish 

cultivars and to evaluate cherimoya germplasm (Ellstrand and Lee, 1987; 

Pascual et al., 1993). This isoenzyme analysis is important, since cultivars 

have been confused and are widely known by the wrong names. For instance, 

there is often confusion with the cherimoya cultivar ‘McPherson’, which is 

incorrectly identified in Spain (Grossberger, 1999). 

In Mexico, Medina et al. (1999) used molecular biological techniques to 

select soursop varieties according to their resistance to fungal diseases. They 

analysed the electrophoretic pattern of peroxidase isozymes and observed the 

variation between isoforms of healthy and infected plants. Since these 

diseases are detrimental to soursop production, the researchers consider these 

isoenzyme patterns as markers to select healthy and infected individuals, and 

potentially to identify resistant and susceptible genotypes. 

Isoenzyme studies are limited because they are carried out using a relatively 

small number of loci. RAPDs offer an enormous number of markers covering 

the whole genome, and is a more powerful technique for genotype 

identification and germplasm evaluation. Ronning et al. (1995) estimated 

variation between cherimoya, sugar apple and atemoya, and determined the 

inheritance of these markers in the F1. All fifteen primers used generated 

repeatable polymorphic patterns, resulting in a very efficient method to 

distinguish genotypes of Annona species. 

In short, the recent biotechnological studies, both cellular and molecular, 

have shown great potential to further annona development, not only to solve 

problems of mass propagation of superior cultivars, through 

micropropagation techniques, but also to identify or fingerprint annona 

cultivars, as well as to determine cultivar parentage, through RAPD markers. 

70

Chapter 10. Agronomy 

10.1 Propagation 


Annona propagation commonly includes sexual or seed propagation, and 

asexual or vegetative propagation. Since both are still important, each will be 

examined carefully, since the quality of planting materials depends upon a 

well-prepared plant. 

10.1.1 Seed propagation 

If seeds are to be used, they should be obtained from selected mother plants, 

whose characteristics should include high fruit yield, excellent fruit quality 

and high resistance to pests and diseases (Torres and Sanchéz, 1992; 

Coronel, 1994; Agustín and Alviter, 1996). Use of seeds bought at market is 

not advisable because these characteristics can not be observed at first hand. 

Annona seeds generally present uneven and irregular germination, which 

occurs over a long time, making sexual propagation difficult. However, since 

seeds lose viability in the field, they should be sown as soon as possible after 

removal from ripe fruits (Coronel, 1994; Nakasone and Paull, 1998). Seed 

storage tolerance and later germination success vary among Annona species 

(Table 10-1). Seeds dried and held at low temperatures provide more leeway 

in time of planting (Torres and Sanchéz, 1992). 

Table 10-1. Time of storage (days) to assure 90% seed viability, time 

(days) for germination, germination percentage, seedling age for 

transplanting and age if used for grafting 


Time of 

Storage 

(days) 

Time of 

Germination 

(days) 

Germination 

(%) 

Age for 

Transplant 

(days) 


Grafting 

(days) 

Cherimoya 50-60 35-45 90-95 70-100 240 

(A. cherimola) 

Custard apple 40-50 30-35 90-95 50-60 180 

(A. reticulata) 

Soursop 30-40 30-40 90-95 60-90 210 

(A. muricata) 

Sugar apple 40-50 35-50 85-95 90-120 220 

71



Time of 

Storage 

(days) 

Time of 

Germination 

(days) 

Germination 

(%) 


Transplant 

(days) 


Grafting 

(days) 

(A. squamosa) 

Source: Hernandez (1983). 

Irregular germination is due to different levels and types of dormancy (Pinto, 

1975 a, b; Ferreira et al., 1997; de Smet et al., 1999; Ferreira et al., 1999; 

Hernández et al., 1999; Moreno Andrade, 1999). Nevertheless, there still 

exists disagreement about the presence of dormancy in Annona seeds and the 

correct treatments to overcome it. Sanewski (1991) claimed that no 

dormancy exists, whereas other authors claim the presence of dormancy 

(Hayat, 1963; Purohit, 1995; Ferreira et al., 1999; Hernández et al., 1999). A 

possible hypothesis to explain this disagreement among authors is the degree 

of seed maturity when tested and the fact that dormancy can be induced by 

environmental factors rather than being innate. Many of the comparisons 

below do not give data on how the seed was handled and what conditions it 

was kept in before testing; hence measures of viability and/or germination 

results may not be truly comparable. 

Pre-treatment of Annona seeds is very important and can be physical, such as 

seed scarification and water immersion to reduce or eliminate the 

impermeability of the seed coat, or chemical, such as gibberellic acid (GA) to 

counteract endogenous germination inhibitors (Campbell and Popenoe, 1968; 

Hartmann et al., 1990; de Smet et al., 1999). Duarte et al. (1974) found that 

dry cherimoya seeds treated with GA at 10,000 ppm significantly increased 

the seed germination to around 70%, compared with 57% for untreated seed, 

while hot water showed an adverse effect on seed germination (28%). 

Pittman (1956) suggested that cherimoya seeds must be soaked in water for 3 

to 4 days and then sown in warm soil, after which they will start to germinate 

in 4 to 5 weeks. 

Castillo Alcopar et al. (1997) found that germination capacity of cherimoya 

seed varied from 66 to 94% and period of germination varied between 58 and 

69 days after planting. They found that seed scarification had only a slight 

influence on germination. Significantly, all physiological responses were 

genotype dependent. De Smet et al. (1999) evaluated germination percentage 

and rate in cherimoya using different pre-treatments, such as soaking in 

different concentrations of GA (extreme values of 500 and 10,000 ppm), 

soaking for different periods (12-72 hours) in distilled water, and chemical 

scarification with sulphuric acid. They found that GA showed a positive 

effect on both physiological parameters. 

72


GA is costly and not affordable for most resource-poor cherimoya growers, 

especially in developing countries. Soaking in distilled water has shown a 

significant effect on germination, although germination was less concentrated 

and more irregular over time; nonetheless, it is a cheaper, though less 

effective, alternative to GA treatment. Another alternative, which also 

resulted in rapid and high germination of cherimoya seeds, is soaking for 48- 

72 h in distilled water, or for a shorter time in hot water (92°C) and gradually 

cooler water (de Smet et al., 1999). Wild soursop (A. senegalensis) produces 

seedlings rapidly when the seeds are scarified (FAO, 1988). 

In general, seedlings show variability in plant growth and fruit yield because 

they are sexually propagated materials. However, soursop seedlings 

cultivated in the Cerrado region of Brazil, which generally start flowering 

and producing at the same age as grafted soursop trees, have similar fruit 

yield and greater longevity (Pinto et al., 2001). In addition, some seed 

propagated orchards are used as seed sources for the establishment of 

rootstocks. 

Seeding substrates, especially those with composts, may contain root rot 

fungi, weed seeds and nematodes, to which most of the annonas species are 

susceptible, and seedlings may not grow well in their presence. The pretreated 

medium should be composed of fine river sand or a mixture of two 

parts fine sand and one part garden soil (Coronel, 1994) or compost. 

Therefore, pre-treatment of seeding compost is strongly recommended 

(Torres and Sánchez, 1992; Junqueira et al., 1996; Kavati and Piza Jr., 1997). 

Recommendations have included methyl chloride, chloropicrin and other 

gaseous chemicals (Hartmann et al., 1990); more recently these chemical 

treatments have been substituted, especially under tropical conditions, by a 

less expensive and safer treatment called the 'solarization system'. 

Solarization involves covering compost (thickness of the compost layer 

should not exceed 30 cm) with a transparent plastic sheet, which allows solar 

rays to penetrate the medium, thereby increasing the temperature high 

enough (> 50°C) to kill most problematic soil microorganisms (Plate 2). 

The treated substrate is transferred to the germination system used in the 

nursery. The seeds can be sown directly in perforated black plastic bags (22 

cm diameter, 25 cm length and 0.2 mm thick; Pinto and Silva, 1996) or can 

be sown in seed boxes, seedbeds or other shallow containers, for later 

transplanting to plastic bags when the seedlings are 8-15 cm tall (Torres and 

Sánchez, 1992; Coronel, 1994; Agustín and Alviter, 1996; Fuentes, 1999). 

73


Seedbeds 1.2 m wide by 4 m long should be prepared, above soil level in 

order to have good drainage (Fuentes, 1999). 

To avoid fungal infection, the seed can be disinfected with a fungicide, such 

as Zineb, if this is available or can be purchased, applied before sowing. 

Seeds should be spaced at 1-3 cm apart and 10 cm between rows. They 

should be sown at about 1-2 cm deep, then covered with a fine layer of soil, 

which should be compacted gently and watered to saturation (Popenoe, 1974; 

Coronel, 1994; Fuentes, 1999). 

Annona species present epigeous germination. The seed coat must be left to 

drop naturally without any interference from the nurseryman, to ensure that 

the plumule and young leaves to emerge without any damage (Fig. 10-1.). 

Figure 10-1. Type and sequence of seed germination of annona species 

After germination, seedlings grow slowly until they are two to three months 

of age. At this stage growth accelerates in some annona species, such as 

soursop, especially during the warmer season. Sugar apple and cherimoya 

seedlings grow more slowly than soursop seedlings. The best time for 

grafting of soursop is around 12-13 months after germination. While for 

budding of sugar apple seedlings, the best time is 15-18 months after 

germination. Cherimoya seedlings reach the appropriate size for budding or 

grafting at around 15 months after germination, when the seedling rootstock 

74


is about 1 m tall (Scheldeman, 2002). However, the trunk diameter of the 

seedling is a more appropriate growth measurement to determine the best 

time for vegetative propagation, which will be explained below. 

10.1.2 Vegetative propagation 

Due to the generally high variability among seedlings, vegetative propagation 

of both scions and rootstocks is desirable. To establish an orchard, the grower 

must evaluate different methods of propagating annonas vegetatively, such as 

by cuttings, layering, inarching, grafting, budding and micro-propagation, as 

different species and varieties react differently in each Annona growing 

region. 

Seedlings for rootstocks are most effectively kept, in large numbers, in a 

protected and well constructed nursery with irrigation (Fig. 10-2 and Fig. 10- 

3), where control of pests, diseases and weeds, as well as seedling 

fertilization, can be planned and carefully executed. Foliar fertilization 

should start on 90-day old seedlings and be repeated monthly, before and 

after grafting, until planting out in the field. A formula that has given good 

results is composed of 5 g of urea and 15 g of triple superphosphate per litre 

of water (Torres and Sánchez, 1992). To avoid herbicide phytotoxicity, most 

annona growers in Brazil control weeds in the nursery by hand once a week, 

while pests and diseases are monitored daily so as to use a minimum of 

pesticides for control. 

75


Figure 10-2. A sketch (plan view) of an irrigated nursery for production 

of grafted soursop and sugar apple trees showing the cement block 

supports, wires, micro-sprinklers and plastic bags, their distances and 

characteristics 

76


Figure 10-3. A sketch (side view) of an irrigated nursery for production 

of grafted soursop and sugar apple trees showing the cement block 

supports, wires, micro-sprinklers and organization of plastic bags, their 

distances and characteristics 

The success of each method varies according to the species grown (Table 10- 

2) and, in addition, rootstock and scion compatibility has a major influence 

on the success of vegetative propagation methods among annona species 

(Table 10-3). Although some species of annonas, such as soursop, can 

successfully be propagated by cuttings, most are difficult to strike, especially 

cherimoya. Cuttings of custard apple and sugar apple succeed only with 

specific cultivars. The experience of the grafter also has a significant 

influence on the success of the propagation procedure. It should be noted that 

wild soursop has not been tested in grafting experiments. 

Table 10-2. Seed and vegetative propagation methods, commercial 

recommendations and success for different Annona species * 

Method 

Annona Species 

Cherimoya Custard Soursop Sugar Apple 

Apple 

Genetically Highly Variable Uniform Low variability 

(seedling) 

variable 

Commercial Use Not 

As 

High; also as Regular to good 

(seedling) 

recommended rootstock rootstock 

Stem and tip 

cuttings 

< 25% Unknown Successful Some cultivars 

only 

Root cuttings Not 

Unknown Successful < 5% 

successful 

Layering Unknown Unknown Unknown High if modified 

technique is used 

Air layering < 5% Unknown Unknown < 8.3% 

77


Method 

Cherimoya 

Annona Species 

Custard Soursop 

Apple 

Sugar Apple 

Budding > 70% > 40% > 40% > 80% 

Grafting > 70% > 70% > 80% > 70% 

Inarching Successful Unknown Successful Unknown 

Topworking Unknown Unknown Successful Unknown 

Micropropagation Successful Unknown Successful Unknown 

Sources: Bourke (1976); Coronel (1994); George and Nissen (1987); 

Nakasone and Paull (1998); Torres and Sánchez (1992). 

* Information not applicable to wild soursop. 

Table 10-3. Rootstock x scion compatibility and the recommended 

vegetative propagation methods for nine annona species 

Rootstock 

Scion Species 

Species A. cherimola A. muricata A. reticulata A. squamosa 

A. cherimola C; budding, NC; none Unknown C; grafting 

grafting 

A. glabra Unknown LC; budding C; budding C; budding, 

grafting 

Atemoya C; budding NC; none NC; none C; budding, 

grafting 

A. montana Unknown C; grafting Unknown Unknown 

A. muricata NC; none C; budding, C; budding LC; none 

grafting 

A. reticulata C; budding C; budding, C; budding C; grafting 

grafting 

A. senegalensis Unknown Unknown Unknown Unknown 

A. squamosa LC; none LC; budding, 

grafting 

C; budding C; budding, 

grafting 

Rollinia spp. C; unknown C; grafting, 

budding 

Unknown NC; none 

C = Compatible; LC = Low compatibility; NC = Not Compatible. 

Sources: Duarte et al. (1974); Popenoe (1974 a, b); Pinto (1975); Hernandez 

(1983); Iglesias and Sanchez (1985); Ferreira et al. (1987); George and 

Nissen (1987); Ledo and Fortes (1991); Singh (1992); Torres and Sánchez 

(1992); Coronel (1994); Pinto and Silva (1996); Bezerra and Lederman 

(1997); Nakasone and Paull (1998). 

Attempts to propagate cherimoya by root cuttings treated with various 

combinations of benzylamino purine (BAP) and indole butyric acid (IBA) 

had no success after 4 months, even though some cuttings produced a few 

roots (George and Nissen, 1987). Hardwood and leafy terminal cuttings of 

cherimoya taken at monthly intervals and placed in an outdoor rooting bed or 

under mist conditions, respectively, were treated with 0, 1250, 2500 and 

5000 ppm of napthalene acetic acid (NAA). None of the cuttings taken from 

78


adult trees rooted, but some leafy terminals taken from one year old plants 

did, with as much as 25% and 20% rooting with 5000 ppm NAA treatment in 

December and January, respectively (Duarte et al., 1974). George and Nissen 

(1987) commented that etiolation of propagation materials has given good 

results with a high percentage of take in a preliminary study of cherimoya 

propagation by cuttings. They said that the extra costs involved in producing 

plants by this method may be compensated for by the higher yields and 

disease resistance of clonal trees. However, no commercially viable method 

for vegetative propagation by cuttings exists to date in cherimoya. 

Tip cutting propagation has been described for atemoya (Annona cherimola x 

A. squamosa hybrid), with greater success when leaves are attached, versus 

without leaves (George and Nissen, 1987; Hartmann et al., 1990). Atemoya 

tip cuttings were propagated in mist beds containing a sterilized 50:50 

sand:perlite mix and bottom heat with temperatures between 25 and 28°C 

(George and Nissen, 1980). An important environmental factor influencing 

tip-cutting success is humidity, since desiccation of cuttings prior to 

placement in mist beds is a common cause of failure. 

In Florida, mature and healthy stem cuttings of sugar apple taken during the 

dormant period were propagated successfully by Noonan (1953). He used 

shoots between 0.5 and 1.0 cm in diameter and cut into 13-15 cm lengths, 

then set them in sand to a depth of 4/5 of their length with one bud exposed 

above the surface. The cuttings produced roots 25-30 days after planting. 

Bourke (1976) evaluated propagation of sugar apple by root cuttings and 

obtained success percentages of less than 5%. 

Layering is a type of vegetative propagation by which adventitious roots are 

produced on a stem while it is still attached to the parent plant (Hartmann et 

al., 1990). A modified layering technique was used by George and Nissen 

(1986) with 100% take. They used one year-old cherimoya seedlings and cut 

them back severely in mid-summer to produce 3-5 juvenile shoots. When the 

new shoots were approximately 15 cm long, metal growth constrictor rings 

were placed over each shoot, then a polyethylene sleeve was placed over 

each shoot and filled with a mixture of 50% sand and 50% sawdust, leaving 

only the growing points exposed. Excellent root systems were produced 4-5 

months later. Marcottage (air-layering) of limbs 1.0-1.5 cm diameter on 

mature trees of cherimoya cultivar ‘Deliciosa’, however, was unsuccessful, 

with less than 5% of marcots rooting (George and Nissen, 1987). 

Orchards established with cuttings are more uniform and less expensive to 

establish than grafted trees, when cutting materials are taken from pruned 

79


branches. However, cutting and air-layering methods do not produce taproots 

and plants are more susceptible to falling over in strong winds, with 

subsequent damage and orchard loss. Hence, these vegetative propagation 

methods are not recommended where strong winds are a problem. 

Cherimoya, soursop and sugar apple can all be successfully inarched with a 

high percentage of success (Morton, 1967; Viñas, 1972; Castillo Alcopar, 

1997). The inarching method is used to join rootstocks to selected scion 

cultivars which are otherwise difficult to root or to graft as detached scions, 

as well as to invigorate weak-growing trees by augmenting their root systems 

(George and Nissen, 1987). However, this method is much more difficult to 

work with and it shows a higher cost/benefit ratio compared with the grafting 

and budding methods. 

The annonas are easily propagated by budding and grafting, although the 

success varies among species. In general, grafted plants grow rapidly but do 

not bear fruit any earlier than seedling stocks. Several authors (Moran et al., 

1972; Duarte et al., 1974; George and Nissen, 1987; Torres and Sánchez, 

1992; Pinto et al. (2001) and Nakasone and Paull, 1998) have described 

splice grafting (Fig. 10-4), also called cleft grafting, as well as shieldbudding 

and inverted T budding (Fig. 10-5), as the most successful methods 

for vegetative propagation of annonas. Generally, annonas have shown 

higher percentage take when propagated by grafting in comparison to other 

methods, except for sugar apple, which shows a greater success with budding 

(Table 10.2). 

80


Figure 10-4. Splice grafting: A) annona rootstock; B) scion from selected 

cultivar and cut rootstock; C) wrapping of scion and rootstock 

producing a humid chamber with a transparent plastic bag; D) new 

grafted plant 

In Ecuador, grafting of cherimoya takes place 15 months after transplanting, 

when the seedling rootstocks are 30-40 cm tall and the stem diameter is 0.5 

cm, and has up to 90% success (Fuentes, 1999). Four types of grafting 

(splice, cleft, whip-and-tongue, and crown) and shield-budding were 

evaluated by Moran et al. (1972) with scions of two diameters (0.8-1.0 cm 

and 1.2-1.5 cm) on cherimoya seedling rootstocks of the same cultivar. They 

found no differences in percentage take between the two sizes of rootstock, 

but subsequent growth was better on the thicker rootstock. Both the budding 

and the four grafting methods led to a success rate of at least 70%. The crown 

grafts took more quickly, but shield-budding led to superior growth 

compared to the other methods. However, splice and whip-and-tongue 

81


graftings on thick rootstocks were considered to be the best as regards 

percentage take, subsequent growth and cost (low). 

Figure 10-5. Steps of the inverted T - budding technique: A and B) 

rootstock cut and prepared for budding; C and D) collecting bud from 

selected cultivar; E and F) insertion of the bud into the rootstock stem; 

G) wrapping of the bud. Adapted from Hartmann et al. (1990) 

In Colombia, Iglesias (1984, cited by Torres and Sánchez, 1992), used three 

budding methods (double bud, patch and shield-budding) and two grafting 

82


methods (whip and veneer) to propagate soursop on several rootstocks 

(commercial soursop, a local soursop type called Chocó soursop, custard 

apple, and sugar apple). The shield-budding method on both soursops as 

rootstocks showed the highest percentage take, with 83% of success. The 

graft or bud union should be placed 15-20 cm above ground level (Pinto and 

Silva, 1994; Nakasone and Paull, 1998) to avoid long trunks and tall 

canopies. 

Both budding and grafting are better carried out in spring with the start of sap 

flow (Wester, 1912; Campbell and Phillips, 1983). According to George and 

Nissen (1986), in the Philippines patch budding of sugar apple is 

recommended prior to leaf abscission, which occurs during the dry season 

(November to February). 

Occasionally an annona grower may establish an orchard with an 

unproductive or unpopular cultivar, whose negative results will only show up 

three years later. He can replace the undesired canopy by using top working 

methods (George and Nissen, 1987), essentially establishing a new 

productive orchard with a high quality cultivar without replanting (Fig. 10-6). 

83


Figure 10-6. Steps of the topworking technique used to regenerate an 

unproductive canopy of an annona tree 

A reliable in vitro method for propagation of Annona species would be of 

considerable benefit to the annona industry because it would allow rapid 

clonal propagation of superior yielding and disease resistant varieties, as well 

as enabling clonal multiplication of superior rootstock material in areas 

where grafted Annona species are required (Rasai et al., 1994). In addition, 

micro-propagation would facilitate the exchanging of germplasm materials 

among research centres. However, rapid multiplication with sustained 

proliferation and in vitro rooting has always proved to be difficult. 

84


10.2 Field establishment 

10.2.1 Orchard location 

The location of an orchard is important as this will influence the quality of 

the fruit and the potential income generated. There are a number of factors to 

be considered before investing in the establishment of an annona orchard, or 

the incorporation of annona trees into existing small holder farming systems. 

The climatic conditions of an existing or potential orchard location will 

determine which species and/or varieties can be grown and their performance 

under these conditions. The ecological requirements of the annona species 

highlighted here are detailed in chapter 5 - Ecology. 

The soil type is also very important. Although most of the Annona species 

grow on a wide range of soil types, from sands to clay loams, higher yields 

are attained from trees grown in sandy loam soils. Soils should be free 

draining as Annona species do not grow well in soils with drainage problems 

and a high water content in the soil can encourage root diseases (Nakasone & 

Paull, 1998). 

Plenty of water for irrigation is essential and proximity to a paved road is 

also important, especially if the grower is concerned about fruit yield and 

quality. In addition, proximity to a large market will allow a higher income, 

principally by saving on transportation costs. 

10.2.2 Land preparation 

The orchard area must be cleared of shrubs and weeds. Four to six months 

before ploughing, a soil sample can be taken to determine the lime 

requirements and soil nutrient levels (Nakasone and Paull, 1998); section 

10.3.5. provides the standard methodology for sampling, however the local 

extension agent should be consulted about sampling methodologies for 

specific locations. Lime and phosphorus are applied before ploughing and 

harrowing (see section 10.3.5 for recommendations), although phosphorus 

can also be applied in the planting pit (Pinto and Silva, 1996; Nakasone and 

Paull, 1998). Drainage should be installed at this time to avoid flooding, with 

either contour or subsurface drains. 

On a medium to light texture oxisol, ploughing to a maximum soil depth of 

30 cm and harrowing twice should be carried out 1 to 2 months before the 

85


wet season to attain the desired soil tillage. This operation will also expose 

the soil seed bank in order to eliminate most of the undesirable weeds (Pinto 

and Ramos, 1997). 

Depending on the degree of slope, the grower can use one of three types of 

planting systems: square (Fig. 10-7 A), rectangular or quincuncial (also 

called triangular) (Fig. 10-7 B). In orchards with slopes greater than 3%, the 

soil should be prepared along contour lines and the quincuncial system 

should be used to minimize soil erosion (Figs. 10-8 A and 10-8 C). On the 

other hand, flat land (with slopes less than 3%) does not need contour line 

practices, and can use a square (Fig. 10-8 B) or rectangular planting system. 

Nakasone and Paull (1998) suggest that minimal tillage can be achieved with 

a 2 m wide cultivated band where the trees are to be planted. 

Figure 10-7. Soursop planting systems: A) square and B) quincuncial. 

Adapted from Torres and Sánchez (1992) 

86

Plate 1. Purple skinned sugar apple is commercialized as an exotic fruit, 

since its colour makes it look somewhat like a rotten fruit thus limiting its 

acceptance at consumer market. 

Plate 2. A solarization system is used as a pre-planting treatment for germination 

and seedling growth media to control fungi and nematode attacks. 

87


Plate 3. An intercropping system using sugar apple (main crop) and papaya 

(secondary crop) can help growers to earn additional income. 

Plate 4. Small plastic containers (e.g. empty film-roll holders) can be used 

to carry pollen . 

88


Plate 5. Small plastic containers to carry pollen should be kept in the operator’s 

pocket, to facilitate hand pollination 

Plate 6. Annona fruit borer (above left), seed borer (also called soursop 

wasp; above right) and trunk borer (below) are the most important annona 

pests. 

89


Plate 7. Soursop damaged by brown rot disease caused by the fungus 

Rhizopus stolonifer. 

Plate 8. Mature sugar apple fruit at its “harvest point”, and fully ripened 

fruit at its “consumption point”. 

90

Figure 10-8. Planting system according to the slope of the land. A) 

Quincuncial system; B) square system; and C) contour line. Adapted from 

Torres and Sánchez (1992) 

10.2.3 Time of planting 

The best time for planting is at the beginning of the wet season, this 

minimizes frequent watering after planting, especially if there are seasonal 

dry periods and no irrigation facilities (Torres and Sánchez, 1992; Pinto and 

Ramos, 1997). When irrigation is provided, planting can be performed in any 

season, although low air relative humidity can cause leaf dryness and 

possible plant death. Hence, even with irrigation, wet season establishment is 

preferable. 

10.2.4 Direct seeding 

Direct seeding in the field is the traditional method and may still be used on 

small holder farms, but is not used commercially any more. Although this 

91


method can, supposedly, save time and money with nursery management of 

the rootstock, its cost-benefit ratio is likely to be very high, considering that 

grafting operations in the field have very low success, obliging the grower to 

replant the orchard later. 

10.2.5 Transplanting and spacing 

In general, annona plants are ready for transplanting into the field or for use 

as rootstocks when they are about 8 to 15 months old, have attained a height 

from 50 to 100 cm and have at least 4 to 6 mature leaves. Before 

transplanting, many growers cut the leaves in half, to reduce transpiration, 

and cut the tip of the main root, apparently to induce production of more 

lateral roots. Both are empirically developed practices and have not been 

validated by research. 

The plastic nursery bags should be removed and the plant's collar placed at 

the ground level or, at the most, a few centimetres above the ground level if a 

planting depression is used to facilitate watering. The young plants should be 

irrigated as soon as possible after transplanting and they should be supported 

to avoid wind damage. Also, a mulch should be provided to avoid soil 

dryness around the newly transplanted materials. Although thick black plastic 

can be used to cover the area under the canopy of the new annona tree to 

avoid water loss through evaporation, dried grass or rice husks are much 

more practical materials; they are also biodegradable and avoid 

environmental problems, as well as being cheaper and easily found in the 

field. 

Current field spacing ranges greatly among commercial orchards, from 6 x 4 

m to 8 x 6 m for cherimoya (George and Nissen, 1992), 4 x 4 m to 8 x 8 m 

for soursop (Torres and Sánchez, 1992; Pinto and Silva, 1996; Pinto and 

Ramos, 1997), 3 x 3 m to 5 x 5 m for sugar apple (Coronel, 1994) or 4 x 4 m 

on poor soil and 5 x 7 m on good soil (Singh, 1992). There is no 

experimental information on the recommended spacing of custard apple, this 

is certainly due to the absence of commercial orchards. For wild soursop, 5 x 

5 m is recommended (FAO, 1988). 

Some authors (Campbell and Phillips, 1983; Nakasone and Paull, 1998) 

comment that annona spacing also depends upon the rootstock and pruning 

management. In Florida, narrow plant (4 - 6 m) and row spacing (6 - 7 m) is 

used for cherimoya; the rows run North-South in a triangular layout, 

whatever the planting distance used. Narrower spacing is also used for 

92


atemoya ‘African Pride’ on sugar apple rootstock and the widest spacing for 

atemoya ‘Pink's Mammoth’ on cherimoya. In Brazil, soursop spacing varies 

from 6 x 6 m to 8 x 8 m (Pinto and Silva, 1996), while sugar apple has a 

wider range of spacing depending on the rootstock vigour, and varies from 6 

x 4 m with sugar apple as a rootstock to 8 x 5 m with custard apple as a 

rootstock (Kavati and Piza Jr., 1997). 

10.3 Orchard management 

Orchard management can be complex and there is a need to experiment to 

ascertain the validity of many aspects. Innovation is also needed, e.g. 

intensive cultural methods, such as trellising or espaliering, require study. 

NRC (1989) reported that espaliering of cherimoya in Madeira (Portugal) 

was extremely successful. 

10.3.1 Windbreaks 

Since most of the Annona species are shallow rooting and consequently very 

susceptible to wind damage, the use of windbreaks is an important orchard 

practice, particularly during the first three years (George et al., 1987). The 

windbreak should be established prior to transplanting the annonas into the 

field. Casuarina (Casuarina equisetifolia) has been used in some Brazilian 

regions as a windbreak, although this species is very competitive for water 

and is susceptible to natural combustion, thus offering a fire risk. The best 

option is to consult the local extension service about windbreak species that 

have proven useful locally, as well as their placement, which depends upon 

their root distribution, plant height and crown density. 

10.3.2 Pruning 

Tustin (1997, cited by Castro et al., 1999) stated that pruning is an important 

cultural practice used to regulate the tree canopy. It influences growth by 

manipulating the balance between vegetative and fruiting growth (Cautin et 

al., 1999). Current pruning methods in cherimoya produce shoots with 

variable levels of vigour. 

The literature mentions several types of pruning of annona trees, such as for 

tree shape, maintenance, rejuvenation and production (also called green 

pruning) (Torres and Sánchez, 1992; Agustín and Alviter, 1996; Pinto and 

Ramos, 1997; Nakasone and Paull, 1998; Bonaventure, 1999). However, 

93


Alvarez et al. (1999) pointed out that there are basically only two types of 

pruning: for plant formation and for plant production. 

Pruning for plant formation begins in the first year at the same time as the 

training operation, though this may vary according to the species, and 

continues until the fifth year after planting out (Agustín and Alviter, 1996). It 

is necessary to begin training cherimoya trees in the nursery (Nakasone and 

Paull, 1998), however, soursop and sugar apple produce their branches close 

to the ground and have a single trunk (Coronel, 1994; Pinto and Ramos, 

1997), therefore, they do not require interference at such an early stage. 

Pruning has several objectives: a) development of good tree architecture in 

order to increase yield; b) acceptable aeration and light penetration; c) ease of 

access for cultural practices, such as artificial pollination, pesticide spraying 

and harvesting; d) removal of lower limbs (especially those touching the 

ground) and branches that are rubbing against each other (Torres and 

Sánchez, 1992; Pinto and Ramos, 1997; Nakasone and Paull, 1998; 

Bonaventure, 1999). 

Anderson and Richardson (1992) described an additional pruning practice 

that should be carried out during the first 4 years. This involves cutting the 

single trunk at 80 cm height, to stimulate production of primary branches. In 

the spring of the second year, the primary branches should be cut at a length 

of 40 cm to stimulate production of secondary branches. Similarly, in the 

third and fourth years, the same pruning procedures should be made to the 

secondary and tertiary branches, however, with 30 cm and 20 cm lengths, 

respectively (Fig. 10-9). It is important that care be taken during this 

procedure to ensure that a large number of internal branches do not remain 

and impede the necessary canopy aeration. Therefore, this type of pruning 

should be supplemented by an annual pruning through which better aeration 

of the internal canopy is maintained (Nakasone and Paull, 1998). 

Agustín and Alviter (1996) described different pruning methodologies for 

cherimoya involving the maintenance of two and three principle branches. 

Pruning using the two-branch system should begin between the fourth and 

the fifth month after transplanting. The two principal branches should be 

selected according to their vigour, and the others should be removed 

establishing a v-shaped training type (resembling two fingers in an opened 

position). The three-branch system described by Nakasone and Paull (1998) 

requires the complete removal of the top part of the tree leaving 90 cm of the 

main trunk, this encourages the production of lateral branches. Agustín and 

Alviter (1996) commented that these lateral branches should be at angles of 

94


approx. 120° from each other. These new branches should be spaced at 15-25 

cm above each other in different directions to develop a good scaffold by the 

fifth year after planting, when the plant is about 2 m tall (Nakasone and 

Paull, 1998). The tree can then be allowed to grow naturally while 

maintaining good aeration. 

Figure 10-9. Stylized diagram of cherimoya tree formation with length of 

the growing branches of subsequent years. Adapted from Anderson and 

Richardson (1990) 

95


Torres and Sánchez (1992) describe two types of pruning for soursop tree 

formation, both called 'free canopy shapes': a) pruning to keep a central axis 

with horizontal lateral branches (Fig. 10-10 A); b) pruning to keep forked 

branches (Fig. 10-10 B). Branch formation should begin between 60 and 80 

cm above the soil, and plant height must be kept to 3.5 m (Fig. 10-10). 

Figure 10-10. Two types of pruning for soursop tree formation: A) by 

keeping a central axis and horizontal lateral branches; B) by keeping 

forked branches. Adapted from Torres and Sanchéz (1992) 

During the pruning for soursop tree formation it is important to break the 

apical dominance of branches, in order to promote the growth of new lateral 

branches; this is followed by another pruning to leave 3 to 4 well distributed 

new branches, with insertion angles of approx. 30° in relation to the trunk 

(Torres and Sánchez, 1992). Soursop trees naturally produce a symmetrical 

canopy well adapted to a central-leader system; therefore, a good alternative 

for soursop tree formation is to develop a mushroom-shaped tree that is 

topped at a height of 2.0-2.6 m (Nakasone and Paull, 1998). When the 

soursop tree is properly trained, little pruning is required, except to thin out 

poorly placed and weak branches, as well as the annual pruning of the 

longest branches extending horizontally and vertically (Nakasone and Paull, 

1998). 

Torres and Sánchez (1992) described pruning for the maintenance and 

rejuvenation of soursop. Pruning for maintenance is required two to three 

times a year, preferably after harvesting. It consists of the elimination of 

undesirable branches, i.e. those which are not productive, dried twigs and 

dead branches, or those which are diseased; it also includes the elimination of 

96


branches above 2.5 m height to avoid excessive growth and alternate yield, 

and to promote better quality fruits. Pruning for rejuvenation is done in old 

soursop orchards, where the semi-abandoned tall trees (Fig. 10-11 A) have 

excessive branching with low yield capacity. All vertical branches are 

eliminated including the thick lateral and terminal ones (Fig. 10-11 B), in 

order to attain a plant height of 2.5-3.0 m, then training the new lateral 

branches into horizontal positions at a later date (Fig. 10-11 C). 

Figure 10-11. Pruning for rejuvenation of a soursop tree: A) canopy of 

an old soursop tree before pruning; B) pruning of lateral and terminal 

branches; C) training of lateral branches into horizontal positions. 

Adapted from Torres and Sánchez (1992) 

Pruning is not usually carried out on sugar apple trees in arid regions, except 

to remove unproductive old branches (Singh, 1992). No information is 

available for custard apple. Methods of pruning have not been worked out for 

wild soursop, but the trees respond well to coppicing (FAO, 1988). 

Pruning for plant production is very common in cherimoya and sugar apple 

(Agustín and Alviter, 1996; Bonaventure, 1999). However, the literature does 

not mention pruning for production in soursop (Torres and Sánchez, 1992; 

Pinto and Ramos, 1997). The position of the bud may be the reason for this, 

since soursop has its lateral buds in the leaf axil while cherimoya and sugar 

have subpetiolar buds ('buried buds') in the base of the swollen leaf petiole 


Leaf shed in cherimoya and sugar apple generally occurs prior to the 

elongation of the 'buried buds', so that mechanical removal of leaves, by 

97


stripping, or chemical removal, with urea or ethephon, will release these buds 

(George and Nissen, 1987). However, this technique is not recommended for 

soursop. Bonaventure (1999) describes this method as 'green pruning' and 

affirms that it is very important in cherimoya. He mentions the two most 

important types of green pruning: the first type to reduce excessive 

vegetation; and the second to separate the two or three new leaves, followed 

by removal of the apical bud with the finger nail. According to this author, it 

is possible to promote late production by using green pruning. 

George et al. (1987) commented that chemical pruning by defoliation of 

cherimoya and sugar apple trees is necessary for cultivars exhibiting strong 

apical dominance, since light fruiting-pruning may not be sufficient to 

release many buds from induced dormancy. They also commented that a 

range of defoliants have been investigated, but a mixture of 250 g urea and 

1 g ethephon, plus wetting agent, in 100 litres of water has been most 

successful. Two defoliation periods are possible: a) at 5 to 10% bud-break, 

when the new shoots are about 3 to 5 cm long; and b) at mid season (first 

week of January in Australia), applied to non-bearing trees, usually 2 to 3 

years of age. 

Pruning for production of sugar apple is very important, since bearing sugar 

apple trees seem to deteriorate, in terms of fruit production, faster than any 

other annona (Coronel, 1994). The heading back of the branches at the onset 

of dormancy to rejuvenate sugar apple trees will result in resumed production 

after 2 years (Coronel, 1994). Dormancy of sugar apple buds depends on the 

climate. In south-eastern Brazil, dormancy is due to low temperatures 

between May and July, while in the semi-arid areas of the north-eastern 

region it is due to strong drought (water stress) during several months (Kavati 

and Piza Jr., 1997). 

In Thailand, sugar apple trees are rejuvenated every year to produce new 

fruiting shoots. This operation consists of removing all small shoots and 

heading back larger shoots to about 10-15 cm long (Coronel, 1994). This 

operation is carried out during the dry season, just prior to shedding of the 

leaves; the plants are then irrigated to promote new shoots and irrigation is 

continued until the rainy season starts. This is the same procedure as in northeastern 

Brazil. 

The pruning for production of sugar apple should begin with 1-year-old 

branches by cutting them back to 10 cm and leaving 120-150 branches per 

tree (Nakasone and Paull, 1998); flower initiation will then begin at the basal 

end of the new growing branch. In China and Taiwan, normal fruit pruning 

98


occurs in January/February, with harvesting from July to September. 

However, a summer pruning with fruit thinning (June-October) can lead to 

harvesting from October to March, effectively increasing the fruit 

availability. The highest winter fruiting occurs when summer sprouts are 

pruned as compared to pruning non-fruiting shoots or pruning in late May. In 

India, light pruning is carried out on budded plants. In São Paulo, Brazil, the 

summer pruning is from January to March with harvest from August to 

October, since the late pruning (after March) can promote flowering during 

periods of low temperature, leading to reduced fruit set (Kavati and Piza Jr., 

1997). There are no such problems in north-eastern Brazil, since high 

temperature, low precipitation and irrigation stimulate successful fruiting of 

sugar apple, which is managed by pruning. 

10.3.3 Orchard maintenance, intercropping and 

cover-cropping 

In most annona orchards, weeds were traditionally eliminated by using hand 

tools, such as mattocks. Nowadays, this practice has been substituted by 

herbicides in commercial orchards. In Spanish cherimoya orchards the 

herbicides most commonly used are simazine or terbumeton mixed with 

terbuthylazine as pre-emergence treatments, and glyphosate or a mix of 

paraquat and diquat as post-emergence treatments (Farré et al., 1999). The 

irrigation system is often used to apply the pre-emergence herbicides. 

However, phytotoxicity may occur after herbicide application. 

Mulching is a very common maintenance system used mainly during the first 

years after planting, since it has a number of positive effects. These include 

improved distribution of humidity in the soil, reduction of evaporation losses, 

avoidance of crust formation on the soil, increased development of healthy 

feeder roots improving efficiency of fertilizer use (Farré et al., 1999; George 

et al. 1987), and reduction of weed infestations. As consumers demand 

chemical-free fruit, mulching will become increasingly more important. In 

practice, dried grass is a practical and cheap material to be used for 

mulching. 

Annonas can be grown as a mono-crop in high density orchards or as an 

intercrop among larger fruit trees, such as mango and citrus (Ochse et al., 

1994), or on small holder farms. Even as the main crop, the space between 

sugar apple trees may be planted with other fruit crops, such as papaya, 

which is a common practice in north-eastern Brazil (Plate 3). It is also 

99


possible to intercrop annonas with annual field or vegetable crops to earn 

additional income from the land during the first 2-3 years before annona 

production. 

The establishment of perennial leguminous or other cover crops, such as 

beans, can provide not only additional income, but also avoid soil erosion 

and improve the physical structure of the soil. Short grass can be used as a 

cover crop in the first 12 months after transplanting (Nakasone and Paull, 

1998), although fertilization needs to be modified to supply two crops instead 

of one. A wild peanut (Arachis pintoi) is currently being tested at Embrapa 

Cerrados Agricultural Research Centre, and has so far been very successful. 

Cover cropping may become more important as consumers demand 

chemical-free fruit. 

10.3.4 Flowering, pollination and fruit set 

Annona flowers are hermaphrodite; both female (carpels) and male (stamens) 

organs are in the same flower. However, the female part matures before the 

male, which is known as dichogamy of the protogynous type. According to 

Mansour (1997), there is a short period of stigma receptivity after anthesis, 

which is 2-3 h in dry weather. Natural pollination is carried out mainly by 

insects, such as Coleoptera (beetles), but is ineffective in several countries 

(Saavedra, 1977; Pinto and Silva, 1996; Grossberger, 1999). In Chile, insects 

rarely visit a cherimoya flower which suggests that entomophilous 

pollination plays only a secondary role in this species (Saavedra, 1977). 

Cherimoya is a native species in Chile and should have co-evolved insect 

pollinators if it were important. 

Although dichogamy and the low population density of pollinator insects are 

important limiting factors to successful natural pollination, the effect of 

climate and pollen viability seem to interfere greatly with the response to 

both natural and artificial (hand) pollination. The effects of these factors may 

result in fertilization failure of all or several ovules, resulting in small or 

asymmetrical fruits, which obviously affects yield and commercialisation 

(Saavedra, 1977). Most annonas flower when atmospheric humidity is low 

(Saavedra, 1977), suggesting the importance of humidity in drying out the 

stigmas and lowering pollen germination. 

Anthesis (flower opening) is most common in the morning or in the evening, 

indicating that it is favoured by low temperatures (Mansour, 1997). Studying 

the problem of fruit set on custard apple in Dhawar, Egypt, Farooqi et al. 

100


(1970) found that flower opening occurs from early morning until noon and 

that the stigma is more receptive at this time on the first day, giving about 

90% fruit set. Thereafter, receptivity decreases gradually, resulting in 

minimal fruit set (8%) on the fifth day. 

Pollen germination of cherimoya has been found to be optimal at 20-25°C, 

which is the same temperature required for good fruit set in the orchard 

(Rosell et al., 1999). Higuchi et al. (1998) studied the effects of warm 

(30/25°C) and cool (20/15°C) day/night temperatures on fruit set and growth 

in potted cherimoya under greenhouse conditions. They found that fruit set at 

warm temperatures was very low, and ascribed this response to both pollen 

and stigmatic damage from heat stress. In Madeira island, the best climatic 

conditions for cherimoya self-pollination are temperatures equal to or above 

22°C, with relative humidity of 70-80%, whereas temperatures below 22°C 

with RH above 90% reduce pollination success (Nunes, 1997). 

The stage of flowering phenology, pollen maturity and viability are factors 

that affect fruit set and yield of Annona species. In New Zealand, Richardson 

and Anderson (1996) compared fruit set at different flowering times by using 

hand pollination methods in individual trees as main plots and time of 

pollination on 20 flowers as sub-plots. They found that cherimoya pollination 

is more successful at the beginning of the flowering period (January), with a 

total yield of 34.8 kg/tree, than at the end of the flowering period (February) 

with a total yield of only 0.1 kg/tree. It was also observed that pollen viability 

varied from 20 to 50% and seediness increased from 11 seeds/fruit at the 

beginning to 59 seeds at the end of the flowering period. In Chile, Saavedra 

(1977) found that the first flowers of cherimoya to open gave poorer fruit set 

with hand pollination than the later ones, probably because the majority of 

the pollen grains at the beginning of flowering were still in the tetrad stage, 

with thick walls and full of starch; at this stage, the pollen has a low viability 

and, consequently, gives a low fruit set. 

Grossberger (1999) commented that when pollen is used within 1-2 h of its 

collection, more than 80% fruit set is generally obtained, as assessed by fruit 

set two weeks after pollination. When pollen was stored for 12 h in a 

standard refrigerator, fruit set fell to about 65% and to 35% after 24 h. This 

would suggest that old pollen results in a low percentage of fruit set, even 

using a brush or blower, which are appropriate tools for hand pollination. 

Fruit set by hand pollination ranges from 44.4 to 60% depending upon the 

species, while fruit set from open pollination is always less than 6% 

(Mansour, 1997). Hand pollination is the only certain strategy to ensure 

101


commercial production, since it guarantees significantly higher production 

and better fruit quality than open pollination. Cogez and Lyannaz (1996) 

compared natural and hand pollination in two sugar apple cultivars: ‘Thai 

Lup’ and ‘New Caledonia’. Natural pollination had 0% and 3.6% success in 

‘Thai Lup’ and ‘New Caledonia’, respectively, while hand pollination 

success was 100% in ‘Thai Lup’ and 90-93% in ‘New Caledonia’. When 

intravarietal and intervarietal pollen was combined with hand pollination, the 

success rate was 90.9% and 92.6% success, respectively, in New Caledonia. 

Pinto and Ramos (1999) obtained 26% fruit set with natural pollination of 

soursop in the Brazilian Cerrado conditions (Brazilian savannah), while fruit 

set was 73% when hand pollination with a paint brush was used. 

Allogamous hand pollination is also effective with cherimoya. Duarte and 

Escobar (1998) applied self pollen on cherimoya cultivar Cumbe in El 

Zamorano, Honduras, as well as cultivar Bronceada, in the morning (6-8 h) 

and in the late afternoon (16-18 h). The highest fruit set (46.4%) was 

obtained with cross pollen, as compared with 30.3% and 23.1% with self 

pollen applied in the morning and in the afternoon, respectively. Fruit set was 

also better at the beginning of the flowering period. 

The success of hand pollination depends on the technique, as well as the 

species and conditions. Economically effective hand pollination of 

cherimoya was carried out for the first time in California by Dr. Schroeder in 

1941 (Grossberger, 1999) and to date the technique has varied little. Several 

authors (Agustín and Alviter, 1996; Mansour, 1997; Bonaventure, 1999; 

Grossberger, 1999) have described the hand pollination technique as follows. 

The cherimoya flowers that will serve as the pollen source are gathered and 

held in a small paper bag until the pollen is shed. The operator should 

remove the 3 petals of the external whorl of freshly opened flowers that will 

be pollinated. The 3 petals of the internal whorl are then held open with one 

hand, while, with the other, the pollen is applied onto the pistils with a small 

camel hair brush, using several back and forth strokes. Pollen can also be 

blown onto the pistils with special blowers (e.g., the Aluminum Hand 

Blower, Technes Industry, São Paulo, Brazil; Bonaventure, 1999). A small 

cylindrical plastic container (such as those used for rolls of film: Plate 3) is 

an important tool used by soursop growers of the Brazilian Cerrados to carry 

pollen, which can be kept in the operator's shirt pocket (Plate 5), leaving the 

grower's hands free to execute the hand pollination. 

Hand pollination of cherimoya is generally practiced in the morning or the 

afternoon as it is too warm and dry at midday and a lower fruit set is obtained 

102


(Grossberger, 1999). Similarly, in soursop, pollination should be practiced in 

the morning, preferably around 9 am and approximately 19 h after the 

collection of the pollen donor flowers (Pinto and Ramos, 1999). The donor 

flowers should be collected from the terminal portion of the branch, since 

these flowers generally present low fruit set themselves (Torres and Sánchez, 

1992; Pinto and Ramos, 1999). 

Hand pollination is considered to be time-consuming and costly (Nakasone 

and Paull, 1998). Attempts have been made to substitute hand pollination by 

growth regulators to enhance fruit set. Experiments carried out by Yang 

(1988, cited by Nakasone and Paull, 1998) indicated that fruits grow very 

slowly and with less fruit drop when indole-acetic acid (IAA) and 

naphthalene acetic acid (NAA) are applied, while gibberellic acid (GA) 

promotes adequate fruit set and growth rate. Saavedra (1979) found that the 

application of either NAA or GA can stimulate some production of seedless 

fruits weighing 200-300 g, but only GA significantly enhanced fruit set and 

stimulated growth of seedless fruit. However, GA can also induce severe 

cracks in fruit rinds before harvest. Therefore, hand pollination is still the 

most effective strategy to increase production and quality of fruit. 

Fruit thinning is necessary to regulate crop load and to maintain fruit size, 

especially for high yielding annona cultivars (George et al., 1987). The 

thinning operation involves the removal of misshapen fruit and thinning of 

fruit clusters, both of which improve the quality of the remaining fruit for 

market. 

10.3.5 Nutrition and fertilization 

Planting and Juvenile Phases 

The establishment of a fertilization programme should be based on the 

nutrient uptake by the target species (Mengel and Kirkby, 1987; Torres and 

Sánchez, 1992; Hermoso and Farré, 1997). In establishing orchards, the 

preliminary fertilization (especially of phosphorous) and soil pH correction 

of the field should be based on soil analysis. Sufficient sub-samples, for 

instance 25 per hectare for phosphate and potassium, should be collected in 

order to obtain a representative sample (Mengel and Kirkby, 1987). Soil subsamples 

should be taken from the 0-40 cm soil layer, by walking a zig-zag in 

the area, a mixture of these sub-samples should then be prepared and a 200- 

300 g composite soil sample sent off for nutrient analysis (Pinto and Silva, 

1996; Bonaventure, 1999). 

103


The fertilization of the planting pit varies according to the species and the 

soil nutrient levels. In Venezuela, 250 g of NPK 10-10-15 or 10-15-15, and 5 

kg of manure is a common recommendation for soursop planting pits 

(Araque, 1971). Annona trees are sensitive to low soil pH and this should be 

adjusted to pH 6.0 if necessary, using either dolomite or calcitic lime before 

planting out. 

For wild soursop, which has not yet reached orchard status, fairly general 

recommendations on fertilizers are available: compound NPK + Mg at rates 

of up to 100 g/tree every 3 months (FAO, 1988). Similarly on sugar apple in 

India, fertilizer can be applied at the start of the monsoon at the rate of 250 g 

N and 125 g P and 125 g K per tree (Anon., 1981). 

Fertilizer rates and timing need to be adjusted according to the tree age and 

the species. Ibar (1979) describes the fertilization schedule for cherimoya 

orchards during the juvenile period (from 1st to 3rd year) according to tree 

age (Table 10-4). However, the schedule for fertilization of the vegetative 

phase would be much more accurate if the amount of fertilizer applied were 

based on the soil nutrient content (Table 10-5 and Table 10-6) (Silva and 

Silva, 1997). 

Table 10-4. Recommendation of NPK fertilization for cherimoya in 

Spain based on tree age after planting during the first three years of 

orchard establishment 

Age after planting 

(years) 

Nitrogen (of 

N/plant) 

Phosphorus (of 

P 2 

O 5 

/plant) 

Potassium (of 

K 2 

O/plant) 

0-1 240 120 120 

1-2 360 180 180 

2-3 480 240 240 

Source: Ibar (1979). 

Table 10-5. Recommendation of P and K for soursop trees of different 

ages according to the amount of N fertilization applied (g/plant) and 

levels of P 2 0 5 and K 2 0 in the soil analysis in the semi-arid region of Brazil 

Tree Age N g of P 2 

O 5 

/plant g of K 2 

O/plant 

Phosphorus in the soil (µg/cm³) Potassium in the soil 

(µg/cm³) 

0-10 11-20 > 20 0-45 46-90 > 90 

0-1 year 40 - - - 60 40 30 

1-2 years 80 80 60 40 80 60 40 

3-4 years 120 120 80 60 120 80 60 

104



O 5 


O/plant 

> 4 years 180 120 80 40 180 120 60 

Source: Silva and Silva (1997). 

Table 10-6. Recommendation of P and K for sugar apple trees of 

different ages according to the amount of N fertilization applied (g/plant) 

and levels of P 2 

0 5 

and K 2 

0 in the soil analysis in the semi-arid region of 

Brazil 


O 5 


O/plant 

Phosphorus in the soil (µg/cm³ Potassium in the soil 

(µg/cm³ 

0-10 11-20 > 20 0-45 46-90 > 90 

0-1 year 50 - - - 70 40 20 

1-2 years 100 80 60 40 60 40 20 

3-4 years 120 120 80 60 120 80 60 

> 4 years 180 160 120 80 200 140 80 

Source: Silva and Silva (1997). 

Besides soil analysis, leaf tissue analysis has been accepted as a way to 

understand nutrient uptake conditions from the soil (Mengel and Kirkby, 

1987) and should be used during the juvenile phase of plant growth. The 

most appropriate methodology for collecting annona leaves for analysis 

depends on the tree's age, the position of the leaves in the canopy and the 

period for sampling. Cherimoya leaves should be collected from intermediate 

branches during the fruiting period or late in the harvest season (Gonzalez 

and Esteban, 1974). Collection of the third and fourth leaf pairs from the 

intermediate branches of the canopy in the four cardinal points is the most 

appropriate for foliar analysis of soursop (Laprode, 1991). Soursop leaves 

should be 8-9 months old and should be collected from intermediate branches 

of healthy plants, free of pesticide residues prior to analysis (Pinto and Silva, 

1996). A sample of 4 leaves per tree from each of 25 randomly selected 

plants in the orchard, totalling 100 leaves, is required for complete tissue 

analysis. Several authors (Avilan, 1975; Sadhu and Ghosh, 1976; Navia and 

Valenzuela, 1978; Silva et al., 1984) have reported a range of nutrients in 

normal and deficient leaves of cherimoya, soursop and sugar apple (Table 

10-7), which serve as a guide for annona plant deficiencies. 

105


Table 10-7. The average normal and deficient levels of macro and micronutrients 

in leaves of cherimoya, soursop and sugar apple 

Species N% P% K% Ca% Mg% S% B 

ppm 

Cherimoya 

(1) 

Fe 

ppm 

Zn 

ppm 

a) Basal 

leaves 

Normal 1.90 0.17 2.00 0.80 0.25 - - 215.0 23.0 

Deficient 0.72 0.09 1.00 0.25 0.04 - 10.0 140.0 12.0 

b) Apical 

leaves 

Normal 2.91 0.17 1.95 0.60 0.26 - - 125.0 29.0 

Deficient 0.90 0.10 1.00 0.15 0.05 - 6.0 40.0 20.0 

Soursop (2) 

Normal 1.76 0.29 2.60 1.76 0.20 - - - - 

leaves 

Deficient 1.10 0.11 1.26 1.08 0.08 - - - - 

Soursop (3) 

Normal 2.5- 

leaves 2.8 

Deficient 1.3- 

1.6 

Sugar 

Apple (4) 

Normal 2.8- 

leaves 3.4 

Deficient 1.9- 

2.8 

0.14- 

0.15 

0.06- 

0.07 

0.34- 

0.34 

0.17- 

0.19 

0.87- 

2.47 

0.75- 

1.66 

2.61 0.82- 

1.68 

2.64 0.45- 

0.81 

0.36- 

0.38 

0.07- 

0.08 

0.15- 

0.17 

0.11- 

0.13 

35.0- 

47.0 

6.0- 

14.0 

- - 

- - 

- - - - - - 

- - - - - - 

Sources: (1) Navia and Valenzuela (1978); (2) Avilan (1975); (3) Silva et al. 

(1984); (4) Sadhu and Ghosh (1976). 

Gazel Filho et al. (1994) found the following variations in the normal 

macronutrient contents (%) of different soursop varieties: N: 1.99 - 2.04; P: 

0.12 - 0.14; K: 1.49 -1.52; Ca: 1.20 - 1.52; Mg: 0.19 - 0.22. Most of these 

macronutrient contents do not fall inside the range presented in Table 10.7, 

which demonstrates that several other factors, such as soil, variety, climate 

etc., influence nutrient contents in living plant tissues (Mengel and Kirkby, 

1987), potentially interfering in the orchard fertilization program. 

Fruit size, colour, shape and taste are quality characteristics affected by 

nutrient deficiencies (Mengel and Kirkby, 1987). Undurraga et al. (1995) 

reported rates of 4.14-6.72 kg of N (as urea) applied to individual cherimoya 

cv. ‘Concha Lisa’ trees in Colombia, resulted in the lowest soluble solids 

contents and fruit pulp firmness values, and the highest titratable acidity 

106


values. This suggests that N application at or above 4.14 kg per cherimoya 

tree impairs the storage quality and organoleptic properties of the fruits. 

Annona fruits have a high demand for potassium, and to avoid serious 

symptoms of potassium deficiency (especially those related to fruit 

production and quality), potassium content in leaves should be maintained 

above the suggested critical level of 1.0% (Torres and Sánchez, 1992; Silva 

and Silva, 1997). Torres and Sánchez (1992) recommended a guide for 

nitrogen, phosphorus and potassium (Tables 10-8, 10-9 and 10-10) for adult 

soursop trees based on the age and the nutrient content of the soil in 

Colombia, for orchard fertilization. 

Table 10-8. A guide for nitrogen fertilization (g of N/tree/year) for adult 

soursop trees in different regions of Colombia, according to age and 

nutrient content in the soil 

Region 

Intermediary Valley 

Atlantic Coast and 

Oriental Plains 

Organic Matter 

Tree Age 

(%) 3 years old 3-6 years old 6 years old 

3 45-70 80-110 110-135 

3-5 30-45 50-80 80-110 

3 50-80 90-120 120-140 

3-5 30-50 60-90 90-120 

5 20-30 30-60 60-90 

Source: Torres and Sánchez (1992). 

Table 10-9. A guide for phosphorus fertilization (g of P 2 

0 5 

/tree/year) for 

adult soursop trees in different regions of Colombia, according to age 

and nutrient content (ppm) in the soil 

Region 

Inter Andean Valley 

Atlantic Coast and 

Oriental Plains 

P ppm 

Tree Age 

3 years old 3-6 years old 6 years old 

20 45-60 60-110 180-240 

20-40 20-45 30-60 120-180 

40 0-20 0-30 60-120 

15 60-80 75-130 140-360 

15-30 30-60 45-75 180-240 

30 0-30 0-45 90-180 


Table 10-10. A guide for potassium fertilization (g of K 2 

O/tree/year) of 

adult soursop trees in Colombia, according to tree age and potassium 

(meq/100 g of soil) content in the soil 

Content of Potassium 

Tree Age 

(meq/100 g of soil) 3 years old 3-6 years old 6 years old 

0.20 40-60 60-90 90-130 

0.20-0.40 20-40 40-60 60-90 

107


Content of Potassium 

Tree Age 

(meq/100 g of soil) 3 years old 3-6 years old 6 years old 

0.40 0-20 0-40 6-60 


The observation of deficiency symptoms, as a field analytical technique, is a 

fast and low cost method. When used with soil and leaf analyses it is very 

important for determining the nutritional status of annonas. The main general 

physiological characteristics and symptoms of deficiencies on annona trees 

are discussed below, following Avilan (1975), Navia and Valenzuela (1978), 

Mengel and Kirkby (1987), Torres and Sánchez (1992) and Silva and Silva 

(1997). 

Nitrogen - When the supply of N from the roots is inadequate, N from the 

older leaves is mobilized to feed the younger parts of the plant. Symptoms of 

nitrogen deficiency on cherimoya seedlings start at 40 days after sowing, 

with reduction of the plant and leaf sizes. The progression of the deficiency 

promotes an intense yellowing of the leaf and then its abscission. In soursop, 

the seedling shows a reduction in height, and leaf yellowing and abscission 

occur more quickly than in cherimoya. The leaf blade also shows a greenyellow 

chlorosis and changes texture. Symptoms of nitrogen deficiency in 

sugar apple seedlings are irregular bud development and lack of branches. 

The leaves are small and show dusty spots; abscission occurs later. 

Phosphorus - In cherimoya seedlings, deficiency symptoms become evident 

60 days after sowing. Irregular chlorosis occurs in the basal leaves and most 

show a darker green colour. As the deficiency develops, the leaves become 

small and irregular in shape, coffee coloured spots develop on the blade, then 

abscission occurs in these affected leaves. Deficient soursop plants only grow 

50% as fast as fertilized plants and the leaves show a necrosis on the blade 

edges. Sugar apple seedlings also show reduced growth with thin shoots and 

their leaves develop abnormal brown spots at their apices and on the blade 

edges. 

Potassium - Due to its great mobility, potassium moves from old organs to 

new ones easily. In general, deficient plants do not have the ability to 

transport carbohydrates produced in the leaves to the other plant organs. 

Potassium deficiency in cherimoya seedlings appears at 50 days after sowing, 

and shows intermediate symptoms that are between phosphorus and nitrogen 

deficiency. Brownish spots start from the apex and basal parts of the leaf 

blade, and gradually coalesce. In soursop, the leaf symptoms appear in the 

8th month after sowing with a reduction in size and an abnormal yellowing, 

then leaf abscission. Sugar apple trees show retarded growth and dryness of 

108


the apical leaves. Deficient plants may flower, but there will be no fruit set. 

A high content of K induces Mg and Zinc deficiencies. 

Calcium - Calcium is an immobile nutrient and its deficiency affects the 

zones of intense growth first. The symptoms of calcium deficiency in leaves 

of cherimoya seedlings appear after 30 days. New leaves show interveinal 

chlorosis. At 70 days, the apical meristem of cherimoya dies and the leaves 

stop growing and become twisted. In the leaves of soursop seedlings, calcium 

deficiency occurs later than in cherimoya (120 days), although the symptoms 

are similar. The best ratio of Ca:Mg is 3:1, a higher ratio can increase Mg 

deficiency. 

Magnesium - The first symptoms of deficiency are generally shown in the 

oldest leaves because Mg is a mobile nutrient. In cherimoya seedlings this 

begins at 50 days. An interveinal chlorosis then starts to become visible in 

the new leaves. In soursop, chlorosis is progressive, turning the leaves 

necrotic. 

Sulphur - This nutrient has low mobility, both in the plant and in the soil, 

especially in soils with low organic matter content and high C/N ratios. 

Therefore, the first symptoms of sulphur deficiency occur in the young 

leaves. The new leaves of cherimoya seedlings are notably smaller than the 

old ones after 75 days of sulphur deficiency, and the new leaves present an 

overall yellowing. Similar symptoms are also detected in the leaves of 

soursop seedlings. 

Boron - Similar to calcium, this nutrient is immobile; therefore, the first 

symptoms of boron deficiency occur in young leaves. In cherimoya seedlings 

the leaf symptoms appear after 70 days. The upper leaves turn intense green, 

with some chlorosis. At 140 days, leaf chlorosis becomes more intense. 

Symptoms are similar on the leaves of soursop seedlings. 

Iron - Like calcium and boron, iron is relatively immobile and the first 

symptoms occur in the young leaves. The initial symptoms are partial 

chlorosis (yellowish green), then the leaf blades become totally yellow, 

except over the veins. 

Zinc - Plants suffering from zinc deficiency often show chlorosis in the 

interveinal areas of the leaf. These areas become pale green, yellow, or white. 

Unevenly distributed clusters of small, stiff leaves are formed at the top of 

the young shoots; this symptom is known as rosette or little-leaf. 

109


Adult Tree Phase 

Physiologically, the fruits act as a sink for nutrients. Nutrient analysis of 

leaves however gives different results from fruits. A new methodology has 

recently been tested using fruits as the source of material for nutrient analysis 

in adult mango trees. Although, in practice, this method is not frequently 

used yet, it would be a useful additional test, together with soil and leaf 

analyses. 

The quantity of nutrients removed from the soil by fruit trees depends on the 

species, the variety and the yield (Kirkby and Mengel, 1987). Guirado (1999) 

observed that a cherimoya orchard with a population density of 156 plants/ha 

and a yield of 89.7 kg of fruits per tree extracted the following amounts of 

nutrients per kg of harvested fruit: 6.8 g N; 0.3 g P; 2.7 g K; 0.6 g Ca; 1.9 g 

Mg. Fruit production of soursop however, is more demanding and extracts 

larger amounts of macronutrients from the soil than cherimoya, except for N, 

Ca and Mg. (Figures are given in Tables 10-11 and 10-12). 

Table 10-11. Removal of macronutrients (kg) per tonne of soursop and 

sugar apple fruits produced 

Macronutrient 

Soursop 

Avilan et al. Silva et al. 

(1980) (1984) 

Sugar Apple 

Silva et al. Silva et al. 

(1984) 

(1991) 

N 2.97 kg/t 2.70 kg/t 7.17 kg/t 4.94 kg/t 

P 0.53 kg/t 0.54 kg/t 0.58 kg/t 0.25 kg/t 

K 2.53 kg/t 3.60 kg/t 5.19 kg/t 5.31 kg/t 

Ca 0.99 kg/t 0.26 kg/t 0.45 kg/t - 

Mg 0.15 kg/t 0.24 kg/t 0.46 kg/t - 

S - 0.27 kg/t 0.27 kg/t - 

Sources: Avilan et al. (1980), Silva et al. (1984). 

Table 10-12. Removal of micronutrients (g) per tonne of soursop and 

sugar apple fruits produced 

Macronutrient Soursop Sugar Apple 

Fe 8.03 18.48 

Cu 1.65 2.68 

Mn 2.71 3.26 

Zn 3.71 6.95 

B 2.75 3.12 

Source: Silva et al. (1984). 

Fertilizer should be applied to adult annona trees in the area under the outer 

third of the canopy (Pinto and Silva, 1996; Pinto and Ramos, 1997). 

Fertilization by foliar spraying is very important to supplement a soil 

110


fertilization programme during the period before flowering and harvesting to 

improve the fruit quality. Commercial liquid fertilizers can be applied 2-3 

times a year (Torres and Sánchez, 1992; Pinto and Silva, 1997). Cherimoya 

trees are sensitive to boron and zinc deficiencies, therefore, boron at 2.0 g/m 2 

should be applied to the ground area below the canopy. Spraying of 0.1% of 

zinc sulphate, applied at monthly intervals, will correct any zinc deficiency. 

In addition, boron and calcium sprays during flowering and early fruit set 

may be beneficial in reducing the incidence of internal fruit browning (Torres 

and Sánchez, 1992; Undurraga et al., 1995; Hermoso and Farré, 1997; 

Bonaventure, 1999). 

Fertilization with easily leachable nutrients, such as N and K, should be split 

into three or more applications during the year. Fertigation techniques 

(application of fertilizers through the irrigation system) is the best 

recommendation for this multiple application practice, providing a quicker 

and more controlled response than soil surface application; this practice also 

promotes a higher yield and fruit quality. However, there are no scientific 

results published on this technical issue which can confirm the 

recommendation for annonas. 

Organic cultivation of annona trees is a recent practice with no conclusive 

studies to identify scientifically acceptable recommendations for production. 

However, an increasing number of growers are reducing the use of 

agrotoxins and increasing the use of organic or biological products to 

improve fruit quality for market. Today, the foliar application of 

microorganism mixes (commercially named as EM-4 and EM-5), as well as a 

bioactivators (commercially named Aminon-25), seem to enhance plant 

metabolism and functions, such as photosynthesis and carbohydrate 

distribution, and give good results in terms of yield and fruit quality of 

organically grown cherimoya (Bonaventure, 1999). Further research results 

are expected to appear on this subject in the coming decade, due to the 

increasing consumer demand of pesticide-free and organic produce. 

10.3.6 Irrigation 

The selection of the appropriate irrigation system is directly related to three 

main factors: technical, economic and human (Silva et al., 1996), within 

which, there are several critical aspects. Water management (availability in 

quantity and quality) and its infiltration, slope of the land, plant phenology 

and climate are technical aspects. For instance, the sprinkler irrigation system 

should be used in areas where water is not a limiting factor and where slopes 

111


are not greater than 16% (Nunes, 1997). The use of sprinkler irrigation also 

has certain restrictions, especially regarding its use during the period of 

pollination and fruit set, since it can impede visits of pollinator insects and 

provoke the abscission of small fruits. Market prices of the irrigation system, 

costs of installation and maintenance, and financial resources are some of the 

economic aspects to be considered. Finally, the quality of the labour which 

will operate the irrigation system is the most important human aspect 

involved (Silva et al., 1996). 

The cost of buying and installing the irrigation system is generally the first 

critical aspect influencing any decision; therefore the cheapest irrigation 

system is often chosen. However, the price is not necessarily related to the 

efficiency. For example, the gravity flow of surface water through furrows 

and flood irrigation are the cheapest systems; however, water distribution is 

seldom uniform, leading to poor efficiency in the orchard as a whole (Santos, 

1997). In Spain, flood irrigation is used only in orchards situated in riverbottoms 

(Farré et al., 1999). In north-eastern Brazil, furrow and flood 

systems are considered old-fashioned, and have been substituted by drip and 

micro-sprinkler irrigation. 

In many areas, water has become very expensive; thus it is necessary to 

minimize costs by not over-irrigating. Drip and micro-sprinkler irrigation 

systems decrease costs and increase the efficiency of water use and 

management (Nunes, 1997), as well as improving fertilizer use and 

management, resulting in higher yields and better fruit quality. These 

localized irrigation systems are, nowadays, the commonest methods used on 

annona trees in north-eastern Brazil. A good irrigation scheme must be 

designed before orchard establishment. Bucks and Davis (1986) outlined a 

typical scheme for establishment of a drip irrigation system in the field (Fig. 

10-12). 

The shape and size of the 'wetted bulb' or area of soil moistened by the 

system, which will be filled with tree roots, depends on the type and size of 

emitter, the volume of water applied, and the structure and texture of the soil 

(Santos, 1997). 

A drip irrigation system should be used in areas where water is very limited, 

e.g., in arid zones (Rungsimanop et al., 1987; Singh 1992). This system has 

several advantages, such as decreasing the loss of water through evaporation, 

reducing weed infestation and the negative effects of increased salt 

concentration (Silva et al., 1996). An enormous disadvantage however, is the 

need for filters, which are sometimes obligatory because of poor water 

112


quality. The filters eliminate, or at least minimize, the possibility of system 

obstruction by suspended materials; without filters there is usually a 

considerable increase in cost of maintenance. 

Figure 10-12. A typical scheme for establishment of a drip irrigation 

system in the field. Adapted from Bucks and Davis (1986) 

Micro-sprinkler irrigation is also recommended for areas where water is 

limited. Like drip irrigation, this system allows more efficient water 

utilization. Micro-sprinkler irrigation is more appropriate for perennial crops, 

like annonas, which are established at low density per hectare and have large 

113


root systems. James (1988) comments that there are several types of microsprinkler 

emitters that are differentiated based on their water distribution 

patterns (Fig. 10-13). Annona growers in Brazil prefer the single microsprinklers 

with 300° of water distribution, or two micro-sprinklers with 180° 

each, so that they don't moisten the tree trunks and encourage infectious 

fungal or bacterial diseases. When selecting a micro-sprinkler system 

additional parameters must be considered, such as the emitter outlet and its 

sprinkling radius, as well as the intensity of water application through the 

radius (Santos, 1997). In general, micro-sprinklers have fewer problems of 

obstruction in the emitters than drip irrigation (Santos, 1997). 

Farré et al. (1999) conducted a study to compare the drip, micro-sprinkler 

and spray irrigation systems with respect to the area moistened under the 

canopy and the planting density. They concluded that with 313 cherimoya 

trees/ha (spacing of 8 x 4 m) 1,800-2,000 drippers/ha (6 drippers/tree) were 

frequently used. With the advent of low-density spray irrigation below the 

canopy, larger areas have been irrigated in Spain with better results. An 

irrigation schedule of 24 to 25 litres/tree and 6 drippers of 4 litres/h results in 

20% of the area moistened to more than 25 cm depth, while one microsprinkler 

of 25 litres/h wets 12 m², resulting in approximately 40% of 

moistened area. Unfortunately the study did not investigate the variation in 

the parameters with different soil types, which is very important in 

understanding the formation of the wetted bulb. 

114


Figure 10-13. Water distribution (degree of opening) of different microsprinklers 

that can be used in annona orchards. A) Type 360°; B) Type 

180°; C) Type 300°; D) Type 280°. Adapted from James (1988) 

Whatever the perennial crop, there is a need to establish a strategy to 

determine when and how much water must be delivered to the tree. To 

determine the need for water, the evaporation of water must be measured and 

then compared to a pre-established evapotranspiration coefficient of the crop 

in that region. The evapotranspiration coefficient is determined by using 

climatic variables, such as solar radiation, temperature, relative humidity 

(RH) and wind velocity (Pinto and Silva, 1994). A formula to calculate the 

irrigation requirement for sugar apple in Brazil's semi-arid northeast is based 

on the averages of air temperature and RH (Santos, 1997). The amount of 

115


water to be applied during each irrigation event depends on the amount of 

water per tree and number of trees per orchard unit. In addition, the time 

taken to apply the required amount of water depends on the number of 

emitters per tree and their outlet volumes. If the watering calculations suggest 

a time that is longer than 3 hours, it is recommended that the irrigation be 

split into 2 or 3 applications to avoid excessive wetness around the root 

system and loss of water due to percolation (Santos, 1997). Irrigation at night 

avoids evaporation losses. Growers should seek expert advice on developing 

any irrigation strategy. 

Water quality is as important as the amount of water and the time of its 

application. The presence of certain nutrients, such as calcium, can 

precipitate phosphates and provoke clogging of the emitters (Pinto and Silva, 

1994). The presence of sodium in irrigation water can cause concern, as this 

can result in salinization, especially in shallow soils without drainage 

systems. Needless to say, salinization is extremely detrimental to plant 

growth and fruit yield. Irrigation water containing NaCl, CaCl 2 

or CaC0 2 

causes depressed growth rate and reduced dry weight of all annona trees, 

especially above 3,000 ppm (Galila et al., 1991, cited by Mansour, 1997), 

leading to leaf burn and defoliation of the seedling leaves. High boron and 

chloride contents in irrigation water also promote phytotoxicity and injury to 

the leaves and fruits (Pinto and Silva, 1994). 

10.3.7 Pest and disease management 

Annona trees are attacked by a large number of insect pests and numerous 

diseases. Peña and Bennet (1995) described 296 species of insects associated 

with annonas, although many of them are not economically important. Nava- 

Díaz et al. (2000) reported that, world-wide, 106 insects, 91 fungi, 5 

nematodes, 2 bacteria and 1 virus have been recorded attacking Annona 

species. In Venezuela, Marin Acosta (1973) described 27 species of pests 

attacking annona trees. A full biological description of each pest, with 

management information, would be a very difficult task with limited 

applicability here, therefore only the major and regionally important 

secondary pests and diseases will be described below. Pest and disease 

control in commercial orchards has traditionally relied on the use of chemical 

products. With increasing awareness of the human and environmental 

hazards in pesticide use, alternative methods of control are suggested. In 

small scale plantings, the potential for rapid increase in pest or disease 

incidence is less, and the capability to monitor any outbreak is easier. 

116


10.3.7.1 Pests 

Major and minor insect pests attacking annonas are described by numerous 

authors (Melo et al., 1983; George et al., 1987; Bustillo and Peña, 1992; 

Oliveira et al., 1992; Torres and Sánchez, 1992; Agustín and Alviter, 1996; 

Junqueira et al., 1996; Pinto and Silva, 1994; Nakasone and Paull, 1998; 

Nava-Díaz et al., 2000) in many countries and ecological regions (Table 10- 

13) with different levels of economic damage and cost of management 

control. 

Table 10-13. Major and selected minor insect pests of Annona species 

Common Name Specific Name Affected Plant Country/Region 

Parts 

Major Insect Pests 

Annona moth Cerconota anonella Seeds and fruits Universal 

Cherimoya seed Talponia batesi Seeds and fruits Mexico, Peru, Spain 

borer 

Atis moth borer Anonaepestis bengalella Fruits Philippines 

Wasps (wasp) Bephratelloides 

maculicollis and 

B. cubensis 

Seeds and fruits Several 

countries/regions 

Borers (trunk 

borer) 

Flies (fruit flies) 

Mealy bugs 

Cratosomus bombina, 

Heilipus catagraphus 

Anastrepha obliqua, 

A. ludens, Ceratitis 

capitata, Bactrocera 

dorsalis and B. tryoni 

Planacoccus citri, 

Dysmicoccus spp., Ferrisia 

virgata, 

P. pacificus (India) 

Trunk and 

branches, base 

of the trunk 

Fruits 

Fruits and 

leaves 

Scale insects Several genera and species Leavesand 

stems 

Spider mites Several genera and species Leaves and 

flowers 

Minor insect pests 

Leaf hopper 

Empoasca fabae, 

Membracis foliata, 

Aethalion spp. 

Leaves and 

stems 

Thecla moths Oenomaus ortygnus Flowers and 

fruits 

Aphids 

Aphis gossypii and Leaves and 

Toxoptera aurantii shoots 

Hemipterous Leptoglossus zonatus and 

insects 

Antiteuchus tripterus 

Fruit spotting 

bug 

Mexico, Brazil 

Peru, Mexico. 

Colombia, Ecuador, 

Spain, Peru, Australia, 

New Zealand 

Several 


Several 


Several 


Amblypelta nitida Young fruits Australia 

Brazil, Colombia, 

Venezuela, American 

tropics 

American tropics, 

Brazil 

USA, Colombia, Peru, 

Brazil and Venezuela 

117


118 


Parts 

Leaf larvae Several genera and species Leaves and 

stems 

American tropics, 

Brazil 

Leaf miners Leucoptera spp. and Leaves 

Cuba and Ecuador 

Phyllocnistis spp. 

Root grubs Anomala Roots Philippines 

Ants ("saúvas") Atta spp. and Acromyrmex 

spp. 

Leaves and 

petioles 

Universal 

There are three important groups of borer insects attacking annona species: 

the trunk borer, the fruit borer and the seed borer. Trunk borers are 

coleopterons, generally weevils, and three species are the most common: 

Cratosomus bombina bombina (Plate 6), Euripages pennatus and Heillipus 

catagraphus (Oliveira et al., 1992). These insects are 2-4 cm long and they 

perforate the trunk, causing plant decline and finally death. The external 

symptom of the attack is a black oxidized sap exudation from the small holes 

where the insects entered the trunk. Injection of liquid pesticides, such as D- 

aletrina and D-tetrametrina into the trunk holes is an effective method of 

control (Oliveira et al., 2001). In the Brazilian Cerrados brushing of the tree 

trunk with a 1% solution of a pesticide, commercially named Creolina, 

together with lime at 10%, has prevented fruit borer attack on soursop. 

The annona moth (Cerconota annonella), commonly known as the 'fruit 

borer', is the most important of the insect pests attacking Annona species 

(Plate 5). Although it is known as the soursop moth in many areas, it has 

been recorded attacking and damaging fruits of several other annonas as well, 

including sugar apple and custard apple (Coronel, 1994; Oliveira et al., 1992; 

Torres and Sánchez, 1992), but has not been reported on cherimoya fruits 

(Bustillo and Peña, 1992). The soursop fruit borer moth has a life cycle from 

egg to adult emergence averaging 36 days. The adult moth is attracted to 

black-light traps, which is an important method for monitoring this insect 

pest (Bustillo and Peña, 1992). The removal of rotted and damaged fruits 

from the ground is also an important cultural control method. Bagging the 

fruits with chemically treated bags (a common type is the chlorpyriphos bag) 

can keep 92% of the fruits totally undamaged before harvesting (Bustillo and 

Peña, 1992). Biological control using two braconids, which parasitize larvae 

of C. annonella, has been successful in Colombia and Ecuador (Bustillo and 

Peña, 1992). In the Cerrado ecosystem of central Brazil, the soursop ecotype 

Morada is less susceptible to the attack of the soursop moth than any other 

ecotypes (Junqueira et al., 1996; Pinto and Silva, 1996), suggesting that 

appropriate cultivar selection can help to minimize the problem. Chemical 

control with triclorphon or fenthion at 0.1%, every 15 days can help to


control this pest. Spraying should be directed at the fruits and started when 

they are still small (Torres and Sánchez, 1992) 

The moth Anonaepestis bengalella is cited as the most destructive pest of 

sugar apple fruit in the Philippines (George and Nissen, 1992; Coronel, 

1994). Another moth Oemanus ortygnus, which is widespread throughout the 

Caribbean region and the American tropics is considered a minor pest 

(Nakasone and Paull, 1998) and attacks the flowers instead of the fruits. In 

addition to the natural control methods described above for the annona moth, 

the removal of damaged and attacked fruits from the ground or even from the 

plant, followed by burial in holes at least 50 cm deep, would be a very 

effective cultural practice. The same chemical control can also be used for 

both species. 

The soursop wasp (Bephratelloides maculicollis or B. cubensis, 

Hymenoptera), also called the annona seed borer (plate 5), is the second most 

important insect pest. Similarly Talponia batesi (Lepidoptera) also attacks 

cherimoya seed in Mexico (Nava-Díaz et al., 2000). All other cultural 

practices for control of Cerconota anonella can be used for Bephratelloides 

spp. and Talponia batesi, except for the use of black-light traps, which are 

ineffective with these species. Chemical control with decamethrin 0.05% 

every 15 days when the fruits are still small (Torres and Sánchez, 1992; 

Junqueira et al., 1996) can reduce infestation. 

Several genera of fruit fly, Anastrepha, Ceratitis and Bactrocera are 

frequently mentioned (George et al., 1987; Peña and Bennet, 1995; Rebollar- 

Alviter et al., 1997; Alvarez et al., 1999; Farré et al., 1999) as important 

insect pests attacking annona fruits in many countries and regions (Table 10- 

13), especially on cherimoya fruits. The infestation occurs with the 

deposition of the eggs by the adult on the fruit skin or through the stem 

cavity (George et al., 1987). By making galleries in the pulp, the larvae 

completely destroy the fruit. The larva starts its pupation phase outside the 

dropped fruit, underground (about 10 cm deep in the soil), from where the 

adults emerge and start a new cycle. The cycle of the fruit fly from egg to 

adult is completed in about 30 days (Nascimento et al., 2000). 

According to Farré et al. (1999), an incidence of fruit fly attack is generally 

due to favourable climatic conditions, high reproductive potential, alimentary 

adaptability and in some circumstances absence of natural enemies, which 

makes them a difficult pest to control. The size of the area under cultivation 

and economic importance of the crop can also add to the impact of an attack 

and the importance of the pest. In Spain, which is one of the most important 

119


cherimoya producers, the Mediterranean fruit fly (Ceratitis capitata) has a 

major economic impact, attacking up to 50% of cherimoya fruits (Farré et al., 

1999), while in Ecuador the incidence of cherimoya fruit fly (Anastrepha 

spp.) is greater than 94% (Alvarez et al., 1999). However, in Brazil, a 

typically tropical country with very small areas producing cherimoya, the 

incidence of fruit flies on this species, or even on its hybrid atemoya, is not 

mentioned in the literature (Kavati et al., 1997; Bonaventure, 1999). Also, 

the incidence of fruit flies on soursop, sugar apple and custard apple fruits is 

negligible or without economic importance, since there is no citation in the 

literature reviewed (Torres and Sánchez, 1992; Junqueira et al., 1996; Pinto 

and Silva, 1994; Kavati and Piza Jr., 1997). The only exception occurs in 

Mexico, where the attack of Anastrepha ludens on soursop is cited by 

Rebollar-Alviter (1987), describing the work of Ponce and Vidal (1981). It is 

not clear why fruit flies are only minor annona pests in this area, because the 

flies attack other fruit species in the same areas where annonas are grown. 

The mining character of fruit fly larvae, together with their underground 

pupation, has led practically all control methods to be directed at the adults, 

by using insecticides (Farré et al., 1999). The spraying of insecticide on 

entire plants is still the most common practice. Distribution of a toxic bait, 

consisting of 4% hydrolyzed protein and 0.15% dimethoate, on the entire tree 

or over the top third only, has also been used (Fuentes et al., 1999) to control 

Mediterranean fly (Ceratitis capitata) on cherimoya fruits. However, besides 

having questionable effectiveness, this method is also questioned by an 

increasingly environmentally sensitive society. 

An integrated control system involving chemical, biological and cultural 

methods should be implemented for management and control of the fruit fly. 

For instance, in Spain, Farré et al. (1999), describing work of Hermoso et al. 

(1994) in the Experimental Station of La Mayora, affirmed that phosphate 

baits or pheromone traps, combined with field hygiene (removal of fruits on 

the ground), can reduce fruit fly attacks to 4 - 7% without any insecticide 

treatment. This percentage of fruit fly attack can be further reduced (to 0.5 - 

2.0%) with immersion of cherimoya fruits in hot water (between 45 and 

47°C) for 60 minutes. Similarly, Rebollar-Alviter et al. (1997) suggest that 

control involving only the removal of dropped over-ripened fruit on the 

ground can reduce fruit fly populations by up to 80%. Fruit bagging also 

provides an adequate protection against attack (Nakasone and Paull, 1998). 

After the removal of the fruit, a cultural practice commonly used in mango 

orchards in north-eastern Brazil, harrowing beneath the canopy, i.e. turning 

over the first soil layer, impedes pupation of fruit flies and interrupts their life 

120


cycle. Given the importance of these pests in many areas, it is curious that 

more complete integrated pest management systems have not been reported. 

In Mexico, the insects Biosteres longicaudatus and Aceratoneuromyia indica 

have been used for biological control of Annona fruit flies (Rebollar-Alviter 

et al., 1997). The use of chemical compounds from seeds or leaves of 

Annona trees, such as wild soursop and sugar apple, have also been tested 

with some success against fruit flies. Extracts from an infusion of 5% (dry 

weight) in water of sugar apple leaves can kill up to 70% of adult fruit flies 

when ingested in the laboratory (Catarino and Ezequiel, 1999) - see Chapter 

6 on the chemical properties of Annona species. 

Mealy bugs, various species of scales or cochineals and spider mites attack 

the stems, leaves, flowers and fruits of Annona species in numerous countries 

and regions (Table 10-13). They are all considered to be sucking insect pests 

and may be considered as economically important, due to their impact when 

they suck the sap of the young vegetative parts and fruits of annona trees. An 

orchard monitoring system is necessary to detect the phenological phase in 

which the attack is occurring at an economic level, so as to make control 

more effective. Mealy bugs are reported to be a major pest on marketable 

fruits in some areas of Australia and red spider mites can become a serious 

problem when attacking growing annona trees in dry areas (Nakasone and 

Paull, 1998). In Colombian and Brazilian soursop orchards, these insect pests 

have been controlled with mineral oil and systemic insecticides (dimethoate) 

(Torres and Sánchez, 1992; Junqueira et al., 1996). It is recommended that 

mites be controlled by spraying specific insecticides (propargite) which can 

kill the eggs and the adult insects. The insecticide and water mixture needs to 

be used with an adherent. The mealy bug Planococcus citri is biologically 

controlled by its predator Cryptolaemus montrouzieri, however, the 

predator's action is hindered when there is a large population of the 

Argentinean ant, Iridomyrmex humilis (Farré et al., 1999). Control of mealy 

bug should focus on biological control where possible, or the use of mineral 

oil. 

Some minor insect pests also have significant economic importance in some 

regions. Aphids can transmit serious viral diseases to annona trees, while 

attacks of aphids and hemipterous bugs on fruits can promote irreversible 

damage to their quality for market. These insect pests are controlled by 

spraying with the pesticides malathion or parathion (Torres and Sánchez, 

1992; Junqueira et al., 1996). Aphids and bugs attacking soursop trees have 

been controlled efficiently in the Brazilian Cerrados by spraying a 20% 

121


solution of macerated leaves of the Neem (Azadirachta indica) tree. The 

fruit-spotting bug (Amblyphelta nitida) is considered a serious annona pest in 

Australia and its damage resembles the symptoms of black canker or diplodia 

rot (Nakasone and Paull, 1998). Where possible, if damage is limited, 

chemical use should be avoided. 

10.3.7.2 Diseases 

Major and minor diseases of annona trees have frequently been described 

(Table 10-14) (George et al., 1987; Junqueira et al., 1996; Pinto and Silva, 

1996; Kavati et al., 1997; Rebollar-Alviter et al., 1997; Nakasone and Paull, 

1998; SPT-TCA, 1999; Nava and Díaz, 2000). The intensity of the damage 

and the control methods differ in a number of ways, according to country and 

region where the attack occurred. 

The most important root diseases caused by fungi are damping-off 

(Rhizoctonia solani and Fusarium spp.) and black root rot (Phytophthora 

spp., Cylindrocladium clavatum and Sclerotium rolfsii), whose attacks occur 

mainly on nursery seedlings, but also occasionally on adult plants (Melo et 

al., 1983; Junqueira et al., 1996). Although these diseases are caused by 

different species, heavy clay soils and high relative humidity (RH) are the 

main contributing factors for attacks. In addition, the symptoms and damage 

(wilting and death of seedlings and adult plants) are similar (Torres and 

Sánchez, 1992; Agustín and Alviter, 1996; Junqueira et al., 1996). To control 

these diseases, the soil can be treated with a solarization system (see topic 

10-1.1 and Plate 2). The use of resistant rootstocks, such as custard apple, 

appears to be an adequate form of management to avoid attack of these fungi 

(Kavati et al., 1997), but is not yet widely used. 

The base of seedlings or adult plants can be sprayed with a fungicide solution 

of benomyl 0.1% (Junqueira et al., 1996) if necessary. Seedlings can also be 

drenched with 0.1% Bavistin at 10-12 day intervals (Singh, 1992). 

Table 10-14. Major and selected minor diseases of Annona species 

Common Name Specific Name Affected Plant 

Parts 

Major Diseases 

Damping-off, Rhizoctonia solani Root and base of 

Black rot (Thanatephorus 

trunk 

cucumeris), 

Phytophthora spp., 

Cylindrocladium 

clavatum, Sclerotium 

rolfsii (Athelia rolfsii) 

Country/Region 

Universal 

122



Parts 

Seedling blight Pithyium spp. Seedlings Universal 

Bacterial wilt Ralstonia solanacearum Roots and canopy Australia and Brazil 

Anthracnose Colletotrichum 

gloeosporioides 

(Glomerella cingulata) 

Leaves, young stems 

and fruits 

Universal 

Black canker 

and Diplodia rot 

Phomopsis spp., 

P. anonacearum, 

Botryodiplodia 

theobromae 

Leaf scorch, twigs 

and fruits, fruits in 

storage 

Universal 

Purple blotch Phytophthora palmivora Fruits Seveal countries 

Brown rot and 

Peduncles and fruit 

fruit rots 

Rhizopus stolonifer, 

Gliocladium roseum, 

Phytophthora spp. 

Brazil, India and 

American countries, 

Universal 

Minor diseases 

Burn of string Corticium koleroga Leaves and twigs Amazon region 

Zoned spot Sclerotium coffeicolum Leaves Amazon region 

Blight Phoma spp. Leaves, stems and Mexico 

twigs 

Black scab Fusarium spp. Trunk, branches and Mexico 

twigs 

Fumagina Stigmella spp. Leaves, stems and Universal 

twigs 

Rust fungus Phakopsora cherimoliae Leaves USA (Florida) 

Rubelose Corticum salmonicolor Branches and twigs Brazil 

Leaf spot Cercospora anonae Leaves Brazil 

Armillaria root Armillaria luteobubalina Roots, base of tree, Ausralia 

rot 

decline 

Nematodes Helicotylenchus spp. and 

Meloidogyne spp. 

Roots 

Universal 

Bacterial wilt is an important root disease, which was responsible for 70% of 

the deaths of atemoya trees established on sugar apple rootstocks in Australia 

(Nakasone and Paull, 1998). This disease is caused by the bacterium 

Ralstonia solanacearum and is manifested by rapid wilting and the death of 

young trees. Collar rots, dark internal discolouration of the root wood tissue, 

tree decline and eventual death are the symptoms on adult trees (George et 

al., 1987). Some cherimoya cultivars are recommended as resistant 

rootstocks, such as ‘White’ in California and ‘Negrito’ and ‘Cristalino’ in 

Spain (George et al., 1987). There is no chemical control for this disease. 

There are several diseases attacking the fruits of Annona species during the 

preharvest and post-harvest phases: anthracnose, black canker, diplodia rot, 

purple blotch and brown rot (Rao et al., 1962; Junqueira et al., 1996; Pinto 

and Silva, 1996; Rebollar-Alviter et al., 1997; Nakasone and Paull, 1998). 

Fruit rot caused by Phytophthora is prevalent on cherimoya, as well as 

123


soursop, custard apple, sugar apple and on related species, e.g., A. 

diversifolia (Weber, 1973). 

Anthracnose (Colletotrichum gloeosporioides) is a cosmopolitan disease 

attacking all annonas. This disease is responsible for 90% of the preharvest 

loss of soursop fruits in Bahia, Brazil (Nieto-Angel, 1999). In Bahia, 

predisposing climatic conditions for anthracnose attacks are highly 

favourable, due to high rainfall and atmospheric humidity, and during wet 

seasons in dry areas (Dhingra et al., 1980). In Mexico, the incidence of 

anthracnose in cherimoya varies from 50 to 70%, although mainly in 

orchards without adequate control (Rebollar-Alviter et al., 1997). This 

disease causes twig dieback, defoliation and dropping of flowers and fruit, 

while in mature fruit its infection causes black lesions (Nakasone and Paull, 

1998). Management and control of anthracnose involves a thorough cleanup 

at the end of the dry season, including the pruning of infected twigs, removal 

of rotting fruits on the ground, and then burning of all waste. Spraying with 

the fungicide benomyl 0.06%, intercalated with copper oxychloride 0.15%, 

every week during the rainy season and every three weeks during the dry 

season, gives adequate control (Junqueira et al., 1996). In India, Singh (1992) 

recommended spraying with 0.05% benomyl or 0.2% mancozeb M43 at 15- 

20 day intervals. 

Black canker (Phomopsis spp.), diplodia rot (Botryodiplodia theobromae) 

and purple blotch (Phytophthora palmivora) are fungal diseases attacking the 

fruits (George and Nissen, 1980; Agustín and Alviter, 1996; Nakasone and 

Paull, 1998). Black canker and diplodia rot occur mainly in neglected 

orchards; they show similar symptoms of purplish to black spots or blotches 

confined to the surface of the fruit. Diplodia rot is distinguished by its darker 

internal discolouration and the extensive corky rotting it produces. Diplodia 

rot has also been described by Junqueira et al. (1996) as attacking the 

junctions between rootstocks and scions of soursop, ultimately killing the 

plant. Purple blotch is distinguished by small spots on immature fruits that 

expand until most of the fruit surface is affected (George and Nissen, 1980). 

There are several management systems to control these diseases, most of 

which should be applied preventively: a) pruning the low branches to avoid 

high humidity under the canopy and brushing with 1% paste of copper 

oxychloride; b) keeping the plants in a good nutritional state; c) avoiding 

physical damage to the fruits, as well as keeping adequate control of fruit and 

seed borers; d) brushing the graft junction with 4% copper oxychloride paste; 

e) spraying with benomyl 0.2%, every 15-20 days during the rainy season 

(Junqueira et al., 1996; Kavati et al., 1997; SPT-TCA, 1999). 

124


Brown rot (Rhizopus stolonifer) is another serious disease which attacks the 

fruits, generally at harvest and during post-harvest periods (Pinto and Silva, 

1996; Rebollar-Alviter et al., 1997) - (Plate 7). The main contributing factors 

for a higher incidence of brown rot are high RH in the orchard and some kind 

of physical damage to the fruit (Torres and Sánchez, 1992). Perforations by 

wasps (soursop seed borers) on the fruit peduncles are probably one of the 

entry points for fungal establishment, the attack of which promotes a brown 

rot and later mummification of the pulp (Torres and Sánchez, 1992; SPT- 

TCA, 1999). A preventive control measure is the elimination of seed borer 

attacks or other physical damage of fruits, as well as removal of damaged 

fruits on the ground (Torres and Sánchez, 1992). 

Like many other tropical fruits, post-harvest rotting is largely responsible for 

the short shelf life of annona fruits and experimental studies have evaluated 

possible solutions to this problem. George et al. (1987) recommended 

dipping annona fruits in either a heated benomyl suspension with 0.5 g a.i./l 

(or the same concentration guazatine solution) at 50° to 52°C for 5 minutes 

for post-harvest control of rots. Dipping the annona fruits in unheated 

prochloraz (0.125 g a.i. per litre) for 1 minute at 25°C also provided a good 

control of post-harvest rotting. However, some treatments, particularly 

prochloraz, can induce skin injury at high concentrations, and should be 

avoided. 

Minor diseases can also be important in some countries or regions (Table 

10.14). Some fungi causing minor diseases develop and attack more severely 

under high relative humidity and hot temperatures. Two soursop diseases 

commonly called 'burning string' (Corticium koleroga) and 'zoned spot' 

(Sclerotium coffeicolum) are examples of these kinds of fungi. In Mexico, 

blight and black scab are also important diseases and the former can even 

show incidences of up to 80% on cherimoya leaves in the high humidity 

period of the year (Nava-Díaz et al., 2000). Fumagina (Stigmella spp.) is a 

cosmopolitan fungus whose attack is aided by certain species of ants, 

although this disease is described as having one of the lowest incidences 

(28.5%) in Mexico (Nava-Díaz et al., 2000). Rubelose (Corticum 

salmonicolor) and cercosporiose (Cercospora annonae) are common 

diseases on twigs and leaves of soursop, although predisposing 

environmental conditions and symptoms are different. Symptoms of rubelose 

are yellowish pink mycelium on the trunk followed by exudation of latex, 

whereas cercosporiose symptoms are distinguished by development of black 

circular lesions on the leaves. The former disease occurs under the high 

humidity and hot temperatures occurring in the Amazon region and in some 

125


north-eastern states of Brazil, while cercosporiose attacks under the lower 

temperatures and dry conditions of Brazil's central region. Spraying with 

copper oxychloride gives adequate control of these diseases (Junqueira et al., 

1996). 

Nematodes of different species, such as Helicotylenchus spp., Meloidogyne 

incognita, Macroposthonia spp., Tylenchorhynchus phaseoli and Xiphinema 

spp. have been described as attacking cherimoya, soursop and sugar apple 

(Sharma, 1973; Sharma, 1977; Monteiro et al., 1978; Ferraz et al., 1989). In 

Brazil, the disease called soursop decline has been associated with the attack 

of nematodes of the Gracilacus species in Ceará State, north-eastern region, 

and in Brasilia (Sharma et al., 1985; Freire and Cardoso, 1997). 

10.3.8 Physiological disorders 

Annona fruits occasionally present abnormalities that are not due to diseases 

or insect pests, but are due to physiological disorders. George et al. (1987) 

described some of them as follows: 

Fruit splitting - probably caused by sudden changes in fruit moisture content 

or temperature, and some species and varieties appear to be less susceptible 

than others. 

Russeting - superficial russeting of the fruit skin. The combination of low 

night temperatures (lower than 13°C) accompanied by low humidity is the 

probable main cause. Nearly mature fruits are more susceptible, and 

cherimoya is less susceptible than sugar apple and their hybrid atemoya. 

Crocodile skin - Fruits show wavy and pointed carpels. Extremely vigorous 

plants show more severe symptoms. No hypotheses of cause have been 

advanced. 

Hard seed casing and brown lumps - There is a suspicion that boron 

deficiency or sudden changes in fruit water content may be the main causes 

of this physiological disorder. In north-eastern Brazil, pulp of sugar apple 

fruit commonly shows hard seed casing disorder when trees are cultivated in 

dry areas without irrigation. 

126

Chapter 11. Harvest, Postharvest 

and Processing 

S. R. M. de Andrade, R. E. Alves, H. A. C. Filgueiras and 


11.1 Introduction 

Cherimoya has been described as the most delicious of the annonas, and as 

the finest dessert fruit in the world, because of its unique flavour (George et 

al., 1987). Soursop produces the largest fruit among the annonas, and 

presents the highest potential for processing (Nakasone and Paull, 1998). The 

pulp flavour of custard apple fruit is inferior to that of cherimoya and 

soursop, and, although some trees produce excellent fruits, they are generally 

not suitable for commercial cultivation (Salunkhe and Desai, 1984; Bora et 

al., 1987; Nakasone and Paull, 1998). Sugar apple is the most widely 

distributed of the Annona species, and is sweeter than soursop, with a high 

soluble solid content (24° Brix) and a low acidity (0.58%) (Rego et al., 

1979), although the acidity recorded by Singh (1992) is lower still (0.19 - 

0.24% depending on cultivar). It is common to find sugar apple in local 

markets, although due to the small size of fruit, large number of seeds and 

poor shelf life, they are seldom cultivated in large commercial orchards 

(Coronel, 1994; Nakasone and Paull, 1998). Wild soursop does not have 

commercial value at this time, but could be transformed by careful strategic 

planning, resources and especially germplasm selection. 

In spite of the potential, the annonas have some limitations, principally 

regarding their resistance to transport to distant markets, because they ripen 

rapidly after harvest and this limits their shelf life. Attempts to enhance the 

post-harvest life of these fruits must take into account ripening physiology, 

physical-chemical aspects of fruit quality, and harvest and post-harvest 

handling (Alves et al., 1997). 

127

Chapter 11. Harvest & Processing 

11.2 Harvest 

Venkataratnam (1959) reported that Annona species usually start flowering 4 

to 5 years after planting, but there is considerable variation; in Latin 

America, soursop usually bears fruit in the third year and cherimoya in the 

third to fourth year. There can be two distinct vegetative flushes, according to 

the season. Therefore, harvest time, also called harvest point, based on 

anthesis is impracticable, because flowering can occur during many months. 

On the other hand, some cultural practices, such as pruning or timing of hand 

pollination, can alter the time of harvesting, as can the use of appropriate 

cultivars. 

Table 11-1. Harvesting season of the four major Annona species in 

different countries and regions 

Countries Cherimoya Custard Soursop Sugar Source 

Apple 

Apple 

Argentina Feb-Jul - - - Nakasone and Paull, 

1998 

Brazil - - Jan-Mar - Lucas, 1994 

Amazonas - - - - Accorsi and Manica, 

1994 

Rio de - - - Dec-May Carvalho et al., 2000 

Janeiro 

São Paulo - - - Feb-Jun TCA, undated 

Pernambuco - - Jul-Sep Jan-Aug - 

Pará - - - - - 

Caribbean - Feb-Apr Yearround 

Jun-Sep Nakasone and Paull, 

1998 

Chile Aug-Dec - - - Nakasone and Paull, 

1998 

China-Taiwan - - - Sep-Oct Tsay and Wu, 1990 

- - - Jul-Sep Nakasone and Paull, 

1998 

- - - Oct-Mar - 

Colombia 

Andes Valley - - Mar-Jun - Torres and Sánchez, 

1992 

Altantic 

Coast 

- - Oct-Dec - Torres and Sánchez, 

1992 

Costa Rica - - Yearround 

- - 

Dominican - - Feb-Mar, - FDA, undated 

Republic 

Jul-Aug 

India 

Sangareddy Nov-Jan Mar-May Apr-Jun Oct-Dec Venkataratnam, 1959 

Poona Nov-Feb - - Aug-Nov Nakasone and Paull, 

128


Countries Cherimoya Custard 

Apple 

Soursop Sugar 

Apple 

Source 

1998 

Indonesia - - Yearround 

- Nakasone and Paull, 

1998 

Mexico - - Jun-Sep - Nakasone and Paull, 

1998 

Peru - - Dec-Mar Nov-Aug Alvarez et al., 1999 

Philippines - - Jun-Aug Jul-Sep Nakasone and Paull, 

1998; Coronel, 1994 

Portugal - - - - Nunes, 1997 

Madeira Nov-Feb - - Oct-Jul Nakasone and Paull, 

1998 

South Africa 

(Natal) - - - - FAO, 1988 

Spain - - - Nov-Feb Accorsi and Manica, 

1994 

Tanzania 

(West) - - - - FAO, 1988 

(East) - - - - 

USA 

California Mar-Apr - - - Nakasone and Paull, 

1998 

Florida Jul-Oct Feb-Apr Jun-Nov Jul-Sep 

Puerto Rico - - Mar-Sep - 

Hawaii - - Jan-Oct - 

Zambia - - - - FAO, 1988 

Zimbabwe - - - - FAO, 1988 

In Taiwan, pruning of sugar apple in January and February leads to harvest 

from July to September, while pruning between June and October leads to 

harvest from October to March (Table 11-1) (Nakasone and Paull, 1998). 

Cherimoya fruit quality attributes, such as total soluble solids (TSS) and 

sugar content, change depending on the pollination time (Nomura et al., 

1997). Temperature has a primary effect on the ripening of annona fruit, with 

low temperature delaying fruit maturation, and high temperatures providing 

premature ripening on the tree, causing fruit fermentation and fruit fall 

(George, 1984; George et al., 1987; Nakasone and Paull, 1998). The harvest 

season of the commercial Annona species differs among countries and 

regions (Table 11-1). 

In Colombia, the soursop harvest occurs in two seasons, according to region 

(Torres and Sánchez, 1992). In the Andean valleys and the lower coffee 

region (altitude 500 to 1250 m), the major production zones; harvesting 

129


occurs between March and June and from October to December, with both 

seasons producing high quality fruits. Along the Atlantic coast, below 500 m, 

harvesting occurs between August and October, and the fruits have a very 

low quality because of the poor genetic resources and the high temperatures 

of the region (Torres and Sánchez, 1992). In Hawaii, soursop production 

occurs during most of the year, with two peaks from January to April and 

from May to August (Nakasone and Paull, 1998). In Brazil, production 

occurs year round, but in some regions, depending on the temperature and 

precipitation, the production shows harvesting peaks. 

All annonas are characterized as climacteric ripening fruits, so fruits are 

harvested when they reach physiological maturity and are still firm, full 

ripening occurs after the climacteric peak. Fruits harvested prematurely will 

soften but have poor quality (Accorsi and Manica, 1994; Coronel, 1994; 

Nakasone and Paull, 1998). The time of harvesting is determined by the fruit 

skin colour, which changes with the proximity of physiological maturity. At 

harvesting time, soursop fruit skin changes from dark green to slightly 

yellowish-green, while the cherimoya and sugar apple fruits change from 

greyish green to yellowish-green, but in all cases their pulp should be firm 


A skin colour index to guide the harvest depends on the location of the 

market. For local markets, fruits must be harvested when mature, with 20 to 

40% yellowish-green skin, and they will ripen in 4 to 6 days; for export 

markets, 10 to 20% yellowish-green skin is satisfactory, as this will provide 

slightly more time before ripening without the loss of quality. When the 

fruits are harvested with more than 75% yellowish-green skin, they will ripen 

in 1 to 3 days, while fruits harvested at less than 5% do not ripen completely 

at all (George et al., 1987). 

Fruit maturation within a plant or orchard is not synchronized, so the harvest 

season can last for 3 to 6 months. Hence, each tree must be inspected 

regularly to collect the fruits at the appropriate harvest point. The most 

suitable time of day to harvest is in the morning just after the evaporation of 

the dew, when the fruits are dry and fungal rot contamination is less likely 

(Accorsi and Manica, 1994). 

Cherimoya fruits are generally harvested when the skin colour changes from 

greyish green to yellow-green, although some cultivars will change to almost 

brown (Accorsi and Manica, 1994). However, sometimes the change of skin 

colour is not very pronounced. Consequently, colour change, pollination time 

and fruit size are not reliable harvest indices. Therefore, the harvest index 

130


needs to be improved for cherimoya to ensure better fruit consistency, 

flavour and quality (Palma et al., 1993; Alves et al., 1997; Nomura et al., 

1997; Nakasone and Paull, 1998). 

Soursop fruits ripen very quickly on the tree. As a consequence, they require 

frequent visits to the orchard during the harvest season. Maturation is 

identified when there is a loss in shine and the skin colour changes from dark 

green to light green (Salunkhe and Desai, 1984; Torres and Sánchez, 1992; 

Accorsi and Manica, 1994). The carpel units spread apart when the fruits are 

mature. In Colombia, growers and wholesalers press the fruit with their 

thumbs to check the fruit maturity (Torres and Sánchez, 1992). It is not 

recommended to leave soursop fruits ripening on the tree, because they fall 

and lose market quality. However, if they are harvested before physiological 

maturity, the fruits do not ripen well and the pulp may become bitter (Torres 

and Sánchez, 1992). 

Sugar apple fruits are considered to be mature and reach their harvesting 

point when the skin changes colour and when the segments spread far apart, 

exposing a creamy yellow skin (Salunkhe and Desai, 1984). At this point 

they have reached their 'consumption point' (Plate 8). They mature at 

irregular intervals over a period of 3 months, so that picking every other day 

or so is obligatory. Premature harvesting can promote poor fruit quality and 

fruits left to ripen on the tree are often eaten by birds and bats, and when 

over-mature they have a tendency to break and decay (Salunkhe and Desai, 

1984; Coronel, 1994; Lucas, 1994; Mosca et al., 1997a). 

Annona fruits must be hand-harvested and put into cushioned boxes or 

baskets to avoid mechanical damage or bruising (Nakasone and Paull, 1998). 

The boxes must remain in the shade and be protected from rain, wind and 

dust (Accorsi and Manica, 1994). The fruits may be cut from the branch with 

pruning scissors, leaving 0.5 to 1 cm of the peduncle to avoid loss in weight 

and fungal diseases (Accorsi and Manica, 1994; Alves et al., 1997). If the 

fruits are pulled from the branch, the floral cushion can be damaged, 

reducing the next harvest. The wounds can also become entry points for 

rotting pathogens (Calzavara and Müller, 1987; Torres and Sánchez, 1992; 

Mosca et al., 1997 b). 

Depending on tree size, some species, such as sugar apple or soursop, are 

harvested by climbing using a ladder, or with a pole with a hook and a basket 

at its end (Torres and Sánchez, 1992; Coronel, 1994). Soursop harvest is 

more difficult and time-consuming than other annonas because the trees are 

usually taller and the fruits are larger (Nakasone and Paull, 1998). Nakasone 

131


and Paull (1998) suggested mechanical harvesting in larger soursop orchards. 

Cherimoya fruits are hand harvested by cutting the peduncle and by using net 

bags to hold or catch the fruit (Accorsi and Manica, 1994; Agustín and 

Angel, 1997). 

11.3 Postharvest handling 

11.3.1 Physiological changes 

Annona fruits have a respiration peak and an increase in ethylene 

concentration after fruit harvest; this is typical of climacteric species. 

Cherimoya, soursop and sugar apple fruits present two successive rises in 

respiration rate, whereas the custard apple presents only one (Brown et al., 

1988). The ripening process occurs during climacteric respiration, with some 

modifications in the chemical composition leading to remarkable changes in 

flavour and a decrease in pulp firmness (Mosca et al., 1997 a). Knowledge 

about this process is very important for post-harvest handling, because 

ripening occurs very quickly after harvest (Torres and Sánchez, 1992). 

Cherimoya presents a climacteric peak 5 days after the harvest point and a 

second one after 10 days, when the fruits soften, and the flavour and aroma 

development are completed (Kosiyachinda and Young, 1975). In the cultivar 

‘Fino de Jete’, Martinez et al. (1993) demonstrated a temporal coincidence 

between ethylene production and physical-chemical alterations. During the 

ripening process at 20°C, the pH dropped to 4.8, total titratable acids 

increased to 0.36 g citric acid/100 g fresh weight, starch content declined to 

20.7 g/100 g fresh weight and Brix increased to 18.7°. 

In soursop fruits, the climacteric peak corresponds to an increase in soluble 

solids content, the pH value decreases and titratable acids rise about 10 fold, 

due to increases in malic and citric acid concentrations (Paull et al., 1983). 

Maximum production of volatile compounds and ethylene diffusion occurs 5 

days after the harvest point. At this time, the highest concentrations of sugars 

and acids are attained; this is the moment of best quality for consumption. 

Fructose and glucose reach their peaks 5 days after the harvest point, while 

sucrose content rises to a maximum concentration 3 days after harvest point 

and then declines. Fructose exceeds sucrose concentration and contributes to 

the sweetness of the fruit (Paull et al., 1983). Starch breakdown by amylases, 

polygalacturonase and cellulase activities increase during ripening 2 days 

after the harvest point (Paull et al., 1983). These changes are ethylene 

132


independent and probably started at fruit detachment (Bruinsma and Paull, 

1984; Paull, 1990). After the climacteric peak, volatile compounds are 

released, sugar and organic acid concentrations decrease, and there is a loss 

of fruit quality. The degree of skin darkening is a useful marker for these 

stages (Paull et al., 1983). The best time for soursop consumption is 6 to 7 

days after harvesting (Paull, 1982). 

Sugar apple fruits soften during their second ethylene peak. The physicalchemical 

properties of sugar apple change very quickly after this peak, and 

abscissic acid increases dramatically and may have a role in fruit ripening 

(Tsay and Wu, 1990). Sugar apple fruits reach physiological maturity 15 to 

17 weeks after pollination, when soluble solids and titratable acids increase 

(Pal and Kumar, 1995; Mosca et al., 1997 a). Mature sugar apple fruits, at 

ambient temperature (28 ± 3°C), ripen 2 to 5 days after the harvest point. 

Ripening is completed and fruits should be consumed when softening is 

apparent, and also when the soluble solids content reaches 28 o Brix and 

titratable acids fall to 0.3% (Pal and Kumar, 1995; Mosca et al., 1997 a). 

11.3.2 Handling 

Annona fruits usually ripen 3 to 7 days after harvesting, thus becoming soft 

and easily injured. Careful, appropriate handling and transportation of fruit is 

necessary to avoid skin bruising. The fruits are very delicate, so one layer of 

fruits per box is recommended for storage and for shipment. If 2 or 3 fruit 

layers are used, fruits must be protected with soft cushioning between them 

(Calzavara and Müller, 1987; FDA, undated). A single layer in trays 

containing 6 to 8 kg of fruits is best (George et al., 1987; Accorsi and 

Manica, 1994). As long as the fruits stay firm it is possible to transport them 

to distant markets, but they should be wrapped individually with soft 

materials, such as paper bags or polystyrene gloves (Salunkhe and Desai, 

1984; Coronel, 1994; Lucas, 1994). A pre-cooling treatment prior to 

shipment improves the post-harvest life (George, 1984), except for sugar 

apple (Singh, 1992). 

Aseptic treatment of tools and containers helps to prevent post-harvest 

infections from pests and diseases. These routines include the immersion of 

pruning scissors in fungicide solution (benomyl 1 g a.i./l) after every fruit 

harvested, to avoid transmission of fungal diseases, mainly Lasiodiplodia 

theobromae (Alves et al., 1997; Mororó et al., 1997). 

133


To control fungal rots, George et al. (1987) suggested dipping the fruits in 

benomyl suspension (0.5 g a.i./L) or guazatine solution (0.5 g a.i./L) at 50 to 

52°C for 5 minutes. Prochloraz solution (0.125 g a.i./L) for 1 minute at 25°C 

also gives good control. Immersion treatments longer than 5 minutes induce 

skin injury, due to the chemical concentrations and interactions. Seemingly, 

anthracnose (Colletotrichum gloesosporoides var. minor), Phomopsis 

annonacearum and Rhizopus stolonifer are also avoided with fungicidal 

treatment, but more studies with these diseases are necessary (George et al., 

1987). A specific treatment for sugar apple consists of rinsing in chlorinated 

water (100 ppm) at 10 to 12°C for 20 minutes, followed by 10 minutes in less 

chlorinated water (20 ppm). To remove chlorine residues, the rinse is 

followed by immersion or aspersion with water containing 2 ppm of chlorine 

and a fungicide (Alves et al., 1997). For transport to distant markets, Babu et 

al. (1990) suggested the immersion of sugar apple fruits in 500 ppm of 

bevestin and placing them in polyethylene bags containing potassium 

permanganate. However, it must be pointed out that no chemical treatment 

can be a general recommendation, as each country has its own regulations 

about the chemicals allowed for each fruit species. Therefore the country's 

agricultural legislation must be clearly understood before the use of any 

product reported in the literature. 

In case of doubt and considering the softness and sensitivity of annona fruits 

to mechanical damage, very careful handling in non-contaminated 

environments can avoid the use of chemical treatments. Gentle cleaning with 

compressed air and wrapping the fruit in paper or plastic film impregnated 

with potassium permanganate are good recommendations for carefully 

handled fruits (Alves et al., 2001). 

11.4 Storage 

Annona fruits are very perishable and have a short post-harvest life; therefore 

they require efficient storage techniques (Coronel, 1994). Optimal storage 

conditions are 15 to 16°C with high relative humidity (RH) and even then 

storage should not exceed about 2 weeks (George et al., 1987). 

Some cherimoya cultivars can be held for 7 to 10 days at 17°C, however, 

normal ripening usually occurs between 15 to 30°C. Fuster and Prestamo 

(1980) suggested 10°C and 85% RH as the best storage conditions for the 

cultivars ‘Campas’ and ‘Fino de Jete’, and the optimal storage time ranges 

from 15 to 21 days. Cherimoya fruits stored at 20°C presented a rapid 

134


decrease in fruit firmness (Fuster and Prestamo, 1980; Lahoz et al., 1993). 

Plaza et al. (1993) suggested storage of the ‘Fino de Jete’ cultivar in 

polyethylene bags at 8.5°C and 98% RH with 3.5 g KMnO4/kg of fruits. Due 

to the difference in temperature sensitivity among cultivars, a general 

recommendation for the storage of cherimoya without risk of chilling damage 

would be 10°C. The use of plastic bags or plastic wrapping film reduces 

water loss due to low relative humidity, and the use of potassium 

permanganate impregnated in the plastic or in sachet inside the pack delays 

softening. 

Mature soursop stored at 22°C and 40-50% RH reaches a climacteric peak in 

4 to 6 days, and is ready to consume 2 days later (Paull et al., 1983; Mosca et 

al., 1997 b). However, under low temperatures the fruits do not ripen 

properly (Livera and Guerra, 1994). Fresh soursop fruits harvested and stored 

at room temperature for 4 to 7 days, will reach optimum quality for 

processing 5 to 6 days after softening begins. Fruits must be stored on racks 

in the shade and inspected daily, by testing the softness with thumb pressure 

(Nakasone and Paull, 1998). Based on current knowledge, the best 

recommendation for the storage of soursop is 15°C and 90% RH. 

Physiologically mature sugar apples stored at 13°C for 12 days, and then 

transferred to room temperature (27.5°C) ripen within 4 days, while at 20°C 

they ripen and soften completely within 6 days; at room temperature ripening 

takes 2 to 4 days (Tsay and Wu, 1989; Coronel, 1994). The ripe fruits may be 

stored for 5 days at 5°C; if they are held 5 to 6 weeks at 4.4°C, the pulp 

remains in good condition, but the skin becomes brown and unattractive 

(Accorsi and Manica, 1994). Ripe fruits can be kept at room temperature for 

one day only (Salunkhe and Desai, 1984; Coronel, 1994), while immature 

fruits stored below 15°C develop chilling injury, resulting in an unpleasant 

appearance (Salunkhe and Desai, 1984; Tsay and Wu, 1989). Broughton and 

Tan (1979) reported that high temperatures (above 20°C) and a dry 

atmosphere accelerated the ripening process, and suggested that the optimal 

conditions to extend sugar apple storage life are temperatures of 15 to 20°C 

and RH of 85 to 90% (Broughton and Tan, 1979; Tsay and Wu, 1990; 

Nakasone and Paull, 1998). Alternatively they can be stored at 15 to 20°C 

with low oxygen and ethylene tensions, combined with 10% CO 2 

and 85 to 

90% RH (Broughton and Tan, 1979). Due to the sugar apple's chilling 

sensitivity, the safest recommended storage temperature is 15°C, with 

relative humidity around 90%. 

135


Sugar apple, as well as cherimoya will benefit from the use of plastic packing 

film and potassium permanganate, since it acts only on the external ethylene 

gas expelled by the fruit during the ripening process by inhibiting this gas 

and prolonging post-harvest life of climacteric fruit (Pinto, 1978). Another 

advantage of potassium permanganate is the possibility of decreasing fruit 

injury by maintaining firmness, as well as being safer for consumption 

compared with other post-harvest preservation products, which may cause 

consumer concerns about allergic problems. 

11.5 Processing 

Annona fruits are mainly consumed as fresh fruit, however, some of them, 

such as soursop, sugar apple and atemoya, can be processed and used in the 

preparation of nectars, drinks, sherbets, ice cream, syrup and cakes. Soursop 

is the most suitable for processing, not only because of its high sugar content 

and delicate, aromatic flavour, but also because its pulp does not oxidize like 

that of sugar apple and cherimoya (George, 1984; Villachica et al., 1996; 

Mororó et al., 1997). 

Figure 11-1. Sketch of a mechanized system to process fruit pulp: A) 

rinse/immersion; B) rinse/aspersion; C) selection; D) endless spiral 

system; E) storage tank; F) pulp extractor; G) pulp storage tank; H) 

pump; I) seal bag 

Adapted from Mororó et al., 1997. 

Before processing, mature soursop fruits should be stored on racks in the 

shade and inspected daily. As the fruits ripen, they should be removed and 

processed for pulp extraction. For hand pulp extraction to produce soursop 

concentrate, Torres and Sánchez (1992) suggested the following steps: 1) 

fruit selection; 2) fresh water rinse; 3) hand peeling; 4) seed removal; 5) pulp 

136


scalding (1 minute); 6) cooling; 7) soluble solids determination; 8) 0.1% 

sodium benzoate addition; 9) blending (10 minutes); 10) sieving; 11) sugar 

addition; 12) air elimination and pulp concentration (60°C); 13) deposition of 

pulp into a container; 14) covering; 15) cooling; 16) labelling; 17) storage. 

The fruits are hand peeled by making 3 to 4 shallow cuts in the skin and 

pulling the skin sections from the apical extremity, being careful to eliminate 

all the skin to avoid depreciation of pulp quality. 

Since soursop hand-peeling is a very difficult process to use commercially, 

Mororó et al. (1997) recommended a more practical processor (Fig. 11-1) for 

pulp extraction. After primary selection of the fruits, they are rinsed by 

immersion in 10 to 15 ppm chlorinated water (Fig. 11-1A), followed by 

rinsing with water (Fig. 11-1B). The aim is to remove the residues on the 

fruit surface, such as dust, soil and pesticides. Following further inspection 

(Fig. 11-1C) any damaged or unripe fruits that could remain from the first 

selection are discarded. After peeling, the fruits are transported by an endless 

spiral system (Fig. 11-1D) to a 'lung container' (Fig. 11-1E). The function of 

the lung container is to equilibrate pulp flux between extraction and 

automatic packing, using a dosimeter pump (Fig. 11-1H). The extraction is 

made by continuously sieving the fruit to separate the pulp from the seeds 

and other residues. Fine extractions can be made by using a small-hole sieve 

to separate the fibres, although the fibres are considered as an important 

dietary component for human health. The pulp is automatically packed into 

polythene bags and sealed (Fig. 11-1G). 

The percentage of pulp recovery ranges from 62 to 85.5%. The variation is 

due to the type of equipment, extraction method, cultivar, cultural practices 

and number of seeds per fruit (Nakasone and Paull, 1998). Soursop pulp is 

viscous, and its dilution produces flat and weak nectar. In addition, the lack 

of uniformity in acidity and soluble solids concentration generally requires 

homogenization (Torres and Sánchez, 1992). In order to obtain a good nectar, 

the pH should be adjusted to 3.7 by the addition of citric acid, and the Brix to 

15° by addition of sugar, thus creating an appropriate balance of acidity, 

sweetness and flavour (Torres and Sánchez, 1992; Nakasone and Paull, 

1998). High starch, polyphenoxidase and peroxidase contents decrease the 

stability of the product's colour and taste (Torres and Sánchez, 1992). To 

produce a high quality product, soursop pulp should be processed below 

93°C and frozen into polythene bags (Pinto, 1991; Torres and Sánchez, 

1992). 

137


After processing, the enriched pulp, sweetened or not, can be further 

processed into various products and puree can be used to prepare ice creams, 

drinks, sherbets and gelatine (Nakasone and Paull, 1998). The optimum 

pasteurisation conditions for soursop natural puree are 69 seconds at 78.8°C 

and pH 3.7 (Umme et al., 1997). It has also been suggested that pulp treated 

at 70°C for 20 minutes, with addition of 0.5% ascorbic acid to avoid 

oxidation, and packed into polythene bags, can be stored for one month at 

5°C. To obtain nectar for juices, marmalade and jams, 17.8% of pulp, 10.7% 

of added sugar, 0.02% of sodium benzoate, 0.02% of sodium metabisulfide 

and water are treated at 100°C for 15 minutes (Villachica et al., 1996). 

The pH of marmalade ranges from 3.1 to 3.3, and contains 60% concentrated 

pulp and 31% added sugar (Villachica et al., 1996). The following steps to 

prepare marmalade are suggested: 1) scalding or boiling the pulp; 2) 

homogenization; 3) water addition; 4) sugar addition; 5) cooking for 30 

minutes; 6) sugar addition; 7) cooking for 45 minutes; 8) fruit piece addition; 

9) final cooking for 10 minutes; 10) placement into a container while warm; 

11) labelling (Torres and Sánchez, 1992). 

Sugar apple pulp mixed with milk results in a delicious drink and can also be 

frozen into ice cream, which is the main type of processing for this fruit. 

138

Chapter 12. Economic Information 

A. C. de Q. Pinto, D. I. Kinpara, S. R. M. de Andrade 

In agro-economic terms, annona species fall into two groups. In the first are 

custard apple and wild soursop, as well as other species grown by subsistence 

farmers under smallholder conditions on a casual basis. Establishment of 

these small holdings is via cultivation of seedlings, even when in small 

orchards, and attention to market demand is a minor concern as long as fruits 

can be sold. In the second are cherimoya, soursop and sugar apple, which are 

often grown on commercial farms, with better technology, numerous inputs, 

such as irrigation and fertilization, have proper commercial organization and 

processing infrastructure, and heed market signals constantly. In many 

regions, however, the species of the second group, e.g., sugar apple and 

soursop, are still cultivated in conditions reminiscent of the species in the 

first group or may even have escaped from cultivation and are treated as an 

extractivist product (as occurs with sugar apple in some parts of India). Two 

factors will be important for expanding production in both groups: first, the 

wider application of existing technologies; and second, intensifying 

commercial production and practices, while heeding market signals. 

It is very difficult to compile reliable statistical data on costs in the annona 

production-to-consumption system, even for cherimoya, which is the most 

important commercial annona fruit. Most of the scattered, available 

information suggests that cherimoya, as well as soursop and sugar apple, are 

highly remunerative crops for both small and medium scale farmers in many 

countries, although the price of annona fruits received by growers has 

decreased in the last ten years, reducing farm incomes. The annona fruit 

market might be strengthened by adopting policies to provide adequate 

institutional support, financial credit (especially lower interest rates), better 

infra-structure (e.g., road and ports), research to breed new cultivars, 

guarantee longer shelf life and develop processed products. Improved access 

to market information may be just as important as other policies, so that 

growers can enter their fruits into new and more demanding markets, 

especially out of season, to obtain better prices. 

139


12.1 Economics of production 

12.1.1 Production cost, price and income 

The cultivation of cherimoya in Latin America is reputed to have a 

comparative advantage over other locations, especially considering the cost 

of production (Van Damme and Scheldeman, 1999). However, these authors 

do not list the costs for one hectare of cherimoya cultivation using standard 

production factors, such as labour, mechanization, fertilizers, transport etc. 

On the other hand, prices of cherimoya have been listed by various authors, 

which allows the estimation of supposed income for cherimoya growers. 

In Spain, cherimoya production has an average yield of 11.8 t/ha during the 

normal harvest period (September to November). According to Requena 

(1998), the mean price of cherimoya in 1996 was about 200 pesetas/kg of 

fruit (at that time US$ 1.00 = pesetas 131.21; hence, the fruit was worth US$ 

1.52/kg). From this we can conclude that a cherimoya grower in Spain could 

expect a gross farm gate income of US$ 17,900 per hectare in 1996. 

However, 15% of this would be subtracted due to fruit perishability between 

harvest and market, leaving a gross income of US$ 15,200. 

The current cost of establishing one hectare of cherimoya in Spain, with a 

density of 357 trees/ha, is US$ 8,000-8,300 (Hermoso González, J.M., La 

Mayora Experimental Station, Spain, personal communication , 2004). This 

price does not include the price of land, which is frequently very expensive in 

the traditional growing areas of southern Spain. At present, land prices vary 

from US$ 185,000 to 190,000/ha and the costs of producing 12-14 t/ha are 

currently about US$ 5,600-6,000/ha. 

Logically, the price of cherimoya depends on a lot of factors, such as size and 

quality of the fruit, place of sale and harvest date. The price of a good 

cherimoya fruit in Spain, for instance, has ranged between US$ 0.20 and 

1.20/kg in the last few years, depending on the fruit size and harvest date. 

During the 2003 Christmas season, a good quality fruit was sold for US$ 

1.50-1.70/kg (Dr. J.M. Hermoso González, personal communication , 2004). 

In Belgium, cherimoya fruits imported from Spain are sold for around US$ 

5.00/kg in supermarkets, while in Ecuador they are sold for less than a US$ 

1.00/kg (Scheldeman, X., personal communication , 2004). In Peru, the price 

of cherimoya fruit varied from US$ 1.00-2.00/kg in 1991 (Tijero, 1992) and 

from US$ 3.00 to 5.00/kg FOB for export in 1996 (INIA, 1997). 

140


In Spain, a cherimoya grower, who owns the land under the orchard, can 

expect an average yield of 13 t/ha and an average price of US$ 0.70/kg of 

fruit, which gives a gross income of US$ 9,100 in 2003. Comparing the gross 

income obtained in 1996 with the income of 2003 there is a decrease of 

49%. However, using a simple analysis, subtracting the cost of production 

(US$ 5,800/ha) from the gross income, there is still a profit of US$ 3,300 per 

hectare, which is an acceptable income per hectare for any fruit grower. 

In Brazil, cherimoya is harvested from February to the end of October, and 

production in 1999 was estimated at 50,000 boxes of 4 to 5 kg, representing a 

total of 200-250 t of fruit. For a seven year-old cherimoya orchard of the 

‘Fino de Jete’ cultivar, with 417 trees/ha and an average yield of 33 kg/tree 

(Richardson and Anderson, 1993; Bonaventure, 1999), a Brazilian grower 

could obtain a gross income of US$ 24,800, when the mean price of fresh 

fruit was US$ 1.80/kg (Bonaventure, 1999). Although there is no up-to-date 

information on cherimoya prices in Brazil, the increase in cultivated area 

surely increases fruit supply and reduces the fruit price in the Brazilian 

market, which is similar to the situation in Spain. 

Today, agricultural production is not discussed in terms of "absolute 

advantage" or "comparative advantage", but "competitive advantage". 

Competitive advantage comes mainly from the creativity of adding "value" to 

the product, for instance, by harvesting fruit out of season when prices are 

higher in the market. Since cherimoya production occurs mainly between 

September and November, Spanish growers should use chemical pruning 

(see Chapter 10) to promote late harvesting (January to February), and get 

better prices for their fruits. Growers can also add value to their products by 

processing. By selling their fruits as frozen pulps, jellies and sweets, the 15% 

due to losses from fruit perishability described by Requena (1998) are 

eliminated. Both of these ideas might help to slow erosion of fruit prices in 

producing countries. 

The available data on production of soursop and sugar apple in the Cauca 

Valley, Colombia, and in the central and north-eastern regions of Brazil 

(Torres and Sánchez, 1992; Pinto and Silva, 1994; Kavati and Piza Jr., 1997) 

suggest that the most important cost in the first year is the purchase of the 

irrigation system (18%). Other inputs, such as the cost of producing or 

buying grafted plants and purchasing fertilizers, are also important during 

this year, which corresponds to the period of orchard establishment (Pinto 

and Silva, 1994). Therefore, soursop growers should expect to have no 

positive net income during the first two years of cultivation; by the third year, 

141


the income starts to cover the cost of establishment and maintenance, and 

provides an economic return (Table 12-1). 

The cost for establishment and maintenance of one hectare of soursop can be 

calculated from a matrix, the units and quantities for each factor of 

production should be multiplied by local prices for each unit (Torres and 

Sánchez, 1992; Pinto and Silva, 1994). Aguiar and Junqueira (2001), in their 

study on costs of establishment and economic returns for soursop, stated that 

the total cost of production varies from US$ 2,485 per hectare in the first 

year to US$ 1,183 in the sixth year (Table 12-1). 

Intercropping with an annual crop (e.g., beans) or another fruit crop, such as 

papaya (Plate 3), could provide additional income and decrease the costs of 

orchard establishment and maintenance during the first unproductive years. 

Generally, the soursop ecotype Morada attains its mature yield of 50-60 kg of 

fruits/plant in the seventh year, when the mean productivity is around 10 t/ha 

and a net annual income of US$ 6,600 is expected. From the second to the 

seventh year, an accumulated net income of approximately US$ 17,000 is 

attained (after subtracting the US$ 4,328 costs of the two first years; Table 

12-1). Of the US$ 2,486 for orchard establishment and maintenance in the 

first year, about 70% is maintenance. These numbers must be used 

cautiously, since mean productivity of soursop in Brazil is only about 4 t/ha 

and local cultivars of the north-eastern region must also be considered. 

Soursop, however, can be sold as a fresh fruit or for processing, the latter 

option adding value to the raw produce, which is a great advantage compared 

with cherimoya and sugar apple, which are sold almost exclusively as fresh 

fruit. 

Table 12-1. Mean costs to establish and maintain one hectare of soursop 

cv Morada, based on 204 plants per hectare, and estimated gross and net 

incomes 

Year 

Establishment/Mai 

ntenance (US$) 

Av. Yield (ton of 

fruitha) 

Gross Income 

(US$) (1) 

Net Income (US$) 

1st 2,486 - 0 0 

2nd 716 2 1,360 644 

3rd 675 4 2,720 2,045 

4th 923 5 3,400 2,477 

5th 996 7 4,760 3,764 

6th 1,184 9 6,120 4,936 

7th 1,212 10 6,800 5,588 

Data from central Brazil (Pinto and Silva, 1994; Aguiar and Junqueira, 2001) 

(1) Price of fresh fruit was US$ 0.68/kg, at an exchange rate of US$ 1.0 = R$ 

2.924 (January 2004). 

142


In south-eastern Brazil, an orchard (417 trees/ha) of 6-year-old sugar apple 

trees can produce 60 fruits per plant (Lucas, 1994). This suggests that it is 

possible for sugar apple growers to obtain yields of 25,000 fruits/ha (12.5 

t/ha). In July 1996, the price of sugar apple was US$ 0.40 per unit of fresh 

fruit (Kavati, 1997), resulting in a gross income of US$ 10,000/ha. However, 

the price of fruit has decreased since then due to a greater supply. In May 

2003, the price of fresh sugar apple fruit (500 g weight) in Brasilia, Brazil's 

capital, was US$ 0.37 per fruit, therefore with the same yield as in 1996, the 

gross income would be US$ 9,250 in 2003. Even with the costs of production 

at about 30% of the income per hectare for productive plants, this profit is 

very good for Brazilian sugar apple growers. 

In the semi-arid tropical São Francisco Valley, at Petrolina, Pernambuco, 

Brazil, sugar apple trees under irrigation can produce two harvests: a main 

harvest during the rainy season with a high yield (80 fruits/plant), and a 

second harvest during the dry season with a lower yield (20 fruits/plant), 

totalling approximately 21 t/ha per year. The semi-arid climatic conditions 

help to guarantee better fruit quality, translating into a better price and 

allowing a gross income at the fresh fruit market in Petrolina, of US$ 7,770, 

even allowing for the lower prices of 2003 (US$ 0.28 per 500 g fruit). 

However, if these semi-arid growers focus on the major Brazilian market in 

São Paulo, via the Central Food Clearing House of São Paulo State 

(CEAGESP), and considering an estimated cost-of-production of 22% of the 

gross income, their profitability will be much lower than that of São Paulo 

growers producing in the same period, due to the cost of transportation, since 

the São Francisco Valley is about 1,200 km from São Paulo. 

12.1.2 Production, productivity and value 

The estimated production area of cherimoya in the world in 1994 was 13,500 

hectares and, considering an average yield of 6 t/ha, the total production was 

estimated at 81,000 t (PROCIANDINO, 1997). Spain, with more than 3,000 

ha, is the most important cherimoya producer in the world, and Peru and 

Chile are the most important producers in South America, with areas larger 

than 1000 ha (Requena, 1998). These three cherimoya producing countries 

account for 46% of this total area. Chile had an average yield in 1998 of 25 

t/ha, which is 4 times higher than the world average and 2.1 times higher than 

Spain (Table 12.2). Chile produced, for the internal market, more than 8,000 

t in 1995 and currently has exported only 3% of its total production (Table 

143


12.2), principally to the USA (70%), Japan (12%), Argentina (10%) and 

Brazil (5%) (Irazabal, 1997). 

Mexico produced more soursop than Brazil in the late 1990s, due both to its 

larger area and to its better productivity. Assuming that the Philippines has 

the same area of sugar apple in the year 2000 as it did in 1978 (2059 ha), 

Brazil had comparatively higher production due to its higher productivity 

(Table 12-2). 

Due to the niche markets of Asians and Latin Americans in the USA market, 

Thailand, the Dominican Republic and Costa Rica are becoming important 

cherimoya exporters to the USA (Crane and Campbell, 1990). Mexico with 

its numerous appropriate microclimate conditions, has a great potential to 

export cherimoya to USA and some growers have already thought of 

replacing their avocado orchards with cherimoya orchards (Agustin and 

Alviter, 1996), suggesting that the USA is still an important "open window" 

for cherimoya exporters. 

Table 12-2. Total area, production, productivity and value of three 

important Annona species in some of the major producing countries 

Species Country Year Area 

(ha) 

Production 

(x 1000-t) 

Productivity 

(t/ha) 

Value 

(x 1000- 

US$) 

Cherimoya (1) Chile 1998 1,152 28.8 25 34,560 

Peru 2000 1,975 14.6 7.4 17,527 

Spain 1998 3,090 36.5 11.8 43,800 

Soursop (2) Brazil 1997 2000 8 4 5,440 

Mexico 1996 5,915 349 5.9 23,732 

Venezuela 1987 3,496 10.1 2.9 6,868 

Sugar apple (3) Brazil 2000 1,294 11.3 8.7 6,328 

Philippines 1978 2,059 6.2 3.0 3,472 

(1) 

Considering a price of US$ 1.20/kg for fresh cherimoya fruit in Spain in 

January 2004. (2) Considering a price of US$ 0.68/kg for fresh soursop fruit in 

Brazil in January 2004. (3) Considering a price of US$ 0.56/kg for fresh sugar 

apple fruit in Brazil in January 2004. 

There is little available production data on soursop except in Mexico, Brazil 

and Venezuela. Due to an increasing demand for soursop, both fresh and 

processed, Mexico increased its soursop production area by 88% between 

1990 and 1996. In Central Brazil, most soursop growers sell their fruits to 

small agroindustries, with prices varying from US$ 0.51-0.56/kg of pulp. The 

price of frozen soursop pulp was the highest (US$ 0.29/100 g in 2001) 

144


among the many fruit pulps sold in important Brazilian markets, such as São 

Paulo, Belo Horizonte, Brasilia and Rio de Janeiro. 

Sugar apple is the most important annona fruit in Alagoas State, northeastern 

Brazil, with production of 7,720 t, which makes Alagoas the most 

important producer in Brazil (Albuquerque, 1997). This production comes 

mainly from the Regional Cooperative of Palmeira dos Indios (CARPIL), in 

Palmeiras dos Indios County, where growers have an average of 1.17 ha of 

sugar apple each. Large sugar apple fresh fruits, produced in north-eastern 

and south-eastern Brazil were sold in May 2001, in Brasilia for US$ 0.56/ kg 

of fruit. In Petrolina, Pernambuco, Brazil, some growers obtained a better 

prices in the supermarket for purple sugar apple fruits (Plate 1) by selling 

them as exotic fruits only, since its fruit colour, which somewhat looks like a 

rotten fruit, is an impediment for better acceptance in the consumer market. 

The price of all annona fruits in the national and international markets 

depends upon the seasonality of production, which interferes in the 

production value at that moment. Therefore, development of more 

technically oriented production systems in both the southern and northern 

hemispheres could expand the availability of fruit and reduce price 

fluctuations. 

12.1.3 Social improvement 

The production, processing, sale and use of annona products can improve 

social conditions in many areas where annonas are grown and processed, 

through the creation of new employment and the encouragement of small 

entrepreneurs. However, most annona growers in Latin America have limited 

knowledge about appropriate technology, so reap few of the possible 

benefits. In addition, few farmers own the appropriate farm implements, and 

most of them are still hand-operated and generally inefficient (Van Damme 

and Scheldeman, 1999). These limitations lessen their chances of competing 

in urban markets. 

Rural people of north-eastern Brazil sell soursop and sugar apple fruits along 

the federal and state highways, contributing to family income. Some 

agroindustries contract small-scale soursop farmers to produce specifically 

for them through a dedicated contract system, which is an especially efficient 

way of increasing rural family incomes. Although the price offered by 

agroindustries is lower than that of urban retailers, the avoidance of 

transportation costs and market competition influences the small growers to 

145


146 

deliver their fruits at the farm gate or along the roads. If these small farmers 

had proper processing infrastructure, they could add value to their raw 

produce themselves. However, limited or complete absence of access to 

capital keeps farmers at the subsistence level, preventing them from making 

decisions for profit maximization (Van Damme and Scheldeman, 1999). 

Organization into associations or cooperatives is a possibility for increasing 

access to capital, as well as producing a significant quantity of fruit that 

could attract larger buyers. 

12.2 Marketing and commercialization 

Marketing involves the policies that provide strategic support to get a product 

or service into the consumer market, helping to guarantee the commercial 

success of the initiative. It also involves the actions taken by individual 

entrepreneurs to sell their products. Marketing is often the weakest part of the 

production-to-commercialisation system in which annona growers must 

participate to sell their products. 

Cherimoya has well established international marketing activities, with Spain 

and Chile as the main producers and marketers. The other annona fruits, 

however, are traded mostly in national markets of the countries where they 

are produced. According to Bandeira and Braga Sobrinho (1997), 

Bonaventure (1999) and Van Damme and Scheldeman (1999), the national 

and international markets for annonas are limited for the following reasons: 

1. Annona species have not received adequate institutional support to be able 

to obtain financial credit with lower interest rates, good paved roads to avoid 

fruit damage, extension services to transfer technology and see good planting 

material is available, or for research to develop new cultivars. 

2. Due to short shelf life and poor postharvest technologies, significant losses 

of fruits in transport impedes their export success. 

3. Processed pulp is sometimes of low quality and does not meet the 

standards of the international markets, principally because adequate 

processing technologies are not readily available to most producers, 

especially small growers and wholesalers. 

4. Lack of international market information, mainly for tropical Annona 

species, restricts crop and product diversification where it might otherwise be 

successful. Success in fruit commercialisation is entirely due to marketing 

policies. These depend on several factors, such as markets with established 

production-to-commercialisation systems, farmers' organisations, transport


and roads, and institutional support, which include access to financial 

services, research and extension. Many of these factors depend upon state 

and national governmental policies and political will. 

Figure 12-1. Commercialization channels for distribution of cherimoya 

in California, USA 

Where there is a market, there is a production-to-commercialisation system, 

varying from a simple system to a more complex one (Fig. 12-1). In a simple 

system, annona growers can sell their fruits in farmers' markets directly to the 

consumer; these are generally small fairs in small cities or villages. A 

complex system leads into larger markets and is constructed around a system 

starting with producers selling to packers and shippers, then a number of 

wholesale companies that send annona fruits to supermarkets, and finally 

cherimoya, soursop or sugar apples are sold to consumers as specialties 

(Grossberger, 1999). 

Prices of annona fruits vary according to their supply and demand. However, 

in general, well packed and shipped fruits are more expensive than fruits 

from a simple marketing system, due to their higher quality (less damage and 

disease, better shape and weight classification, and uniformity of external 

colour). Bonaventure (1999) commented that in São Paulo, Brazil, a carton 

with four high quality cherimoya fruits attained a price of R$ 35.00 (US$ 

147


14.46/carton at US $1.00 = R $2.42) in 1998. According to this author, 500 g 

is the ideal weight for a cherimoya fruit in the Brazilian markets. 

Fruit packing has a pattern of classification, which influences prices 

significantly. Grossberger (1999) commented that packing of cherimoya fruit 

for the USA market is generally in boxes of 4.5 kg, with plastic insertions 

that hold 6 to 16 fruits in appropriate positions. Lower quality fruit is 

generally sold in a loose pack, which is commonly packed in 18 kg boxes, as 

is seen in California. Fruits sold to supermarkets are considered specialties 

and only small amounts are displayed at any one time. 

Soursop growers pack their fruits in single layer trays with 9 or 12 fruits per 

tray to be sold at supermarkets or they sell individual fruits in the local fresh 

fruit markets. Because soursop is a large fruit, especially that of the ecotype 

Morada, it is difficult to sell to an individual consumer. Some retailers in 

Brasilia, the capital of Brazil, slice the ripe soursop fruits and pack the slices 

(around 1 kg per slice) in trays with thin plastic covers, displaying them in 

freezers of large supermarkets in order to meet their consumers' requirements 

and facilitate commercialisation. 

In Australia, sugar apple is commonly packed in single layer trays of 6 to 8 

kg (George et al., 1987). In Brazil, sugar apples are often packed in wooden 

boxes with weight varying according to the state or local market; smaller 

fruits are often sold in packages with 12 to 24 fruits (Lucas, 1994). Yokota 

(1986) commented that the types of classifications for sugar apple in São 

Paulo, Brazil, are mainly based on length, diameter and weight of the fruits. 

The fruit arrangement in the box varies according to the fruit size and 

number of fruits per box; a type 9 carton has fruits with average weights of 

500 g, which is the best commercial weight, and the package has 9 fruits 

(Table 12-3). 

Table 12-3. Carton types for classification and packing of sugar apple 

fruit in the São Paulo market, Brazil 

Carton 

Fruit Characteristics 

Fruit Arrangement in the 

Package 

Length (cm) Diameter (cm) Weight (g) Columns Rows 

8 >10.5 10.5-11.5 600-620 2 4 

9 9.5-10.5 10.0-10.5 480-520 3 3 

12 8.5-9.5 9.0-10.0 360-390 3 4 

15 8.0-8.5 8.5-9.0 280-320 3 5 

18 7.5-8.0 7.5-8.5 210-215 3 6 

Source: Yokota (1986). 

148


Packing represents 8-10% of the gross value reflected in the market price and 

poor packing can decrease the fruit price by up to 30% (Kavati, 1997). Sugar 

apple packing carton type 9 is the most commonly commercialised in southeastern 

Brazil. However, in Brasilia, a smaller and cheaper wooden box with 

six fruits is the most common type of packaging, with a retail price of R$ 

4.00 per box (US$ 1.65/box) in May 2001, while the supermarket price was 

generally twice as high. 

To provide the consumer with a good quality product on a timely basis with 

affordable prices, a well-constructed farmers' organization is necessary. 

Close linkages between growers and retailers can be established through 

producer cooperatives. The development of this type of organization is 

extremely important for the success of most modern agricultural initiatives, 

but is beyond the scope of this book. 

Looking at the participants of the market chain of annona fruits, growers are 

the ones who have the highest risk and the lowest profit, whereas consumers, 

at the end of the chain, pay for all the growers', wholesaler's and retailers' 

profits. The best example of this common economic equation is given by 

Alvarez et al. (1999) in Ecuador. Cherimoya prices in Ecuador in 1999 

ranged from US$ 0.01 to 0.05 per fruit of 600-800 g at the farm gate. 

Wholesale buyers sold cherimoya to the supermarkets at the price of US$ 

0.075 per fruit and supermarkets sold the fruit to the final consumer for US$ 

0.25, which is an increase of 80 to 96% over the price received by the 

growers compared with the price received by the supermarket owners. 

However, it is necessary to remember that most of this price composition is 

aggregated by services needed to get the final product into market due to its 

perishability. 

149

Chapter 13. Conclusions and 

Research Needs 


Several of the Annona species discussed here, especially cherimoya, soursop 

and sugar apple, have a great potential for an expanded world-wide market 

for fresh fruit consumption and industrially processed products. The lesser 

known species, such as custard apple, wild soursop, and other species not 

discussed, have limited importance for consumption as fresh fruit, although 

the uses of custard apple as a rootstock and of wild soursop for insecticidal 

formulations have demonstrated their relevance in many countries. Selection 

and agronomic development could change this in the mid-term future. 

Important agronomic advances have occurred with Annona species. New 

cherimoya, soursop and sugar apple varieties offer better fruit quality for 

consumer markets. Even the purple sugar apple, which is offered in the 

market as a rarity or curiosity, has obtained double the price of a normal fruit. 

The hybrid atemoya produces better in inhospitable environments, and offers 

better fruit resistance to transport and post-harvest management, compared 

with either of its parents. 

Relevant field technologies, such as the use of artificial pollination to 

increase fruit set and the development of new pruning and training 

techniques to improve plant architecture have both had positive effects on 

yield and fruit quality. Also, intercropping techniques now allow the growers 

to obtain additional income during the first 2-3 years in the field. Postharvest 

technologies for the soft and difficult to handle fruits have been 

improved and new methods of fruit packing, with their systems of 

classification, have significantly influenced fruit quality and prices. 

Packaging of frozen soursop pulp in an appropriate size and format is a 

recent marketing strategy. 

A major breakthrough in recent years has been the better identification and 

isolation of important industrial and medicinal compounds in Annona 

species. Almost without exception, Annona species have bioactive 

compounds in their roots, leaves, bark, fruits and seeds that have great 

potential for use in industry and medicine. 

150

Chapter 13. Conclusions 

However, the expansion of the cultivated areas of annonas is still limited, 

except for cherimoya in China, Taiwan, Spain and Chile, soursop in Brazil 

and Mexico, and sugar apple in India. This is due partially to ecological, but 

principally to political and social factors. Field labour qualified to use new 

technologies, reduction of input costs and better fruit prices are the most 

important factors limiting cultivation and marketing today. Lack of 

government support for financial credit, research and extension services, 

roads and tax structures are policies that negatively influence the expansion 

of cultivation. 

This review has highlighted intensive research on cherimoya in Spain. This 

has been wide-ranging and has covered studies on germplasm, agronomy and 

processing. Other research has focused on soursop in Mexico, Brazil and 

Colombia, which has had positive impacts on increasing commercialisation 

and demand. The research on soursop has been more fragmented than that on 

cherimoya, and there has been less successful dissemination and adoption of 

research findings. The application of new technologies by growers in Latin 

America is still limited, due mainly to poor education levels and low 

technical qualification. Additionally, few farmers have the appropriate 

implements and orchard management is still manual, with consequent 

impacts on fruit quality limiting commercialisation. 

There has been other significant research, summarized in the body of the text. 

The basic research in India on sugar apple has been noteworthy, and basic 

and applied research in Australia, the Philippines, China and Taiwan on 

several species likewise. 

Two important conclusions emerge from this review. First, all too often the 

adoption of modern agronomic practices is slow. This is a complex issue and 

requires targeted down-streaming of technologies and support from extension 

services. Second, too little attention has been given to the better exploitation 

of the Annona gene pool; current use of germplasm collections is woefully 

inadequate in all countries holding them. This means that selection of new 

cultivars has been almost a random process, when there is the urgent need to 

have a range of cultivars readily available for the agro-ecological niches 

where growers live and produce. 

There are also socio-economic factors relevant to the production of annonas. 

For instance, most Latin American banks charge high interest rates for the 

financial credit they offer to small growers - a strong barrier against 

introduction and use of new technologies. Also, governments have given 

poor support to the construction and maintenance of paved roads, resulting in 

151


fruits of poor quality and lower prices in the market, a disincentive to both 

growers and consumers. Prices of annona fruits vary according to supply and 

demand, and small farmers sell their fruits mostly into simplified market 

channels in which low quality and price are demanded, resulting in low 

profits. In Brazil, fruit growers must pay high export taxes, representing up to 

25% of the product's free on board (FOB) price, which negatively influences 

competitiveness in world markets. 

Consumption habits are important social factors restricting expansion of 

annonas in various parts of the world. Cherimoya is not known by people 

living in many tropical regions of the world, although it can be found on 

supermarket shelves of big cities. Soursop and sugar apple are similarly 

poorly or unknown in temperate countries. Although wild soursop adapts 

very well to different altitudes and climate conditions, its presence is 

essentially restricted to Africa and its potential for expanded use beyond that 

continent is currently unknown. 

The following section highlights those aspects which require development to 

allow utilisation of Annona species to their fullest potential for improvement 

of grower livelihoods, both at small and large scale. Adequate training in the 

growing, processing and marketing of Annona species is urgently needed. 

13.1 Research requirements and technology 

transfer 

The most important research requirements are listed below, although 

priorities will surely vary among institutions and countries that study and 

cultivate annonas. 

13.1.1 Genetic resources and genetic improvement 

a) Conservation of Annona germplasm is important to avoid excessive 

genetic erosion. Loss of diversity among traditional farmers is likely because 

the economic environment of the farm household strongly influences the 

extent of the diversity maintained on farm. Conservation of genetic resources 

should be carried out both in situ and ex situ. Developing a conservation 

strategy for the various Annona genepools also requires enhanced basic 

research on the species relationships and the patterns of variation within the 

genepools (the use of DNA markers is a particularly useful tool for this). 

152


b) Better methods and enhanced attention to methods of characterizing 

germplasm are needed. This also depends on implementing the basic research 

in the above point. 

c) Accessions in germplasm collections are by no means representative of the 

patterns of variation in wild and cultivated species. The current collections 

need urgent assessment and they should be nationalized and enriched. The 

collections are limited to a few Annona species (especially cherimoya and 

soursop) and to a few countries. 

d) Variability in seedling rootstock performance, due to genetic diversity, is a 

major cause of low scion yield and poor fruit quality. Therefore, research on 

selection of vigorous and genetically stable rootstocks is very important. 

e) Development of new scion cultivars, and crosses with cultivars with 

superior agronomic and yield traits is essential to provide high quality fruits 

(among other selected traits, cultivars should have high natural fruit set, out 

of season harvest, fruits with symmetrical form, excellent taste and a hard, 

resistant rind to improve pest and disease resistance, as well as to prolong 

post-harvest life). 

f) During selection and breeding there are opportunities to find fruits with 

odd characteristics that might attract consumers and better prices. Skin colour 

is the obvious trait of interest, but others may exist. 

g) For specific pharmacological purposes, evaluation and selection of wild 

and cultivated species with important medicinal and insecticidal uses should 

be pursued, and supported by field surveys and laboratory analysis 

(Abubakar and Abdurahman, 1998; Farrera Villanueva et al., 1999). 

h) Biotechnology needs to be much more widely applied. Studies, such as 

those of Encina et al. (1999) on genetic transformation to improve the control 

of the ripening, to change the post-harvest characteristics of fruits, and to 

provide pest and disease resistance should continue. 

13.1.2 Propagation studies 

a) Vegetative propagation by budding or grafting is slow, often costly and in 

some cases, inefficient. Tissue culture research should be accelerated to 

provide reliable micro-propagation of desirable planting materials. 

153


13.1.3 Studies on crop management 

a) Pruning for increased yield should be better developed for each species in 

a range of environmental conditions. These studies would look for better fruit 

distribution and supply, and help to stabilize prices in the markets and 

profitability for growers. 

b) Water management and modern irrigation techniques have not been fully 

investigated or developed and need more attention. Intercropping and covercropping 

systems should be better studied, both to support water 

management and to improve fruit production. 

c) A full range of organic production techniques and practices should be 

examined and adapted to production of the major export annonas, especially 

cherimoya. Locally, both sugar apple and soursop might benefit in specific 

markets. 

13.1.4 Postharvest and processing 

a) Annona fruits do not ripen adequately when harvested at the immature 

stage, hence identification of the best stage for picking is essential. On the 

other hand, ripe fruits are particularly vulnerable to rot and decay, thus 

quickly losing their shelf life and quality. This raises the need for adequate 

on and off-farm facilities and technology to store and/or process these fruits. 

b) Appropriate packaging techniques for long-distance transport are 

necessary, especially for markets that demand better flavour and appearance, 

due to the fact that fruits are nearer maturity when picked. 

c) Currently, adding value through product diversification is a priority area 

for research attention, although this should be done by the private sector. 

However, in most less developed countries, the public sector must get 

involved, preferably in partnership with the private sector. 

13.1.5 Industrialisation and marketing 

a) Better technology for pre-prepared juices is an urgent requirement, since 

many annona flavour components are extremely volatile and are lost with 

current preparation technologies. Capture and return of these volatile 

components may be the key to preparing bottled and tetra-packed juices for 

many markets, including export. 

154


b) Annona species are already used for the preparation of medicinal products 

with high values. Soursop acetogenins have potential anticancer uses, but no 

information is available about which is the best plant part for extraction of 

these secondary metabolites. Rain Tree, USA, is the only enterprise so far 

that has shown an interest in the industrialization of secondary metabolites. 

Interest by the public sector should be mobilized to accelerate development 

of these opportunities, always in accordance with the Convention on 

Biological Diversity. 

c) Extract of wild soursop is prepared domestically and used with success as 

an insecticide in Africa. Studies on methods for more efficient small-scale 

industrialization of the extract should be pursued. 

d) Traditional marketing systems are still used by the majority of growers 

and, consequently, prices received by them are still low, especially for small 

growers. It is extremely important that government agencies and institutions 

carry out policy research and stimulate organization of small growers into 

associations or co-operatives, so that they can better compete in national 

markets and reach international markets. 

155

References 

Abubakar M. S. and Aburahman E. M. (1998) “Useful Plants in Traditional 

Control of Insect Pests.” Journal of Herbs, Spices and Medicinal 

Plants, 6 (2) : 49-54. 

Accorsi M. R. and Manica I. (1994) “Colheita, Armazenamento e 

Utilização.” [Portuguese] In: Fruticultura-Cultivo das Anonáceas: Ata- 

Cherimólia-Graviola. Edited by I. Manica Evangraf, Porto Alegre. : 

pp. 92-116 

Adeogan E. K. and Durodola J. I. (1976) “Antitumor and Antibiotic 

Principles of Annona senegalensis.” Phytochemistry, 15 (8) : 1311- 

1312. 

Aguiar J. L. P. and Junqueira N. T. V. (2001) “Custo de implantação e 

retornos econômicos. ” [Portuguese] In: Graviola Produção: Aspectos 

Técnicos. Embrapa Informação Tecnológica, Brasilia, DF. : pp. 61-78. 

Agustín J. A. (1997) “El Cultivo de la Cherimoya (Annona cherimola 

Mill.).” [Spanish] In: Memorias del Congreso Internacional de 

Anonaceas Universidad Autonoma Chapingo (UAC), Chapingo, 

México : pp. 110-123. 

Agustín J. A. (1999) “Advances in Research on Genetic Resources of 

Cherimoya (Annona cherimola Mill.) in Michoacán State, Mexico.” In: 

Proceedings of the First International Symposium on Cherimoya. 

Edited by V. Van Damme, Van Damme P. and Scheldeman X. 

Evangraf, Porto Alegre. Acta Horticulturae, 497 : 189-200. 

Agustín J. A. and Alviter A. R. (1996) El Cultivo de la Chirimoya (Annona 

cherimola Mill.) en el Estado de Michoacán. [Spanish] Universidad 

Autonoma Chapingo (UAC), Chapingo, México : 61 pp. 

Agustín J. A. and Angel D. N. (1997) “El Cultivo de la Chirimoya (Annona 

cherimola Mill.) en México.” [Spanish] In: Anonaceas: Produção e 

Mercado (Pinha, Graviola, Atemóia de Cherimólia) Edited by A. R. 

São José, Vilas Boas I., Morais O. M. and Rebouças T. N. H. 

Universidade Estadual do Sudoeste da Bahia, Depto de Fitotecnia e 

Zootenia. Vitória da Conauista, Bahia, Brasil : pp. 7-19. 

156

References 

Aiyelaagbe I. O. O. (1994) “Fruit Crops in the Cashew-Coconut System of 

Kenya: Their Use, Management and Agroforestry Potential.” 

Agroforestry Systems, 27 : 1-16. 

Albuquerque H. C. (1997) “Situação Atual e Perspectivas para Anonáceas no 

Estado de Alagoas.” [Portuguese] In: Proceedings of the I Brasilian 

Symposium on Annonaceae. Edited by A. R. São José, Souza I. V. B., 

Morais O. M. and Rebouças T. N. H. Universidade Estadual do 

Sudoeste da Bahia, Vitória da Conauista, Bahia, Brasil : pp. 150-155. 

Almeida S. P., Proença C. E. B., Sano S. M. and Ribeiro J. F. (1998) 

Cerrado, Espécies Vegetais Úteis. [Portuguese] Embrapa. Centro de 

Pesquisa Agropecuária dos Cerrados. Planaltina, Federal District, 

Brazil. 

Alvarez J. V. U., Motoche J. P. R. and Scheldeman X. (1999) “Estudio del 

Cultivo de la Chirimoya (Annona cherimola Mill.) en las Provincia de 

Loja, Ecuador.” [Spanish] II Congresso International de Anonáceas 

Universad de Ciencias y Artes del Estado de Chiapas, Tuxtla Gutiérrez, 

Chiapas, México. : pp. 230-237. 

Alves R. E., Filgueiras H. A. C. and Mosca J. L. (1997) “Colheita e Póscolheita 

de Anonáceas.” [Spanish] In: Anonáceas: Produção e 

Mercado (Pinha, Graviola, Atemóia de Cherimólia). Edited by A. R. 

São José, Vilas Boas I. and Rebouças T. N. H. Universidade Estadual 

do Sudoeste da Bahia, Depto de Fitotecnia e Zootenia. Vitória da 

Conauista, Bahia, Brasil : pp. 240-256. 

Anderson P. and Richardson A. (1990) Which Cherimoya Cultivar is Best? 

The Orchadist, Keri Keri Research Report. December. : pp. 17-19. 

Anderson P. and Richardson A. (1992) “Cherimoya Pruning Essential for 

High Quality Fruit.” The Orchadist of New Zealand, : 32-34. 

Anjaria J. V. (1989) “Herbal Drugs: Potential for Industry and Cash.” In: 

New Crops for Food and Industry. Edited by G. Wickens, Haq N. and 

Day P. Chapman and Hall, London, UK : pp. 84-92. 

Anonymous (1981) Annual Report of the All India Coordinator Fruits 

Improvement Project. ICAR, India. 

Araque R. (1971) “La Guanábana.” Semán, 2 : 23-29. 

Arckoll D. B. and Clement C. R. (1989) “Potential New Food Crops from the 

Amazon.” In: New Crops for Food and Industry. Edited by G. 

157

References 

Wickens, Haq N. and Day P. Chapman and Hall, London, UK : pp. 

150-165. 

Ashmore S. E. (1997) Status Report on the Development and Application of 

In Vitro Techniques for the Conservation and Use of Plant Genetic 

Resources. IPGRI, Rome, Italy. 

Asolkar L. V., Kakkar K. K. and Chakre O. J. (1992) Second Supplement to 

Glossary of Indian Medicinal Plants with Active Principles Part I. 

CSIR, New Delhi, India. 

Avilan L. R. (1975) “Efecto de la Omisson de los Macronutrientes em el 

Desarrolo y Composición Química de la Guanábana (Annona muricata 

L.) Cultivada en Soluciones Nutritivas.” [Spanish] Agronomia Tropical 

(Maracay), 25 (1) : 73-79. 

Ayoade J. O. (1991) Introdução á Climatologia para os Trópicos. 

[Portuguese] Edited by A. S. Bertrand, Brazil. 3, II, : 232-233. 

Azcón-Aguilar C., Encina C. L., Azcón R. and Barea J. M. (1994) “Effect of 

Arbuscular Mycorrhiza on the Growth and Development of 

Micropropagated Annona cherimola Plants.” Agricultural Science in 

Finland, 3 : 281-288. 

Babu K. H., Zaheeruddin Md., Prasad P. K., (1990) “Studies on Postharvest 

Storage of Custard Apple.” Acta Horticulturae, 269 : 299. 

Bailey L. H. (1949) Manual of Cultivated Plants. Macmillan, New York. 

Bajaj M. and Williams J. T. (1995) Healing Forests: Healing people. IDRC, 

New Delhi, India. 

Belotto F. A. and Manica I. (1994) “Clima e Solo.” [Spanish] In: 

Fruticultura-cultivo das Anonáceas (Ata-Cherimólia-Graviola). Edited 

by I. Manica. Porto Alegre. Chapter 3 : pp. 13-17. 

Bandeira C. T. and Sobrinho R. B. (1997) “Situação Atual e Perspectivas da 

Pesquisa e da Agroindústria da Anonaceas no Estado do Ceará.” 

[Portuguese] In: Proceedings of I Brazilian Symposium on 

Annonáceous Edited by A. R. São José, Souza I. V. B., Morais O. M. 

and Rebouças T. N. H. Universidade Estadual do Sudoeste da Bahia, 

Depto de Fitotecnia e Zootenia. Vitória da Conauista, Bahia, Brasil : 

pp. 156-160. 

158

References 

Beneto J. R., Rodriguez A. J. and Raman de Sandoval A. (1971) “A Soursop 

Pulp Extraction Procedure.” Journal of the Agricultural University of 

Puerto Rico, 55 : 518-519. 

Benjoy M. and Hariharam M. (1992) “In Vitro Plantlet Differentiation in 

Annona muricata.” Plant Cell, Tissue and Organ Culture, 31 : 245- 

247. 

Benza J. C. (1980) “Annonaceas.” [Spanish] In: 143 Frutales Nativos. 

Libreria “El Estudiante”. Lima, Peru. : pp. 33-39. 

Bezerra J. E. F. and Lederman I. E (1997) “Propagação Vegetativa de 

Anonáceas por Enxertia.” [Portuguese] In: Anonáceas, Produção e 

Mercado. Edited by A. R. São José, Souza I. V. B., Morais O. M. and 

Rebouças T. N. H. Universidade Estadual do Sudoeste da Bahia, 

Vitória da Conauista, Bahia, Brasil : pp. 61-67. 

Bhakuni D. S., Tewari S. and Dhar M. M. (1972) “Aporphine Alkaloids of 

Annona squamosa.” Phytochemistry, 11 (5) : 1819-1822. 

Bonaventure L. (1999 a) “The Cultivation of the Cherimoya and of its 

Hybrid Atemoya in Brazil.” In: Proceedings of the First International 

Symposium on Cherimoya. Edited by V. Van Damme, Van Damme P. 

and Scheldeman X. Evangraf, Porto Alegre. Acta Horticulturae, 497 : 

143-146. 

Bonaventure L. (1999 b) “Ecologia Arborícola.” [Portuguese] In: A Cultura 

da Cherimóia e de seu Híbrido a Atemóia. Edited by Nobel. São Paulo. 

Chapter 5 : pp. 31-36. 

Bonaventure L. (1999 c) “Dicas Gerais e Receitas com Cherimóia e 

Atemóia.” [Portuguese] In: A Cultura da Cherimóia e de seu Híbrido a 

Atemóia. Edited by Nobel. São Paulo. Chapter 24 : pp. 160-182. 

Bora P. S., Narain N., Holschuh H. J., Vasconcelos M. A. S. and Santos C. 

M. G. (1987) “Caracterização Física dos Frutos da Gravioleira 

Oriundos do Trópico Semi-árido da Paraíba.” [Portuguese] In: 9° 

Congresso Brasileiro de Fruticultura. 2 : 487-491. 

Bories C., Loiseau P., Cortes D., Myint S. H., Hocquemiller R., Gayral P., 

Cavé A. and Laurens A. (1991) “Antiparasitic Activity of Annona 

muricata and Annona cherimola Seeds.” Planta Medica, 57 : 434-436. 

Bourke D. O. D. (1976) “Annona spp.” The Propagation of Tropical Fruit 

Trees. Edited by R. J. Garner and Chaudhri S. A. Horticultural Review 

159

References 

N. 4. Commonwealth Bureau of Horticulture and Plantation Crops, 

East Malling, Kent, UK : pp. 223-248. 

Bridg H. (1984) Micropropagation and Determination in the In Vitro 

Stability of Annona cherimola and Annona muricata. Dissertation von 

Dipl. Biol. 25, 01, 64, Stantafe de Bogota, Kolumbien. 

Broughton W. J. and Tan G. (1979) “Storage Conditions and Ripening of the 

Custard Apple Annona squamosa.” Scientia Horticulturae, 10 : 73-82. 

Brown B. I., Wong L. S., George A. P. and Nisse R. J. (1988) “Comparative 

Studies on the Postahrvest Physiology of Fruit from Different Species 

of Annona (Custard Apple).” Journal of Horticultural Science, 63 : 

521-528. 

Bruinsma J. and Paull R. E. (1984) “Respiration During Postharvest 

Development of Soursop Fruit, Annona muricata L.” Plant Physiology, 

76 : 131-138. 

Bucks D. A. and Davis S. (1986) “Introduction to Historical Development.” 

In: Trickle Irrigation for Crop Production: Design, Operation and 

Management. Edited by F. S. Nakayama and Bucks D. A. Elsevier, 

Amsterdam, The Netherlands : pp. 1-21. 

Bueso C. E. (1980) “Soursop, Tamarind and Cherimoya.” Tropical and 

Subtropical Fruits - Composition, Properties and Uses. Edited by S. 

Nagy and Shaw P. E. Avi Publishing Inc., Westport, Connecticut, USA 

: pp. 375-387. 

Bustillo A. E. and Peñame J. E. (1992) “Biology and Control of the Annona 

Fruit Borer Cerconota annonella (Lepidoptera:Oecophoridae).” Fruits, 

47 (1) : 81-84. 

Calzavara B. B. G. and Müller C. H. (1987) Fruticultura Tropical: A 

Gravioleira (Annona muricata L.). [Portuguese] Belém, EMBRAPA- 

CPATU, Documentos 47 : 36 pp. 

Campbell C. W. and Phillips R. L. (1983) The Sugar Apple. Fruit Crops 

Factsheet, Florida Cooperative Extension Service, University of 

Florida, USA : pp. 1-3. 

Campbell C. W. and Popenoe J. (1968) “Effect of Gibberelic Acid on Seed 

Dormancy of Annona diversifolia Saff.” Proceedings of Tropical 

Region American Society of Horticultural Science, 11 : 31-36. 

160

References 

Caparros-Lefebvre D., Elbaz A. and the Caribbean Parkinsonism Study 

Group. (1999) “Possible Relation of Atypical Parkinsonism in the 

French West Indies with Consumption of Tropical Plants: A Casecontrol 

Study.” The Lancet, 354 : 281-286. 

Carvalho P. S. de, Bezerra J. E. F., Lederman I. E., Alves M. A. and Melo 

Neto M. L. (2000) “Avaliaçao de Genotipos de Pinheira (Annona 

squamosa L.) no Vale do Rio Moxoto III - Caracteristicas de 

Crescimento e Produçao, 1993 a 1997.” [Portuguese] Revista 

Brasileira de Fruticultura, 22 (1) : 27-30. 

Cassady J. M. (1990) “Natural Products as a Source of Potential Cancer 

Chemotherapeutic and Chemopreventative Agents.” Journal of Natural 

Products, 53 (1) : 23-41. 

Castillo Alcopar P., Muno Perez R. B., Rubi Arriaga M. and Cruz Castillo J. 

G. (1997) “Metodos de Propagacion del Chirimoyo (Annona cherimola 

Mill.) en Catepec Harinas, México.” [Spanish] In: Memorias del 

Congreso Internacional de Anonaceas Universidad Autonoma 

Chapingo (UAC), Chapingo, México : pp. 195-203. 

Castro M., Cautin R. and Biancani L. (1999) “Evaluation of Three 

Disinfection Protocols and Three Protocols for the Use of Antioxidants 

in In Vitro Cultivation of Cherimoya (Annona cherimola 

Mill.) and Three Quantitative Determination of Branch Phenolic 

Content.” Acta Horticulturae, 497 : 303-307. 

Castro F. A., Maia G. A., Holanda L. F. F., Guedes Z. B. L. and Moura Fe J. 

A. (1984) “Caracteristicas Fisicas e Quimicas da Graviola.” 

[Portuguese] Pesq. Agrop. Bras, Brasilia. 19 (3) : 361-365. 

Catarino P. S. and Ezequiel M. D. (1999) “Control de Moscas de la Fruta con 

Extractos de Anona (Annona squamosa, L.), Annonaceae.” [Spanish] II 

Congreso International de Anonáceas. Universad de Ciencias y Artes 

del Estado de Chiapas, Tuxtla Gutiérrez, Chiapas, México : pp. 163- 

165. 

Cautin R., Fassio C. and Ovalle A. (1999) “Productive Behaviour of Fruiting 

Wood in Three Trellis Systems in Cherimoya.” In: Proceedings of the 

First International Symposium on Cherimoya. Edited by V. Van 

Damme Damme P. Van Scheldeman X. ISHS, Loja, Ecuador. Acta 

Horticulturae, 497 : 315-322. 

Cavalcante P. B. (1976) Frutas Comestíveis da Amazônia. [Portuguese] 

INPA, Belém do Pará, Brasil. 

161

References 

Chang F. R., Chen J. L., Chiu H. F., Wu M. J. and Wu Y. C. (1998) 

“Acetogenins from Seeds of Annona reticulata L.” Phytochemistry, 47 

(6) : 1057-1061. 

Chang F. R., Wu Y. C. and Duh C. Y. (1993) “Studies on the Acetogenins of 

Formosan Annonaceous Plants, II. Cytotoxic Acetogenins from 

Annona reticulata.” Journal of Natural Products, 56 (10) : 1688-1694. 

Chao-Ming L., Ning-Hua T., Hui-Lan Z., Qing M., Xiao-Jiang H., Yi-Neng 

H. and Jun Z. (1998) “Cyclopeptide from the Seeds of Annona 

muricata.” Phytochemistry, 48 (3) : 555-556. 

Chao-Ming L., Ning-Hua T., Qing M., Hui-Lan Z., Xiao-Jiang H., Yu W. 

and Jun Z. (1997) “Cyclopeptide from the Seeds of Annona squamosa 

L.” Phytochemistry, 45 (3) : 521-523. 

Chen C. Y., Chang F. R., Chiu H. F., Wu M. J. and Wu Y. C. (1999) 

“Aromin-A, an Annonaceous Acetogenin from Annona cherimola.” 

Phytochemistry, 51 : 429-433. 

Chen C. Y., Chang F. R., Yen H. F. and Wu Y. C. (1998) “Amides from 

Stems of Annona cherimola.” Phytochemistry, 49 (5) : 1443-1447. 

Chuliá S., Ivorra M. D., Cavé A., Cortés D., Nogueira M.ª and D´ocón M. P. 

(1995) “Relaxant Activity of Three Aporphine Alkaloids from Annona 

cherimolia on Isolated Aorta of Rat.” Journal of Pharmaceutical 

Pharmacology, 47 : 647-650. 

Clement C. R. (1999) “1492 and the Loss of Amazonian Crop Genetic 

Resources. I The Relation Between Domestication and Human 

Population Decline.” Economic Botany, 53 (2) : 188-202. 

Cogez X. and Lyannaz J. P. (1996) “Manual Pollination of Sugar Apple 

(Annona squamosa L.).” Tropical Fruits Newsletter, 19 : 5-6. 

Coppens d'Eeckembrugge G., Ferla D. L. and Ferreira F. R. (1998) 

“Diversidade e Potencial das Fruteiras Neotropicais.” [Portuguese] In: 

15º Congresso Brasileiro de Fruticultura. Lavras-UFLA : pp. 19-47. 

Coronel R. E. (1994) “Atis.” In: Promising Fruits of the Philippines. Edited 

by R. E. Coronel, Zuno J. C. and Sotto R. C. College of Agriculture, 

University of the Philippines at Los Baños, Laguna, Philippines. : pp. 

1-18. 

Cortés D., Figadere B. and Cavé A. (1993 a) “a) Bis-Tetrahydrofuran 

Acetogenins from Annonaceae.” Phytochemistry, 32 (6) : 1467-1473. 

162

References 

Cortés D., Myint S. H., Dupont B. and Davoust D. (1993 b) “Bioactive 

Acetogenins from Seeds of Annona cherimolia.” Phytochemistry, 32 

(6) : 1475-1482. 

Couceiro E. M. (1973) “Pinha, Fruta do Conde ou Ata, sua Cultura e 

Origem.” [Portuguese] Publicação da CEASA-PE, Recife, 1 (8) : 7 

Crane J. H. and Campbell C. W. (1990) “Origin and Distribution of Tropical 

and Subtropical Fruits.” In: Fruits of Tropical and Subtropical Origin. 

FSS, Florida, USA : pp.1-65. 

Croat J. B. (1978) Flora of Barro Colorado Island. Stanford University 

Press, Stanford, USA. 

de Oliveira S. L., Guerra N. B., Maciel M. I. S. and Livera A. V. S. (1994) 

“Polyphenoloxidase Activity, Polyphenols Concentration and 

Browning Intensity During Soursop (Annona muricata L.) 

Maturation.” Journal of Food Science, 59 (5) : 1050-1052. 

de Smet S., Van Damme P., Scheldeman X. and Romero J. (1999) “Seed 

Structure and Germination of Cherimoya (Annona cherimola Mill.).” 

In: Proceedings of the First International Symposium on Cherimoya, 

Loja, Ecuador. Edited by V. Van Damme, Van Damme P. and 

Scheldeman X. ISHS, Loja, Ecuador. Acta Horticulturae, 497 : 269- 

278. 

Dhingra J., Mehrotra R. S. and Aneja I. R. (1980) “A New Postharvest 

Disease of Annona squamosa L.” Current Science, 49 : 477-478. 

Diego S. (1989) Lista Parcial de Especies Frutales en Venezuela. I Jornadas 

Sobre Producción y Exportación de Frutas. [Spanish] Maturin, 

Venezuela. 

Donadio L. C. (1997) “Situação Atual e Perspectivas das Anonáceas” 

[Portuguese] In: Anonáceas: Produção e Mercado (Pinha, Graviola, 

Atemóia e Cherimólia). Edited by A. R. São José, Souza I. V. B., 


Sudoeste da Bahia, Depto de Fitotecnia e Zootecnia. Vitória da 

Conquista, Bahia, Brasil : pp. 1-4. 

Duarte O. and Escobar O. (1998) “Improving Fruit Set of Cherimoya 

(Annona cherimola Mill.) cv. Cumbe, by Autogamous and Allogamous 

Hand Pollination.” Proceedings of the Interamerican Society for 

Tropical Horticulture, 41 : 162-165. 

163

References 

Duarte O., Villagarcia J. and Franciosi R. (1974) “Efecto de Algunos 

Tratamientos en la Propagación del Chirimoyo, por Semillas, Estacas e 

Injertos.” [Spanish] Proceedings of the Tropical Region of American 

Society Horticulture Science, 18 : 41-48. 

Duke J. A. (1970) “Ethnobotanical Observations on the Choco Indian.” 

Economic Botany, 24 (3) : 344-366. 

Duret P., Gromek D., Hocquemiller R. and Cavé A. (1994) “Isolation and 

Structure of Three New Bis-Tetrahydrofuran Acetogenins from the 

Roots of Annona cherimolia.” Journal of Natural Products, 57 (7) : 

911-916. 

Ellstrand N. C. and Lee J. M. (1987) “Cultivar Identification of Cherimoya 

(Annona cherimola Mill.) Using Isozymes Markers.” Scientia 


Encina C. L., Barceló-Muñoz A., Herrero-Castaño A. and Pliego-Alfaro F. 

(1994) “In Vitro Morphogenesis of Annona cherimola Mill. Bud 

Explants.” Journal of Horticultural Science, 69 : 1053-1059. 

Encina C. L., Padilla I. M. G., Carzola J. M. and Caro E. (1999) “Tissue 

Culture in Cherimoya.” In: Proceedings of the First International 

Symposium on Cherimoya. Edited by V. Van Damme Van Damme P. 

Scheldeman X. ISHS, Loja, Ecuador. Acta Horticulturae, 497 : 289- 

294. 

Evangelista J. (1992) “Alimentos - Um Estudo Abrangente.” [Portuguese] 

Atheneu, Série Enfermagem, Nutrição, Módulo, 2 : 33. 

Fang X. P., Gu Z. M., Rieser M. J., Hui Y. H. and McLaughlin J. L. (1993) 

“Structural Revisions of Some Non-Adjacent Bis-Tetrahydrofuran 

Annonaceous Acetogenins.” Journal of Natural Products, 56 (7) : 

1095-1100. 

FAO (1983) “Food and Fruit-Bearing Forest Species, 1: Examples from 

Eastern Africa.” FAO Forestry Paper, 4 (1) : 11-14. 

FAO (1988) “Traditional Food Plants - A Resource Book for Promoting the 

Exploitation and Consumption of Food Plants in Arid, Semi-Arid and 

Sub-Humid Lands of Eastern Africa.” FAO Food and Nutrition Paper, 

42. 

FAO (1990) “Utilization of Tropical Food, Fruits and Leaves.” FAO Food 

and Nutrition Paper, 47 (7) : 10-14. 

164

References 

Farooqi A. A., Parvatikar S. R. and Nalawadi U. G. (1970) “Preliminary 

Studies on the Problem of Fruit Set in Annona reticulata L.” The 

Mysore Journal of Agricultural Sciences, 4 (1) : 44-53. 

Farré J. M. and Hermoso J. H. (1997) “El Chirimoyo en España.” In: 

Anonáceas: Produção e Mercado (Pinha, Graviola, Atemóia e 

Cherimólia). Edited by A. R. São José, Boas I. V., Morais O. M. and 

Rebouças T. N. H. Universidade Estadual do Sudoeste da Bahia, Depto 

de Fitotecnia e Zootecnia, Vitória da Conquista, Bahia, Brasil : pp. 84- 

87. 

Farré J. M., Hermoso J. M., Guirado E. and Garcia-Tapia J. (1999) 

“Techniques of Cherimoya Cultivation in Spain.” In: Proceedings of 

the First International Symposium on Cherimoya. Edited by V. Van 

Damme, Van Damme P. and Scheldeman X. ISHS, Loja, Ecuador. 

Acta Horticulturae, 497 : 91-103. 

Farrera Villanueva S. B., De La Cruz Chacón I., Luna Cazáres L. M., Pérez 

Amador M. C., González Esquinca A. R. and Martínez Vásquez M. 

(1999) “Citoxicidad Diferencial en Estructuras de Annona diversifolia 

Saff. Y Annona lutescens Saff.” [Spanish] In: Memórias del Congresso 

Internacional de Anonáceas. Universad de Ciencias y Artes del Estado 

de Chiapas, Tuxtla Guttiérrez, Chiapas, México: pp. 170-173. 

Fatope M. O., Audu O. T., Takeda Y., Zeng L., Shi G., Shimada H. and 

McLaughlin J. L. (1996) “Bioactive Ent-Kaurene Diterpenoids from 

Annona senegalensis.” Journal of Natural Products, 59 : 301-303. 

FDA, Fundacion de Desarrolo Agropecuario. (undated) “Cultivo de 

Guanabana.” [Spanish] Boletin Tecnico. 12 : 11p. 

Fehr W. R. (1987 a) Principles of Cultivar Development, Theory and 

Technique. Vol. 1. Macmillan Publishing Company, New York. 

Fehr W. R. (1987 b) Principles of Cultivar Development, Crop Species. Vol. 

2. Macmillan Publishing Company, New York. 

Ferraz L. C. C. B., Monteiro A. R. and Silva G. S. (1989) “Sobre a 

Ocorrência de Espécies de Xiphinema no Estado do Maranhão.” 

[Portuguese] Nematologia Brasileira, 13 : 185-188. 

Ferreira F. R. (2001) “Conservação de Germoplasma de Espécies Frutíferas 

no Campo.” [Portuguese] In: III SIRGEALC – Simpósio de Recursos 

Genéticos para a América Latina e Carib. Londrina, IAPAR. : pp. 38- 

40. 

165

References 

Ferreira F. R. and Bustamante P. G. (2000) “Etnobotânica: a História do 

Curauá.” [Spanish] Genebio, 2 (6) : 16. 

Ferreira G., Cereda E., Silva C. de P., Cunha R. J. P. and Cataneo A. (1997) 

“Imbibition Studies of Sugar Apple (Annona squamosa L.) and 

Atemoya (Annona Hibrid) Seeds.” In: Memorias del Congreso 

Internacional de Anonáceas. Universad Autonoma Chapingo (UAC), 

Chapingo, México : pp. 210-225. 

Ferreira G. Z., Fogaça L. A. and Malavasi M. M. (1999) “Germinación de 

Semillas de Annona squamosa L. Sometidas a Diferentes Tiempos y 

Concentraciones de ácido Giberélico.” [Spanish] In: Memorias del II 

Congreso Internacional de Anonáceas. Universad de Ciencias y Artes 

del Estado de Chiapas, Tuxtla Gutiérrez, Chiapas, México : pp. 79. 

Ferreira S. A. N. and Clement C. R. (1987) “Avaliação de Diferentes Porta- 

Enxertos para Gravioleira na Amazónia Central I. Métodos de 

Enxertia.” In: IX Congresso Brasileiro de Fruticultura. Campinas, 

Brasil : pp. 475-479. 

Fouqué A. (1972) “Espéces Fruitiéres Dámérique Tropicale.” [French] 

Fruits, 27 (1) : 62-72. 

Freire F. de C. O. and Cardoso J. E. (1997) “Doenças das Anonáceas.” 

[Portuguese] In: Proceedings of I Brazilian Symposium on 

Annonáceous. Edited by A. R. São José, Souza I. V. B., Morais O. M. 

and Rebouças T. N. H. Universidade Estadual do Sudoeste da Bahia, 

Vitória da Conquista, Bahia, Brasil : pp. 196-213. 

Fresno A. Villar del and Cañavate J. L. R. (1983) “Alkaloids from Annona 

cherimolia Seed.” Journal of Natural Products, 46 (3) : 438. 

Fries R. E. (1959) “Annonaceae.” In: Die Natürlichen Pflanzenfamilien 2 e . 

Edited by A. Engler and Prantl K. Aufl., Band 17a II: 1-171, Berlin, 

Germany. 

Fuentes L. J. (1999) “Production of Cherimoya (Annona cherimola Mill.) in 

Ecuador.” In: Proceedings of the First International Symposium on 

Cherimoya. Edited by V. Van Damme, Van Damme P. and 

Scheldeman X. ISHS, Loja, Ecuador. Acta Horticulturae, 497 : 59-63. 

Fusagri A. A. F. G. (1982) “Estudios de Dinamica de Maduracion en 

Guanabana.” [Spanish] Proceedings of the Tropical Region, American 

Soceity for Horticultural Science, 25 : 267-274. 

166

References 

Fuster C. and Prestamo G. (1980) “Variation of Cherimoya (Annona 

cherimola) Texture During Storage, as Determined with An Instron 

Food Testing Instruments.” Journal of Food Science, 45 : 142-143. 

Galila A. S., El Masry H. M. and Hagag L. F. (1991) “Effect of Saline Water 

Irrigation on Growth and Mineral Content of Annona Plants.” Journal 

of Applied Science, 6 (8) : 304-315. 

Gardiazabal I. F. and Cano G. L. (1999) “Caracterización de 10 Cultivares de 

Chirimoyo (Annona cherimola Mill.) y su Respuesta a la Polinización 

Artificial en Quillota, Chile.” [Spanish] In: Proceedings of the First 

International Symposium on Cherimoya. Edited by V. Van Damme, 

Van Damme P. and Scheldeman X. ISHS, Loja, Ecuador Acta 


Gazel Filho A. B., Carvalho A. C. A. and Menezes A. J. E. A. (1994) 

“Teores de Macronutrientes em Folhas de Graviola.” [Portuguese] 

Revista Brasileira de Fruticultura, Cruz das Almas, Bahia. 16 (2) : 

121-124. 

George A. P. (1984) “Annonaceae.” In: Tropical Tree Fruit for Australia. 

Edited by P. E. Page. Queensland Department of Primary Industries, 

Brisbane, Australia: pp. 35-41. 

George A. P., Broadley R. H., Nissen R. J., Hamil S. D. and Topp B. L. 

(1999) “Breeding Strategies for Atemoya and Cherimoya.” In: 


Edited by V. Van Damme, Van Damme P. and Scheldeman X. ISHS, 

Loja, Ecuador. Acta Horticulturae 497 : 255-260. 

George A. P. and Nissen R. J. (1980) “Custard Apple Propagation.” Biennial 

Report Maroochy Horticulture Research Station, 2 : 19-20. 

George A. P. and Nissen R. J. (1986) “Custard Apple.” Biennial Report 

Maroochy Horticulture Research Station, 4 : 46-68. 

George A. P. and Nissen R. J. (1987) “Propagation of Annona Species: A 

Review.” Scientia Horticulturae, 33 : 75-85. 

George A. P. and Nissen R. J. (1992) “Edible Fruits and Nuts: Annona 

cherimola Mill. and Annona squamosa L.” In: Plant Resources of 

South-East Asia, 2. Edited by E. W. M. Verheij and Coronel R. E. 

Prosea, Bogor, Indonesia. : pp. 71-75. 

George A. P., Nissen R. J. and Brown B. I. (1987) “The Custard Apple.” 

Queensland Agricultural Journal, 113 (5) : 287-297. 

167

References 

Geurts F. (1981) Annonaceous Fruits. Royal Tropical Institute, Amsterdam, 

the Netherlands.: 16 pp. 

Gleye C., Duret P., Laurens A., Hocquemiller R. and Cavé A. (1998) “cis- 

Monotetrahydrofuran Acetogenins from the Roots of Annona 

muricata.” Journal of Natural Products, 61 : 576-579. 

Gleye C., Laurens A., Hocquemiller R., Laprévote O., Serani L. and Cavé A. 

(1997) “Cohibins A and B Acetogenins from Roots of Annona 


Gleye C., Laurens A., Laprévote O., Serani L. and Hocquemiller R. (1999) 

“Isolation and Structure Elucidation of Sabadelin, an Acetogenin from 

the Roots of Annona muricata.” Phytochemistry, 52 (8) : 1403-1408. 

Goeschl T. (1998) “In Situ Conservation On Farm - Data Collecting and 

Analysis: Market/Non-Market Incentives and Agricultural Policy for 

On Farm Conservation.” In: Proceedings of a Workshop to Develop 

Tools and Procedures for In Situ Conservation On Farm. Edited by D. 

I. Jarvis and Hodgkin T. IPGRI, Rome, Italy: pp. 22-23. 

Gonzalez C. and Esteban E. (1974) “Nutrición del Chirimoya: Ciclo Anual.” 

[Spanish] Anales de Edafologia e Agrobiologia, Madrid, España. 33 : 

371-380. 

Grossberger D. (1999) “La Industria de Chirimoya en California.” [Spanish] 

In: Proceedings of the First International Symposium on Cherimoya. 

Edited by V. Van Damme, Van Damme P. and Scheldeman X. ISHS, 

Loja, Ecuador. Acta Horticulturae, 497 : 131-142. 

Guirado E. (1999) “El Chirimoyo en España: História, Producción 

Comercial, Técnicas de Cultivo, Investigación en Marcha.” [Spanish] 

In: II Congreso International de Anonáceas. Universad de Ciencias y 

Artes del Estado de Chiapas, Tuxtla Gutiérrez, Chiapas, México: pp. 

43-57. 

Guirado E., Hermoso G. J. M., Pérez de Ortega M. A., Garcia-Tapia Bello J. 

and Farré M. J. M. (2001) Polinización del Chirimoyo. [Spanish] Junta 

de Andaucia, Consejería de Agricultura y Pesca, Sevilla, Spain. : 45 pp. 

Haise R. H. and Hagan R. M. (1967) “Soil, Plant and Evaporative 

Measurements as Criteria for Scheduling Irrigation.” In: Irrigation of 

Agricultural Lands. Edited by R. M. Hagan, Haise H. R. and Edminster 

T. W. Madison : pp. 577-604 (Agronomy Series, 11). 

168

References 

Hartmann H. T., Kester D. E. and Davies Jr. F. T. (1990) “Principles of Seed 

Selection.” In: Plant Propagation, Principles and Practices. Prentice 

Hall, New Jersey, USA. Chapter 5 : pp. 94-103. 

Hartshorn G. S. and Proveda L. J. (1983) “Checklist of Trees.” In: Costa 

Rican Natural History. Edited by D. H. Janzen. University of Chicago 

Press, Chicago, USA : pp. 158-183. 

Hayat M. A. (1963) “Morphology of Seed Germination and Seedling in 

Annona squamosa.” Botanical Gazette, 124 : 360-362. 

Hermoso G. J. M., Pérez de Oteyza M. A., Ruiz Nieto A. and Farré J. M. 

(1999) “The Spanish Germplasm Bank of Cherimoya (Anonna 

cherimola Mill.).” In: Proceedings of the First International 


and Scheldeman X. ISHS, Loja, Ecuador. Acta Horticulturae, 497 : 

201-212. 

Hermoso J. M. and Farré J. M. (1994) “Experimentación en Fruticultura 

Tropical de Utilidad para la Agricultura Ecológica.” [Spanish] In: 

Actas I Congreso S.E.A.E. Toledo, España : pp. 194-201. 

Hermoso J. M. and Farré J. M. (1997) “Cherimoya Growing in Spain.” 

Mesfin Newsletter, 3 (1) : 5-10. 

Hernández C. R. and Angel D. N. (1997) “Anonáceas com Propriedades 

Insecticidas.” [Portuguese] Anonáceas: Produção e Mercado (Pinha, 

Graviola, Atemóia e Cherimólia). Edited by A. R. São José Boas I. V., 


Sudoeste da Bahia, Depto de Fitotecnia e Zootecnia, Vitória da 

Conquista, Bahia, Brasil : pp. 229-239. 

Hernández D. C. M., Rios M. J. A., Vidal-Lezama E. and Marroquin A. L. 

M. (1999) “Efectos de Giberelinas y Sustrato Sobre la Germinación de 

Semillas de Chincuya (Annona purpurea Moc & Sesse).” [Spanish] In: 

Memorias del II Congreso Internacional de Anonáceas. Universad de 

Ciencias y Artes del Estado de Chiapas, Tuxtla Gutiérrez, Chiapas, 

México : pp. 81-82. 

Hernandez L. V. (1983) La Reproducción Sexual y Multiplicación Vegetativa 

de las Anonáceas [Spanish] Universidad Veracruzana, Vera Cruz, 

México : pp. 102-122. 

Hernandez M. C. L. V. and Nieto Angel D. (1997) Diagnostico Técnico y 

Commercial de la Guanabana en México. [Spanish] Memorias del 

169

References 

170 

Congreso Internacional de Anonaceas Universidad Autonoma 

Chapingo (UAC), Chapingo, México : pp. 1-18. 

Herrera A. L. (1999) “Biodiversidad, Bioseguridad y Propriedad Intelectual 

de los Recursos Geneticos.” [Spanish] In: Memórias del II Congresso 


de Chiapas, Tuxtla Gutiérrez, Chiapas, México : pp. 31-42. 

Higuchi H., Utsunomiya N. and Sakutani T. (1998) “Effects of Temperature 

on Growth, Dry Matter Production and CO 2 

Assimilation in Cherimoya 

(Annona cherimola Mill.) and Sugar Apple (Annona squamosa L.) 

Seedlings.” Scientia Horticulture, 73 : 89-97. 

Hisham A., Sunitha C., Sreekala U., Pieters L., De Bruyne T., Van den 

Heuvel H. and Claeys M. (1994) “Reticulacinone, an Acetogenin from 

Annona reticulata.” Phytochemistry, 35 (5) : 1325-1329. 

Holanda L. F. F., Maia G. A., Martins C. B. and Fe J. de A. M. (1980) 

“Estudo do Processamento e Estabilidade da Polpa e Néctar da 

Graviola (Annona muricata L.).” [Spanish] Ciéncias Agronomy, 10 (1) 

: 103-107. 

Hong T. D., Linington S. and Ellis R. H. (1996) Seed Storage Behaviour: A 

Compendium. IPGRI, Rome, Italy. : 656 pp. 

Hopp D. C., Alali F. Q., Gu Z. M. and McLaughlin J. L. (1998) “Mono-THF 

Ring Annonaceous Acetogenins from Annona squamosa L.” 

Phytochemistry, 47 (5) : 803-809. 

Hopp D. C., Zeng L., Gu Z. M., Kozlowski J. F. and McLaughlin J. L. (1997) 

“Novel Mono-tetrahydrofuran Ring Acetogenins, from the Bark of 

Annona squamosa, Showing Cytotoxic Selectivities for the Human 

Pancreatic Carcinoma Cell Line, PACA-2.” Journal of Natural 

Products, 60 : 581-586. 

Hopp D. C., Zeng L., Gu Z. M. and McLaughlin J. L. (1996) “Squamotacin: 

An Annonaceous Acetogenin with Cytotoxic Selectivity for the Human 

Prostate Tumor Cell Line (PC-3).” Journal of Natural Products, 59 (2) 

: 97-99. 

Ibar L. (1979) “El Chirimoya.” [Spanish] In: Cultivo del Aguacate, 

Chirimoya, Manga, Papaya. Editorial Aedos, Barcelona, España. : pp. 

121-144. 

Ibar L. (1986) “El Chirimollo.” [Spanish] In: Aguacate, Chirimollo, Mango y 

Papaya. Editorial Aedos, Barcelona, España : pp. 23-147.

References 

Idstein H., Herres W. and Schreier P. (1984) “High-Resolution Gas 

Chromatography-Mass Spectrometry and Fourier Transform Infrared 

Analysis of Cherimoya (Annona cherimolia Mill.) Volatiles.” Journal 

of Agricultural Food Chemistry, 32 : 383-389. 

Iglesias A. A. (1984) Propagación del Guanábano (Annona muricata L.) por 

Medio de Injerto sobre Diferentes Patrones de Anonáceas. [Spanish] 

Master Thesis, Universidad Nacional de Colombia, Faculdad de 

Ciencias Agropecuarias de Palmira, Colombia : 73 pp. 

Iglesias A. A. and Sanchez L. A. (1985) “Propagation of Soursop, Annona 

muricata L. by Grafting on Different Annonaceous Rootstocks.” 

[Spanish] Acta Agronomica, 35 (3) : 53-58. 

INIA (1997) El Cultivo del Chirimoyo. Instituto Nacional de Investigación 

Agraria. Boletim Técnico 11, Lima, Peru. : 28 pp. 

IPGRI (2000) Directory of Germplasm Collections. IPGRI, Rome, Italy. 

Irazabal F.G. (1997) “Cherimoya Cultivation in Chile.” Mesfin Newsletter, 1 

: 2-4. 

James L.G. (1988) Principles of Farm Irrigation System Design. Wiley, New 

York, USA. 

Jirovetz L., Buchbauer G. and Ngassoum M. B. (1998) “Essential Oil 

Compounds of the Annona muricata Fresh Fruit Pulp from Cameroon.” 

Journal of Agricultural Chemistry, 46 : 3719-3720. 

Johnson S. (1990) “Eating Anona.” Tropical Garden Fruit World. 1 (4) : 

128. 

Junqueira N. T. V., Cunha M. M. da, Oliveira M. A. S. and Pinto A. C. de Q 

(1996) Graviola para Exportação: Aspectos Fitossanitários. 

[Portuguese] Embrapa-SPI, Brasília: 67 pp. 

Kalil A. C., Lerner B. R., Kuroba C. H., Boog M. C. F., Oliveira M. C. M., 

Stefarini M. L. R., Sobrinho O. N. N., Lepper R. M., Philippi S. T. and 

Faria Z. (1979) Manual Básico de Nutrição. [Portuguese] 2 a Ediãço 

Estado de São Paulo, Sec.do Estado de Saúde, Inst.de Saúde, Seção 

Nutrição no 5, publição no.34, série E: 11-13, 69-70. 

Kavati R. (1997) “Embalagem e Comercialização.” [Portuguese] In: 

Anonáceas, Produção e Mercado. Edited by A. R. São José, Souza I. 

V. B., Morais O. M. and Rebouças T. N. H. Universidad Estadual do 

Sudoeste da Bahia, Vitória da Conquista, Bahia, Brasil : pp. 257-262. 

171

References 

Kavati R., Bueno S. C., Sales Tokunaga T., Nogueira E. M. C., Takassaki J. 

and Perioto N. W. (1997) A Cultura da Atemoia. [Portuguese] CATI, 

Campinas, São Paulo, Brasil: 22 pp. 

Kavati R. and Piza Jr. C. T. (1997) “Formação e Manejo do Pomar de Frutado-Conde, 

Atemoia e Cherimoia.” [Portuguese] In: Anonáceas, 

Produção e Mercado. Edited by A. R. São José, Souza I. V. B., Morais 

O. M. and Rebouças T. N. H. Universidade Estadual do Sudoeste da 

Bahia, Vitória da Conquista, Bahia, Brasil : pp. 76-83. 

Kessler P. J. A. (1993) “Annonaceae.” In: The Families and Genera of 

Vascular Plants. Vol.II, Flowering Dicotyledons. Edited by K. Kubitzi, 

Rohwer J. G. and Brittrich V. Springer-Verlag, Berlin, Germany. : pp. 

93-104. 

Kerharo J. and Adam J. G. (1974) La Pharmacopée Sénégalaise 

Traditionelle, Plantes Médicinales et Toxiques. [French] Vigot Fréres, 

Paris. 

Khan M. R., Kornine K. and Omoloso A. D. (1997) “Antibacterial Activity 

of Some Annonaceae Part I.” Fitoterapia, 69 (4) : 367-369. 

Kim D. H., Ma E. S., Suk K. D., Son J. K., Lee J. S. and Woo M. H. (2001) 

“Annomolin and Annocherimolin, New Cytotoxic Annonaceous 

Acetogenins from Annona cherimola Seeds.” Journal of Natural 

Products, 64 (4) : 502-506. 

Kim G. S., Zeng L., Alali F., Rogers L. L., Wu F. E., McLaughlin J. L. and 

Sastrodihardjo S. (1998 a) “Two New Mono-Tetrahydrofuran Ring 

Acetogenins, Annomuricin E and Muricapentocin, from the Leaves of 

Annona muricata.” Journal of the Natural Products, 61 : 432-436. 

Kim G. S., Zeng L., Alali F., Rogers L. L., Wu F. E., Sastrodihardjo S. and 

McLaughlin J. L. (1998 b) “Muricoreacin and Murihexocin C, 

Monotetrahydrofuran Acetogenins, from the Leaves of Annona 


Koesriharti (1992) “Edible Fruits and Nuts: Annona cherimola Mill. and 

Annona squamosa L.” In: Plant Resources of South-East Asia, 2. 

Edited by E. W. M. Verheij and Coronel R. E. Prosea, Bogor, 

Indonesia. : pp. 75-78. 

Kooiman P. (1967) “The Constitution of the Amyloid from Seeds of Annona 

muricata L.” Phytochemistry, 6 (12) : 1665-1673. 

172

References 

Kosiyachinda S. and Young R. E. (1975) “Ethylene Production in Relation to 

the Initiation of Respiratory Climacteric in Fruit.” Plant and Cell 

Physiology, 16 : 595-602. 

Kowalska M. T. and Puett D. (1990) “Potential Biomedical Applications for 

Tropical Fruit Products.” Tropical Garden Fruit World, 1 (4) : 126- 

127. 

Lahoz J. M., Gutierrez M., Sola M. M., Salto R., Pascual L., Martinez- 

Cayuela M. and Vargas A. M. (1993) “Ethylene in Cherimoya Fruit 

(Annona cherimola Mill.) Under Different Storage Conditions.” 

Journal of Agricultural Food Chemistry, 41 (5) : 721-23. 

Langason R. B. F., Akunyili D. N. and Akubue P. I. (1994) “A Preliminary 

Study of the Gastrointestinal Effects of Some Nigerian Medicinal 

Plants.” Fitoterapia, 65 (3) : 235-241. 

Laprode S. C. (1991) “Variación Estacional de Nutrimentos Foliares em 

Guanabana (Annona muricata L.).” [Spanish] Corbana (Costa Rica), 

15 (35) : 6-10. 

Leal F. (1990) “Sugar Apple.” In: Fruits of Tropical and Subtropical Origin 

Composition, Properties and Uses. Edited by S. Nagy, Shaw P. E. and 

Wardowski W. F. Florida Science Source, Inc., Lake Alfred, Fla. : pp. 

149-158. 

Lebeouf M., Legueut C., Cave A., Desconclois J. F., Foracs P. J. and 

Jacquemin H. (1981) “Alkaloids of the Annonaceae XIX. Alkaloids of 

Annona muricata.” [French] Planta Medica, 42 (1) : 37-44. 

Lederman I. E. And Bezerra J. E. (1997) “Situação Atual e Perspectivas de 

Anonáceas no Estado de Pernambuco.” In: Produção e Mercado. 

Edited by A. R. São José, Souza I. V. B., Morais O. M. and Rebouças 

T. N. H. Universidade Estadual do Sudoeste da Bahia, Vitória da 

Conauista, Bahia, Brasil. : pp. 173-177. 

Ledo A. S. (1992) Recomendações Básicas para o Cultivo da Gravioleira 

(Annona muricata L.). [Portuguese] Min. Agric. Ref. Agrar., 

EMBRAPA, CPAF - Acre, Rio Branco, AC : 10 pp. 

Ledo A. da S. and Fortes J. M. (1991) “Avaliação de Métodos de Enxertia 

para a Gravioleira em Viçosa-MG.” [Portuguese] Revista Brasileira de 

Fruticultura, 13 (1) : 63-66. 

173

References 

Lemos E. E. P. (2000 a) “Organogênese e Micropropagação em Anonáceas.” 

[Portuguese] In: Palestras - III Workshop Sobre Avanços na 

Propagação de Plantas Lenhosas. UFLA, Lavras-MG: pp. 4-21. 

Lemos E. E. P. (2000 b) “Graviola 'Gigante das Alagoas'.” [Portuguese] In: 

Novas Variedades Brasileiras de Frutas. Edited by L. C. Donadio 

Sociedade Brasileira de Fruticultura-SBF, Jaboticabal-SP : pp.96-97. 

Lemos E. E. P. and Baker D. A. (1998) “Shoot Regeneration in Response to 

Carbon Source on Internodas Explants of Annona muricata L.” Plant 

Growth Regulation, 25 : 105-112. 

Lemos E. E. P. and Blake J. (1994) “Leaf Abscission in Micropropagated 

Sugar Apple (Annona squamosa L.).” Edited by P. J. Lumsden, 

Nicholas J. R. and Davies W. J. Kluwer Academic Publisher, 

Dordrecht, the Netherlands : pp. 227-233. 

Lemos E. E. P. and Blake J. (1996 a) “Control of Leaf Abscission in Nodal 

Cultures of Annona squamosa L.” Journal of Horticultural Science, 71 

: 721-728. 

Lemos E. E. P. and Blake J. (1996 b) “Micropropagation of Juvenile and 

Adult Annona squamosa.” Plant Cell, Tissue and Organ Culutre, 46 : 

77-79. 

Lemos E. E. P. and Blake J. (1996 c) “Micropropagation of Juvenile and 

Mature Annona muricata L.” Journal of Horticulural Science, 71 : 

395-403. 

León J. (1987) Botánica de los Cultivos Tropicales. [Spanish] IICA, San 

José, Costa Rica. 

Li X. H., Hui Y. H., Rupprecht J. K., Liu Y. M., Wood K. V., Smith D. L., 

Chang C. J. and McLaughlin J. L. (1990) “Bullatacin, Bullatacinone, 

and Squamone, a New Bioactive Acetogenin, from the Bark of Annona 

squamosa.” Journal of Natural Products, 53 (1) : 81-86. 

Livera A. V. S. (1992) Desenvolvimento e Maturidade Fisiológica da 

Graviola (Annona muricata L.). [Portuguese] Tese de Mestrado, Univ. 

Fed. Pernambuco, Recife, Brasil : 80 pp. 

Livera A. V. S. and Guerra B. (1994) “Determinação da Maturidade 

Comercial da Graviola (Annona muricata) Através do Disco de 

Coleta.” [Portuguese] In: 13° Congresso Brasileiro de Fruticultura. 

Vol. 2. : pp. 603-604. 

174

References 

Lizana L. A. and Reginato G. (1990) “Cherimoya” In: Fruits of Tropical and 

Subtropical Origin: Composition, Properties and Uses. Edited by S. 

Nagy, Shaw P. E. and Wardowski W. F. Florida Science Source, Lake 

Alfred, Florida, USA : pp. 131-148. 

Lloyd B. F. and Jackson M. (1986) Plant Genetic Resources: An Introduction 

to their Conservation and Use. Edward Arnold, Baltimore, Maryland, 

USA : 146 pp. 

Loh C. L. (1975) “Fruits in Penninsular Malaysia.” Edited by J. T. Williams, 

Lamoureux C. H. and Wulijani S. South East Asian Plant Genetic 

Resources, LIPI, Bogor, Indonesia. 

Londershausen M., Leicht W., Lieb F. and Moeschler H. (1991 b) 

“Molecular Mode of Action of Annonins.” Pesticide Science, 33 : 427- 

438. 

Londershausen M., Leicht W., Weiss H. and Lieb F. (1991 a) “Annonins - 

Mode of Action of Acetogenins from Annona squamosa.” Pesticide 

Science, 30 (4) : 443-445. 

Lucas A. P. (1994) “O Cultivo da Pinha Traz Lucro em Dólar.” [Portuguese] 

Manchete Rural, 82 : 19-21. 

MacLeod A. J. and Pieris N. M. (1981) “Volatile Flavour Components of 

Soursop (Annona muricata).” Journal of Agricultural and Food 

Chemistry, 29 (3) : 488-490. 

Maeda U., Hara N., Fujimoto Y., Srivastava A., Gupta Y. K. and Sahai M. 

(1993) “N-Fatty Acyl Tryptamines from Annona reticulata.” 


Mahdeem H. (1990) “Best of Annonas.” Tropical Fruit World, 1 (4) : 110- 

114. 

Manica I. (1997) “Taxionomia, Morfología and Anatomia.” [Portuguese] In: 

Proceedings of I Brazilian Symposium on Annonaceous. Edited by A. 

R. São José, Souza I. V. B., Morais O. M. and Rebouças T. N. H. 

Universidade Estadual do Sudoeste da Bahia, Depto de Fitotecnia e 

Zootenia. Vitória da Conauista, Bahia, Brasil : pp. 20-35. 

Mansour K. M. (1997) “Current Status of Annonaceae in Egypt.” Mesfin 

Newsletter, 3 (1) : 5-10. 

175

References 

Marin Acosta J. C. (1973) “Lista Preliminar de Plagas de Annonaceae, 

níspero (Achras zapota L.), Guajava (Psidium guajava L.) en 

Venezuela.” [Spanish] Agronomia Tropical, 23 : 205-216. 

Marler J. E., George A. P., Nissen R. J. and Andersen P. J. (1994) 

“Miscellaneous Tropical Fruits - Annonas.” In: Handbook of 

Environmental Physiology of Fruit Crops. Vol. II. Subtropical and 

Tropical Crops. Edited by B. C. Schaffer and Andersen P. C. CRC 

Press, Boca Raton, Florida, USA : pp. 200-206. 

Martinez G., Serrano M., Pretel M. T., Riqueline F. and Romojas F. (1993) 

“Ethylene Biosynthesis and Physico-chemical Changes During Fruit 

Ripening of Cherimoya (Annona cherimola Mill).” Journal of 

Horticultural Science, 68 : 477-483. 

Medina N., Del Rocio M., Cruz Castillo G., Guerra Bustos, Luna Rodriguez 

M. and Dominguez Martinez V. (1999) “Peroxidases as Variation 

Tracers Within Annona muricata Individuals.” In: Memórias del II 

Congreso Internacional de Anonáceas. Universad de Ciencias y Artes 

del Estado de Chiapas, Tuxtla Gutiérrez, Chiapas, México : pp. 81. 

Melo G. S., Gonzaga Neto L. and Moura R. J. M. (1983) “Cultivo da 

Gravioleira.” Empresa Pernambucana de Pesquisa Agropecuária-IPA, 

Recife, Pernambuco State, Brasil : 4 pp. (Instruçães Técnica, 13) 

Mengel K. and Kirkby E. A. (1987) Principles of Planta Nutrition. 

International Potash Institute, Bern, Switzerland : 687 pp. 

Mishra A., Dogra J. V. V., Singh J. N. and Jha O. P. (1979) “Post Coital 

Antifertility Activity of Annona squamosa and Ipomoea fistula.” 

Planta Medica, 35 (3) : 283-285. 

Monastra F. (1997) “Current Status of Annonaceae in Italy.” Mesfin 

Newsletter, 3 (1) : 11-12. 

Monteiro A. R., Martinelli N. M., Ferraz L. C. C. B. and Lordello R. R. A. 

(1978) “Nematóides Parasitos de Plantas na Região de Ilha Solteira, 

São Paulo.” [Portuguese] In: III Reunião da Sociedade Brasileira de 

Nematologia. Mossoró, Rio Grande do Norte : pp. 35-38. 

Moran R., Bautista C., Bermudez R., Calzada B. and Chavez F. (1972) 

“Comparativo de Dos Diametros de Patron y Cinco Tipos de Injertos 

en la Propagación del Chirimoyo (Annona cherimola Mill.).” [Spanish] 

Anuario Cientifico La Molina, 10 (3/4) : 158-176. 

176

References 

Moreno Andrade R., Luna Cazáres L. and González Esquinca A. R. (1999) 

“Estudios Sobre la Germinación de Annona lutescens.” [Spanish] In: 

Memorias del II Congreso Internacional de Anonáceas. Universad de 

Ciencias y Artes del Estado de Chiapas, Tuxtla Gutiérrez, Chiapas, 

México : 82 pp. 

Mororó R. C., Freire E. S. and Sacramento C. K. (1997) “Processamento da 

Graviola para Obtenção de Polpa.” [Portuguese] In: Anonáceas: 

Poduoção e Mercado (Pinha, Graviola, Atemóia e Cherimólia). Edited 

by A. R. São José Boas I. V., Morais O. M. and Rebouças T. N. H. 

Universidade Estadula do Sudoeste da Bahia, Vitória da Conquista, 

Brasil : pp. 263-274. 

Morton J. F. (1987) Fruits of Warm Climates. J. F. Morton Publisher, Miami, 

Florida, USA. 

Mosca J. L., Alves R. E., Filgueiras H. A. C. and Oliveira J. F. (1997 b) 

“Determination of Harvest Index for Soursop Fruits (Annona muricata 

L.).” In: Memorias, Congresso Internacional de Anonaceaes. 

Universidad Autonoma Chapingo (UAC), Chapingo, México : pp. 315- 

322. 

Mosca J. L., Assis J. S., Alves R. E., Filgueiras H. A. C. and Batista A. F. 

(1997 a) “Physical, Physical-Chemical and Chemical Changes During 

Growth and Maturation of Sugar Apple (Annona squamosa L.).” In: 

Memorias, Congresso Internacional de Anonaceaes. Universidad 

Autonoma Chapingo (UAC), Chapingo, México : pp. 304-314. 

Moura J. V. (1988) “A Cultura da Graviola em Áreas Irrigadas.” 

[Portuguese] In: Uma nova opção. Fortaleza, DNOCS : 42 pp. 

Mowry H., Toy L. R. and Wolfe H. S. (1941) “Miscellaneous Tropical and 

Sub-Tropical Florida Fruits.” Agriculture Extension Service, 

Gainesville, Florida, Bulletin, 109 : 11-21. 

Mukhopadhhyay G., Mukherjee B., Patra A., Ghosh R., Roychowdhury P. 

and Lowe P. (1993) “Refined NMR and x-Ray Crystallographic 

Studies with a Diterpene from Annona squamosa.” Fitoterapia, 64 (1) : 

7-10. 

Myint S. H., Cortes D., Laurens A., Hocquemiller R., Leboeuf M., Cavé A., 

Cotte J. and Quéro A. M. (1991) “Solamin, a Citotoxic Mono- 

Tetrahydrofueranic γ-Lactone Acetogenin from Annona muricata 

Seeds.” Phytochemistry, 30 (10) : 3335-3338. 

177

References 

Nair S., Gupta P. K. and Mascarenhas A. F. (1983) “Haploid Plants from In 

Vitro Anther Culture of Annona squamosa L.” Plant Cell Reports, 2 : 

198-200. 

Nair S., Shirgurkar M.V. and Mascarenhas A. F. (1986) “Studies on 

Endosperm Culture of Annona squamosa L.” Plant Cell Reports, 5 : 

132-135. 

Nakasone H. Y. and Paull R. E. (1998) “Annonas.” In: Tropical Fruits. 

Edited by H. Y. Nakasone and Paull R. E. CAB International, 

London,UK. : pp. 45-75. 

NAS (1975) Underexploited Tropical Plants with Promising Economic 

Value. National Academy of Sciences, Washington DC, USA. 

Nascimento A. S., Coutinho C. de C., Ferreira F. R., Santos Filho H. P., 

Cunha M. M. and Junqueira N. T.V. (2000) Manga Fitossanidade. 

[Portuguese] Embrapa-CTT, Brasília : 104 pp. 

Nava-Díaz C., Osada-Kawasoe S., Rendon-Sánchez G. and Ayala-Escobar 

V. (2000) “Organismos Asociados a Cherimoyo (Annona cherimola 

Mill.) en Michoacán, México.” [Spanish] Agrociencia, 34 : 217-226. 

Navia V. M. G. and Valenzuela J. B. (1978) “Sintomatologia de Deficiências 

en Chirimoya (Annona cherimola Mill.) cv. Bronceada.” [Spanish] 

Agricultura Técnica (Santiago), 38 : 9-14. 

Ndambuki B. M. (1991) “In Situ Conservation and Monitoring of Rangeland 

Genetic Resources.” In: Crop Genetic Resources of Africa. Edited by 

F. Attere, Zedan H., Ng N. Q. and Perrino P. 

IBPGR/UNEP/IITA/CNR, Rome, Italy : 656 pp. 

Ng N. Q. (1991) “Long-term Seed Conservation.” In: Crop Genetic 

Resources of Africa. Edited by F. Attere Zedan H. Ng N. Q. Perrino P. 

IBPGR/UNEP/IITA/CNR, Rome, Italy : pp. 135-148 

Nieto-Angel D., São José A. R., Rebouças T. N. H., Bonfim M. P. Y. and 

Brandão A. L. S. (1999) “Hongos Asociados a Frutos de Guanábana 

(Annona muricata L.) in Bahia, Brazil.” [Portuguese] In: II 

International Congress of Annonaceae. Universad de Ciencias y Artes 

del Estado de Chiapas, Tuxtla Gutiérrez, Chiapas, México : pp. 91-92. 

Nomura K., Terai H., Yabe K., Maeda M., Rahman M. S. M., Yoshida M. 

and Yonemoto J. Y. (1997) “Comparison of Changes in Sugar and 

Ethylene Production of Cherimoya Fruit Grown at Different Seasons.” 

Journal of Horticulural Science 72 : 617-622. 

178

References 

Nonfon M., Lieb F., Moeschler H and Wendisch D (1990) “Four Annonins 

from Annona squamosa.” Phytochemistry, 29 (6) : 1951-1954. 

Noonan J. C. (1953) Review of Investigations on the Annona species. 

Proceedings Florida State Horticulture Society : pp. 205-210. 

NRC (1989) Lost Crops of the Incas. Little Known Plants of the Andes with 

Promise for Worldwide Cultivation. National Academy Press, 

Washington DC, USA. 

Nunes R. E. de F. (1997) “The Actual Status of Cherimoya Cultivation in 

Madeira Island.” In: Second MESFIN on Plant Genetic Resources. 

Madeira, Portugal. : pp. 135-151. 

Ochse J. J., Soule Jr. M. J., Dijkman M. J. and Wehlburg C. (1974) “Otros 

Cultivos Frutales.” [Spanish] In: Cultivo y Mejoramiento de Plantas 

Tropicales y Subtropicales. Editorial Limusa, México. : pp.587-818. 

Oliver-Bever B. (1986) Medicinal Plants in Tropical West Africa. Cambridge 

University Press, Cambridge, UK. 

Oliveira M. A. S., Genú P. J. de C., Junqueira N. T. V. and Pinto A. C. de Q. 

(1992) Pragas da Gravioleira no Cerrado. [Portuguese] Embrapa 

Cerrado, Brasília : 11 pp. (Serie Documentos, 41). 

Oliveira M. A. S., Icuma I. M., Alves R. T., Junqueira N. T. V. And Pinto A. 

C. de Q. (2001) “Insectos-praga e seu Controle.” In: Graviola, 

Produção. Embrapa Informação Tecnológica, Brasilia, DF. : pp. 34-38. 

Padilla I. M. G. (1997) Micropropagation of Cherimoya (Annona cherimola 

Mill.) cv. Fino de Jete. University of Malaga, Spain : 334 pp. 

Page P. E. (1984) Tropical Tree Fruits for Australia. Information Series 

QI83018. Queensland Department of Primary Industries, Brisbane, 

Australia. 

Pal D. K. and Kumar P. S. (1995) “Changes in the Physio-chemical and 

Biochemical Compositions of Custard Apple (Annona squamosa L.) 

Fruits During Growth, Development and Ripening.” Journal of 

Horticultural Science, 70 (4) : 569-572. 

Palacios Rangel M. I. and Cano Gracia G. V. (1997) “La Comercialización 

de Anonáceas en México.” [Spanish] In: Memorias del Congreso 

Internacional de Anonáceas. Universad Autonoma Chapingo (UAC), 

Chapingo, México : pp. 68-91. 

179

References 

Palma T., Aguilera J. M., Stanley D. W. (1993) “A Review of Postharvest 

Events in Cherimoya.” Postharvest Biology and Technology, 2 : 187- 

208. 

Pandey G. P. and Varma B. K. (1977) “Annona Seed Powder as a Protectant 

of Mung Against Pulse Beetle, Callosobruchus maculatus (Fabr.).” 

Bulletin of Grain Technology, 15 (2) : 100-104. 

Pareek O. P. (1985) “Fruit Crops.” In: Efficient Management of Dryland 

Crops. Research Institute for Dryland Agriculature, Hyderbad, India. : 

pp. 293-408. 

Pascual L., Perfectti F., Gutierrez M. and Varga A. M. (1993) 

“Characterizating Isozymes of Spanish Cherimoya Cultivars.” 

HortScience, 28 (8) : 845-847. 

Paull R. E. (1982) “Postharvest Variation in Composition of Soursop 

(Annona muricata L.) Fruit in Relation to Respiration and Ethylene 

Production.” Journal of American Society of Horticultural Science, 107 

: 582-585. 

Paull R. E. (1990) “Soursop Fruit Ripening - Starch Breakdown.” Acta 


Paull R. E., Deputy J. and Chen N. J. (1983) “Changes in Organic Acids, 

Sugars, and Headspace Volatile During Fruit Ripening of Soursop 

(Annona muricata L.).” Journal of American Society of Horticultural 

Science, 108 : 931-934. 

Pawshe Y. H., Patil B. N. and Patil L. P. (1997) “Effect of Pre-germination 

Seed Treatment on the Germination and Vigour of Seedlings in Custard 

Apple (Annona squamosa L.).” Annals of Plant Physiology, 2 (11) : 

150-154. 

Payumo E. M., Pilac L. M. and Maniquis P. L. (1965) “The Preparation and 

Storage Properties of Canned Guaybano (Annona muricata L.) 

Concentrate.” Philippine Journal of Science, 94 : 161-169. 

Peña J. E. and Bennett F. D. (1995) “Arthropods Associated with Annona 

spp. in the Neotropics.” Florida Entomologist, 78 (2) : 329-349. 

Pérez de Oteyza M. A. and Farré J. M. (1999) “El Banco Español de 

Germoplasma de Chirimoyo.” [Spanish] In: Parámetros Estudiados y 

Variabilidad. VIII Congreso Nacional de Ciencias Hortícolas, Murcia. 

: pp. 7-12. 

180

References 

Perfectti F. (1995) Estudio de Marcadores Genéticos en Chirimoyo, con 

Aplicación a al Identificación Varietal. Evaluación de los Recursos 

Genéticos. [Spanish] Tesis doctoral. Universidad de Granada. 

Philipov S., Kandé K. M. and Machev K. K. (1995) “Alkaloids of Annona 

sengelensis.” Fitoterapia, 66 (3) : 275-276. 

Philipov S., Machev K. and Tsankova E. (1994) “Liriodenine from Annona 

muricata Seeds.” Fitoterapia, 65 (6) : 555. 

Pinto A. C. de Q. (1975 a) “Produção e Utilização da Graviola e Pinha.” 

[Portuguese] In: Semi-Anual Field Progress Report. Centro de 

Pesquisa Desenvolvimento CEPED, Bahia, Brazil : pp. 1-18. 

Pinto A. C. de Q. (1975 b) “Influência de Hormônio Sobre o Poder 

Germinativo de Sementes de Graviola (Annona muricata L.).” 

[Portuguese] In: Anais do III Congresso Brasileiro de Fruticultura, V. 

II. Sociedade Brasileira de Fruticultura, Rio de Janeiro, Brasil : pp. 

415-421. 

Pinto A. C. De Q. (1978) Influência do ácido giberélico, do Permanganato 

de Potássio e da Embalagem de Polietileno na Conservação e 

Qualidade da Banana ‘Prata’. [Portuguese] Ms. Thesis, Escola 

Superior de Agricultura de Lavras, Lavras, Minas Gerais State, Brazil. : 

80 pp. 

Pinto A. C. de Q. and Ramos V. H. V. (1997 a) “Melhoramento Genético da 

Graviola.” [Portuguese] In: Proceedings of the I Brazilian Symposium 

on Annonaceous. Edited by A. R. São José, Souza I. V. B., Morais O. 

M. and Rebouças T. N. H. UESB, Vitória da Conquista, Brazil : pp.55- 

60. 

Pinto A. C. de Q. and Ramos V. H. V. (1997 b) “Graviola: Formação do 

Pomar e Tratos Culturais. Anonáceas, Produção e Mercado.” In: 

Anonáceas, Produção e Mercado. [Portuguese] Edited by A. R. São 

José, Souza I. V. B., Morais O. M. and Rebouças T. N. H. 

Universidade Estadual do Sudoeste da Bahia, Vitória da Conquista, 

Bahia, Brasil : pp. 94-104. 

Pinto A. C. de Q. and Ramos V. H. V. (1999) Polinização Artificial da 

Graviola. [Portuguese] Guia Técnico do Produtor Rural, N° 20. 

Embrapa Cerrados, Brasília, Brasil. 

181

References 

Pinto A. C. de Q., Ramos V. H. V. and Rodriques A. A. (2001) “Formação 

do Pomar.” In: Graviola, Produção. [Portuguese] Edited by Oliveira 

M. A. A. Embrapa Informação Tecnológica, Brasilia, DF. : pp. 22-25. 

Pinto A. C. de Q. and Silva E. M. (1994) Graviola para Exportação: 

Aspectos Técnico da Produção [Portuguese] FRUPEX, Min. 

Agricultura, do Abastecimento e da Reforma Agrária, Sec. de 

Desenvolvimento Rural SDR, Prog. De Apoio à Prod. E Export. De 

Frutas, Hortaliças, Flores e Plantas Ornamentais. : 41 pp. 

Pinto A. C. de Q. and Silva E. M. (1996) Graviola para Exportação, 

Aspectos Técnicos da Produção. [Portuguese] Embrapa/SPI, Brasília, 

Brasil 

Pinto E. T. (1991) Recomendaciones Tecnicas para el Cultivo del 

Guanabano en las Region Altantica de Costa Rica. [Portuguese] 

Ministerio da Agricultura y Ganadeira, Siquirres. : 22 pp. 

Pittman E. C. (1956) La Chirimoya. [Spanish] Circ. Est. Exp. Agric. La 

Molina, Peru. 71 : pp. 26. 

Piza Jr. C. T. and Kavati R. (1997) “Situação Atual e Perspectivas da Cultura 

de Anonáceas no Estado de São Paulo.” [Portuguese] In: Proceedings 

of the I Brazilian Symposium on Annonaceous. Edited by A. R. São 

José Souza I. V. B., Morais O. M. and Rebnouças T. N. H. UESB, 

Vitória da Conquista, Brazil : pp. 184-195. 

Plaza J. L., Rossi S. and Calvo M. L. (1993) “Inhibitory Effects of the 

Ethylene Chemisorption on the Climateric of Cherimoya Fruit in 

Modified Atmosphere.” Acta Horticulturae, 343 : 181-183. 

Ponce H. L. and Vidal H. (1981) “Asistência Eécnica Sobre o Cultivo de la 

Guanábana.” [Spanish] In: Memória de III Congreso Nacional de 

Fruticultura. Guadalajara, México : 171 pp. 

Popenoe W. (1939) Importantes Frutas Tropicais. [Portuguese] Washington, 

DC, União Panamericana, Depto de Cooperação Agrícola. : 29 pp. 

(série Agricultura, 81-82). 

Popenoe W. (1952) “Central American Fruit Culture.” CEIBA, 1 (5) : 299- 

304. 

Popenoe W. (1974 a) “The Anonaceous Fruits; The Cherimoya.” In: Manual 

of Tropical and Subtropical Fruits, Facsmile of the 1920 ed. Hafner 

Press, a Division of Macmillan Publishing Co, Inc., New York, Collier- 

Macmillan Publishers, London, Chapter 5 : 161-189. 

182

References 

Popenoe W. (1974 b) “The Anonaceous Fruits; The Soursop.” In: Manual of 

Tropical and Subtropical Fruits, Facsmile of the 1920 ed. Hafner 

Press, a Division of Macmillan Publishing Co, Inc., New York, Collier- 

Macmillan Publishers, London, Chapter 5 : 182-186. 

Prasada R. R. and Rao S. D. T. (1984) “Oil from Custard Apple (Annona 

squamosa) Seed.” Indian Food Industry, 3 : 163-4. 

PROCIANDINO (1997) Estudio Global para Identificar Oportunidades de 

Mercado de Frutas y Hortalizas de la Región Andina-PROCIANDINO, 

Fruthex. [Spanish] Quito, Equador : 120 pp. 

Purohit A. (1995) “Annonaceous Fruits.” In: Handbook of Fruit Science and 

Technology: Production, Composition, Storage and Processing. Edited 

by D. K. Salunkle and Kadam S. S. M. Dekker, New York, USA : pp. 

377-385. 

Purseglove J. W. (1968) “Other Useful Products: Annonaceae.” Tropical 

Crops, Dicotyledons. Longman, London, UK. : pp.624-625. 

Qadri S. S. H. and Rao B. B. (1977) “Effect of Combining some Indigenous 

Plant Seed Extracts Against House-hold Insects.” Pestiscides, 11 (12) : 

21-23. 

Rao V. G., Desai M. K. and Kulkarni N. B. (1962?) “A New Phytophthera 

Fruit Rot of Annona senegalensis from India.” Plant Disease Reporter, 

46 : 874-876. 

Rasai S., George A. P., Kantharajah A. S. (1995) “Tissue Culture of Annona 

spp. (Cherimoya, Atemoya, Sugar Apple and Soursop): A Review.” 

Scientia Horticulturae, 62 : 1-14. 

Rasai S., Kantharajah A. S. and Dodd W. A. (1994) “The Effect of Growth 

Regulators, Source of Explants and Irradiance on In Vitro Regeneration 

Atemoya.” Australian Journal of Botany, 42 : 333-340. 

Rebollar-Alviter A., Domínguez Alvarez J. L., Nieto-Angel D., Nava-Diaz 

C. and Delgadillo Sánchez F. (1997) “Principales Plagas y 

Enfermedades de las Anonaceas en México.” [Spanish] In: 

Proceedings of the I Brazilian Symposium on Annonaceous. Edited by 

A. R. São José, Souza I. V. B., Morais O. M. and Rebnouças T. N. H. 

UESB, Vitória da Conquista, Brazil : pp. 222-228. 

Rego F. A. O., Alves R. E., Lima E. D. P. A., Lima C. A. A., Silva H. and 

Silva A. Q. (1979) “Caracterização Física e Química de Diferentes 

183

References 

Frutos da Família Annonaceae.” [Portuguese] In: Anais do 10° 

Congresso Brasileiro de Fruticultura. : pp. 493-497. 

Requena J.C. (1998) El Mercado Español de la Cherimoya: Situación Atual 

y Perspectivas. [Spanish] Departamento de Economia y Sociología 

Agraria, CIDA, Granada, España. : pp. 79-105. 

Richardson A. C. and Anderson P. (1993) “A Detailed Evaluation of 

Cherimoya Cultivars.” The Orchadist, 66 (6) : 28-31. 

Richardson A. C. and Anderson P. A. (1996) “Flowering Date Affects 

Pollination of Cherimoya.” Orchadist, 69 (1) : 49-52. 

Rieser M. J., Gu Z. M., Fang X. P., Zeng L., Wood K. V. and McLaughlin J. 

L (1996) “Five Novel Mono-tetrahydrofuran Ring Acetogenins from 

the Seeds of Annona muricata.” Journal of Natural Products, 59 : 100- 

108. 

Roblot F., Laugel T., Leboeuf M., Cavé A. and Laprévote O. (1993) “Two 

Acetogenins from Annona muricata Seeds.” Phytochemistry, 34 (1) : 

281-285. 

Román E. and Damián R. (1999) “Collección y Evaluación de Selecciones y 

Cultivares de Chirimoyo.” [Spanish] Memórias del Congreso 


de Chiapas, Tuxtla Gutiérrez, Chiapas, Méxo : pp. 188-192. 

Ronning C. M., Schnell R. J., Gazit S. (1995) “Using Randomly Amplified 

Polymorphic DNA (RAPD) Markers to Identify Annona Cultivars.” 

Journal of the American Society for Horticultural Science, 120 (5) : 

726-729. 

Rosell P., Herrero M. and Galan Saúco V. (1999) “Pollen Germination of 

Cherimoya (Annona cherimola Mill.), In Vivo Characterization and 

Optimization of In Vitro Germination.” Scientia Horticulturae 81 (3) : 

251-165. 

Rungsimanop C., Suksri A. and Srinukul S. (1987) “Some Irrigation Methods 

which Influence the Growth of Custard Apple and Papaya when 

Intercropped in Northeast Thailand.” Phytochemistry, 30 (10) : 3335- 

3338. 

Rupprecht J. K., Hui Y. H. and McLaughlin J. L. (1990) “Annonaceous 

Acetogenins: A Review.” Journal of Natural Products, 53 (2) : 237- 

278. 

184

References 

Saavedra E. (1977) “Influence of Pollen Grain Stage at the Time of Hand- 

Pollination as a Factor of Fruit Set of Cherimoya.” HortScience, 12 (2) 

: 117-119. 

Saavedra E. (1979) “Set and Growth of Annona cherimola Mill. Fruit 

Obtained by Hand-Pollination and Chemical Treatments.” Journal of 

American Horticultural Science, 104 (5) : 668-673. 

Sadhu M. K. and Ghosh S. K. (1976) “Effects of Different Levels of 

Nitrogen, Phosphorus and Potassium on Growth, Flowering, Fruiting 

and Tissue Composition of Custard Apple (Annona squamosa L.).” 

Indian Agricultural, 20 (4) : 297-301. 

Sahpaz S., González M. C., Hocquemiller, R., Zafra-Polo M. C. and Cortés 

D. (1996) “Annosenegalin and Annogalene: Two Cytotoxic Mono- 

Tetrahydrofuran Acetogenins from Annona senegalensis and Annona 

cherimolia.” Phytochemistry, 42 (1) : 103-107. 

Salluja A. K. and Santani D. D. (1990) “Phytochemical Study of Annona 

squamosa.” Fitoterapia, 61 (4) : 359-360. 

Salunkhe D. K. and Desai B. B. (1984) “Custard Apple and Jujube.” In: 

Postharvest Biotechnology of Fruits. II. CRS Press, Boca Raton : pp. 

133-135. 

Samuel R., Pinsker W., Balasubhramaniam S. and Morawetz W. (1991) 

“Allozyme Diversity and Systematics in Annonaceae - A Pilot 

Project.” Plant Systematics and Evolution, 178 : 125-134. 

Sánchez-Nieva F. (1953) “In: Bueso, CE (1980) Soursoup, Tamarind and 

Chironja.” In: Tropical and Subtropical Fruits Composition, Properties 

and Uses. Edited by S. Nagy and Shaw F. Avi Publishing Inc, 

Wetsport, Connecticut, USA : pp. 375-387. 

Sánchez-Nieva F. (1970) “Frozen Soursop Puree.” Journal of Agricultural 

University of Puerto Rico, 54 : 220-236. 

Sanewski G. M. (1991) Custard apple - Cultivation and Crop Protection. 

Information Séries QI90031. Queensland Department of Primary 

Industry, Brisbane, Australia 

Santos C. R. (1997) “Irrigação em Anonáceas.” [Portuguese] In: Proceedings 

of the I Brazilian Symposium on Annonaceous. Edited by A. R. São 

José, Souza I. V. B., Morais O. M. and Rebouças T. N. H. UESB, 

Vitória da Conquista, Bahia, Brazil : pp. 105-117. 

185

References 

Sanyal D. (1924) Vegetable Drugs of India. Dehra Dun, India. 

Sarabia E. R. and Alatorre L. M. (1997) “Las Annonas en el México 

Prehispanico.” [Spanish] In: Memorias del Congreso Internacional de 

Anonáceas. Universidad Autonoma Chapingo (UAC), México : pp. 

169-186. 

Scheldeman X. (2002) “Literature Review.” In: Distribution and Potential of 

Cherimoya (Annona cherimola Mill.) and Highland Papayas 

(Vasconcellea spp.) in Ecuador. Thesis, Faculty of Agricultural and 

Applied Biological Sciences, University of Ghent, Belgium. : pp. 5-62. 

Scheldeman X. and Van Damme P. (1999) “Promising Cherimoya (Annona 

cherimola Mill.).” In: Proceedings of the First International 


and Scheldeman X. Evangraf, Porto Alegre. Acta Horticulturae, 497 : 

173-180. 

Schnell R. J., Knight R. J., Harkins D. M. and Zill G. (1994) “Eliminating 

Zygotic Seedlings in 'Turpentine' Mango Rootstock Populations by 

Visual Roguing.” HortScience, 29 : 319-320. 

Sharma R. D. (1973) “Plant Parasitic Nematodes in the São Francisco Valley, 

Pernambuco, Brazil.” Nematropica, 3 (2) : 51-54. 

Sharma R. D. (1977) “Nematodes of the Cocoa Region of Bahia, Brazil; 

Nematodes Associated with Tropical Fruit Trees.” In: II Reunião da 

Sociedade Brasileira de Nematologia. Piracicaba, São Paulo, Brazil : 

pp. 109-113. 

Sharma R. D., Pinto A. C. de Q. and Loof P. A. A. (1985) “Declínio da 

Gravioleira (Annona muricata L.) Sob Condições de Cerrados do 

Distrito Federal.” [Portuguese] Nematologia Brasileira, 9 : 38. 

Silva A. Q. and Silva H. (1997) “Nutrição e Adubação em Anonáceas.” 

[Portuguese] In: Anonáceas, Produção e Mercado (Pinha, Graviola, 

Atemó e Cherimólia). Edited by A. R. São José, Souza I. V. B., Morais 

O. M. and Rebouças T. N. H. Universidade Estadual do Sudoeste da 

Bahia, Vitória da Conquista, Bahia, Brasil : pp. 118-137. 

Silva E. M. da, Pinto A. C. Q. and Azevedo J. A. (1996) Manejo da 

Irrigação e Fertirrigação na Cultura da Mangueira. [Portuguese] 

Embrapa Cerrados, Brasília : 77 pp. (Serie Documentos 61). 

Silva H., Silva A. Q. da, Cavalcante A. T. and Malavolta E. (1984) 

“Composição Mineral das Folhas de Algumas Fruteiras do Nordeste.” 

186

References 

[Portuguese] In: Anais do VII Congresso Brasileiro de Fruticultura. 

Sociedade Brasileira de Fruticultura, Florian´polis, Brazil 1 : 320-325. 

Singh S., Krishnamurthy S. and Katyol S. L. (1967) Fruit Culture in India. 

ICAR, New Delhi, India. 

Singh S. P. (1992 a) Fruit Crops for Wasteland. Scientific Publisher, 

Jodhpur, India. 

Smith N. J. H., Williams J. T., Plucknett D. L. and Talbot J. P. (1992) 

Tropical Forests and Their Crops. Cornell University Press, Ithaca, 

NY, USA. 

Smith R. W. (1997) “ICUC's Role in Developing an Action Plan for 

Promoting Under-utilised Crops through Co-operation amongst 

National Institutions.” 2nd MESFIN Meeting on Plant Genetic 

Resources and Fruit Production. 5-8 August, 1997. Madeira. 

SPT-TCA (1999) Impactos Actuales y Potenciales de las Enfermedades de 

los Cultivos Perennes de la Amazonia y Posibilidades de Control para 

el Desarrollo Sostenible de la Region. [Spanish] Tratado de 

Cooperación Amazonica, Venezuela. : pp. 79-84. 

Standley P. C. (1930) “A Second List of the Trees of Honduras.” Tropical 

Woods, 21 : 9-41. 

Sturrock D. (1959) Fruits for Southern Florida. South Eastern Printing Co., 

Stuart, FL, USA. 

Sundararaj D. D. and Balasubramanyam G. (1959) Guide to the Economic 

Plants of South India. Madras, India. 

TCA, Tratado de Cooperacion Amazonica (undated) “Guanabana Annona 

muricata L.” [Spanish] In: Fructales y Hortalizas Promisorios de la 

Amazonia. Lima, Peru : pp. 130-136. 

Thakur D. R. and Singh R. N. (1965) “Studies on Pollen Morphology, 

Pollination and Fruit Set in Some Annonas.” Indian Journal of 

Horticulture, 22 (1) : 10-18. 

Tijero R. F. (1992) El Cultivo del Chirimoyo en el Peru. [Spanish] Ediciones 

Fundeagro, Lima, Peru. .108 pp. 

Torres W. E. and Sánchez L. A. (1992) Fruticultura Colombiana, 

Guanábano. [Spanish] Instituto Colombiano Agropecuario. ICA, 

Manual de Asistencia Técnica 57, Bogotá. 

187

References 

Tsay L. M. and Wu M. C. (1989) “Studies on the Postharvest of Sugar 

Apple.” Acta Horticulturae, 258 : 287-294. 

Tsay L. M. and Wu M. C. (1990) “Studies on the Physio-chemical Properties 

of Postharvest Sugar Apple.” Acta Horticulturae, 269 : 241-247. 

Tustin S. (1997) “Fisiología della Potatura e dell`allevamento come Regolare 

lo Sviluppo Vegeto-Produttivo di Rami e Branche.” [Portuguese] 

Rivista de Frutticoltura e di Ortofloricoltura, 3 : 53-55. 

Umme. A., Aubi. B. A., Salmah Y., Junainah A. H. and Jamilah B. (1997) 

“Characteristics of Soursop Natural Puree and Determination of 

Optimum Conditions for Pasteurization.” Food Chemistry, 58 (1-2) : 

119-124. 

Undurraga P. L., Olaeta J. A. and Marnich M. L. (1995) “Effect of Increasing 

Nitrogen Fertilizer Rates on Cherimoya (Annona cherimola Mill.) cv. 

Concha Lisa Fruits in Refrigerated Storage. Harvest and Postharvest 

Technologies for Fresh Fruits and Vegetables.” In: Proceedings of 

American Society of Agricultural Engineers (ASAE). Guanajuanto, 

Mexico : pp. 405-413. 

Uphof J. C. Th. (1959) Dictionary of Economic Plants. H. R. Englemann, 

New York, USA. 

Van Damme P. and Scheldeman X. (1999) “Commercial Development of 

Cherimoya (Annona cherimola Mill.) in Latin America.” In: 


Edited by V. Van Damme Van Damme P. Scheldeman X. ISHS, Loja, 

Ecuador. Acta Horticulturae, 497 : 17-28. 

Venkataratnam L. (1959) “Floral Morphology and Blossom Biology Studies 

in Some Annonaceae.” Indian Journal of Agronomy Science, 29 : 69- 

75. 

Villachica H., Carvalho J. E. U., Müller C. H., Díaz S. C. and Almanza M. 

(1996) Frutales y Hortalizas Promisorios de la Amazonia. [Spanish] 

Tratado de Cooperación Amazonica, Secretaria Pro-Tempore, Lima, 

Peru. [Guanabana, Annona muricata L., pp. 130-136.] 

Villachica V. (1996) Frutales y Hortalizas Promisorios de la Amazonia. 

[Spanish] Tratado de Cooperación Amazonica, Secretaria Pro- 

Tempore, Lima, Peru: 367 pp. 

188

References 

Viñas R. C. (1972) “Atis (Annona squamosa L., Annonaceae). Cultural 

Directions for Philippine Agricultural Crops.” Publ. Aff. Off. Press, 

Bur. Plant Industry, Manila. Fruits, 1 : 31-36. 

Viteri P. F., León J. F. And Ortega C. (1999) “Evaluation of Shoot Inducers 

in Cherimoya (Annona cherimola Mill.).” In: Proceedings of the First 

International Symposium on Cherimoya. Edited by V. Van Damme, 

Van Damme P. and Scheldeman X. Evangraf, Porto Alegre. Acta 


Vogt K. (1995) A Field Worker´s Guide to the Identification, Propagation 

and Uses of Common Trees and Shrubs of Dryland Sudan. SOS Sahel 

International, UK. 

Von Maydell H. J. (1986) Trees and Shrubs of the Sahel, Their 

Characteristics and Uses. Russdorf: TZ-Verlagsgellschaft, Germany. 

Wagner H., Reiter M. and Fersti W. (1980) “New Drugs with Cardiotonic 

Activity. I Chemistry and Pharmacology of the Cardiotonic Active 

Principle of Annona squamosa.” [German] Planta Medica, 40 (1) : 77- 

85. 

Weber G. F. (1973) Bacterial and Fungal Diseases of Plants in the Tropics. 

University of Florida Press, Gainesville, FL, USA. 

Wester P. J. (1912) “Annonaceous Fruits and Their Propagation.” Philippine 

Agriculture Review, 5 : 298-304. 

Wester P. J. (1913) “Annonaceous Possibilities for the Plant Breeder.” 

Philippine Agricultural Review, 6 : 312-321. 

Williamson J. (1974) The Useful Plants of Malawi. Montford Press, Limbe, 

Malawi. 

Woo M. H., Chung S. O. and Kim D. H. (1999) “Cis-annonacin and (2,4) Cis 

and Trans Isoannonacins: Cytotoxic Monotetrahydrofuran 

Annonaceous Acetogenins from the Seeds of Annona cherimola.” 

Arch. Pharm. Research, 22 (5) : 524-528. 

Woo M. H., Kim D. H., Fotopoulos S. S. and McLaughlin J. L. (2001) 

“Annocherin and (2,4) Cis and Trans Annocherimomes, Monotetrahydrofuran 

Annonaceous Acetogeninns with a C-7 Carbonyl 

Group from Annona cherimola Seeds.” Journal of Natural Products, 

63 (2) : 294. 

189

References 

Wu Leung W. T. and Flores M. (1961) Tabla de composición de alimentos 

para uso en America Latina. [Spanish] INCAP-ICNND, Ciudad de 

Guatemala, Guatemala : 132 pp. 

Wu F. E., Gu Z. M., Zeng L., Zhao G. X., Zhang Y. and McLaughlin J. L. 

(1995 a) “Two New Cytotoxic Monotetrahydrofuran Annonaceous 

Aetogenins, Annomuricins A and B, from the Leaves of Annona 

muricata.” Journal of Natural Products, 58 (6) : 830-836. 

Wu F. E., Zeng L., Gu Z. M., Zhao G. X., Zhang Y., Schwedler J. T. and 

McLaughlin J. L. (1995 b) “Muricatocins A and B, Two New Bioactive 

Monotetrahydrofuran Annonaceous Acetogenins from the Leaves of 

Annona muricata.” Journal of Natural Products, 58 (6) : 902-908. 

Wu F. E., Zeng L., Gu Z. M., Zhao G. X., Zhang Y., Schwedler J. T. and 

McLaughlin J. L. (1995 c) “New Bioactive Monotetrahydrofuran 

Annonaceous Acetogenins, Annomuricin C and Muricatocin C, from 

the Leaves of Annona muricata.” Journal of Natural Products, 58 (6) : 

909-915. 

Wu Y. C., Hung Y. C., Chang F. R., Cosentino M., Wang H. K. and Lee K. 

H. (1996) “Identification of Ent-16?,17-Dihydroxykauran-19-oic Acid 

as an Anti-HIV Principle and Isolation of the New Diterpenoids 

Annosquamosins A and B from Annona squamosa.” Journal of 

Natural Products 59 (6) : 635-637. 

Yang C. S. (1998) “Application of Plant Growth Regulators on Annona 

culture.” In: The Application of Plant Growth Regulators on 

Horticultural Crops, Symposium Proceedings. Edited by H. S. Lin, 

Chang L. R. and Lin J. H. Changhua, Taiwan : pp. 305-320 (Special 

Publication, 12). 

Yang T. H. and Chen C. M. (1970) “Studies on the Consitituents of Annona 

squamosa L.” Journal of the Chinese Chemical Society (Tapei), 17 (4) : 

243. 

Yokota T. (1986) Incremento na Produção de Pinhas. [Portuguese] Anuário 

do Fórum Paulista de Fruticultura. : pp. 59-66. 

You M., Wickramaratne D. B. M., Silva G. L., Chai H., Chagwedera T. E., 

Farnsworth N. R., Cordell G. A., Kinghorn A. D. and Pezzuto J. M. 

(1995) “(-) Roemerine, an Aprophine Alkaloid from Annona 

senegalensis that Reverses the Multidrug-Resistance Phenotype with 

Cultured Cells.” Journal of Natural Products, 58 (4) : 598-604. 

190

References 

Yu Jing-Guang, Gui Hua-Qing, Luo Xiu-Zhen and Sun L. (1998) 

“Murihexol, A Linear Acetogenin from Annona muricata.” 


Zayas J. C. (1966) Las Frutas Anonáceas. Ediciones Fruticuba : pp. 5-17. 

Zeng L., Wu F. E., Oberlies N. H., McLaughlin J. L. and Sastrodihadjo S. 

(1996) “Five New Monotetrahydrofuran Ring Acetogenins from the 

Leaves of Annona muricata.” Journal of Natural Products, 59 : 1035- 

1042. 

191

Appendix A. Common chemical 

compounds found in annonas 

Table A-1. Some of the Most Common Chemical Compounds Found in 

Different Parts of Cherimoya (Annona cherimolia), Custard apple (A. 

reticulata), Soursop (A. muricata), Wild soursop (A. senegalensis) and 

Sugar Apple (A. squamosa)*. 

Compounds References A.c. A.r. A.m. A.se. A.sq. 

Acetogenins 

4 deoxyannoreticuin Hopp et al., 1998 b 

Alumunequin Cortés et al., 1993b; r,s 

Duret et al., 1994 

Annocherin Woo et al., 2000 S 

Annogalene Sahpaz et al., 1996 S 

Annoncherimolin Kim et al., 2001 S 

Annomolin Kim et al., 2001 S 

Annomonicin Chang et al., 1993 l 

Annomuracins A, B Wu et al., 1995 a c l 

Annomuracin E Kim et al., 1998 l 

Annonacin 10 one Wu et al., 1995 a c; 

l,s 

Rieser, 1996 

Annonacin A Wu et al., 1995 a c s 

Annomuricatin B Vhao-Ming et al., 

s 

cyclopeptide 

1998 

Annonastatin Nonfon et al., 1990 s 

Annopentocin A, B, C Zeng et al., 1996 l 

Annoreticuin Chang et al., 1993; 

l,s 

Chang et al., 1998 

Annoreticuin 9 Hopp et al., 1997 b 

Annoreticuin 9 one Change et al., 1993; 

l, s 

Change et al., 1998 

Annosenegalin Sahpaz et al., 1996 s 

Annosquamosins A, B Chao-Ming et al., 

b 

cyclopeptides 

1997 

Anonacins Nonfon et al., 1990 s 

Anonins Nonfon et al., 1990; 

s l,s s 

Pinto & Silva, 1994 

Arianacin Reiser, 1996 s 

192

Appendix A. 


Aromin A Chen et al., 1999 st,l,s 

Asimicin Nonfon et al., 1990 s 

Bullatacin Li et al., 1990; 

Hernández & Angel, 

1997; hang et al., 

1999 

Bullatacinone Li et al., 1990; r 

Duret et al., 1994; 


1997 

Chermolin 1 Cortés et al., 1993 b s 

Cherimolin 2 Cortés et al., 1993 b s 

Cis 4 deoxyannoreticuin Hopp et al., 1998 B 

Cis annonacin Reiser, 1996; 

s 

Woo et al., 1999 a 

Cis annonarin-10-one Reiser, 1996 s 

Cis and trans 

Kim et al., 1998 a 

annomuricin C,D,E 

Cis goniothalamicin Wu et al., 1995 c; 

Rieser, 1996 

Cis trans bullatacinone Chang et al., 1998 s 

Cis trans isomurisolenin Chang et al., 1998 s 

Cis trans murisolinone Chag eet al., 1998 s 

Cyclopeptide 

Wu et al., 1995 c 

s 

muricatonins A,B 

Donhexocin Yu et al., 1998 s 

Epomuricenins A,B Roblot et al., 1993 s 

Gigantetrocin A Wu et al., 1995 c l,s 

Isoannoreticuin Chang et al., 1993 l 

Isocherimolin 1 Duret et al., 1994 r 

Isomolvizarin 1 Duret et al., 1994 r 

Isomolvizarin 2 Duret et al., 1994 r 

Javoricin Reiser, 1996 s 

Liriodenine Philipov et al., 1994 s 

Laherradurin Cortés et al., 1993 b s 

Molvizarin Hisham et al., 1994 stb 

Muricapentocin Kim et al., 1998 a l 

Muricatocins A,B,C Wu et al., 1995 c l 

Muricin Pinto & Silva, 1994 st,l,s 

Muricinin Pinto & Silva, 1994 s 

s 

l 

l,s 

b,s 

b,s 

193

Appendix A. 


Muricatetrocins A,B Wu et al., 1995 b l,s 

Muricoreacin Kim et al., 1998 b l 

Murihexocin C Kim et al., 1998 b l 

Murihexol Yu et al., 1998 s 

Otivarin Cortés et al., 1993 a S 

Panatellin Gleye et al., 1998 r 

Reticulatacin Gleye et al., 1998 r 

Reticulatacin-10-one Gleye et al. 1998 r 

Reticulacinone Hisham et al., 1994 stb 

Rolliniastatin Chang et al., 1993 l 

Rolliniastatin 2 Hisham et al., 1994 stb 

Sabedelin Gleye et al., 1999 r 

Solamin Myint et al., 1991; 

l,s r,s 

Chang et al., 1993; 

Gleye et al., 1998 

Squamocin Duret et al., 1994; r s S 


1997 

Squamone Li et al., 1990; 

l 

B 

Chang et al., 1993 

Squamotacin Hopp et al., 1996 B 

Uvariamicin I Gleye et al., 1998 r 

Uvariamicin IV Gleye et al., 1998 r 

2,4 cis and trans 

annocherinone 


isoannonacin 


squamoxinone 


mosinone A 

Alkaloids 

Woo et al., 2000 

Wu et al., 1995 b; 

Woo et al., 1999 a 

Hopp et al., 1998 

Hopp et al., 1997 

(-)Roemerine 

Bhakuni et al., 

1972; Oliver-Bever, 

1986; Cassady, 

1990; 

You et al., 1995; 

Chuliá et al., 1995; 

Fatope et al., 1996 

Anonaine Yang & Chen, 1970; 

Bhakuni et al., 

1972; Fresno & 

Cañavate, 1983; 

Bridg, 1984; 

s 

s 

l 

r l,s l L 

r,stb, 

l,p,b, 

s 

B 

B 

r,b l r,l,b r,l,b 

194

Appendix A. 


Philipov et al., 1995 

Aporphine Cassady, 1990; r s l L 

Salluja et al., 1990; 

Maeda et al., 1993; 

Chuliá et al., 1995; 

You et al., 1995; 

Fatope et al., 1996 

Benzyllioquinoline Maeda et al., 1993 s 

Caffeine 

Lizana & Reginato, s 

1990 

Corydine Oliver-Bever, 1986 l,s b,r,st 

,s 

Dehydroroemerine Chuliá et al., 1995 r 

α-Glaucine Oliver-Bever, 1986 b,r,st 

,s 

Isoboldine Philipov et al., 1995 r,l,b 

Isocorydine Oliver-Bever, 1986 b,r,st 

,s 

Isoquinoline Leboeuf et al., 1981; 

Maeda et al., 1993 

s l,s,r, 

b 

Lanuginosine 

Fresno & Cañavate, 

1983 

stb,l, 

p,b,s 

Liriodenin 

Fresno & Cañavate, stb,l, r,b s r,l,b 

1983; Morton, 1987; 

Philipov et al., 1995 

p,b,s 

Norcorydine Oliver-Bever, 1986 b,r,st 

,s 

Norisocoryline Bhakuni et al., 1972 L 

Pyrimidine β carboline Maeda et al., 1993 s 

Other Compounds 

β farnesene essential oil Leal, 1990 S 

α pirene essential oil Leal, 1990 S 

β pirene essential oil Leal, 1990 S 

Aliphatic ketone You et al., 1995 l 

Alkanes You et al., 1995 l 

Alkanols You et al., 1995 l 

Aminoacids You et al., 1995 b 

Amyl caproic acid Pinto & Silva, 1994 p 

Amyloids Kooiman, 1967 s 

Annosquamosins A,B Wu et al., 1996 s f,b 

diterpenoids 

Cherimoline Chen et al., 1998 st,l 

Cherinonaine Chen et al., 1998 st,l 

195

Appendix A. 


Coclamine Khan et al., 1997 p 

Couximine Khan et al., 1997 p 

Cohibins A,B Gleye et al., 1997 r 

Dihydroferuloyltyramine Chen et al., 1998 st,l 

Diterpenes 

Mukhophadhyay et 

r,stb 

al., 1993 

Ent-kaurenoids Fatope et al., 1996 rb,st, 

s,tb 

Essential oils (others) Bridg, 1964; l,s l,r f f S,l 

MacLeod & Pieris, 

1981; Jivoretz et al., 

1998 

Esthers Idstein et al., 1984; p 

You et al., 1995 

Flavonoids Cassady, 1990; 

l 

Langanson et al., 

1994 

Geranyl caproic acid Pinto & Silva, 1994 l 

Kaurane diterpenoids Adeogan & 

st,l s b f,r 

Durodola, 1976; 

Maeda et al., 1993; 

Wu et al., 1996 

Lactam amide Chen et al., 1998 st,l 

Limorene essential oil Leal, 1990 S 

Monoterpenoids You et al., 1995 l 

Murisolin Khan et al., 1997 p 

n-trans caffeoyltyramine Chen et al., 1998 st,l 

n-cis caffeoyltyramine Chen et al., 1998 st,l 

n-cis 

Chen et al., 1998 st,l 

feruloymethoxytyramine 

n-cysferuloyltyramin Chen et al., 1998 st,l 

n-fatty acyl tryptamine Maeda et al., 1993 s 

n-p-coumaroyltyramine Chen et al., 1998 st 

n-trans 

Chen et al., 1998 st,l 

feruloylmethoxytyramine 

n-trans feruloyltyramine Chen et al., 1998 st,l 

Purine Cehn et al., 1998 st 

Reticulin Khan et al., 1997 p 

Rhamnoside Salluja et al., 1990 l,s 

Saponins Salluja et al., 1990; 


1994 

l 

S 

196

Appendix A. 


Sesquiterpenoids You et al., 1995 l 

Stepharine Khan et al., 1997 p 

Steroids Chen et al., 1998 st 

Sterols You et al., 1995 l 

Tannins Burkill, 1966; 


1994 

Trans orimene essential Leal, 1990 

oils 

Volatiles (terpenes 

hydrocarbons, esters, 

carbonyls) 

Idstein et al., 1984 

l l p l 

Key: r-root; rb-root bark; st-stem; stb-stem bark; l-leaf; f-fruit; p-pulp; b-bark; s- 

seed. 

* This is not a comprehensive list of compounds. 

p 

S 

197

Appendix B. Uses of Annona species in medicine 

Table B-1. Some Uses of Annona Species in Medicine, Cherimoya (Annona cherimolia), Custard apple (A. reticulata), 

Soursop (A. muricata), Wild soursop (A. senegalensis) and Sugar Apple (A. squamosa) 

Human Bioactive Effects Compound Reference 

Abortion Unkown Salluja & Santini, 1990; Asolkar et al., 1992 

Anal prolapse Unknown Chao-Ming et al., 1997 

Antibacterial Murisolin, couxine, couclamine, 

stepharine, reticulin 

Asolkar et al., 1992; Khan et al., 1997 

Anti-HIV principle 16β17 dihydroxy kauran19oic Wu et al., 1996 

acid 

Antineuralgic properties Essential oils Calzavara et al., 1987; Moura, 1988 

Antiparasitic and protozoa activity 

Probably acetogenins and essential oils 

Calzavara et al., 1987; Bories et al., 1991; 

Philipov et al., 1994 s s,l 

Antispasmodic Probably flavonoids alkaloids tannins saponins 

Moura, 1988; Philipov et al., 1994; Chuliá et al., 

1995 r l,s 

Antiulcer Flavonoids Langanson et al., 1994 

Astringent Unknown Calzavara et al., 1987; Asolkar et al., 1992; Khan 

et al., 1997; f b l,f 

A.c. 

A.r. A.m. A.se. A.sq. 

s s 

l l 

l + 

l 

l 

f 

198 

198

Appendix B. 

Appendix B. 


Atonic dyspepsia Unknown Calzavara et al., 1987; Khan et al., 1997 

Bed bugs/lice Acetogenins, alkaloids flavonoids 

tannins saponins 

Rupprecht et al., 1990; Bories et al., 1991; 

Asolkar et al., 1992; Langanson et al., 

1994;Hernández & Angel, 1997; Abubaka & 

Abdurham, 1998 

Burning of the throat Resin Lizana & Reginato, 1990 

Cancer treatment Kaurene, diterpenoids, 

acetogenins 

Cassady, 1990; Asolkar et al., 1992; Chang et 

al., 1993; Hopp et al., 1994; Wu et al., 1995; 

You et al., 1995; Fatope et al., 1996; Hopp et al., 

1996; Reiser et al., 1996; Sahpaz et al., 1996; 

Zeng et al., 1996; Hopp et al., 1997; Kim et al., 

1998 a 

Cathartic Unknown Lizana & Reginato, 1990; Asolkar et al., 1992 

Cemopreventative agents Flavonoids Cassady, 1990 

Chest pain - You et al., 1995 

Colic Murisolin, couxine, couclamine, 


Khan et al., 1997 

Convulsions Unknown You et al., 1995 

Depression (sedative, 

anxyolitic) 

Alkaloids, others Bories et al., 1991; Chao-Ming et al., 1997 

199 

A.c. 


f 

s l,s,p s l,s,dri 

ed f 

S 

s l 

s s,l,b s,r,stb 

,l,b 

B 

s 

l,b,f, 

r,s 

l,b,f, 

r,s 

l,b,f, 

r,s 

l,b,f,r 

,s 

l,b,f, 

r,s 

r,l 

f 

r,l 

l l R 

199

Appendix B. 

Appendix B. 



A.c. 

200 

Diabetes Unknown Calzavara et al., 1987 

l 

b l,f,r r,b,l + 

Calzavara et al., 1987; Leal, 1990; Asolkar et al., 

1992; Philipov et al., 1995; You et al., 1995; 


Diarrhoea, dystentery Alkaloids, murisolin, couxine, 

couclamine, stepharine, reticulin 

Diuretic Unknown Calzavara et al., 1987; Khan et al., 1997 

f 

Dryness of the mouth Lizana & Reginato, 1990 

Emetic Resin Calzavara et al., 1987; Lizana & Reginato, 1990; 

Asolkar et al., 1992 s s + 

Calzavara et al., 1987; Philipov et al., 1995 

fw l 

Eye inflammation Alkaloids, flavonoids tannins 

saponins 

l,f r 

Calzavara et al., 1987; Asolkar et al., 1992; You 

et al., 1995; Fatope et al., 1996; Khan et al., 

1997 

Febrifuge Murisolin, couxine, couclamine, 


Filariosis Unknown You et al., 1995; Fatope et al., 1996 

r,l 

l l r,b,l l,r 

Leal, 1990; Langason et al. 1994; Philipov et al., 

1995; Voigt et al., 1995; You et al., 1995; Khan 

et al., 1997 

Alkaloids, flavonoids tannins 

saponins 

Gastric & digestive 

processes, intestine diseases 

in general, as a tonic or 

S 

laxative 

Haemolysis of red blood Saponins Salluja & Santani, 1990 

cells 

Immunosupressant Acetogenins Rupprecht et al., 1990 

r,s s r,l,s l,s S 

Intense photophobia Resin Lizana & Reginato, 1990 

s 

200

Appendix B. 

Appendix B. 


Liver disease, jaundice Acids Calzavara et al., 1987 

Malaria Murisolin, couxine, couclamine, 



Male impotency Alkaloids You et al., 1995 

Nausea & vomiting Resin Calzavara et al., 1987; Lizana & Reginato, 1990; 

Khan et al., 1997 s f,s 

Oedema Murisolin, couxine, couclamine, 



Parasites of the feet Acids Calzavara et al., 1987; Asolkar et al., 1992; 

Philipov et al., 1994 s p 

Peptic ulcers Murisolin, couxine, couclamine, 



Pneumonia Unknown Williamson, 1974; Vogt, 1995 

Pupil dilation Resin Lizana & Reginato, 1990 

Purgative Alkaloids Leal, 1990 

Rash, skin diseases, sores Murisolin, couxine, couclamine, 


Rheumatological problems Essential oils Calzavara et al., 1987; Moura, 1988 

Scorbutic Probably vitamin C Calzavara et al., 1987, Khan et al., 1997 

A.c. 

s 


p 

f 

f 

f 

r,l 

b,l + 

Asolkar et al., 1992; Vogt, 1995; Khan et al., 

1997; Chen et al., 1998 s,t f r,l,b L 

l 

f 

R 

201 

201

Appendix B. 

Appendix B. 



A.c. 

202 

Snake bite Alkaloids Philipov et al., 1995; Vogt, 1995; You et al., 

1995 b,l 

Spinal marrow disease Alkaloids Chao-Ming et al., 1997 

r 

Swelling Unknown Chao-Ming et al., 1997 

L 

Trypanosomiasis Alkaloids, flavonoids others You et al., 1995; Fatope et al., 1996 l 

Venereal disease Alkaloids Asolkar et al., 1992; Philipov et al., 1995; Vogt, 

1995, You et al., 1995 r,l + 

s s l,s r,b 

Nonfon et al., 1990; Bories et al., 1991;Asolkar 

et al., 1992; Cortés et al., 1993; Philipov et al., 

1994; Vogt, 1995; You et al., 1995 

Worm infestations Essential oils alkaloids flavonoids 

acetogenins 

Key: r-root; rb-root bark; st-stem; stb-stem bark; l-leaf; f-fruit; fw-flower; p-pulp; b-bark; s-seed; +-whole plant. 

202

Appendix C. Institutions and 

Individuals Engaged in Annona 

Research and Development 

INTERNATIONAL INSTITUTIONS 

International Centre for Underutilised Crops - 

ICUC 

International Water Mangaement Institute 

127 Sunil Mawatha 

Pelawatte 

Battaramulla 

Sri Lanka 

International Plant Genetic Resource Institute 

- IPGRI 

Via dei Tre Denari 472/a 

00057 Maccarese 

Rome 

Italy 

Food and Agriculture Organization of the 

United Nations - FAO 

Via delle Terme di Caracalla 

00100 Rome 

Italy 

COUNTRY, INDIVIDUALS AND NATIONAL 

INSTITUTIONS 

AUSTRALIA 

Broadley R. 

Maroochy Research Centre, Department of 

Primary Industries 

P.O. Box 5083 

Sunshine Coast Mail Center 

Nambour-Queensland 4560 

Australia 

Campbell J. A. 

Maroochy Horticultural Research Station, 

Department of Primary Industries 

P.O. Box 5083 


Nambour, Queensland 4560 

Australia 

George A. P. 



P.O. Box 5083 


Hamill S. D. 



P.O. Box 5083 


203

Appendix C. 


Australia 

Howitt C. 

Biometry Branch, Department of Primary 

Industries 

P.O. Box 46 

Brisbane, Qld. 4000 

Australia 


Australia 

Nissen R. J. 



P.O. Box 5083 



Australia 

Topp B. L. 

Granite Belt Research Station 

PO Box 501 

Stanthorpe, Queensland 4380 

Australia 

BANGLADESH 

Malek A. 

BCSIR Laboratories 

1205 Dhaka 

Bangladesh 

BARBADOS 

Carrington C. M. S. 

Biology Department, University of the West 

Indies 

P.O. Box 64 

Bridgetown 

Barbados 

Worrell D. B. 

Biology Department, University of the West 

Indies 

P.O. Box 64 

Bridgetown 

Barbados 

BELGIUM 

de Smet S. 

Laboratory for Tropical and Subtropical 

Agronomy and Ethnobotany, 

University of Ghent 

Coupure Links 653 

9000 Ghent 

Belgium 

Scheldeman X. 





9000 Ghent 

Belgium 

Snellings M. 

Bergstraat 7, 

Van Damme P. 


204

Appendix C. 

3740 Bilzen 

Belgium 




9000 Ghent 

Belgium 

Van Damme V. 

Clos de la Sucrerie 6, 

7760 Escanaffles 

Belgium 

Van Ranst E. 

Laboratory for Soil Science, 

University of GhentKrijgslaan S8 

9000 Ghent 

Belgium 

Vandersmissen M. 

Korsele 48 

9667 Horebeke 

Belgium 

BRAZIL 

Andrade, S. R. M. de 

EMBRAPA Cerrados 

Km 18 da Br 020, P.O. Box 08223 

73301-970 Planaltina-DF 

Brazil 

Bonaventure, L. E. 

Rua Bennett, 727 

05464-010 São Paulo 

Brazil 

Bonfim, M. P. 

Departamento de Fitotecnia e Zootecnia, 

Universidade Estadual do Sudoeste da Bahia 

Cx. 95 cep. 45000 000 

Vitória da Conquista, Bahia 

Brazil 

Calzavara, G. B. B. 

EMBRAPA Trópicos Úmidos - CPATUP.O. 

Box 48 

66240 Belém PA 

Brazil 

Castro, F. A. de 

Nucleo de Tecnologia Industrial do Estado do 

Ceara- NUTEC 

60000 Fortaleza CE 

Brazil 

Cataneo, A. 

Centro de Ciências Agrárias, 

Universidade Federal de Alagoas 

Rua Judson Roosevelt Cabral, 34 Quadra "d" 

CEP 57061 330, Tabuleiro Maceió 

Brazil 

205

Appendix C. 

Cavalcante, R. L. R. R. 

Departamento de Fitotecnia e Fitossanidade, 



57072 970, Maceió AL 

Brazil 

Cavalcante, T. R. M. 

Universidade Federal de Vicosa 

Departamento de Fitotecnia 

36571 000, Vicosa, Minas Gerais 

Brazil 

Clement, C. R. 

Instituto Nacional de Pesquisa da Amazônia – 

INPA 

CP478 69000, Manaus AM 

Brazil 

Cordeiro, M. C. R. 


Km 18 da Br 020 

P.O. Box 08223 

73301-970 Planaltina DF 

Brazil 

Ferreira Filho, W. C. 

Instituto Nacional de Pesquisa da Amazônia 

69000, Manaus AM 

Brazil 

Ferreira, F. R. 

EMBRAPA Recursos Genéticos e Biotecnologia 

SAIN Parque Rural 

Brasília DF 

Brazil 

Ferreira, G. 

Universidade Estadual do Oeste do Paraná-R 

Marechal Cândido Rondom, 1777 

859609, Pernambuco 

Brazil 

Flores, W. B. C. 

Instituto Nacional de Pesquisa da Amazônia 

CP478 69000, Manaus AM 

Brazil 

Fogaça, L. A. 

Universidade Estadual do Oeste do Paraná-R 

Marechal Cândido Rondom, 1777 

859609, Pernambuco 

Brazil 

Freitas, G. B. 

Departamento de Fitotecnia, 

Universidade Federal de Viçosa 

36571 000, Viçosa, Minas Gerais 

Brazil 

Guedes, Z. B. L. 

Universidade Federal do Ceara 

P.O. Box 3038 

60000, Fortaleza CE 

Brazil 

Hojo Rebouças, T. N. 



Vitória da Conquista, Bahia 

Brazil 

206

Appendix C. 

Lemos, E. E.P. 

Departamento Agronomia, 


BR 104 Norte, Km 14 

57072970 Maceió AL 

Brazil 

Lima, C. L. C. De 





Brazil 

Lima, G. P. P. 





Brazil 

Maia, G. A. 

Universidade Federal do Ceara 

P.O. Box 3038 


Brazil 

Marcondes, M. A. 





Brazil 

Marinho, G.A. 

Departamento de Fitotecnia e Fitossanidade, 



57072 970, Maceió 

Brazil 

Melo, M. R. 


Avenida Santa Rita 304-A 

Viçosa MG 

Brazil 

Moura Fé, J. A. 

Universidade Federal do Ceará 

P.O. Box 3038 


Brazil 

Nascimento, R. J. 

116 Várzea 

50741 390, Recife 

Brazil 

Pinto, A. C. de Q. 


Km 18 da Br 020 

P.O. Box 08223 

73301-970 Planaltina-DF 

Brazil 

Rua, I. B. B. 

116 Várzea 

50741 390, Recife 

Brazil 

Santiago, A. D. 





Brazil 

São José, A. R. 



Silva, A. C. da 

Universidade Estadual do Sudoeste da Bahia, 

Departamento de Fitotecnia e Zootecnia 

207

Appendix C. 

45000-000 Vitória da Conquista, Bahia 

Brazil 

P.O. Box 95 

45000-000 Vitória da Conquista, Bahia 

Brazil 

Vieira, M. F. 

Departamento de Biologia Vegetal, 


36571 000, Vicosa, Minas Gerais 

Brazil 

Zucarelli, C. 

Universidade estadual do Oeste do Paraná 

Rua Pernambuco 1777, marechal Cândido 

Rondon, 859609 Paraná 

Brazil 

CHILE 

Cano G. L. 

Universidad Católica de Valparaiso 

Casilla 476 

Quillota 

Chile 

Castro M. 

Laboratorio de Micropropagación, 

Facultad de Agronomia, 

Universidad Católica de Valparaíso 

Cassilas 4 - D, Quilhota 

Chile 

Fassio Ortiz C. 

Departamento de Fruticultura, 

Universidad Católica de Valparaíso 

Casilla 4-D, Quillota V Región 

Chile 

Gardiazabal Irazabal F. 

Universidad Católica de Valparaiso 

Avenida Brasil 2950 

Casillas 4059, Valparaiso 

Chile 

Ibacache A. 

Instituto de Investigaciones Agropecuarias - 

INIA 

Casilla 73 

Vicuña 

Chile 

Montiel Moreno M. 

Villa La Cruz, Pje 1 

5e Región, La Cruz (Quillota) 

Chile 

Morales R. C. 


Universidad Catolica de Valparaíso 

Casilla 4-D, Quillota, V Region 

Chile 

Navia V. M. G. 

Subestación Experimental Cauquenes, 

Instituto de Investigaciones Agropecuarias 

Casilla 165 

Cauquenes, Maule 

Chile 

Ovalle A. 


Universidad Catolica de Valparaíso 

Casilla 4-D, Quillota, V Región 

Chile 

Razeto B. 

Facultad de Ciencias Agrarias e Forestales, 

Universidad de Chile 

Casilla 1004, Santiago 

Chile 

208

Appendix C. 

Saavedra E. 

Faculdad de Agronomia, 

Universidad de Chile 


Chile 

Undurraga Martinez P. L. 


Universidad de Valparaiso 

Casilla 4-D, Quillota, V Región 

Chile 

Valenzuela J. 

Programa frutales y Viñas, 

Estación Experimental La Platina, 

Instituto de Investigaciones Agropecuarias - 

INIA 


Chile 

CHINA 

Li Chao-Ming 

Kunming Institute of Botany, 

Chinese Academy of Sciences 

650204, Heilongtang, Kunming, Yunnan 

China 

Mu Quing 




China 

Sun Han-Dong 




China 

Tan Ning-Hua 




China 

Weidong Tang 

Medicine Material Institute of Shanghai, 

The Chinese Academy of Sciences 

Shanghai 200031 

China 

Xiao-Jiang Hao 



Heilongtang, Kunming 650204, Yunnan 

China 

Xu Bin 

Medicine Material Institute of Shanghai, 


Shanghai, 200031 

China 

Yu-Ping Lu 



650204, Heilongtang, Kuming, Yunan 

China 

Yang Lu 

Medicine Material Institute of Beijing, 

The Chinese Academy of Medical Sciences 

Beijing 100050 

China 

Zhen Hui-Lan 




China 

209

Appendix C. 

Zhou Jun 




China 

COLOMBIA 

Escobar Torres W. 

Instituto Colombiano de Agricultura - ICA 

Apartado Aéreo 233 

Palmira 

Colombia 

Lastra R. 

International Plant Genetic Resources Institute 

- IPGRI 

Apartado Postal 6713, Cali 

Colombia 

Rios Castano D. 

M.S. Frutales Ltda 

Apartado Aereo 6282, Cali 

Colombia 

Salazar Castro R. 



Palmira 

Colombia 

Sánchez López L. A. 



Palmira 

Colombia 

Torres Monedero R. 



Palmira 

Colombia 

COSTA RICA 

León J. 

Instituto Interamericano de Cooperacion para 

la Agricultura 

San José 

Costa Rica 

CUBA 

Dominicis Maria E. 

Departamento de Bioproductos, 

Instituto de Ecología y Sistematica 

Finca la Chata, carretera de Verona, km 3.5 

10800, Capdevila, Boyeros 

Oviedo R. 





210

Appendix C. 

Cuba 

Cuba 

Payo A. 





Cuba 

Sandoval D. 



Finca La Chata 

Carretera de Verona, km 3.5 


Cuba 

CYPRUS 

Gregoriou C. 

Agricultural Research Institute, 

Ministry of Agriculture, Natural Resources 

and Environment 

P.O. Box 2016 

1516 Nicosia 

Cyprus 

ECUADOR 

Agila Palacios E. P. 

Ave. De los Paltas y Paraguay 

Loja 

Ecuador 

Aguirre R. Q. 

CATER 

Casilla 399, Loja 

Ecuador 

Aguirre Mendoza Z. H. 

Herbario "Reinaldo Espinosa", 

Universidad Nacional de Loja 

Casilla "B", Loja 

Ecuador 

Apolo V. 

Proyecto VLIR 


Ecuador 

Armijos Tandajo J. E. 

CATER 


Ecuador 

Briceño Ortiz J. I. 

CATER 


Ecuador 

Briceño Ortiz V. A. 

Proyecto Europeo 


Ecuador 

Coronel Benitez S. B. 

Quintas Experimentales, 


Casilla "S" 656, Loja 

211

Appendix C. 

Ecuador 

Cuenca Ojeda R. A. 

Piura y Rosario 

Loja 

Ecuador 

Cuenca Ortiz K. I. 



Ecuador 

Delgado Cueva T. 

Herbario "Reinaldo Espinosa" 

Casilla "B" Loja 

Ecuador 

Eguiguren G. 

Loja Flor 

Casilla 11-01-142, Loja 

Ecuador 

Honorio Rivera R. 



Ecuador 

Espinosa González C. A. 

CATER 


Ecuador 

Feijó Cisneros N. G. 



Ecuador 

Feijó Cisneros T. F. 



Ecuador 

González M. 

Univesidad Nacional de Loja 

Casilla "S", 656 Loja 

Ecuador 

Guamán Diaz F. 



Ecuador 

León Chimbo F. S. 

PREDESUR 

Av. Orillas Del Zamora 

Sector Norte, Loja 

Ecuador 

León Fuentes J. F. 

Instituto Nacional Autónomo de Investigaciones 

Agropecuarias - INIAP 

Casilla 17-01-340, Quito 

Ecuador 

Maldonado Astudillo N. P. 

Facultad de Ciencias Agrícolas, 

Universidad Nacional de LojaCasilla "S" 656, 

Loja 

Ecuador 

Mariaca J. 

EcoCiencia 

P.O. Box 17-12-257 

Quito 

Ecuador 

Moreira Palacios M. O. 

Universidad Técnica Particular de Loja 

Casilla 11.01.608, Loja 

Ecuador 

Morocho Pesantez J. G. 

Proyecto VLIR 


Ecuador 

212

Appendix C. 

Ortega C. 


Agropecuarias - INIAP 


Ecuador 

Pazmiño Pineda E. M. 



Ecuador 

Quishpe W. 

Herbario "Reinaldo Espinosa", 


Casilla "B", Loja 

Ecuador 

Romero Motoche J. P. 

Centro Andino de Tecnología Rural, 



Ecuador 

Rivadeneira Arqudo L. A. 

Calle Novena 109 y Avenida Domingo 

Comín, Oficina Fadesa 

Guayaquil 

Ecuador 

Serrano Armijos T. 



Ecuador 

Solórzano L. V. 

Proyecto Bosque Seco 

Calle Garcia Moreno 410 

Celica, Loja 

Ecuador 

Su´rez Chacón J. P. 

Universidad Técnica Particular de Loja 

Casilla 11.01.608, Loja 

Ecuador 

Suárez L. 

EcoCiencia 

P.O. Box 17-12-257 

Quito 

Ecuador 

Ureña Alvarez J. V. 



Ecuador 

Ureña V. 

Centro Andino de Tecnología Rural, 



Ecuador 

Valdivieso Caraguay E. S. 

CATER 


Ecuador 

Viteri P. F. 


Agropecuarias – INIAP 

Granja Experimental Tumbaco 


Ecuador 

213

Appendix C. 

EGYPT 

Mansour K. M. 

Agricultural Research Center, Horticultural 

Research Institute 

9 Gamma Street, Giza, Orman 

Egypt 

FRANCE 

Pallares P. 

25, Chemin de Clapiers 34 730 

St. Vincent de Barbeyrargues 

France 

GERMANY 

Ebert G. 

Institut fur Gartenbauwissenschaften, 

Fachgebiet Obstbau der Humboldt, 

Universitat zu Berlin 

Albrecht Thaerweg 14195 

Germany 

GREECE 

Lionakis S. 

Department of Subtropical Plants, 

National Agricultural Research Foundation, 

Subtropical Plants and Olive Trees Institute 

Khania, Crete, 73100 

Greece 

ITALY 

Fiorino P. 

Dipartimento Ortoflorofrutticoltura Universita 

Firenze 

Italy 

Monastra F. 

Instituto Sperimentale per la Fruticoltura 

Via Fioranello, 52 

00040 Ciampino, Rome 

214

Appendix C. 

Italy 

Parri G. 

Dipartimento Ortoflorofrutticoltura Universita 

Firenze 

Italy 

Pestelli P. 

Instituto Propagazione Specie Legnose 

C.N.R.Firenze 

Italy 

Tazzari L. 

Instituto Propagazione Specie Legnose C.N.R. 

Firenze 

Italy 

INDIA 

Bejoy M. 

Tropical Botanic Garden and Research 

Institute 

Palode 695562 

Kerala 

India 

Farooqi A. A. 

College of Agriculture 

Dwarwar 

India 

Gupta P. K. 

Division of Biochemical Sciences, 

National Chemical Laboratory 

Pune 411.008 

India 

Hariharan M. 

Department of Postgraduate Studies and 

Research in Botany, 

University of Calicut, 673635 

Kerala 

India 

Mascarenhas A. F. 



Pune 411.008 

India 

Nalawadi U. G. 


Dwarwar 

India 

Nair S. 



Pune 411.008 

India 

Parvatikar S. R. 


Dwarwar 

India 

Shirgurkar M. V. 



Pune 411.008 

215

Appendix C. 

India 

ISRAEL 

Blumenfeld A. 

Institute of Horticulture, 

The Volcani Center 

P.O. Box 6 

Bet Dagan, 50-250 

Israel 

Gazit S. 

Faculty of Agriculture, 

The Hebrew University of Jerusalem 

P.O. Box 12 

Rehovot 76100 

Israel 

JAPAN 

Higuchi H. 

Graduate School of Agriculture, 

Kyoto University 

Kitashirakawa Sakyo-ku, Kyoto 606-8502 

Japan 

Sakuratani T. 

Laboratory of Tropical Agriculture, 

Faculty of Agriculture, Kyoto University 

Kitashirakawa Sakyo-ku, Kyoto 60601 

Japan 

Utsunomiya N. 


Kinki University 

Nakamachi, Nara 631-8505 

Japan 

Nakanishi T. 


Kobe University 

Rokkoudai, Nada-ku, Kobe 657-8501 

Japan 

Tomita E. 

Wakayama Prefecture Fruit Tree Experiment 

Station 

Kibi-cho, Arida-gun, Wakayama 643-0022 

Japan 

Yonemoto Y. 

Wakayama Prefecture Experiment Station for 

Primary Industry 

Mountainous Regions, Kozagawa, 

Higashimuro 

Wakayama 649-4222 

Japan 

KENYA 

Bydekerke L. 

Div. Environmental Assessment and Early 

Warning, Database and Atlas Project, 

UNEP 

PO Box 30552 

Nairobi 

Kenya 

216

Appendix C. 

MÉXICO 

Abraján Hernández P. 

Laboratorio de Fisiología y Química Vegetal, 

Universidad de Ciencias y Artes del Estado de 

Chiapas 

C.P 29000, Tuxtla Gutiérrez Chiapas 

México 

Aldana Llanos L. 

Centro de Desarrollo de Productos Bióticos, 

Instituto Politécnico Nacional 

Km. 8.5 de la carretera Yautepec- Jojutla 

C.P 62731, Colonia San Isidro 

C.P 24, Yautepec, Morelos 

México 

Alfaro Romero T. de J. 



Chiapas 

C.P 29000, Tuxtla Gutiérrez, Chiapas 

México 

Agustín J. A. 

Universidad Autônoma Chapingo en el Centro 

Regional Universitario Centro-Occidente 

Morelia, Michoacan 

México 

Avilez Terán F. 

Escuela de Biología, 

Universidad Autónoma de Sinaloa 

Apdo. Postal 264 Culiacán, Sinaloa 

México 

Ayala Escobar V. 

Secretaria de Agricultura, 

Ganaderia y Desarrollo Rural 

Guilhermo Pérez Valenzuela 127 

Col. Del Carmen, 04100 Coyoacán 

México 

Becerril R.A.E. 

Colegio de Postgraduados, Especialidad de 

Fruticultura 

Km 38.5 Carretera México- Texcoco 

C.P 56230, Montecillos 

México 

Beltran Magallanes J. A. 




México 

Benitez Morteo Y. 

Instituto de Salud Pública y Facultad de Biología, 

Universidad Veracruzana 

Jalapa, Veracruz 

México 

Bolóvar Fernández N. 

Instituto Tecnológico de Mérida 

Av. Tecnológico s/n. km 5 

Mérida, Yucatan 

México 

Castañeda Vildozola A. 

Departamento de Fitotecnia, Fundación Salvador 

Sanches Colín, CICTAMEX S.c 

C.P 51700, Coatepec Harinas 

México 

Cedillo Portugal E. 


Universidad Autónoma Chapingo 

Carretera México- Texcoco, km 36.5 

C.P 56230, Chapingo 

México 

217

Appendix C. 

Cervera Backhauss E. 





México 

Coeto Juárez L. P. 

Laboratorio de Productos Naturales Área de 

Química, Universidad Autônoma Chapingo 



México 

Cortés Popoca R. 



Km 8.5 de la carretera Yautepec- Jojutla 

C.P 62731, Colonia San Isidro, Morelos 

México 

Chávez Primitivo E. V. 





México 

Colar Gómez E. 




México 

Cruz Castillo J. G. 

Centro Regional Univeritario Oriente, 


Apartado 65, Huatusco, Veracruz 

México 

De la Cruz Chacón I. 



Chiapas 


México 

Díaz Camacho A. 


Chiapas 


México 

Domínguez Martínez V. 




México 

Edith A. 

Centro de Investigaciones Biológicas, 

Universidad Autónoma del Estado de Morelos 

Av. Universidad No. 1001 Chamilpa 

México 

Estrada Reyes R. 



Km 8.5 de la carretera Yautepec - Jojutla 


México 

Evangelista Lozano S. 



Km 8.5 de la carretera Yautepec- Jojutla 


México 

Farrera Villanueva S. B. 



Chiapas 


México 

Ferrera Cerrato R. 

Seccion de Microbiologia, 

Colegio de Postgraduados 

Km 38.5, Carretera México - Texcoco 

56230, Montecillos 

México 

García V. E. González Chávez M. C. 

218

Appendix C. 

Especialidad de Fruticultura, 

Colegio de Postgraduados 

Km 38.5, Carretera México - Texcoco 

56230, Montecillos 

México 

González Esquinca A. R. 



Chiapas 


México 

Instituto Tecnológico Agropecuario No. 2 

Km 16.3, Antigua Carretera México - Motul 

Municipio de Congal, 97100 Mérida 

México 

Guerra Bustos O. 




México 

Heinze G. 



Carretera México - Texcoco km 36.5 

CP 56230, Chapingo 

México 

Heredia J. B. 

Centro de Investigación en Alimentación y 

Daserrollo, A. C - Unidad Culiacan 

Pascual Orozco y Topolobampo 1603, 

Culiacán 

México 

Hernández Domínguez C. 





México 

Ibarra Salazar M. A. 




México 

Jesús Peralta M. 

Depto. de Preparatoria Agrícola, 




México 

Nathan P. J. 

Centro de Investigación y de Estudios Avanzdos, 


D. F 07000, México 

México 

Nieto A. D. 

Colegio de Postgraduados, Área: Enfermedades 

de Frutos en Postcosecha, 

Instituto de Fitossanidad 

CP 56230, Montecillo, Texcoco 

México 

López L. L. 

Departamento de Fitotecnia, Fundación Salvador 

Sanches Colín, CICTAMEX S.c 

Calle Ignácio Zaragoza 


México 

López Herrera A. 




México 

Luna Cazáres L. M. 


Escuela de Biología, Universidad de Ciencias 

y Artes del Estado de Chiapas 


México 

219

Appendix C. 

Luna Rodríguez M. 

Instituto de Salud Pública, 


Ernesto Ortiz Medina No. 3 

91020, Xalapa, Veracruz 

México 

Marroquín Andrade L. M. 

Dpto. de Fitotecnia, 

Universidad Autônoma Chapingo 

Chapingo 

México 

Martinez Díaz E. 

CBTA 17 de Ú:rsulo Galvan Veracruz 

Apartado Postal 1439 

Central Camionera Aguascalientes 

CP 20170, Aguascalientes 

México 

Martinez Mata N. L. 






México 

Martinez Santiago M. G. 



Chiapas 


México 

Monroy R. 

Centro de Investigaciones Biológicas, 

Universidad Autónoma del Estado de Morelos 

Av. Universidad No. 1001 

México 

Morales Franco L. 






México 

Moreno Andrade E. R. 


Universidad de Ciencias y Artes Del Estado 

de Chiapas 

CP 29000, Tuxtla Gutiérrez, Chiapas 

México 

Mosqueda V. R. 

Especialización de Fruticultura, 


91020, Jalapa, Veracruz 

México 

Nadal Medina R. 



91020, Jalapa, Veracruz 

México 

Nava-Díaz C. 





México 

Navarrete Castro A. 

Departamento de Farmácia, 

Universidad Nacional Autónoma de México 

CP 56230, México D.F 

México 

Orozco M. R. 

Facultad de Ciencias Agropecuarias UAEM 

Av. Universidad 1 Colonia Chamilpa 

Osada Kawasoe S. 



220

Appendix C. 

Cuernavaca, Morelos 

México 



México 

Perales C. 

Casa Blanca 1003, Fracc. Casa Blanca 

CP 20270, Aguascalientes 

México 

Pizón López L. L. 

Instituto Tecnológico Agropecuario 

Antigua Carretera México - Motul 

97100, Mérida, Yucatán 

México 

Regollar Alviter A. 

Universidad Autônoma Chapingo en el Centro 

Regional Universitario 

Centro-Occidente 

Morelia, Michoacan 

México 

Reyes Trejo B. 

Laboratorio de Productos Naturales Área de 

Química, Universidad Autonoma Chapingo 

Carretera México, Texcoco km 36.5 

P.O. Box 56-230 

Chapingo, Edo. De México 

México 

Saucedo Veloz C. 

Instituto Tecnológico de Mérida 

Av. Tecnológico s/n, km 5 

Mérida, Yucatán 

México 

Rendón Sánchez G. 





México 

Román Rubén D. E. 


Fundación Salvador Sanches Colín, 

CICTAMEX S.c 


México 

Sauri Duch E. 

Instituo Tecnológico de Mérida 

Av. Tecnológico s/n, km 5 

Mérida, Yucatán 

México 

Tun Suárez J. M. 

Instituto Tecnológico Agropecuário 

N° 2, km 16,3 

Antigua Carretera Mérida-Motul 

Município de Conkal, Yucatán 

México 

Valdés Estrada M. E. 

Centro de Desarrolo de Productos Bióticos, 


Km 8,5 de la Carretera Yautepec-Jojutla 

Colonia San Isidro, P.O. Box 62731, Morelos 

México 

Vásquez Aguilar R. F. 

Instituto Tecnológico Agropecuário 

N° 2, km 16.3, 

Antigua Carretera Mérida-Motul 

Município de Conkal 

97100 Mérida-Yucatán 

México 

Verdugo Munguía H. 

Escuela de Biología, Universidad Autonoma 

de Sinaloa 

Aptdo. Postal 264 

Culiacán, Sinalo 

México 

Vidal Hernández L. 

Especialización en Fruticultura, 

Vidal Lezama E. 


221

Appendix C. 


Ernesto Ortiz Medina N° 3, 

91020 Jalapa, Veracruz 

México 




México 

Villegas M. A. 

Colegio de Postgraduados, 

Especialidad de Fruticultura 

Km 38.5 Carretera México-Texcoco 

Montecillos, Estado do México, P.O. Box 

56230 

México 

NEW ZEALAND 

Anderson P.T. 

The Horticulture and Food Research Institute, 

Kerikeri Research Centre 

PO.Box 23 

Kerikeri 

New Zealand 

Richardson A. C. 

Kerikeri Research Centre, 

The Horticulture and Food Research Institute 

P.O.Box 23 

Kerikeri 

New Zealand 

NIGERIA 

Abdurahman E. M. 

Department of Pharmacognosy and Drug 

Development, Ahmadu Bello University 

Zaria 

Nigeria 

Abubakar M. S. 

Department of Pharmacognosy and Drug 

Development, Ahmadu Belo University 

Zaria 

Nigeria 

Aku A. A. 

Department of Crop Production, University of 

Agriculture 

P.M.B. 2373, Makurdi, Benue State 

Nigeria 

Attah J. A. 

Department of Crop Production, University of 

Agriculture 


Nigeria 

Ogunwolu E. O. 

Department of Crop Production, 

University of Agriculture 


Nigeria 

222

Appendix C. 

PERÚ 

Duarte O. 

Departamento de Horticultura, 

Universidad Nacional Agrária 

La Molina, Lima 

Perú 

Franciosi R. 


Estación Experimental Agrícola 


Perú 

Rios Lobo M. L. 

Instituto Nacional de Investigación Agraria 

INIA 

La Molina, Lima 12, Lima 

Perú 

Riveros L. 

Faculdad de Ciências Forestales, 


12C Apartado 404 


Perú 

Villagarcia J. 




Perú 

PHILIPPINES 

Coronel R. E. 

University of Philippines, 


Los Baños, Laguna 

Philippines 

PORTUGAL 

de Freitas Nunes R. M. 

Direcção Regional de Agricultura 

Madeira 

Portugal 

Leca da Silva J. J. 

Secretaria Regional de Agricultura, Florestas e 

Pescas, Região Autônoma da Madeira 

São Martinho - 9000- 254, Funchal, Madeira 

Portugal 

SPAIN 

Alique R. 

Dpto. Ciencia y Tecnología de Productos 

Azcón-Aguilar C. 

Estación Experimental del Zaidin 

223

Appendix C. 

Vegetales, Instituto del Frio (C.S.I.C.) 

Ciudad Universitaria s/n 

28040 Madrid 

Spain 

C.S.I.C. 18008 Granada 

Spain 

Barcelo Muñoz A. 

Centro de Investigacion y Desarrollo Agrario 

29140 Churriana, Málaga 

Spain 

Caro E. 

Department of Hortofruticultura Subtropical, 

Estación Experimental "La Mayora" 

29750 Algarrobo, Costa, Málaga 

Spain 

Casorla J. M. 




Spain 

Cortés Martínez D. 

Departamento de Farmacología, 

Farmacognosía y Farmacodinamia, 

Facultad de Farmacia 

46100, Burjassot, Valencia 

Spain 

del Mar Sola M. 

Departamento de Bioquímica y Biología 

Molecular, 

Universidad de Granada 

18071 Granada 

Spain 

Farre Massip J. M. 

Consejo Superior de Investigaciones 

Científicas, 



Spain 

Galán Saúco V. 

ICIA, Departamento de Fruticultura Tropical 

Apartado 60, La Laguna, 38200 Tenerife 

Canary Islands 

Spain 

Garcia-Tapia J. 

Caja Rural Granada 

Fina "La Nacla", Mtril, Granada 

Spain 

Guirado E. 

Estacián Experimental "La Mayora" 


Spain 

Herrero M. 




Spain 

Hermoso González J. M. 

Consejo Superior de Investigaciones 

Científicas, 



Spain 

Herrero Castano A. 



Spain 

Lahoz J. M. 


López Encina C. 


224

Appendix C. 

Molecular, Universidad de Granada 

18071 Granada 

Spain 

Estación Experimental "La Mayora" (C.S.I.C) 

29750 Algarrobo Costa, Málaga 

Spain 

Martinez G. 

Centro de Edafología y Biologia Aplicada del 

Segura - CEBAS 

Avda. de la Fama, 130003 Murcia 

Spain 

Martinez Cayuela M. 



18071 Granada 

Spain 

Oliveira G. S. 

Dpto. Ciencia y Tecnología de Productos 

Vegetales, Ciudad Universitaria s/n 

28040 Madrid 

Spain 

Padilla I. M. G. 




Spain 

Pascual L. 


Molecular, 


18071 Granada 

Spain 

Majada J. P. 

Unidad de Fisiología Vegetal, 

Departamento B.O.S, 

Universidad de Oviedo 

33071 Oviedo 

Spain 

Perez M. A. 



Spain 

Pliego Alfaro F. 

Departamento de Biología Vegetal, 

Universidad de Málaga 

Campus de Teatinos s/n 

29071 Málaga 

Spain 

Perfectti F. 

Departamento de Genetica, 


Granada, 18071 

Spain 

Pretel M. T. 




Spain 

Riquelme E. 




Spain 

Romojaro F. 




Spain 

Rosell P. 




Ruiz Nieto A. 



Spain 

225

Appendix C. 

Spain 

Salto R. 



18071 Granada 

Spain 

Serrano M. 




Spain 

SOUTH AFRICA 

Conradie W. 

Citrus and Subtropical Fruit Research Institute 

Private Bag x11208, Nelspruit 1200 

South Africa 

du Preez R. J. 



South Africa 

Welgemoed C. P. 



South Africa 

TURKEY 

Tuzcu O. 

Department of Horticulture, 

University of Çukurova 

Adana - 01330 

Turkey 

UNITED KINGDOM 

Chatrou L. W. 

Molecular Systematic Section, 

Royal Botanic Gardens, Kew 

Richmond, Surrey, TW9 3DS 

United Kingdom 

Gold C. 

2 Hillside, Newbury Road, East Hendred 

Wantage, Oxon, OX12 8LE 


Haq N. 

Centre for Under-utilised Crops, University of 

Southampton 

Southampton, SO17 1BJ 

Wilkins R. M. 

Department of Agricultural and 

Environmental Science, 

University of Newcastle 

Newcastle-on-Tyne, NE1 7RU 

226

Appendix C. 



USA 

Bennett F. D. 

Tropical Research and Education Center, 

University of Florida 

18905 S.W. 280th Street 

Homestead, FL 33031 

USA 

Bustillo A. E. 

Tropical Research and Education Center 

18905 SW 280th Street 

Homestead FL 

USA 

Ellstrand N. C. 

Department of Botany and Plant Sciences, 

University of California 

Riverside, CA 92521-0124 

USA 

Grossberger D. 

Condor Growers 

6301 Worth Way 

93012, Camarillo 

USA 

Grumet R. 

Horticulture Department, 

Michigan State University 

East Lansing, MI 48824 

USA 

Huber D. J. 

Horticultural Science Department, 


P.O. Box 110690 

Gainesville, FL 32611 

USA 

Ismail M. A. 

Agricultural Research and Education Center, 

IFAS, University of Florida 

FL33850 Lake Alfred 

USA 

Lee J. M. 

Department of Botany and Plant Sciences, 

University of California 

Riverside, CA 92521-0124 

USA 

Nadel H. 





USA 

Nakasone H. Y. 

University of Hawaii at Manoa 

Honolulu, Him 

USA 

Patty L R. 

Department of Biochemistry, 

University of California, Riverside 

California 92521 

USA 

Pena J. E. 


Paull R. E. 

University of Hawaii at Manoa 

Honolulu, HI 

USA 

Ronning C. M. 

National Clonal Germplasm Repository, 

227

Appendix C. 




USA 

U.S. Dept. of Agriculture 

13601 Old Cutler Road 

Miami 

USA 

Subtropical Horticultural Research Unit, 

USDA 

13601 Old Cutler Road 

Miami FL 33158 

USA 

VENEZUELA 

Avilan Rovira L. 

FONAIAP - Centro Nacional de Investigaciones 

Agropecuarias 

Aptdo. 4653, Maracay 2101 

Venezuela 

León de Sierralta S. 

Departamento de Química, 

Facultad de Agronomía, 

La Universidad de Zulia 

Apartado 15205 

4005, Maracaibo 

Venezuela 

Martínez Vázquez M. 






Venezuela 

Medina D. 






Venezuela 

Ramírez M. 

Departamento de Botánica, 




4005, Macaraibo 

Venezuela 

228

Appendix D. Countries and 

Institutions with Collections of 

Germplasm 

Legend Headings: For the germplasm descriptions, the following legend was 

used: 

Taxon sample : taxon specific name 

Sample Type: AC – advanced cultivar; BL – breeding and inbred lines; CU – 

cultivated; GS – genetic stocks; IF – introgressed forms; LR – landrace or traditional 

cultivar; MT – mutants; WS – Wild/Weedy species; UN – unknown; no 

description 

Geogr. Origin: UN – Unknown; ISO code for the country where the sample was 

originally collected or bred; No description. 

N°: Number of accessions per taxon. 

Updated: date when record was last updated. 

AUSTRALIA 

1. Tropical Fruit Research Station, Department of Agriculture 

P.O. Box 72, Alstonville, New South Wales 2477 

Details of Holdings 

Taxon Sample Sample Type Geogr. Origin N° Updated 

A. cherimola No description Aus 2 31-12-1991 

A. hybrid (atemoya) AC Aus 2 31-12-1991 

A. diversifolia No description UN 1 31-12-1991 

A. reticulata No description UN 1 31-12-1991 

A. squamosa No description UN 1 31-12-1991 

2. Northern Territory Department of Primary Production 

P.O. Box 5160, Darwin, Northern Territory, 5794 Australia 



A. muricata BL UN 4 31-12-1991 

A. reticulata AC AUS 2 31-12-1991 

229

Appendix D. 

3. Maroochy Horticulture Research Station 

Queensland Department of Primary Industries 

P.O. Box 5083 SCMC, Nambour, Queensland, 4560 Australia 



A. cherimola AC AUS, ISR, USA 8 31-12-1991 

A. hybrid (atemoya) AC AUS., ISR, USA 20 31-12-1991 

BRAZIL 

1. Departamento de Genética da Universidade de Brasilia 

Caixa Postal 04477, 70919-000 Brasilia-DF 



A. coriacea No description BRA 1 07-05-1999 

A. crassiflora No description BRA 1 07-05-1999 

2. Laboratórios de Recursos Genéticos (CCTA) 

Universidade Estadual do Norte Fluminense 

Av. Alberto Lamego 2000, Bairro Horto 

28015-620 Campos dos Goytacazes-RJ 



A. muricata LR BRA 28 14-05-1999 

A. reticulata LR BRA 6 14-05-1999 

A. squamosa LR BRA 8 14-05-1999 

3. Estação Experimental de Fruticultura Tropical (EBDA) 

Via Conceição do Almeida-São Felipe, km 4 

Conceição do Almeida, Bahia State 


230

Appendix D. 


A. glabra WS BRA 1 30-04-1999 

A. muricata AC BRA (8), MEX 12 30-04-1999 

(1), VEN (3) 

A. squamosa WS BRA 4 30-04-1999 

4. Estação Experimental de Itajaí 

Empresa de Pesquisa Agropecuária de Santa Catarina - EPAGRI 

Via Antonio Heil, km 6 s/n; Caixa Postal 277, 88301-970 Santa Catarina 

State, e-mail: eeitajai@melim.com.br 



Annona cacaus WS BRA 1 13-05-1999 

5. Dept. de Horticultura, FCAVJ/UNESP 

Rodovia Carlos Tonanni, km 5 

Caixa Postal 145, 14870-000 Jaboticabal, São Paulo State 



A. cacaus WS UN 1 07-05-1999 

A. cherimola No description UN 2 07-05-1999 

A. cher. x .A squam. No description UN 5 07-05-1999 

A. coriacea WS UN 1 07-05-1999 

A. glabra WS UN 2 07-05-1999 

A. muricata LR UN 5 07-05-1999 

A. purpurea WS UN 1 07-05-1999 

A. reticulata No description UN 3 07-05-1999 

A. squamosa No description UN 2 07-05-1999 

6. Instituto Nacional de Pesquisa da Amazonia – INPA 

Ministério de Ciência e Tecnologia, Alameda Cosme Ferreira 1756, 

Caixa Postal 478, Manaus, Amazonas State 



A. montana LR BRA 1 28-04-1999 

7. Empresa Pernambucana de Pesquisa Agropecuária – IPA 

231

Appendix D. 

Av. General San Martin 1371, Bonji 

50761-000 Recife, Pernambuco State 



A. muricata LR BRA 18 06-10-1999 

A. squamosa LR/WS BRA 85 06-10-1999 

CAMEROON 

Center de Recherches Agronomiques de Njombe (IRA/CRA) 

P.O. Box 13, Nkongsamba 



A. muricata LR UN 4 18-03-1991 

A. reticulata LR UN 4 18-03-1991 

A. squamosa LR UN 1 18-03-1991 

COSTA RICA 

1. Estación Experimental “Fabio Baudrit Moreno” 

Universidad de Costa Rica (UCR), Apartado 183-4050, 

4050 Alajuela, Costa Rica, e-mail: eefbm@cariari.ucr.ac.cr 




A. muricata No description UN 10 19-04-1999 

2. Escuela de Ciencias Agrarias, Universidad Nacional 

Apartado 86, 3000 Heredia, Costa Rica 




3. Corporación Bananera Nacional S.A. (CORBANA) 

Apartado postal 390-7210, La Rita, Pococi, Limón, Costa Rica 

232

Appendix D. 



A. muricata LR, AC, GS No description 6 15-09-1993 

4. Asociation ANAI 

Apartado 170, 2070 Sabanilla, Montes de Oca, Costa Rica 



A. cherimola No description No description 1 17-09-1993 

A. diversifolia No description No description 1 17-09-1993 

A. montana No description No description 1 17-09-1993 

A. muricata No description No description 1 17-09-1993 

A. purpurea No description No description 1 17-09-1993 

A. reticulata No description No description 1 17-09-1993 

A. spp. No description No description 2 17-09-1993 

A. squamosa No description No description 1 17-09-1993 

A. hybrid (atemoya) No description No description 2 17-09-1993 

5. Centro Agronomico Tropical de Investigación y Ensenanza (CATIE) 

Apartado 7170, e-mail: name@catie.ac.cr 

7170 Turrialba, Cartago, Costa Rica. 



A. glabra No description BRA 2 15-04-1999 

A. muricata No description AC, COL, CRI, 50 15-04-1999 

ECU, HND, SLV, 

MEX, PAN, PRI 

A. pittieri No description CRI 1 15-04-1999 

A. purpurea No description PAN, CRI, SLV, 4 15-04-1999 

GTM, MEX 

A. reticulata No description CRI, SLV, GTM, 3 15-04-1999 

HND, MEX, PAN 

PER, USA 

A. spp. No description CRI, HND 2 15-04-1999 

233

Appendix D. 

CUBA 

1. Instituto de Investigaciones Fundamentales en Agricultura Tropical 

(INIFAT), Calle 1, esq. 2, Stgo. de las Vegas; e-mail: inifat@cenial.inf.cu, 

17200 Boyeros, Ciudad de la Habana 



A. cherimola AC No description 1 30-07-1999 

A. cherimola x AC No description 1 30-07-1999 

A. purpurea 

A. cinerea No description No description 2 30-07-1999 

A. glabra No description No description 1 30-07-1999 





A. salzmannii No description No description 1 30-07-1999 


2. Dirección de Investigaciones de Citros y Otros Frutales 

Calles 7ª y 42 Miramar 

La Habana, Cuba 



A. cherimola BL No description 1 31-12-1991 

A. muricata BL No description 3 31-12-1991 

A. reticulata BL No description 4 31-12-1991 

A. squamosa BL No description 2 31-12-1991 

CYPRUS 

Plant Genetic Research and Herbarium Agricultural Research Institute 

P.O. Box 2016, e-mail: ari@athena.cc.ucy.ac.cy 

Nicosia, Cyprus 



A. cherimola AC USA 4 11-01-1999 

234

Appendix D. 

ECUADOR 

1. Centro Andino de Tecnologia Rural (CATER) 


Casilla 399, Loja, Ecuador 



A. cherimola WS ECU 150 19-04-1999 

2. Estación experimental Napo Payamino (INIAP) 

km 5 via Coca-Lago Agrio, El Coca Napo, Ecuador 



Annona glabra LR ECU 20 14-04-1999 

3. Estación Experimental Portoviejo (INIAP) 

km 12 Carretera Santa Ana, Apartado 13-01-100 

e-mail: iniapo@po.iniap-ecuador.gov.ec 

Portoviejo, Manabi 



A. cherimola LR ECU 1 10-06-1999 

A. muricata LR ECU 1 10-06-1999 

A. reticulata LR ECU 1 10-06-1999 

4. Estacion Experimental Tropical Pichilingue (INIAP) 

km 5 via Quevedo-El Empalme, C.P. 24; 

Quevedo, Los Rios 



A. cherimola LR ECU 1 19-04-1999 

A. muricata LR ECU 4 19-04-1999 

A. squamosa LR ECU 3 21-04-1999 

235

Appendix D. 

5. Granja Experimental de Tumbaco (INIAP) 

Casilla 2600, Tumbaco, Pichincha 



A. cherimola LR ECU, ESP, USA, 66 14-04-1999 

PER, CRI, AUS 

EL SALVADOR 

Centro Nacional de Tecnología Agropecuaria y Forestal (CENTA) 

Km 33,5 Carretera Santa Ana a Cantón; C.P. 885; 

e-mail: cdtzal@es.com.sv 

Arce, San Andrés, Dept. La Libertad 



A. diversifolia WS SLV 23 10-06-1999 

A. glabra WS HND 2 10-06-1999 

A. muricata WS SLV 30 10-06-1999 

A. purpurea WS SLV 2 10-06-1999 

A. reticulata WS SLV 4 10-06-1999 

FRANCE 

CIRAD-FLHOR Station de la Guadeloupe 

Neufchateau-Sainte Marie 

97130 Capesterre Belle-Eau 



A. cherimola No description CRI, ESP, AUS 4 03-01-1992 

A. hybrid (atemoya) No description USA, AUS 2 03-01-1992 

A. muricata No description CRI, BRA, AUS 13 03-01-1992 

A. reticulata No description GLP, GTM 2 03-01-1992 

A. squamosa No description USA, AUS, THA, 4 

NCL 

03-01-1992 

236

Appendix D. 

GERMANY 

Greenhouse for Tropical Crops 

Inst. Prod./Nutr. World Crops, Un. Kassel 

Steinstrasse 19, 37213 Witzenhausen 


Taxon Sample Sample Type Geogr. Origin N o Updated 

A. cherimola No description ESP 1 17-08-1994 

A. muricata No description IDN 1 17-08-1994 

A. squamosa No description TWN 1 17-08-1994 

GHANA 

Crop Research Institute Plant Genetic Resources Unit 

P.O. Box 7, Bunso 



A. hybrid (atemoya) AC No description 1 11-07-1994 

A. muricata AC No description 1 11-07-1994 

A. reticulata AC No description 1 11-07-1994 

A. squamosa AC No description 1 11-07-1994 

GRENADA 

Caribbean Agricultural Research & Development Institute (CARDI) 

Westerhall, St. David´s, P.O. Box 270 

e-mail: cardignd@caribsurf.com 

St. George´s 



A. hybrid (atemoya) AC USA 2 12-04-1999 

A. squamosa AC, LR GRD, CUB 2 12-04-1999 

GUATEMALA 

237

Appendix D. 

Centro Universitario de Sur Occidente (CUNSUROC) 

Universidad de San Carlos 

Apartado Postal 606, Mazatenango, Suchitepequez 



A. diversifolia WS GTM 3 14-05-1999 

A. glabra WS GTM 8 14-05-1999 

A. muricata WS GTM 4 14-05-1999 

A. primigenia WS GTM 1 14-05-1999 

A. purpurea WS GTM 5 14-05-1999 

A. reticulata WS GTM 6 14-05-1999 

A. scleroderma WS GTM 2 14-05-1999 

HONDURAS 

1. Centro Universitario Regional del Litoral Atlantico (CURLA) 

Km 8, Carretera La Ceiba-Tela, Apartado 89 

La Ceiba, Dept. de Atlantida 



A. scleroderma No description No description 1 24-05-1999 

2. Jardin Botanico Wilson Popenoe de Lancetilla 

Apartado Postal 49, Tela, Atlantida 







Annona spp. No description No description 1 13-07-1999 

INDIA 

Indian Institute of Horticultural Research 

Hessaraghatta Lake Post, 560089 Bangalore, Karnataka 

238

Appendix D. 




ISRAEL 

Horticultural Institute Volcani Center 

P.O. Box 6 

e-mail: vhwisma@volcani.bitnet 

50250 Bet Dagan 




JAMAICA 

1. Research Development Division Ministry of Agriculture & Mining 

Hope Gardens, P.O. Box 480. 

Kingston 



Annona spp. AC No description 9 31-12-1991 

2. College of Agriculture, Science and Education 

Passley Gardens, Portland, Post Box 170. 

Port Antonio, Portland 



A. muricata WS JAM 1 09-04-1999 

A. squamosa WS JAM 1 24-01-2000 

MALAWI 

Bvumbwe Agricultural Research Station 

239

Appendix D. 

P.O. Box 5748, Limbe 



Annona spp. AC BRU, MOZ 3 31-12-1991 

MEXICO 

Instituto Nacional de Investigaciones Forestales, Agricolas y Pecuarias 

(INIFAP), Serapio Rendon 83, 06470 Col. San Rafael, México-D.F. 




A. cherimola AC No description 2 19-04-1999 


PANAMÁ 

1. División de Mejoramiento Genético (INDIAP) 

Apartado 6-4391. El Dorado 

Panamá 




2. Facultad de Ciencias Agropecuarias, Universidad de Panamá 

Apartado Estafeta Universitad, Panamá City 



A. cinerea No description No description 1 14-06-1999 


PAPUA NEW GUINEA 

240

Appendix D. 

Lowlands Agriculture Experimental Station (LAES) 

P.O. Box 204, Kokopo, East New Britain Province 




PERU 

1. Universidad Nacional Hermilio Valdizan (UNHEVAL) 

Giron 2 de Mayo 680; Apartado 278 

Huánuco 



A. cherimola LR PER 11 19-05-1999 

2. Instituto de Desarrollo del Medio Ambiente 

Jr. Junin 459 

e-mail: ldmahua@net.cosapidata.com.pe 

Huánuco 




3. Universidad Agraria La Molina 

Av. La Universidad, Apartado 456 




A. cherimola AC PER 4 02-01-1992 


A. muricata WS PER 6 02-01-1992 

4. Estacion Experimental Pucallpa – Ucayali (INIA) 

241

Appendix D. 

Av. Centenario km 4, Apartado 203 

Pucallpa, Coronel Portillo 



A. muricata LR PER 2 21-05-1999 

PHILIPPINES 

1. Institute of Plant Breeding, College of Agriculture (UPLB) 

e-mail: opd@ipb.uplb.edu.ph 

4031 College, Laguna 



A. hybrid (atemoya) AC No description 8 02-01-1992 

A. muricata WS No description 7 02-01-1992 

A. reticulata WS No description 11 02-01-1992 

Annona spp. WS No description 5 02-01-1992 

A. squamosa WS No description 43 02-01-1992 

2. National Plant Genetic Resources Laboratory (IPB/UPLB) 

College, 4031 Laguna 




PORTUGAL 

Departamento de Fitotecnia Estação Agronomica Nacional 

2780 Oeiras 



Annona cherimola AC No description 7 13-02-1995 

242

Appendix D. 

PUERTO RICO 

Agricultural Experiment Station, University of Puerto Rico 

HC-01, P.O. Box 1165; e-mail: eealajas@caribe.net 

00667 Lajas 



A. muricata WS PRI 7 19-05-1999 

SAINT LUCIA 

Caribbean Agricultural Research & Development Institute (CARDI) 

La Ressource, Dennery, Post Box 971 

e-mail: cardi@candw.lc 

Castries 




SEYCHELLES 

Grand Anse Experimental Centre Ministry of Agric. & Fisheries 

P.O. Box 166 

Mahe 




SOUTH AFRICA 

Institute for Tropical and Subtropical Crops 

Private Bag X11208, 1200 Nelspruit, Transvaal 



243

Appendix D. 

A. cherimola AC CHL 4 11-11-1991 

A. cherimola WS USA 7 11-11-1991 

SUDAN 

Horticultural Research Section, Agricultural Research Corporation 

P.O. Box 126, Wad Medani 



A. squamosa AC SDN 7 02-01-1992 

SURINAME 

1. Agricultural Experimental Station, Ministry of Agriculture 

L. Vriesdelaan 10, Post Box 160, Paramaribo 




2. STIPRIS (Foundation for Experimental Gardens), Tijgerkreek-West and 

Boma, L. Vriesdelaan 9, Post Box 160. 

Paramaribo 





TAIWAN 

Chiayi Agricultural Experiment Station (TARI) 

2, Min-cheng Road 

60014 Chia-yi 


244

Appendix D. 







A. hybrid (atemoya) No description No description 3 10-10-1991 

TANZANIA 

Tropical Pesticides Research Institute 

POB 3024, Arusha 





UNITED STATES OF AMERICA (USA) 

1. Agricultural Experiment Station 

University of the Virgin Islands 

R.R. # 02 Box 10000, Kingshill, St. Croix, USVI 00850 



Annona spp. AC USA 9 07-04-1999 

2. Tropical Agricultural Research Station 

Clonal Repository USDA-ARS, P.O. Box 70, 00709-0070 Mayaguez 




3. Department of Horticulture, College of Agriculture Science 

University of Puerto Rico, Mayaguez, Puerto Rico 


245

Appendix D. 


A. muricata WS No description 250 02-01-1992 

4. Subtropical Horticultural Research Unit (USDA) 

13601 Old Cutler Road, Miami-FL 33158 

http://www.ars-grin.gov/ars/SoAtlantic/Miami/homes 



A. bullata No description No description 2 12-08-1988 



A. hybrids No description No description 21 12-08-1988 






VENEZUELA 

Centro Nacional de Investigaciones Agropecuarias (CENIAP-FONAIAP) 

Apartado 4653, 2101 Maracay, Estado de Aragua 

e-mail: fonaiap@conicit.ve 



A. muricata BL VEN 7 14-04-1998 

Source: IPGRI, 2001. 

246

Glossary 

A 

abcission - The normal shedding of leaves, flowers or fruits from a plant at a 

special separation layer. 

abiotic - not living, as opposed to biological. 

acute - terminating with a sharp or well defined angle. 

actinomorphic - having radially symmetric shape, usually refers to the petals 

of a flower. 

acuminate - the shape of a tip or base of a leaf or perianth segment where the 

part tapers gradually and usually in a concave manner. 

adnate - joined to or attached to. 

adventitious - describes an organ growing where it is not normally expected. 

albumen - Starchy and other nutritive material in a seed, stored as 

endosperm inside the embryo sac, or as perisperm in the surrounding nucellar 

cells; any deposit of nutritive material accompanying the embryo. 

alternate - describes leaves that are not opposite to each other on the axis, 

but arranged singly and at different heights. 

anatropous - bent over through 180 degrees to lie alongside the stalk. 

androecium - all the male reproductive organs of a flower; the stamens. cf. 

gynoecium. 

angiosperm - a plant producing seed enclosed in an ovary. A flowering 

plant. 

annual - a plant that completes its life cycle from germination to death 

within one year. 

anterior - front; on the front side; away from the axis. 

anther - the pollen-bearing (terminal) part of the male organs (stamen), 

borne at the top of a stalk (filament). 

247

Glossary 

anthesis - flower bud opening; strictly, the time of expansion of a flower 

when pollination takes place, but often used to designate the flowering 

period; the act of flower bud opening. 

apex - the tip of an organ, the growing point. 

apical - pertaining to the apex. 

apiculate - having a short point at the tip. 

arcuate - bow-shaped. 

areole - a small pit or cavity marked out upon a surface. 

asexual - lacking sexual characteristics, or when referring to reproduction, 

occurring without the fusion of egg and sperm. 

autotrophy - refers to a process by which an organism that can process 

inorganic materials in to organic by using energy outside the organism such 

as sunshine on chlorophyll. 

axil - the upper angle formed by the union of a leaf with the stem. 

axillary - pertaining to the organs in the axil, e.g. buds flowers or 

inflorescence. 

axis - the main or central stem of a herbaceous plant or of an inflorescence. 

B 

basal - borne on or near the base. 

biotic - refers to any aspect of life, but especially to characteristics of entire 

populations or ecosystems. 

bipinnate - (of leaves) a pinnate leaf with primary leaflets themselves 

divided in a pinnate manner; cf pinnate. 

blade - the flattened part of a leaf; the lamina. 

budding - method of propagating woody plants. A cutting of one variety, 

called the scion with buds attached, is joined onto another related species or 

variety called the rootstock. As the plant grows, the two parts graft together 

to form one plant. 

248

Glossary 

C 

caducous - falling off early, or prematurely, as the sepals in some plants. 

calyx - the outer whorl of floral envelopes, composed of the sepals. 

carpel - one of the flowers' female reproductive organs, comprising an ovary 

and a stigma, and containing one or more ovules. 

chlorosis - a yellowing of the leaves, reflecting a deficiency of chlorophyll 

and caused by waterlogging or a lack of nutrients, often iron. 

clone - a group of plants that have arisen by vegetative reproduction from a 

single parent, and which therefore all have identical genetic material. 

cordate - heart-shaped, often restricted to the basal portion rather to the 

outline of the entire organ. 

coriaceous - of leathery texture. 

cotyledon - seed leaf; the primary leaf or leaves in the embryo. 

crenate - shallowly round-toothed, scalloped. 

crenulate - finely crenate. 

cross pollination - the transfer of pollen from the anther of the flower of one 

plant to the flowers of a different plant. 

cultivar - a race or variety of a plant that has been created or selected 

intentionally and maintained through cultivation. 

cuneate - wedge-shaped; triangular with the narrow end at point of 

attachment, as in the bases of leaves or petals. 

cuspidate - with an apex abruptly and sharply constricted into an elongated, 

sharp-pointed tip. 

cyme - a broad, more or less flat-topped, determinate flower cluster, with 

central flowers opening first. 

cymose - inflorescence showing the cyme arrangement. 

249

Glossary 

D 

deciduous - falling at the end of one season of growth or life, as the leaves of 

non-evergreen trees. 

decoction - herbal preparation made by boiling a plant part in water. 

deflexed - bent abruptly downward; deflected. 

dehiscence - the method or process of opening a seed pod or anther. 

dentate - with sharp, spreading, course indentations or teeth, perpendicular 

to the margin. 

denticulate - minutely or finely dentate. 

derived - originating from an earlier form or group. 

dichogamy - the differing times of maturation of stamens and pistils in a 

flower. 

dichotomous - forked, in 1 or 2 pairs. 

dicotyledon - a flowering plant with two cotyledons. 

diploid - having two sets of chromosomes. 

dorsal - upon or relating to the back or outer surface of an organ. 

downy - covered with short and weak soft hairs. 

E 

ecosystem - an interacting complex of a community, consisting of plants 

and/or animals and functioning as an ecological unit. 

elliptic - oval in outline. 

emarginate - having a shallow notch at the extremity. 

endocarp - the inner layer of the pericarp or fruit wall. 

endosperm - the starch and oil-containing tissue of many seeds. 

entomophilous -insect pollinated. 

250

Glossary 

epigynous - borne on or arising from the ovary; used of floral parts when the 

ovary is inferior and flower not perigynous. 

explant - a plant part asceptically excised and prepared for culture in a 

culture medium. 

exocarp - the outer layer of the pericarp or fruit wall. 

F 

flacate - scythe-shaped; curved and flat, tapering gradually. 

fasicle - a condensed or close cluster. 

faveolate - honey-combed. 

ferruginous - pertaining to or coloured like iron rust. 

filament - thread; particularly the stalk of the stamen, terminated by the 

anther. 

filiform - thread-shaped, long, slender and terete. 

flexuose - zig-zig; bending from side to side; wavy. 

fulvous - dull, brownish-yellow. 

fuscous - dusky, greyish-brown. 

G 

genotype - the genetic constitution of an organism, acquired from its parents 

and available for transmission to its offspring. 

genus - a group of related species, the taxonomic category ranking above a 

species and below a family. 

glabrous - not hairy. 

glaucous - bluish white; covered or whitened with a very fine, powdery 

substance. 

globose - globe-shaped. 

glabrescent - becoming glabrous with age. 

251

Glossary 

grafting - a method of propagation, by inserting a section of one plant, 

usually a shoot, into another, so that they cam grow together into a single 

plant. 

gynoecium - all the female parts of a flower. 

H 

haploid - half the full set of genetic material found in a chromosome. 

hermaphrodite - bisexual - with both male and female reproductive parts in 

the same flower. 

homonym - a scientific name given two or more times to plants of the same 

taxonomic rank but which are quite distinct from each other. 

hybrid - a cross-breed of two species, usually having some characteristics of 

both parents. 

hypocotyl - the axis of an embryo below the cotyledons which on seed 

germination develops into the radicle. 

I 

indehiscent - not regularly opening , as a seed pod or anther. 

indigenous - native and original to the region. 

inflorescence - the flowering part of a plant and especially the mode of its 

arrangement. 

integuments - an outer covering or coat. 

K 

karyotype - characterization of a chromosome set of an individual or group. 

L 

lamina - a blade, the leafy portion of a frond or leaf. 

252

Glossary 

lanceolate - shaped like a lance head, several times longer than wide, 

broadest above the base and narrowed toward the apex. 

lateral - side shoot, bud etc. 

lamellae - a thin, flat plate or laterally flattened ridge. 

lenticellate - having a body of cells as a pore, formed on the periderm of a 

stem, and appearing on the surface of the plant as a lens-shaped spot. 

locular - having a cavity or chamber inside the ovary, anther or fruit. 

M 

membranous - thin in texture, soft and pliable. 

mesocarp - the fleshy middle portion of the wall of a succulent fruit between 

the skin and the stony layer. 

micro-propagation - propagation of plants thought tissue culture. 

monophyletic - descended from a single ancestral line. See Also: 

polyphyletic. 

mucronate - terminated abruptly by a distinct and obvious spur or spiny tip. 

N 

naturalized - to cause a plant to become established and grow undisturbed as 

if native. 

nectar - sweet secretion of glands in many kinds of flower. 

nectrotic - Death of cells or tissues through injury or disease. 

nectiferous - producing nectar. 

O 

oblique - slanting, unequal sided. 

obovate - inverted ovate; egg-shaped, with the broadest part above. 

obtuse - blunt or rounded at the end. 

253

Glossary 

octaploid - having 8 times the basic number of chromosomes. 

orbicular - circular. 

ovary inferior - with the flower-parts growing from above the ovary. 

ovary superior - with the flower-parts growing from below the ovary. 

ovate - egg-shaped, with the broader end at the base. 

ovule - the body which after fertilization becomes the seed. 

P 

parthenocarpic - refers to the fruiting of plants which have not been 

pollinated or otherwise fertilized. 

pedicel - a tiny stalk; the support of a single flower. 

pendulous - more or less hanging or declined. 

peduncle - a primary flower stalk supporting either a cluster or a solitary 

flower. 

perianth - the floral envelope consisting of the calyx and corolla. 

pericycle - the tissue of the stele lying just inside the endodermis. 

perigynous - adnate to the perianth, and therefore around the ovary and not 

at its base. 

petal - a division of the corolla; one of a circle of modified leaves 

immediately outside the reproductive organs, usually brightly coloured. 

petiole - the stalk of a leaf that attaches it to the stem. 

phenology - the science of the relations between and periodic biological 

phenomena 

phenotype - the morphological, physiological, behavioural, and other 

outwardly recognizable forms of an organism that develop through the 

interaction of genes and environment. 

pilose - hairy, especially with soft hairs. 

pilosulous - minutely pilose. 

pinnate - a compound leaf consisting of several leaflets arranged on each 

side of a common petiole. 

254

Glossary 

pistil - the seed-bearing organ of the flower, consisting of the ovary, stigma 

and style when present. 

polygamous - bearing male and female flowers on the same plant. 

polyphyletic - having members that originated, independently, from more 

than one evolutionary line. 

polyploidy - having more than two sets of chromosomes. 

polyporate - pollen grain with many apertures. 

prolate - having flattened sides due to lengthwise elongation. 

propagate - to produce new plants, either by vegetative means involving the 

rooting or grafting of pieces of a plant, or sexually by sowing seeds. 

protandrous - refers to a flower, when the shedding of the pollen occurs 

before the stigma is receptive. 

protogynous - referring to a flower where the shedding of the pollen occurs 

after the stigma has ceased to be receptive. 

psilate - referring to a pollen grain having a smooth surface. 

pubescent - covered with hairs, especially short, soft and down-like. 

R 

raceme - a simple inflorescence of pediceled flowers upon a common more 

or less elongated axis. 

rachis - the main stalk of a flower cluster or the main leafstalk of a 

compound leaf. 

radicle - the portion of the embryo below the cotyledons that will form the 

roots. 

ramification - branching. 

reticulate - in the form of a network, net-veined. 

retuse - with a shallow notch at a rounded apex. 

rootstock - the root system and lower portion of a woody plant to which a 

graft of a more desirable plant is attached. 

rotundate - nearly circular; orbicular to oblong. 

rugose - wrinkled. 

255

Glossary 

rugulose - covered with minute wrinkles. 

S 

scandent - climbing but not self-supporting. 

scarify - to scar or nick the seed coat to enhance germination. 

scion - a cutting from the upper portion of a plant that is grafted onto the 

rootstock of another plant, usually a related species. 

self pollination - the transfer of pollen from the anther of a flower to the 

stigma of the same flower, or different flowers on the same plant. 

sepal - a division of a calyx; one of the outermost circle of modified leaves 

surrounding the reproductive organs of the flower. 

serrate - having sharp teeth pointing forward. 

serrulate - finely serrate. 

sessile - without a stalk. 

sheath - a tubular envelop. 

spinescent - 1. having spines 2. terminating in a spine 3. modified to form 

a spine. 

stamen - one of the male pollen-bearing organs of the flower. 

stigma - that part of a pistil through which fertilization by the pollen is 

effected. 

stipule - an appendage at the base of a petiole, often appearing in pairs, one 

on each side, as in roses. 

style - the usually attenuated portion of the pistil connecting the stigma and 

ovary. 

subulate - awl-shaped. 

sulcate - grooved or furrowed. 

syncarp - an aggregate or multiple fruit produced from coherent or fused 

pistils, the small single fruits massing and growing together into a single 

fruit. 

256

Glossary 

T 

tetraploid - having 4 sets of chromosomes (twice the normal number of 

chromosomes). 

testa - the outer seed coat. 

tomentose - covered with a thick felt of radicles; densely pubescent with 

matted wool. 

tomentulose - rather tomentose. 

tomentum - closely matted, woolly hairs. 

transverse - cross-wise in position. 

triploid - having three sets of chromosomes. 

tropism - the movement of an organism in response to an external source of 

stimulus, usually toward or away from it. 

truncate - ending abruptly, as if cut off transversely. 

tuberculate - bearing tubercles, covered with warty lumps. 

U 

unguiculate - narrowed, clawed. 

V 

valvate - open by valves. 

Z 

zygomorphic - capable of division by only one plane of symmetry. 

257

Index 

accessions, 52, 53, 153, 229 

acetogenins, 35, 36, 44, 45, 46, 

155, 198, 199, 202 

agroforestry, 47 

cover crops, 99, 100 

intercropping, 88, 99, 150 

windbreak, 93 

agronomy 

direct seeding, 91 

fertilisation See fertilisation, 151 

field establishment, 85 

irrigation See irrigation, 151 

land preparation, 85 

mulching, 92, 99 

planting systems, 86 

pruning, 28, 92, 93, 94, 96, 97, 

98, 124, 128, 129, 131, 133, 

141, 150, 154 

spacing, 28, 92, 114 

topworking, 78, 83, 84 

training, 94, 97, 150 

transplanting, 71, 73, 81, 92, 93, 

94, 100 

Angola, 21, 24 

Annona arenaria, 5 

Annona asiatica, 5 

Annona bonplandiana, 4 

Annona cearensis, 4 

Annona chrysophylla, 5 

Annona chrysophylla var. 

porpetac, 5 

Annona cinerea, 5, 54, 234, 240 

Annona diversifolia, 47, 48, 49, 54, 

124, 229, 233, 236, 238, 246 

Annona excelsa, 4 

Annona glabra, 1, 3, 10, 13, 48, 49, 

54, 78, 231, 233, 234, 235, 236, 

238, 246 

Annona laevis, 4 

Annona longifolia, 4, 49 

Annona longipes, 49 

Annona macrocarpa, 4 

Annona montana, 3, 11, 48, 49, 54, 

78, 231, 233, 234, 245, 246 

Annona porpetac, 5 

Annona pubescens, 4 

Annona purpurea, 1, 48, 49, 54, 

233, 234, 236, 238 

Annona riparia, 4 

Annona scleroderma, 1, 48, 49, 54, 

238 

Annona senegalensis var. porpetac, 

5 

Annona stenophylla, 49 

Annona tripetala, 4 

Antilles, 19, 21, 48, 49, 52 

Argentina, 27, 28, 49, 128, 144 

atemoya, 3, 22, 25, 26, 66, 79, 93, 

120, 123, 126, 136, 150, 229, 

230, 233, 236, 237, 242, 245 

Australia, 21, 22, 23, 24, 53, 98, 

117, 121, 122, 123, 148, 151, 

230 

Bolivia, 19, 24 

Botswana, 49 

Brazil, 1, 5, 6, 21, 24, 25, 26, 29, 

31, 32, 33, 34, 44, 48, 49, 50, 53, 

65, 66, 73, 75, 93, 98, 99, 102, 

104, 105, 112, 114, 115, 117, 

118, 120, 123, 124, 126, 128, 

130, 141, 142, 143, 144, 145, 

147, 148, 149, 151, 152 

breeding, 48, 153 

budding, 49, 74, 75, 78, 80, 81, 82, 

83, 153 

double bud, 82 

inverted T, 80, 82 

patch, 82, 83 

shield, 82 

258

Index 

Chile, 4, 19, 23, 24, 26, 27, 28, 38, 

49, 100, 101, 128, 143, 146, 151 

chilling injury, 135 

China, 21, 22, 29, 54, 98, 128, 151 

China-Taiwan, 128 

chromosome number, 48 

cimarrona, 48 

climate, 12, 28, 29, 32, 38, 98, 100, 

106, 111, 152 

altitude, 19, 21, 26, 28, 29, 30, 

32, 129, 152 

rainfall, 21, 28, 29, 30, 31, 34, 

124 

relative humidity, 30, 91, 101, 

115, 125 

shade, 28, 30, 131 

temperature, 21, 28, 29, 30, 31, 

32, 98, 99, 100, 101, 115, 125, 

126, 129, 130 

wind, 28 

Colombia, 24, 82, 106, 107, 117, 

118, 128, 129, 131, 141, 151 

common names, 3, 4, 22 

composition 

acids, 35, 36, 39, 46 

alkaloids, 35, 36, 44, 45, 46 

amino acids, 36, 38 

carbohydrates, 37, 38, 39 

falvonoids, 35, 36, 44, 45 

fats, 35, 36 

fatty acids, 36, 37 

fibre, 13, 37, 38 

minerals, 37, 38 

oils, 35, 45 

proteins, 37, 38 

sugars, 38, 39 

tannins, 39, 45 

terpenes, 35 

vitamins, 37, 38, 39, 44 

water, 39 

conservation, 48, 49, 50, 51, 152 

ex situ, 49, 51, 152 

in situ, 49, 50, 152 

in vitro, 51 

on farm, 50, 152 

Costa Rica, 10, 21, 24, 53, 128, 

144 

Cuba, 21, 22, 24, 29, 30, 39, 53, 

118 

cultivars, 10, 13, 17, 66, 77, 80, 81, 

82, 83, 98, 102, 103, 118, 123, 

127, 128, 130, 132, 134, 137, 

139, 141, 142, 146, 148, 151, 

153 

cuttings See vegetative 

propagation, 75, 77, 78, 79 

demand See economics, 1, 25, 26, 

27, 41, 99, 100, 107, 111, 139, 

144, 147, 151, 152, 154 

description of genus, 6 

description of species, 7 

direct seeding See agronomy, 91 

diseases, 26, 71, 75, 116, 122, 133 

anthracnose, 123, 124 

Armillaria luteobubalina, 123 

armillaria root rot, 123 

Athelia rolfsii, 122 

bacterial wilt, 123 

black canker, 123, 124 

black scab, 123 

blight, 123 

Botryodiplodia theobromae, 123, 

124 

burn of string, 123 

Cercospora anonae, 123 

Colletotrichum gloeosporioides, 

123, 124 

Corticium koleroga, 123, 125 

Corticum salmonicolor, 123, 125 

Cylindrocladium clavatum, 122 

damping-off, 122 

diplodia rot, 123, 124 

fumagina, 123, 125 

Fusarium spp., 122, 123 

Gliocladium roseum, 123 

Glomerella cingulata, 123 

Helicotylenchus spp., 123, 126 

leaf spot, 123 

Meloidogyne spp., 123 

nematodes, 73, 116, 126 

Phakopsora cherimoliae, 123 

259

Index 

Phoma spp, 123 

Phomopsis spp, 123, 124 

Phytophthora palmivora, 123, 

124 

Phytophthora spp, 122, 123 

Pithyium spp, 123 

purple blotch, 123, 124 

Ralstonia solanacearum, 123 

Rhizoctonia solani, 122 

Rhizopus stolonifer, 90, 123, 

125, 134 

rubelose, 123, 125 

rust fungus, 123 

Sclerotium coffeicolum, 123, 125 

Sclerotium rolfsii, 122 

seedling blight, 123 

Stigmella spp., 123, 125 

Thanatephorus cucumeris, 122 

zoned spot, 123 

diversity, 19, 21, 22, 48, 49, 50, 52, 

152 

Dominican Republic, 24, 25, 128, 

144 

economics, 2, 50, 52, 111, 117, 

119, 121, 139, 142, 149, 151, 

152 

costs, 79, 80, 82, 85, 92, 108, 

112, 113, 117, 139, 140, 141, 

142, 143, 145, 151 

demand, 1, 25, 26, 27, 41, 99, 

100, 107, 111, 139, 144, 147, 

151, 152, 154 

employment, 145 

income, 85, 88, 100, 140, 141, 

142, 143, 145, 150 

yield, 23, 24, 25, 26, 65, 71, 73, 

85, 94, 97, 100, 101, 110, 111, 

116, 140, 141, 142, 143, 150, 

153, 154 

Ecuador, 19, 24, 34, 49, 52, 53, 81, 

117, 118, 120, 140, 149 

Egypt, 26, 28, 100 

endemism, 50 

evaluation, 153 

export See marketing, 23, 24, 26, 

130, 140, 144, 146, 152, 154 

fertigation, 111 

fertilization, 75, 103, 104, 110, 111 

flower 

anthesis, 7, 12, 100, 128 

description, 7, 11, 13, 16, 17 

phenology, 28, 29, 32, 33, 73, 

98, 99, 101, 102, 128 

pollination, 6, 7, 13, 17, 22, 26, 

28, 32, 94, 100, 101, 102, 103, 

112, 129, 130, 133, 150 

uses, 47 

flowers 

uses, 45 

France, 27, 53 

fruit, 1, 100, 102, 117, 124, 138, 

142, 144, 148 

description, 7, 13, 14, 17 

form(cherimoya), 9, 10 

fruit-set, 13, 17, 101, 102, 103, 

109, 111, 112, 150, 153 

maturation, 32, 33, 129, 133 

phenolgy, 32, 33 

phenology, 32, 33 

properties, 37, 38, 39, 45 

thinning, 103 

uses, 41, 42, 43, 44, 45 

fruit-set, 100, 101 

genebank, 51, 52 

genetic, 51, 152 

diversity, 50, 153 

erosion, 48, 152 

improvement, 152 

transformation, 153 

variability, 48 

genetic resources, 48, 49, 50, 130 

germplasm, 10, 19, 21, 51, 52, 53, 

84, 127, 151, 152, 153, 229 

collection, 53, 151, 153 

grafting, 49, 71, 74, 75, 77, 78, 80, 

81, 82, 83, 92, 153 

cleft, 80 

splice, 80 

veneer, 83 

260

Index 

whip-and-tongue, 82, 83 

Guanabanus muricatus, 4 

Guanabanus squamosus, 5 

Guatemala, 4, 48, 49, 53 

harvesting, 2, 26, 32, 94, 96, 99, 

103, 105, 111, 118, 125, 127, 

128, 129, 130, 131, 132, 133, 

140, 141, 143, 153 

harvesting period, 140 

harvesting point, 90, 128, 130, 

131, 132, 133 

Honduras, 48, 49, 52, 53, 102 

hybridization, 3, 22, 52, 79 

ilama, 48 

inarching See vegetative 

propagation, 78 

India, 4, 10, 17, 21, 24, 25, 27, 33, 

34, 46, 47, 53, 99, 104, 117, 

123, 124, 128, 139, 151 

intercropping See agroforestry, 88, 

99, 150 

irrigation, 24, 26, 75, 85, 91, 98, 

99, 111, 112, 115, 116, 126, 

139, 141, 143, 154 

drip, 112, 113, 114 

micro-sprinkler, 112, 113, 114 

salinization, 116 

water quality, 113 

Israel, 22, 23, 26, 28, 53, 66 

Jamaica, 24, 45, 53 

Japan, 27, 144 

Kenya, 5, 21, 30, 47 

layering See vegetative 

propagation, 75, 77, 79, 80 

leaves, 79 

description, 7, 11, 13, 16, 17 

properties, 35, 36, 45, 46 

uses, 44, 45, 46, 47 

Malawi, 17, 49, 53 

marketing, 2, 23, 25, 26, 27, 50, 

65, 71, 85, 103, 111, 121, 130, 

131, 139, 140, 141, 143, 144, 

145, 146, 147, 148, 149, 150, 

151, 152, 154, 155 

domestic, 23, 25, 143 

export, 23, 26, 130, 140, 144, 

146, 152, 154 

international, 23, 145, 146, 155 

Mauritania, 21 

Mexico, 1, 5, 10, 19, 21, 22, 23, 

24, 26, 29, 31, 32, 33, 34, 44, 

48, 49, 52, 53, 65, 117, 119, 

120, 121, 123, 124, 125, 129, 

144, 151 

micropropagation See vegetative 

propagation, 75, 84, 153 

molecular markers, 19, 152 

Mozambique, 21 

mulching See agronomy, 99 

Namibia, 49 

nursery, 73, 75, 76, 77, 92, 94, 122 

origin, 2, 10, 19, 21, 22, 26, 49, 66 

packaging, 148, 149, 150, 154 

Panama, 19, 24, 49, 53 

Pemba, 21 

Peru, 19, 23, 24, 25, 28, 34, 49, 52, 

54, 117, 129, 140, 143, 144 

pests, 26, 37, 45, 71, 75, 89, 116, 

117, 118, 119, 120, 121, 126, 

133 

Acromyrmex spp, 118 

Aethalion spp, 117 

Amblypelta nitida, 117 

Anastrepha ludens, 120 

Anastrepha obliqua, 117 

annona moth, 117 

Anonaepestis bengalella, 117, 

119 

Antiteuchus tripterus, 117 

aphids, 117, 121 

Aphis gossypii, 117 

atis moth borer, 117 

Atta spp, 118 

Bactrocera dorsalis, 117 

Bephratelloides maculicollis, 

117, 119 

Ceratitis capitata, 117, 120 

Cerconota anonella, 117, 119 

cherimoya seed borer, 117 

Cratosomus bombina, 117, 118 

Dysmicoccus spp, 117 

261

Index 

Empoasca fabae, 117 

Ferrisia virgata, 117 

fruit borer, 66, 118 

fruit fly, 119, 120, 121 

fruit borer, 89 

fruit-spotting bug, 122 

Heilipus catagraphus, 117 

hemipterous insects, 117 

leaf hopper, 117 

leaf larvae, 118 

leaf miners, 118 

Leptoglossus zonatus, 117 

Leucoptera spp, 118 

mealy bugs, 117, 121 

Membracis foliata, 117 

Oenomaus ortygnus, 117 

Phyllocnistis spp, 118 

Planococcus citri, 121 

root grubs, 118 

scale insects, 117 

seed borer, 89, 118, 119, 124, 

125 

spider mites, 117 

Talponia batesi, 117, 119 

thecla moths, 117 

Toxoptera aurantii, 117 

trunk borer, 118 

wasps, 117 

phenology, 101, 111 

Philippines, 4, 21, 22, 25, 33, 54, 

83, 117, 118, 119, 129, 144, 151 

pollen, 7, 88, 101, 102, 103 

germination, 17, 100, 101 

maturity, 101 

viability, 100, 101 

pollination, 6, 7, 13, 17, 22, 26, 28, 

32, 94, 100, 101, 102, 103, 112, 

129, 130, 133, 150 

hand, 7, 13, 88, 101, 102, 103, 

128 

insect, 7, 100, 112 

Polynesia, 21 

pond apple, 1, 48 

Portugal, 26, 27, 28, 54, 93, 129, 

223 

postharvest handling, 2, 123, 125, 

127, 132, 133, 134, 136, 146, 

150, 153, 154 

processing, 2, 23, 25, 39, 41, 42, 

43, 65, 127, 135, 136, 137, 138, 

139, 141, 142, 144, 145, 146, 

150, 151, 154 

canning, 42 

freezing, 25, 41, 42 

pasteurization, 41, 43 

pulp preservation, 42 

production, 1, 2, 22, 23, 24, 25, 26, 

27, 29, 34, 47, 49, 93, 98, 102, 

107, 111, 129, 139, 140, 141, 

142, 143, 144, 145, 146, 147, 

151, 154 

areas, 22, 23, 24, 25, 26, 34, 

143, 144 

rootstock See vegetative 

propagation, 31, 47, 49, 73, 74, 

75, 77, 80, 81, 82, 83, 84, 92, 

122, 123, 124, 150, 153 

seed 

dormancy, 72, 98 

germination, 71, 72, 73, 74, 87 

pre-treatment, 72 

propagation, 65, 71, 73 

properties, 46 

sowing, 74, 108 

storage, 51, 71 

selection, 22, 65, 118, 127, 150, 

151, 153 

Senegal, 21 

shelf life, 26, 125, 127, 139, 146, 

154 

soil requirements, 10, 31, 73, 85, 

114 

solarization, 73, 87 

soncoya, 1, 48 

Spain, 10, 19, 23, 27, 28, 32, 34, 

54, 104, 112, 114, 117, 119, 

120, 123, 129, 140, 141, 143, 

144, 146, 151 

Sri Lanka, 21 

262

Index 

storage, 42, 44, 51, 107, 123, 133, 

134, 135, 136, 137 

Sudan, 21, 54 

Tanzania, 5, 21, 30, 33, 34, 54, 129 

Thailand, 24, 25, 98, 144 

topworking See agronomy, 78, 83, 

84 

transportation, 85, 133, 143, 145 

tree management See agronomy, 

13, 32, 92, 93, 112, 116, 117, 

124, 150, 151, 154 

United Kingdom, 27 

United States of America, 19, 21, 

22, 23, 24, 25, 27, 29, 54, 117, 

123, 129, 144, 147, 148, 155, 

227 

uses, 2, 42, 44, 45, 46, 150, 153, 

155 

antibacterial, 44 

anti-bacterial, 35 

anti-cancer, 35, 44 

anti-helminthic, 35 

cytotoxic, 35, 36, 37, 44, 45 

drinks, 39, 41, 43, 136, 138 

essential oils, 36, 43, 45 

food, 37, 42 

industrial, 42, 43, 150 

insecticide, 35, 46, 94, 105, 118 

medicinal, 1, 35, 44, 45, 46, 48, 

150, 153, 155, 198 

rootstocks, 47, 49, 73, 74, 75, 

77, 80, 81, 82, 83, 84, 92, 

122, 123, 124, 150, 153 

vegetative propagation, 71, 75, 77, 

78, 79, 80 

budding See budding, 49, 74, 

75, 78, 80, 81, 82, 83, 153 

cuttings, 75, 77, 78, 79, 94, 98, 

132 

grafting See grafting, 49, 71, 74, 

75, 77, 78, 80, 81, 82, 83, 92, 

153 

inarching, 75, 80 

layering, 75, 77, 79, 80 

micro-propagation, 51, 75, 84, 

153 

rootstocks, 31, 47, 49, 73, 74, 

75, 77, 80, 81, 82, 83, 84, 92, 

122, 123, 124, 150, 153 

Venezuela, 24, 25, 54, 104, 116, 

117, 144 

windbreak See agroforestry, 93 

yield, 23, 24, 25, 26, 65, 71, 73, 

85, 94, 97, 100, 101, 110, 111, 

116, 140, 141, 142, 143, 150, 

153, 154 

Zanzibar, 21 

Zimbabwe, 49, 129 

263

Annona Species Monograph.pdf - Crops for the Future

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?