Tropical Plant Biol. (2011) 4:62–89
DOI 10.1007/s12042-011-9068-3
Sugarcane (Saccharum X officinarum): A Reference Study
for the Regulation of Genetically Modified Cultivars
in Brazil
Adriana Cheavegatti-Gianotto & Hellen Marília Couto de Abreu & Paulo Arruda &
João Carlos Bespalhok Filho & William Lee Burnquist & Silvana Creste &
Luciana di Ciero & Jesus Aparecido Ferro & Antônio Vargas de Oliveira Figueira &
Tarciso de Sousa Filgueiras & Mária de Fátima Grossi-de-Sá & Elio Cesar Guzzo &
Hermann Paulo Hoffmann & Marcos Guimarães de Andrade Landell &
Newton Macedo & Sizuo Matsuoka & Fernando de Castro Reinach & Eduardo Romano &
William José da Silva & Márcio de Castro Silva Filho & Eugenio César Ulian
Received: 6 October 2010 / Accepted: 13 January 2011 / Published online: 22 February 2011
# The Author(s) 2011. This article is published with open access at Springerlink.com
Abstract Global interest in sugarcane has increased significantly in recent years due to its economic impact on
sustainable energy production. Sugarcane breeding and
better agronomic practices have contributed to a huge
increase in sugarcane yield in the last 30 years. Additional
increases in sugarcane yield are expected to result from the
use of biotechnology tools in the near future. Genetically
modified (GM) sugarcane that incorporates genes to
increase resistance to biotic and abiotic stresses could play
a major role in achieving this goal. However, to bring GM
sugarcane to the market, it is necessary to follow a
regulatory process that will evaluate the environmental
and health impacts of this crop. The regulatory review
process is usually accomplished through a comparison of
the biology and composition of the GM cultivar and a nonGM counterpart. This review intends to provide informa-
Communicated by: Marcelo C. Dornelas
A. Cheavegatti-Gianotto : H. M. C. de Abreu : S. Matsuoka :
E. César Ulian (*)
CanaVialis/Alellyx S.A., Rua James Clerk Maxwell,
320, 13069-380 Campinas, São Paulo, Brasil
e-mail: eugenio.c.ulian@monsanto.com
A. Cheavegatti-Gianotto
e-mail: adriana.cheavegatti@alellyx.com.br
S. Matsuoka
e-mail: sizuo.matsuoka@gmail.com
S. Creste : M. G. de Andrade Landell
IAC/APTA - Centro de Cana, Instituto Agronômico de Campinas,
Rodovia Antonio Duarte Nogueira, Km 321,
CP 206, 14032-800 Ribeirão Preto, São Paulo, Brazil
L. di Ciero
Amyris Crystalsev Biocombustíveis Ltda.,
Rua James Clerk Maxwell,
315, 13069-380 Campinas, São Paulo, Brasil
P. Arruda
Centro de Biologia Molecular e Engenharia Genética,
Universidade Estadual de Campinas,
13083-875 Campinas, São Paulo, Brasil
F. de Castro Reinach
Amyris Crystalsev Biocombustíveis Ltda., Amyris Inc,
5885 Hollis St, Ste 100,
Emeryville, CA 94608, USA
e-mail: fernando.reinach@gmail.com
J. C. Bespalhok Filho
Universidade Federal do Paraná,
Rua dos Funcionários, 1540, Cabral,
80035-050 Curitiba, Paraná, Brasil
J. A. Ferro
Campus de Jaboticabal, Departamento de Tecnologia,
Universidade Estadual Paulista,
14884-900 Jaboticabal, SP, Brasil
W. L. Burnquist
Centro de Tecnologia Canavieira,
CP 162, 13400-970 Piracicaba, São Paulo, Brasil
A. V. de Oliveira Figueira
CENA/USP - Laboratório de Melhoramento de Plantas,
CP 96, 13400-970 Piracicaba, São Paulo, Brasil
Tropical Plant Biol. (2011) 4:62–89
tion on non-GM sugarcane biology, genetics, breeding,
agronomic management, processing, products and byproducts, as well as the current technologies used to develop
GM sugarcane, with the aim of assisting regulators in the
decision-making process regarding the commercial release
of GM sugarcane cultivars.
Keywords Biosafety . Ethanol . Biofuel . Saccharum .
Sugarcane
Introduction
Economic interest in sugarcane has increased significantly in recent years due to the increased worldwide
demand for sustainable energy production. The Brazilian
experience in sugarcane ethanol production has paved the
way for the establishment of a consolidated world supply
to meet the demand of a proposed ethanol addition of
approximately 10% to gasoline worldwide. It is estimated
that the Brazilian production of sugarcane must double in
the next decade to meet this goal. In the future,
biotechnological advances might help to reduce the
environmental impacts of increased sugarcane production
by developing solutions that would produce more
sugarcane with decreased requirements for fertilizers
and water. Among these solutions, GM sugarcane will
play a key role in providing growers with more
productive and resistant varieties. In Brazil, the National
Biosafety Technical Commission (CTNBio) has already
approved field trials with genetically modified sugarcane,
incorporating traits such as increased yield, drought
tolerance, insect resistance and herbicide tolerance. It is
expected that the enormous research and development
efforts being conducted by government and private
T. de Sousa Filgueiras
Reserva Ecológica do IBGE,
CP 08770, 70312-970 Brasília, Distrito Federal, Brazil
M. d. F. Grossi-de-Sá
Embrapa Recursos Genéticos e Biotecnologia,
Parque Estação Biológica,
Av. W5 Norte, 70770-900,
CP 02372 Brasília, Distrito Federal, Brasil
63
institutions will in the medium-term result in commercial
release in Brazil of genetically modified sugarcane.
This review was produced to serve as a source of
baseline information of conventional sugarcane to assist in
the decision-making process related to possible future
commercial release of GM sugarcane cultivars. Here, we
will cover the origin of the species that have contributed to
the genetic makeup of modern sugarcane cultivars, the
agronomic behavior and growth habit of sugarcane, the
botanical aspects of its reproductive biology, its genetic
composition, potential for lateral gene transfer by pollen, its
model of seed dispersion, and allergenicity of its derived
products. This review also addresses the utilization of
sugarcane and the byproducts generated by the sugarcane
agribusiness.
Economic Importance
The sugarcane crop has been relevant to the Brazilian
economy since the beginning of the 16th century. The first
sugarcane plants were brought from Madeira Island and
established in Brazil around 1515; the first sugar mill was
established in 1532. Currently, Brazil is the world’s largest
sugarcane producer, with approximately 7.5 million cultivated hectares, which produced approximately 612 million
tons in the 2009/2010 crop season. Approximately half of
the sugarcane was used to produce sugar, and the remainder
was used to produce 25 billion liters of ethanol (CONAB
2009). Brazil also produces sugarcane for animal feed,
cachaça (sugarcane spirit), and sugarcane syrup, among
other products.
In 2009, Brazil’s sugar and ethanol exports generated
approximately US$ 9.9 billion in revenue, ranking sugarcane third among its exports (CONAB 2009).
H. P. Hoffmann
Campus de Araras, Universidade Federal de São Carlos,
Rodovia Anhanguera, Km 174,
CP 153, 13600-970 Araras, São Paulo, Brasil
N. Macedo
Araujo & Macedo Ltda.,
Rua Oswaldo Cruz, 205, Jardim Santa Cruz,
13601-252 Araras, São Paulo, Brasil
e-mail: newton_macedo@yahoo.com.br
E. Romano
Embrapa Recursos Genéticos e Biotecnologia,
Empresa Brasileira de Agropecuária,
Av. W5 Norte, 70770-900,
CP 02372 Brasília, Distrito Federal, Brasil
W. J. da Silva
Dow Agroscience, Rodovia Anhanguera,
Km 344,
Jardinópolis 14680-000 São Paulo, Brasil
E. C. Guzzo
Embrapa Tabuleiros Costeiros, Unidade de Execucao de Pesquisa,
BR 104 Norte, Km 85, 57061-970,
CP 2013 Maceió, Alagoas, Brasil
M. de Castro Silva Filho
Departamento de Genética, Escola Superior de Agricultura Luiz
de Queiroz, Universidade de São Paulo,
Avenida Pádua Dias, 11,
CP 83, 13400-970 Piracicaba, São Paulo, Brasil
64
Currently, the Brazilian sugar-ethanol agribusiness is
experiencing a period of excitement due to the combination
of extremely favorable factors, including prospects for the
growth of internal and external markets, a long-term trend of
rising international oil prices, possession of the world’s lowest
ethanol production costs, growth of the flex fuel fleet, and a
universal concern for sustainable energy alternatives. These
conditions have attracted global attention to the Brazilian fuel
ethanol program and the country’s potential to supply a
significant part of the world’s demand for renewable fuel. The
result has been a substantial increase in sugarcane production
over the last few years. In 2007, the planted area grew by 12%
over the previous year and the same level of growth is
expected for the next several decades (FNP 2009).
Geographical Distribution
Sugarcane is grown in all tropical and subtropical regions of
the world, on both sides of the equator, up to approximately
35° N and 35° S (van Dillewijn 1952; Gomes and Lima
1964). In 2007, the main sugarcane-producing countries
were Brazil (33% of the world’s production), India (23%),
China (7%), Thailand (4%), Pakistan (4%), Mexico (3%),
Colombia (3%), Australia (2%), the United States (2%) and
the Philippines (2%) (FNP 2009).
In Brazil, sugarcane cultivation is concentrated in the
southeastern region, which is responsible for approximately
70% of the national sugarcane production (Fig. 1). Northeastern Brazil, another traditional producing area, is
responsible for 14% of sugarcane production, and midwestern
Brazil, where the crop is rapidly advancing, represents 13% of
the national production (CONAB 2009). It should be noted
that, despite the concerns about the expansion of sugarcane
Fig. 1 Sugarcane production in
regions of Brazil (Source:
Conab 2009)
Tropical Plant Biol. (2011) 4:62–89
cultivation toward the Amazon region, the actual cultivation
in this region is minimal and decreasing.
Classification and Nomenclature
The genus Saccharum was first described by Linnaeus
(1753) in his book Species Plantarum. The generic name is
derived from the Greek word sakcharon, which means sugar
and was duly Latinized by the author. The book described
two species: Saccharum officinarum L. and S. spicatum L.,
which is currently classified under the genus Perotis (P.
spicata (L.) T. Durand and H. Durand) (Dillon et al. 2007).
The taxonomy and nomenclature of the genus has always
been challenging (Bentham 1883; Hackel 1883; Pilger 1940;
Hitchcock 1951; Bor 1960; Almaraj and Balasundaram 2006).
The genus has two known synonymous names, Saccharophorum and Saccharifera. When it was initially described, the
genus consisted only of five to ten species from the Old
World, including S. officinarum, S. spontaneum, S. sinense, S.
edule and S. barberi. Later, several species that were allocated
in other genera, including Andropogon, Anthoxanthum,
Eriochrysis, and Erianthus, were transferred to Saccharum.
The case of Erianthus is particularly interesting.
Currently, the genus Erianthus comprises species from the
Old World and New World that were previously separated
into two different genera since the Old World species were
firstly placed under the genus Ripidium (Grassl 1971). Later,
authors classified the Old World species under Erianthus,
section Ripidium (Almaraj and Balasundaram 2006). This
distinction is supported by the fact that the Old World
species contain a flavonoid (di-C-glycoside) that is absent in
New World species (Williams et al. 1974). Cordeiro et al.
(2003), analyzing these two groups by microsatellites
Tropical Plant Biol. (2011) 4:62–89
markers also found that Erianthus accessions clustered
separated in accordance to their geographical origin.
Erianthus is considered to be closely related to Saccharum
and many species have been assigned to either of these
genera, depending on the criteria used. Several botanists,
however, have considered that they are distinct genera
(Hooker 1896; Haines 1921, Jeswiet 1925; Grassl 1946;
Dutt and Rao 1950; Mukherjee 1958). This distinction has
been reinforced by studies using the presence of root tip
tannin (Rao et al. 1957), leaf lipoid (Vijayalakshmi and Rao
1963), esterase isozyme alleles (Waldron and Glasziou
1972), and flavonoid composition (Williams et al. 1974)
and microsatellites markers (Cordeiro et al. 2003).
Despite these data, the genera and species are more often
separated and identified primarily on the basis of floral
characteristics, which are considered by botanists to be
more stable than vegetative morphological characters and
the basic criterion used to differentiate both genera is the
presence (in Erianthus) or absence (in Saccharum) of a
floret structure called an awn, which is an extension of the
mid-rib of the floral bract, on the top of lemma II or the
upper lemma. The lemma in which the awn is absent is
referred to as awnless. The current trend regards Erianthus
as synonymous with Saccharum because this commonly
used criterion to differentiate both genera (presence/absence
of awn on the lemma) is variable and is not a consistent
characteristic in the complex (Bor 1960; Renvoize 1984;
Clayton and Renvoize 1986). Therefore, the genus Saccharum
currently comprises all species that were previously described
under Erianthus. The current sugarcane botanical classification is shown in Table 1.
The tribe Andropogoneae consists of tropical and subtropical grass species that are grown in the Old World and New
World. The most important cultivated members of the tribe are
corn (Zea mays) and sorghum (Sorghum bicolor) (Daniels
and Roach 1987). Saccharum and Sorghum share many
similarities in their genetic composition, as they originated
from a common ancestral lineage that diverged approximately
65
5–8 million years ago (Al-Janabi et al. 1994; Guimarães et al.
1997; Figueira et al. 2008).
Botanical Description
The currently cultivated sugarcane plants are hybrids
derived from crossings mainly between plants of S.
officinarum and S. spontaneum (Dillon et al. 2007). The
plants are perennial grasses that form stools of stalks or
culms that can be several meters in length and are juicy,
with high concentrations of sucrose.
The sugarcane root system consists of adventitious and
permanent shoot root types. Adventitious roots emerge
from the culm root zone and are responsible for water
uptake during bud sprouting and plant support until the
permanent roots develop. Permanent roots are fasciculated
at the base of growing shoots and are classified into support
or anchor roots and the more network like absorption roots.
(Mongelard 1968; Thompson 1964; Moore and Nuss
1987). The ratio between one root type and another are
somewhat species specific. Saccharum officinarum generally contains fewer support roots than does S. spontaneum
(Moore 1987a), which could explain the increased vigor
and resistance to environmental stresses characteristic of S.
spontaneum.
The stalk or culm consists of alternating nodes and
internodes. On the node, there is a leaf scar, an axillary bud
and a circumferential band of axillary root primordia. Stalk
morphology is highly variable from one genotype to
another and represents an important element for varietal
characterization (Martin 1961). Sugarcane leaves are
alternate and are attached to the stalk, with one leaf per
internode. Sheathes consist of the sheath proper and the
much smaller acropetal blade joint consisting of a leaf
collar, dewlap, ligule and auricles. The shape, size and
Table 1 Current sugarcane classification
Order:
Poales
Family:
Subfamily:
Tribe:
Poaceae
Panicoideae
Sub
Tribe:
Genus:
Species:
Saccharinae
Andropogoneae
Saccharum
Saccharum officinarum; Saccharum spontaneum;
Saccharum sinense; Saccharum barberi; Saccharum
robustum; Saccharum edule (Old World). Saccharum
villosum; Saccharum asperum (New World) and others.
Fig. 2 Kuijper’s leaf numbering system (1915)
66
Tropical Plant Biol. (2011) 4:62–89
Fig. 3 Diagram of a sugarcane
propagule: a evident propagule,
cupulate coma, empty pedicel
and rachilla plus sessile spikelet.
The other figures depict the
different parts of the spikelet. b
Glume I. c Glume II. d Palea. e
Flower with two lodicules, three
stamens, and gynoecium with
ovary and two feathered stigmas
(Illustration: Klei Sousa)
distribution of trichomes and the shape of the ligule and
auricles are traits of taxonomic importance for varietal
identification. Sugarcane leaves are numbered from top to
bottom starting with the uppermost leaf showing a visible
dewlap designated as leaf +1 (Fig. 2) (Moore 1987a).
The inflorescence of sugarcane is a ramified, conoidal
panicle with a main stem, called the rachis, which is the
continuation of the last stalk internode. The rachis holds
secondary branches which in turn hold tertiary branches.
The spikelets are located at the base of the tertiary branches
and on the top of the secondary branches. Each spikelet has
one flower, which is disposed alternately along the
inflorescence secondary and tertiary branches. At the base
of the spikelet, there is a ring of silky, colorless trichomes
(‘coma’) that covers the spikelet (Fig. 3a) and help with
spikelet dispersion. Next, there is a series of bracts called
glumes (‘glume I’ and ‘glume II’), both glabrous (Fig. 3b
and c); upper lemma (or fertile lemma); and palea, which is
hyaline and without veins and may be either rudimentary or
absent (Fig. 3d). When the inflorescence matures, an
anemochoric (by wind) dispersion of the propagules
begins. The propagules consist of the coma, some floral
clusters and the spikelet (Fig. 3a). The flowers (Fig. 3e)
consist of two lodicules, the androecium and the gynoe-
cium. The pollen grains are spherical when fertile and
prismatic when sterile. Sugarcane fruit, called the caryopsis (Fig. 4), is dry, indehiscent and one-seeded, and it
cannot be separated from the seed. The fruit can only be
distinguished from the seed when viewed with scanning
electronic microscopy.
Fig. 4 Sugarcane caryopses. Note the presence of the remains of
styles at the tip of the caryopsis and the differentiated embryo region
(at the opposite extremity of the style remains) (Photograph: Alellyx)
Tropical Plant Biol. (2011) 4:62–89
Hybridization and Introgression
Modern sugarcane cultivars are the products of crosses
between species of the genus Saccharum that were made by
breeders in the late 19th century (Matsuoka et al. 1999).
The most important species contributing to modern sugarcane varieties were S. officinarum, which was widely
cultivated for its ability to accumulate sucrose in its stalks,
and S. spontaneum, which is a vigorous, widely adapted
wild species which contributed genes for disease and stress
resistance. The species S. sinense, S. barberi and S.
robustum also provided minor contributions toward the
development of some modern sugarcane varieties.
S. officinarum L. is generally known as the noble cane
because it is stout and produces abundant sweet juice.
Culms are thick (normally over 3.5 cm in diameter) and
soft due to low fiber content. Assessions of this species
display long, wide leaf blades (1 m long×5 cm wide)
and a relatively small, shallow root system (Scarpari and
Beauclair 2008). S. officinarum is highly demanding in
specific climate conditions, high soil fertility and water
supply. S. officinarum accessions are generally susceptible
to diseases such as mosaic, gummosis, leaf scorch, root rot
and Fiji disease (Martin 1961; Ricaud and Autrey 1989;
Ricaud and Ryan 1989), but they tend to be resistant to
sugarcane smut (Segalla 1964). S. officinarum includes all
old traditional sugarcane varieties that were cultivated
throughout the world prior to the introduction of hybrid
varieties (Segalla 1964).
Known as wild sugarcane, S. spontaneum L. is highly
polymorphic, with plant stature ranging from small grasslike plants without stalks to plants over 5 m high with long
stalks. Leaf blades vary in width from very narrow mostly
restricted to the mid-rib or up to a width of 4 cm
(Matsuoka et al. 1999). Plants show highly adaptive
plasticity and are found in different environments. S.
spontaneum is the species that has contributed to the
improvement in sugarcane vigor, hardness, tillering,
ratooning ability and resistance to biotic stresses (Mohan
Naidu and Sreenivasan 1987). These plants tend to be
immune to most diseases, including ‘Sereh’ and mosaic,
but they are susceptible to sugarcane smut (Segalla 1964).
In some regions of the world, such as the United States, S.
spontaneum is considered a harmful invasive species
(USDA 2008). In Brazil, it is considered an exotic and
non-invasive plant (Instituto Horus 2008; Global Invasive
Species Database 2008).
Saccharum sinense Roxb. and Saccharum barberi Jesw.
are known as Chinese or Indian canes, as they were initially
grown in China and India before the spread of modern
varieties (Mohan Naidu and Sreenivasan 1987). Some
taxonomists consider them a single species (Matsuoka et
al. 1999), and according to more modern studies, they are
67
natural hybrids between S. officinarum and S. spontaneum
(Irvine 1999; D’Hont et al. 2002). Stems from both species
are long (up to 5 m), thin (approximately 2 cm in diameter)
and fibrous, presenting long, fusiform internodes. The
plants have a vigorous, well-developed root system and
good tillering, which enables adaptation to poor and dry
soil and allows the production of large volumes of biomass.
However, both species have medium sugar content and
early maturation. S. sinense and S. barberi tend to exhibit
resistance to root diseases; some individuals are resistant to
mosaic, immune to ‘Sereh’ disease, resistant to sugarcane
borers and susceptible to sugarcane smut (Segalla 1964). Due
to poor flowering and sterility of most genotypes of these
species, they are rarely used in sugarcane breeding programs
(Roach and Daniels 1987), although they may have
contributed to the development of some modern varieties
(Mohan Naidu and Sreenivasan 1987; Dillon et al. 2007).
S. robustum Brandes and Jeswiet ex Grassl represents
wild sugarcanes adapted to broad environmental conditions. It possesses a high fiber content and vigorous
stalks that are 2.0–4.4 cm in diameter and up to 10 m
high, but like S. officinarum, it does not have rhizomes.
The culms are hard and have little juice, are poor in sugar
content and have a hard rind, a characteristic that is
exploited to build hedges (Matsuoka et al. 2005; Mozambani
et al. 2006). S. robustum tends to be highly susceptible to
mosaic (Segalla 1964). Few current commercial cultivars
have received a contribution from this species. However,
there are reports of a successful program to broaden the
genetic basis of the current varieties by introgressing S.
robustum into Hawaiian sugarcane varieties (Mohan Naidu
and Sreenivasan 1987).
Centers of Origin and Diversity
The genus Saccharum probably originated before the
continents assumed their current shapes and locations. The
genus consists of 35–40 species and has two centers of
diversity: the Old World (Asia and Africa) and the New
World (North, Central and South America). Asia has
approximately 25 native species, North America six native
species and four or five introduced species, and Central
America has three or four native and some introduced
species (Webster and Shaw 1995). Africa has two native
and Australia have one naturalized species (Darke 1999;
Bonnett et al. 2008).
The Brazilian Saccharum species have not been well
characterized. Only regional floristic surveys have reported
the presence of these species. One study described the
native species S. asperum, S.angustifolium, S. purpureum,
S. biaristatum, S. glabrinodis, S. clandestinus and S.
villosum, but the authors commented that these species
68
were poorly defined so that it is possible that they all might
be variations of a single species (Smith et al. 1982). In fact,
from the species listed on this work, only S. asperum, S.
angustifolium and S. villosum are currently accepted
scientific names (The Plant List 2010). In another study,
the native species were identified as S. villosum, S. asperum
and S. baldwinii (Filgueiras and Lerina 2001).
The Saccharum species involved in the development of
modern sugarcane cultivars originated from Southeast Asia
(Roach and Daniels 1987). Because S. officinarum and S.
spontaneum are the major contributors to the genomes of
modern varieties, the geographical origins of these species
will be described in more detail.
S. officinarum has been cultivated since prehistoric times
(Sreenivasan et al. 1987). It is believed that its center of origin
is Polynesia and that the species was disseminated throughout
Southeast Asia, where a modern center of diversity was
created in Papua New Guinea and Java (Indonesia); this is the
region where the majority of specimens were collected in the
late 19th century (Roach and Daniels 1987).
The center of origin and diversity of S. spontaneum is
the more temperate regions of subtropical India. However,
because S. spontaneum can be grown in a wide range of
habitats and altitudes (in both tropical and temperate
regions), it is currently spread over latitudes ranging from
8°S to 40°N in three geographic zones: a) east, in the South
Pacific Islands, Philippines, Taiwan, Japan, China, Vietnam,
Thailand, Malaysia and Myanmar; b) central, in India, Nepal,
Bangladesh, Sri Lanka, Pakistan, Afghanistan, Iran and the
Middle East; and c) west, in Egypt, Kenya, Sudan, Uganda,
Tanzania, and other Mediterranean countries. These zones
roughly represent natural cytogeographical clusters
because S. spontaneum tends to present a different number
of chromosomes in each of these locations (Daniels and
Roach 1987).
Genetic Constitution
Saccharum species present high ploidy levels. S. officinarum
is octoploid (2n=80) having x=10 chromosomes, which is
the basic chromosome number of members of the Andropogoneae tribe (D’Hont et al. 1995; Cesnik and Miocque 2004;
Nobrega and Dornelas 2006). S. spontaneum has x=
8 chromosomes (D’Hont et al. 1996) but presents great
variation in chromosome numbers with five main cytotypes:
2n=62, 80, 96, 112 or 128 (Daniels and Roach 1987;
Sreenivasan et al. 1987).
Modern sugarcane cultivars, which were derived from the
hybridization between these two species, are considered
allopolyploid hybrids (Daniels and Roach 1987), with most
exhibiting a 2n+n constitution, representing two copies of
the S. officinarum genome plus one copy of the S.
Tropical Plant Biol. (2011) 4:62–89
spontaneum genome (Cesnik and Miocque, 2004). The S.
officinarum genome usually duplicates when it is hybridized
with S. spontaneum. This phenomenon facilitated the work
of the first breeders because nobilization consisted of
increasing the ratio of the S. officinarum to that of the S.
spontaneum genome (Bremer, 1961). In situ hybridization
studies have shown that the genomes of modern hybrids are
composed of 10–20% of S. spontaneum chromosomes, 5–
17% of recombinant chromosomes containing part of S.
officinarum and part of S. spontaneum chromosomes and
the remainder composed of S. officinarum chromosomes
(Piperidis and D’Hont, 2001; D’Hont 2005).
The hybrids are usually aneuploid, with a prevalence of
bivalents, a significant proportion of univalents and rare
multivalent associations during meiosis (Daniels and Roach,
1987). Despite this genome complexity, evidence suggests a
diploid-like mode of inheritance (Hogarth, 1987).
The “Saccharum Complex” Theory
It has been hypothesized that an intercrossing group named
the “Saccharum complex” consisting of the genera Saccharum (including species previously classified under
Erianthus sect. Ripidium), Sclerostachya, Narenga and
Miscanthus sect. Diandra provided the basis for modern
sugarcane varieties (Mukherjee, 1957; Roach and Daniels
1987; Daniels and Daniels, 1975). Saccharum officinarum
was likely derived from crosses involving S. spontaneum,
Miscanthus, S. arundinaceus (Syn: Erianthus arundinaceus)
and S. robustum (Roach and Daniels, 1987). The presence of
whole S. officinarum chromosomes, homologous to chromosomes from Miscanthus and from some Saccharum species
previously classified as belonging to Erianthus sect. Ripidium,
supports the hypothesis of hybridization among these species
giving rise to Saccharum officinarum (Daniels and Roach,
1987; Besse et al., 1997).
Despite the fact that the aforementioned Saccharum
complex hypothesis is currently broadly accepted, particularly by sugarcane breeders, who consider species within
the “Saccharum Complex” as the primary gene pool for
sugarcane breeding, recent molecular data do not support
this theory (D’Hont et al. 2008). Thus, there is a
suggestion that Saccharum is a well-defined lineage that
diverged over a long evolutionary period from the
lineages leading to the Erianthus and Miscanthus genera
(Grivet et al. 2004).
Sugarcane Breeding
Sugarcane breeding is based on the selection and cloning of
superior genotypes from segregating populations that was
Tropical Plant Biol. (2011) 4:62–89
69
obtained by crossing contrasting individuals. To maximize
the efficiency of this rather long process, it is divided into
various phases, including choosing suitable parentals and
quantifying environmental effects on the expression of the
selection characters.
The first step of a sugarcane breeding program consists
of establishing a germplasm collection in a heavy flowering
area where the flowering time of the parental lines can be
synchronized. To meet sugarcane flowering requirements,
the breeding stations operating in Brazil are all located in
the heavy flowering northeastern region and include the
breeding programs of CanaVialis at Maceió (AL), Rede
Interuniversitária para o Desenvolvimento do Setor
Sucroalcooleiro - RIDESA at Murici (AL), Centro de
Tecnologia Canavieira – CTC at Camamú (BA) and
Instituto Agronômico de Campinas - IAC at Uruca (BA).
Other sugarcane breeding programs have been active in
Brazil in the past and the codes for identifying varieties
developed in those programs are listed in Table 2.
A typical sugarcane variety development program
(Fig. 5) begins by making a large number of crosses among
selected parental genotypes. The resulting seeds give rise to
a large number of progeny (seedlings), which are needed to
increase the chance of obtaining improved cultivars from
superior genetic combinations. The selection process is
conducted in distinct locations to identify superior genotypes with improved agronomical performance and tolerance to biotic and abiotic stresses. On average, one
commercial variety can be obtained for every 250,000
seedlings evaluated in the first stage of the breeding
program (T1). The selection process continues in the
second and third phases, which are evaluated under
different environmental conditions (Fig. 5).
Table 2 Main Brazilian sugarcane breeding programs
(*) Still in operation
Reproductive Biology
Sugarcane flowering is regulated by day length known as
the photoperiod. Flowering induction and development
occur when the hours of light decreases from 12.5 h to
11.5 h. Panicle emergence occurs with an additional
decrease to approximately 11 h. Under these conditions in
the southern hemisphere, flowering occurs close to the
autumnal equinox (March 21st) and is delayed by approximately 2 days for each additional degree of latitude (Brett,
1951; Moore and Nuss, 1987). Flowering induction is only
effective after the juvenile period has been completed, i.e.,
when at least two to four internodes have matured
(Clements and Awada, 1967; Coleman, 1969, Julien,
1973). Adequate water and temperatures above 18°C are
also necessary (Barbieri et al., 1984; Coleman, 1969).
S. officinarum is refractory to flowering, which effectively
occurs only at low latitudes. The low flowering percentage of
S. officinarum is exploited as a beneficial agronomic trait of
hybrid cultivars once flowering is undesirable since sugar
yield decreases during flower development as flowering
culms stop to grow, become diseased and, ultimately,
senesce. (Moore and Osgood, 1989).
The optimal temperatures for panicle development and
pollen fertility are 28°C during the day and 23°C at night.
Temperatures below 23°C delay panicle development and
reduce pollen fertility (Brett and Harding, 1974; Berding,
1981). Daytime temperatures above 31°C and nighttime
temperatures below 18°C are detrimental (Clements and
Awada, 1967; Moore and Nuss, 1987).
Sugarcane pollen is small, ca. 50 μM, with a honeycombed exine and is dispersed primarily by wind. Sugarcane pollen grains dry rapidly after dehiscence with an
Breeding Programs
Period
Abbreviation
Escada, PE
Campos, RJ
1913–1924
1916–1972
EB
CB
Barreiros, PE
São Bento, Tapera, PE
Curado, Recife, PE
EECAPO, Piracicaba, SP
Agronomic Institute of Campinas, Campinas, SP
COPERESTE, Sertãozinho, SP
EECA, Rio Largo, AL
COPERSUCAR, Piracicaba, SP
CTC, Piracicaba, SP (former COPERSUCAR)
PLANALSUCAR, Brazil
Barra Plant, Barra Bonita, SP
Federal Universities, Brazil
CanaVialis, Campinas, SP
1924–1933
1928-?
1933–1974
1928–1935
1935-+(*)
1963-1969
1968-1971
1968-2004
2004-+(*)
1971-1990
1975-1996
1991-+(*)
2003-+(*)
EB
SBP
(PB) – IANE
IAC
COP
SP
CTC
RB
PO
RB
CV
70
Tropical Plant Biol. (2011) 4:62–89
Fig. 5 Flowchart of a sugarcane
breeding program. T1: seedling
selection. T2: clone selection.
T3: local trial. T4: regional
trial. Source: CanaVialis
estimated half-life of approximately 12 min. After 35 min at
26.5°C and 67% relative air humidity, the pollen has lost
viability (Moore 1976; Venkatraman, 1922). Therefore,
pollen dispersed over large distances is not expected to be
viable. In addition, even under ideal conditions, some
sugarcane varieties show poor pollen fertility or even male
sterility due to the cytogenetic abnormalities that occur
during meiosis that are associated with sugarcane’s high
polyploid multispecies genetic complex (Ethirajan, 1987).
The conditions required for sugarcane flowering can be
summarized as follows:
&
&
&
Latitudes between 5° and 15°: a region with a gradual
reduction of photoperiod is an essential factor for flowering induction and panicle development (Midmore, 1980).
High temperatures, mainly during the night. Temperatures below 18.2°C are considered non-inductive.
Sugarcane requires at least 10 inductive nights for
flowering, but 15 nights are ideal. Non-inductive nights
delay panicle development and reduce pollen fertility
(Berding, 1981).
High relative humidity is critical not only for the
induction and development of the panicle and pollen
fertility, but also for anthesis and seed formation.
Flower opening and anthesis, which are both affected
by relative air humidity (RAH), only occur a few hours
before sunrise, when the plant is totally hydrated and
RAH is high (Moore and Nuss, 1987).
Because sugarcane is predominantly an outcrossing large
stature plant, the first breeding programs used seeds from
the open pollination of field grown plants, with no control
over the male parent. Currently, crosses are conducted in a
controlled manner under plastic domes or lanterns, where
parentage can be guarenteed (Fig. 6). For specific crosses,
the flowering dates of desired parents are synchronized by
manipulating the photoperiod (Moore, 1987b). Stalks with
inflorescences from selected parents are harvested and kept
in an acidic solution to keep the stalks fresh which
facilitates successful fertilization and seed maturation
(Heinz and Tew, 1987). Panicles from both parents are
placed under a dome for 12–15 days, with the male parent
positioned slightly above the female parent. Normally,
crosses are conducted in protected locations, such as in
the middle of the forest or in covered sheds containing
isolation cells, to avoid undesired cross-contamination.
After 3–4 days for hybridization, fertilized panicles are
kept in a shed for 1 week to promote seed maturation.
Panicles are then harvested and placed in a heated chamber
to dry the seeds (Ethirajan, 1987).
In a variety development program, sugarcane cultivars
are selected for low flowering. However, because flowering
is influenced by environmental conditions, flowering in
cultivation fields may still occur in a given location or in a
given year. If seeds are produced and fall onto the soil,
germination only occurs under conditions of high temperature and humidity. Therefore, sexual reproduction is
strongly compromised in locations that have a dry, cold
autumn, such as southern Brazil and the southern parts of
the southeastern and midwestern regions of Brazil. In the
northern parts of the southeastern and midwestern regions,
relative humidity but not nighttime temperature is normally
restrictive. In general, some flowering may occur in the
southern, southeastern and midwestern regions of Brazil,
and in some years, it may even be intense. Nevertheless,
because RAH is low heavy flowering does not mean that
seedings are produced; even if seeds are formed, field
conditions are very unfavorable for germination because of
low soil humidity, since it is well knwn that sugarcane
seeds lose approximately 90% of their viability after 80 days
at 28°C (Rao, 1980). However, in the northeastern region,
conditions are favorable for both flowering and seed
Fig. 6 Sugarcane crossing conducted under domes (lanterns). Source:
CanaVialis
Tropical Plant Biol. (2011) 4:62–89
formation. Seed dehiscence occurs during the wet season,
which favors seed germination under field conditions.
Potential for Lateral Gene Transfer
Commercial sugarcane production is performed exclusively
using vegetatively propagated material of comercial
hybrids. Under ideal flowering conditions, sugarcane pollen
is dispersed by wind, with no participation of animal or
insect vectors (McIntyre and Jackson, 2001). Because
sugarcane pollen has low viability, natural hybridization
can only occur close to the pollen-supplying plant (Moore,
1976; Venkatraman, 1922). Thus, little seed set is expected
since pollen rapidly loses its viability.
Although crossings between species of the genus
Saccharum with other closely related species have been
suggested to occur in the wild (Grassl 1980; Daniels and
Roach, 1987), wild hybridization has not been reported
with current sugarcane varieties.
Species belonging to the “Saccharum Complex” exhibit
different levels of sexual compatibility with S. officinarum
and S. spontaneum under artificial controlled crossings
(Bonnett et al., 2008). Hybridization among sugarcane
species and Erianthus sect. Ripidium and Miscanthus
species are more probable than with Narenga and Sclerostachya under breeder’s intervention. However, genetic
transfer among commercial hybrids and these ancestral
species, if existent, are much lower under natural conditions
(Bonnett et al., 2008). It is important to note that there are
no members of the “Saccharum Complex” species native in
Brazil. In addition, there is no data on the biology of the
wild Brazilian Saccharum species such as S. villosum, S.
asperum, S. angustifolius and S. baldwinii (Filgueiras and
Lerina, 2001; Kameyama, 2006; Carporal and Eggers,
2005) nor on the possibility of gene flow occurring between
them and commercial sugarcane hybrids.
The Saccharum species that gave rise to commercial
sugarcane varieties (S. officinarum and S. spontaneum, with
minor contributions of S. robustum, S. barberi and S. sinense)
are not native to Brazil. In Brazil, these species exist only in
germplasm collections which are used in sugarcane breeding
programs. Under breeding station conditions, they can flower
synchronously and successfully hybridize with modern
varieties. However, lateral transfer of genes among modern
sugarcane hybrids and those species is not expected to occur
under natural Brazilian environmental conditions.
Commercial Sugarcane Cultivation
The first step towards establishing a commercial sugarcane field is the production of vegetative planting
71
material from the desired commercial variety under
approved sanitary conditions at nurseries. To assure the
starting material is disease-free, it is common practice
that the stalks to be used as planting material are either
exposed to thermotherapy (a hot water treatment to
control systemic bacterial infections such as ratoon
stunting disease), or they are obtained aseptically through
meristem culture (free of bacteria and viruses), or from a
combination of methods. Essentially, there are three
types of nurseries differing primarily in size and
generations removed from initial asepsis:
&
&
&
Basic Nursery or Pre-Primary Nursery originates from
buds of aforementioned treated stalks or meristem
propagated plants.
Primary Nursery originates from the Basic Nursery but
is roughly ten times larger than that source. The first
ratoon of the Basic Nursery is also considered a Primary
Nursery.
Secondary Nursery originates from the Primary Nursery
and is 10–15 times larger than the previous nursery. The
second ratoon of the Basic Nursery and the first ratoon
of the Primary Nursery can also be considered
Secondary Nurseries (Xavier et al., 2008).
Commercial plantations are generally established
using time proven conventional methods. Plowing is
30 cm deep, and the furrows cut to a depth of 25–30 cm.
Rows are spaced at distances varying from 0.8 to 1.5 m
and are planted with 8–12 tons of planting material per
hectare. Stalks are distributed in furrows in pairs with the
base of one stalk paired against the upper part of the
other, i.e., two stalks are laid in opposite directions
because the buds from the upper part of the stalk tend to
germinate better than those of the base. After the stalks
are distributed in the furrow, they are sectioned into 2 to
3-node seed pieces to interrupt apical dominance that
exists in the intact stalk. In soils known to be infested by
insect pests or nematodes, pesticides are applied over the
cuttings in the furrows. The last step of the planting
operation is to cover the cuttings in the furrows with 10–
15 cm of soil (dos Anjos and Figueiredo, 2008).
In Brazil, the use of irrigation in commercial sugarcane
fields is generally not necessary, contributing to low
production costs. Currently, as marginal production areas,
primarily drier areas with inadequate rainfall, are added to
the sugarcane industry through crop expansion, drought
tolerance is seen as an increasingly important trait for
sugarcane varieties (Pires et al., 2008).
When the crop begins to grow, the most important
agronomic practice is weed control. Once an optimal plant
stand is established, the major concern is to employ
72
practices to insure good crop development to achieve good
maturation, i.e. accumulation of sugar, in an optimal time
span. Proper practices assure optimization of the
harvesting-milling operations and, consequently, overall
economic return. This aim is achieved by mills cultivating
a range of varieties having different soil nutrient requirements, rates of maturation and reliable disease resistance.
In Brazil, sugarcane harvesting is either semi-mechanized
or completely mechanized. In the first case, the cane is
harvested manually, but it is loaded onto trucks mechanically;
in the second case, the cane is harvested by machines that load
it directly onto trucks. Although the fully mechanized harvest
system has the advantage of not requiring a prior burning step,
it cannot be adopted everywhere because current harvesting
machines cannot operate in areas where the slope exceeds 15–
17% (Ripoli and Ripoli, 2008).
Commercial Sugarcane Crop Cycle
Sugarcane is a semiperennial crop in commercial fields. It has
to be replanted approximately every three to six harvests when
grown under the rainfed conditions of Brazil. Replanting is
required because of declining yields due to crop and soil
damage caused by the heavy traffic of machines and trucks
over the stumps during harvesting. In addition, there could be
a progressive accumulation over time of pathogens in the
sugarcane crop, some of which reduce stand population while
others impair plant growth. There may also be a genetic
component contributing to yield decline because most of the
commercial cultivars have been selected to produce well only
for the first three to four cultivation cycles. The overall result
is a decrease in year-over-year productivity, which can reach
economically unfeasible levels and the need to replant the
field (Matsuoka et al., 1999).
There are two basic sugarcane production cycles. The
plant-cane cycle starts with planting and ends after the first
harvest. The ratoon, or ratoon-cane, cycle starts after the
harvest of the plant cane and continues with successive ratoon
crops until field renewal (Fig. 7). The complete cycle of a
sugarcane field lasts either four or five seasons, after which
time the crop is renewed. Eradication of the crop after it has
become economically unfeasible is performed by ploughing
it under and harrowing the soil, which is often preceded by
the application of a systemic herbicide.
Sugarcane-Associated Insects
The most important sugarcane insect pests in Brazil are the
sugarcane borers (Diatraea saccharalis and Diatraea
flavipennella), the giant sugarcane borer (Telchin licus),
spittlebugs (Mahanarva fimbriolata and Mahanarva posticata),
Tropical Plant Biol. (2011) 4:62–89
termites (different genera), the migdolus beetle (Migdolus
fryanus), the sugarcane weevil (Sphenophorus levis) and
herbivorous ants (Atta spp. and Acromyrmex spp.).
D. saccharalis is widely distributed in Brazil, while D.
flavipenella is restricted to the northeastern region of the
country. Both species construct galleries in the stalks,
leading to less biomass and sugar production, and an
increase in fungal infections and juice contamination. These
sugarcane borers are mainly controlled by massive release
of the parasitoid Cotesia flavipes, which is normally reared
in labs at the various mills. Sugarcane borers are also
controlled, although by a lesser extent, by the release of the
parasitic wasp Trichogramma galloi. Currently there is
strong evidence that the sugarcane borer population has
been increasing due to the expansion of cane into new
areas, the cultivation of susceptible varieties, and the failure
to use biological control. The increase in borer populations
has been causing an increase in the use of pesticides to
control these insects (Dinardo-Miranda, 2008a). The giant
sugarcane borer (T. licus) is another lepidopteran pest that
attacks the crop. T. licus was considered to be restricted to
northeastern Brazil, but there have been recent reports of its
occurrence in São Paulo State, Brazil’s main sugarcane
producing state. T. licus also construct galleries in the
stalks, but they more easily outright kill the ratoons due to
the extensive damage caused by their large size (DinardoMiranda, 2008a). Biological and chemical control mechanisms against T. licus are not effective, and the economic
impact of the pest, if it spreads throughout Brazilian
sugarcane-producing regions, has yet to be assessed.
The root froghopper (M. fimbriolata) has become an
important sugarcane pest since Brazil started to abolish crop
burning. Crop damage is caused by the young insect
(nymph), which sucks water and nutrients from plant roots
and injects toxins into them, leading to a decrease in root
function and, consequently, a loss of productivity. Release
of the fungus Metarhizium anisopliae results in good
biological control, which can be complemented or replaced
by pesticide spraying (Dinardo-Miranda, 2008a). The leaf
spittlebug (M. posticata) is more predominant in northeastern Brazil; this insect sucks leaf sap, causing leaf drying; it
can be controlled in the same way as M. fimbriolata.
There are many species of termites that attack sugarcane
in the the country. Among these species, H. tenuis is the
most harmful. These underground insects attack the stalks
used for planting, leading to low bud germination and the
need for replanting (Dinardo-Miranda, 2008a). Farmers
control termites by spraying pesticides over the stalks in the
furrow during planting.
The migdolus beetle Migdolus fryanus is a native Brazilian
insect that attacks the roots of many crops, including
sugarcane, coffee, eucalyptus and beans (Bento et al. 1985).
The insect can destroy the root system, leading to an early
Tropical Plant Biol. (2011) 4:62–89
Fig. 7 Sugarcane phenological cycle. a Stalk pieces used in planting;
b Beginning of bud sprouting and rooting; c Tillering initiation; d
Intense tillering; e Beginning of maturation; f Manufacturable stalks in
73
optimal sucrose concentration; g Harvesting; h Ratoon sprouting.
Illustration: Rogério Lupo
74
Tropical Plant Biol. (2011) 4:62–89
need for field replanting. Control of M. fryanus is difficult
because the larvae live deep within the soil so that pesticide
application during planting is not very effective. Recently, the
use of pheromone traps has been shown to be very promising
for controlling this pest (Nakano et al., 2002). The sugarcane
weevil (Sphenophorus levis) is another beetle that attacks the
sugarcane root system, leading to damage similar to that
caused by the migdolus beetle. S. levis has low dissemination
ability so its spread is linked to human activities (DinardoMiranda, 2008a). Consequently, one of the more effective
control practices for this pest is to avoid planting cane that is
harvested from infected areas.
Ants that behave as pests in the sugarcane crop belong to
the genera Atta and Acromyrmex. The species Atta bisphaerica
and Atta capiguara cause most of the losses, but Atta sexdens
and Atta laevigata also cause damage. Studies have shown
that the ants of one anthill can reduce sugarcane productivity
by 3.2 ton ha−1 (Dinardo-Miranda, 2008a). Control of these
pests is accomplished using pesticides that must be applied
very carefully because the pesticides could kill predator ants
that are beneficial to the crop.
Many species of nematodes are found in association
with sugarcane, but in Brazil, most of the damage is
caused by five species: Meloidogyne incognita, Meloidogyne
javanica, Pratylenchus zeae, Pratylenchus brachyurus and
Helycotylenchus dihystera. These nematodes are mainly
controlled with chemical pesticides because nematoderesistant varieties have not been developed in the Brazilian
sugarcane breeding programs (Dinardo-Miranda, 2008b).
Nematicide application during sugarcane planting can
increase productivity up to 30% in some infested areas
(Copersucar, 1982). It is also possible to use nematocides
Table 3 Registered products to control insects and nematodes in sugarcane fields in Brazil. Source: AGROFIT (2010)
Common name
Chemical group
Commercial name
Target organism²
(Z)-11Hexadecenyl
acetate
(Z)-7-dodecenyl
acetate
(Z)-9tetradecenyl
acetate
Aldicarb
unsaturated acetate
Bio Spodoptera
Spodoptera frugiperda
unsaturated acetate
Bio Spodoptera
S. frugiperda
unsaturated acetate
Bio Spodoptera
S. frugiperda
oxime
methylcarbamate
biological
Temik 150
Mahanarva fimbriolata, Meloidogyne incognita,
Pratylenchus zeae
S. frugiperda, Mocis latipes
pyrethroid
Bistar 100 EC, Brigada EC, Brigade 100 EC,
Capture 100 EC, Capture 400 EC, Talstar 100
EC
Carboran Fersol 350 SC, Diafuran 50, Furacarb
100 GR, Furadan 100 G, Furadan 350 SC,
Furadan 50 GR, Ralzer 50 GR
Bacillus
thuringiensis
Bifentrin
Carbofuran
benzofuranyl
methylcarbamate
Endosulphan
cycledienochloride
Ethiprole
Fipronil
phenylpyrazole
pyrazole
Imidacloprid
neonicotinoid
Metarhizium
anisopliae
N-2'Smethylbutyl2-methyl
butylamide
Terbufos
Thiametoxam
Trichlorfon
biological
Triflumuron
Bac-Control WP, Dipel WP, Thuricide
Dissulfan EC, Endosulfan Nortox 350 EC,
Endosulfan 350 EC Milenia, Endozol, Thiodan
EC
Curbix 200 SC
Regent 20 GR, Regent 800 WG
Confidor 700 WG, Evidence, Nuprid 700 WG,
Warrant
Metarril Wp E9
Heterotermes tenuis, Migdolus fryanus,
Procornitermes triacifer
Meloidogyne. javanica, M. incognita,
Helicotylenchus dihystera, P. zeae, M.
fimbriolata, Diatraea saccharalis, M. fryanus,
H. tenuis
H. tenuis, M. fryanus, Cornitermes cumulans
H. tenuis, M. fimbriolata
Neocapritermes opacus, H. tenuis, P. triacifer, C.
cumulans, D. saccharalis, M. fryanus
H. tenuis, M. fryanus, N. opacus, M. fimbriolata
M. fimbriolata
amide
(pheromone)
Migdo
M. fryanus
organophosphorate
neonicotinoid
organophosphorate
Counter 150 G
Actara 10 GR, Actara 250 WG
Dipterex 500
benzoylurea
Certero
P. zeae, M. incognita, H. tenuis, M. javanica
M. fimbriolata, H. tenuis
M. latipes, S. frugiperda, Tomaspis furcata, M.
fimbriolata
D. saccharalis
Tropical Plant Biol. (2011) 4:62–89
in ratoon cycles, but the control of nematodes on ratoons
is not as effective (Dinardo-Miranda, 2008b). The
pesticides that are registered in Brazil are displayed in
Table 3.
In addition to insects that act as pests, sugarcane fields
contain other associated insects with different biological
functions, which comprise the sugarcane insect fauna (entomofauna). There have been several sugarcane entomofauna
studies aimed at identifying the impact of sugarcane field
burning on the associated insect crop population. These
studies revealed a species rich system (Macedo and
Araújo, 2000a; Macedo and Araújo, 2000b; Araújo et al.,
2004; Araújo et al., 2005). However, identifying the risks
to the insect fauna of adopting specific technologies,
such as new agrochemicals and genetically modified
cultivars for crop improvement requires detailed analyses
of the individual species that are most important and
meaningful for monitoring (Romeis et al., 2008). Species
selection must be based on the biological functions of the
insects, their abundance and their economic importance.
Other, less objective criteria such aesthetic value, cultural
value and species at risk, may also be used (Romeis et al.,
2008).
Table 4 displays a no exhaustive list of insects and
nematodes that are most relevant to the sugarcane
agroecosystem.
Sugarcane Weed Control
Although sugarcane is a vigorous plant, the crop suffers
from weed competition in its initial development stages.
The most detrimental weed species are in the Poaceae
and Cyperaceae families, but morning glory species may
also interfere by coiling around the sugarcane plants,
reducing leaf unfurling to decrease the photosynthetic
area and slowing mechanical harvesting. In most sugarcane production areas of the world, herbicide use
(chemical control) is the most common way to control
sugarcane weeds. Table 5 displays a non exhaustive list of
the most common sugarcane weeds occurring in sugarcane
fields and Table 6 displays the registered products used to
control them.
Sugarcane Diseases
Most commercial sugarcane varieties are genetically resistant to most sugarcane diseases. In addition to genetic
resistance, use of pathogen-free planting material is
commonly used to avoid spreading of diseases. The
Brazilian sugarcane industry does not usually control
sugarcane diseases in commercial fields, but recently,
75
coinciding with the first detection of the Orange rust
(Puccinia kuehnii) in the country, the Agriculture Department has registered a product (azoxystrobin and ketoconazole) to control fungal disesases at sugarcane fields (Santos,
2008; Agrofit, 2010). Additionally, the Triazole fungicides
triadimefon and triadimenol are registered to treat sugarcane stalks before planting to prevent smut contamination
caused by Ustilago scitaminea (AGROFIT 2010). Sugarcane smut disease is also contolled by destroying contaminated plants in the field (roguing) when varieties having
intermediate resistance to the pathogen are planted.
Table 7 displays a list of the most common sugarcane
diseases in Brazil.
Environmental Impacts
Sugarcane’s high efficiency in fixing CO2 into carbohydrates
for conversion into fuel has awakened the world’s interest in
the crop. Emerging data indicates that sugarcane could be the
best crop for the production of renewable energy, which could
reduce some effects of global warming caused by the use of
fossil fuels (Buckeridge, 2007). The impact of sugarcane on
the environment might be reduced by adopting environmentally friendly agricultural practices such as the elimination of
burning before harvest, modifying other practices for a
reduction in diesel-driven transportation and a reduction in
the use of oil-based fertilizers (Ometto et al., 2005).
Brazil’s land area currently occupied by sugarcane is mainly
the result of the large expansion of the sugarcane industry that
occurred in the 1970s, when the Pró-Álcool (pro-ethanol)
program was created. During this period, sugarcane expansion
occurred in areas that were originally covered with Atlantic
Rain Forest, but which were already being used for pastures
and annual crops. The current rapid expansion of sugarcane
into new areas that have never been used to grow the crop raises
questions about sustainability, particularly in those areas of the
cerrado biome that are already threatened by the expansion of
other crops (Rodrigues and Ortiz, 2006). This recent concern
about the potential environmental impacts of crops has
encouraged government agencies to promote studies to
establish zones for planting sugarcane and to regulate how
expansion will take place to avoid expansion into protected
areas. This zoning proposal was recently approved and will
allow the Brazilian Government to use Climatic Risk Zoning
as a tool for the establishment of a sustainable sugarcane
agribusiness in the country (Embrapa, 2009).
Hitorically, the sugarcane has been burned prior to
harvest as a means to facilitate and thus reduce the costs
for harvest and hauling of cane, whether harvested by
hand or by machine. In addition, burning of the crop
generally increased recovery of the sucrose contained in
the plant. However, in the process of burning, carbon
76
Tropical Plant Biol. (2011) 4:62–89
Table 4 Insects and nematodes there are most relevant to sugarcane agrosystem in Brazil
Specie
Order
Common name english (Portuguese)
Leaf/shoot apex/stalk pests
Diatraea saccharalis (Fabricius, 1794)
Diatraea flavipennella (Box, 1931)
Mocis latipes (Guenée, 1852)
Spodoptera frugiperda (J. E. Smith, 1797)
Mahanarva posticata (Stal, 1855)
Elasmopalpus lignosellus (Zeller, 1848)
Aclerda campinensis (Hempel, 1934)
Lepidoptera
Lepidoptera
Lepidoptera
Lepidoptera
Hemiptera
Hemiptera
Hemiptera
Sugarcane borer (Broca da cana-de-açúcar)
Sugarcane borer (Broca da cana-de-açúcar)
Striped grassworm (Curuquerê-dos-capinzais)
Fall armyworm (Lagarta-do-cartucho)
Spittlebug (Cigarrinha da folha)
Lesser cornstalk borer (Lagarta elasmo)
Sugarcane mealybug (Cochonilha parda)
Hemiptera
Hemiptera
Hemiptera
Hymenoptera
Hymenoptera
Coleoptera
Hemiptera
Pink sugarcane mealybug (Cochonilha rosada)
Sugarcane aphid (Pulgão)
Corn leaf aphid (Pulgão)
Giant leaf-cutting ant (Saúva mata-pasto)
Grass cutting ant (Saúva parda)
West Indian sugarcane weevil (Besouro-rajado-da-cana)
Spittlebug (Cigarrinha-das-pastagens)
Hemiptera
Lepidoptera
Lepidoptera
Root froghopper (Cigarrinha da raíz)
Giant sugarcane borer (Broca gigante)
Borer (Broca peluda)
Isoptera
Isoptera
Isoptera
Isoptera
Isoptera
Isoptera
Isoptera
Termite
Termite
Termite
Termite
Termite
Termite
Termite
Isoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Termite (Cupim)
Migdolus beetle (migdolus)
Sugarcane weevil (Bicudo-da-cana)
Sugarcane beetle (Pão-de-galinha)
Banana beetle (Pão-de-galinha)
Scarab beetle (Pão-de-galinha)
Masked chafers (Besouro)
Hymenoptera
Hymenoptera
Diptera
Diptera
Larval parasitoid (Cotesia)
Egg parasitoid (Tricograma)
Larval parasitoid (Mosca parasitóide)
Larval parasitoid (Mosca parasitóide)
Hymenoptera
Hymenoptera
Hymenoptera
Hymenoptera
Dermaptera
Fire ant (Formiga lava-pé)
Ant (Formiga)
Ant (Formiga)
Ant (Formiga)
Linear earwig (Tesourinha)
Coleoptera
Ladybird (Joaninha)
Hymenoptera
Honeybee (Abelha)
Saccharicoccus sacchari (Cockerell, 1895)
Melanaphis sacchari (Zehnter, 1897)
Rhopalosiphum maidis (Fitch, 1856)
Atta bisphaerica (Forel, 1908)
Atta capiguara (Gonçalves, 1944)
Metamasius hemipterus (Linnaeus, 1765)
Tomaspis furcata (Germar, 1821)
Stool pests
Mahanarva fimbriolata (Stal, 1854)
Telchin licus (Drury, 1773)
Hyponeuma taltula (Schaus, 1904)
Underground pests
Heterotermes tenuis (Hagen, 1858)
Heterotermes longipes (Snyder, 1924)
Procornitermes triacifer (Silvestri, 1901)
Neocapritermes opacus (Hagen, 1858)
Neocapritermes parvus (Silvestri, 1901)
Syntermes molestus (Burmeister, 1839)
Cornitermes spp.
Rhynchotermes spp.
Migdolus fryanus (Westwood, 1863)
Sphenophorus levis (Vaurie, 1978)
Euetheola humilis (Burmeister, 1847)
Ligyrus bituberculatus (Beauvois, 1805)
Stenocrates laborator (Fabricius., 1775)
Cyclocephala spp.
Parasitoids
Cotesia flavipes (Cameron, 1891)
Trichogramma galloi (Zucchi, 1988)
Lydella minense (Townsend, 1927)
Billaea claripalpis (Wulp, 1896)
Predators
Solenopsis saevissima (F. Smith, 1855)
Crematogaster sp.
Dorymyrmex sp.
Pheidole sp.
Doru lineare (Eschscholtz, 1822)
Cycloneda sanguinea (Linnaeus, 1763)
Other associated insects
Apis mellifera (Linnaeus, 1758)
(Cupim)
(Cupim)
(Cupim)
(Cupim)
(Cupim)
(Cupim)
(Cupim)
Tropical Plant Biol. (2011) 4:62–89
77
Table 4 (continued)
Specie
Order
Common name english (Portuguese)
Nematodes
Meloidogyne incognita (Kofoid and White, 1919)
Meloidogyne javanica (Treub, 1885)
Pratylenchus zeae (Graham, 1951)
Pratylenchus brachyurus (Godfrey 1929)
Helicotylenchus dihystera (Cobb, 1893)
Tylenchida
Tylenchida
Pratylenchidae
Pratylenchidae
Tylenchida
Root knot nematode (Nematóide das galhas)
Root knot nematode (Nematóide das galhas)
Dagger nematode (Nematóide-das-lesões)
Dagger nematode (Nematóide-das-lesões)
Spiral nematode (Nematóide-espiralado)
dioxide and other greenhouse gases, and soot or ash are
released (Cançado et al., 2006). Brazilian law No. 11.241
(Sept. 19, 2002) in São Paulo State established a goal for
the gradual elimination of sugarcane burning and associated measures by 2021, with all harvesting being entirely
mechanized on non-burned cane by 2031. On June 4, 2007
in anticipation of this law, UNICA (União da Indústria de
Cana-de-açúcar) which is the Brazilian Sugarcane Industry Association representing the sugar, ethanol and
bioelectricity producing industry of the State of São
Paulo, and the state government signed the Protocolo
Agroambiental do Setor Sucroalcooleiro [Sugar and
Alcohol Industry Agro-Environmental Protocol]. This
protocol established several environmental principles and
technical guidelines to be observed by the sugarcane
industry. Perhaps the most important guidelines are those
addressing the anticipation of the legal deadline for the
end of sugarcane harvesting within 14 years.
Table 5 List of common weeds occurring in Brazilian sugarcane fields. Source: Azania et al. (2008)
Species
Family
Common name english (Portuguese)
Annual
Acanthospermum australe (Loefl.) Kuntze
Acanthospermum hispidum DC.
Ageratum conyzoides (L.) L.
Alternanthera ficoidea (L.) Sm.
Amaranthus spp.
Bidens pilosa L.
Brachiaria plantaginea (Link) Hitchc.
Cenchrus echinatus L.
Commelina spp.
Croton lobatus L.
Digitaria horizontalis Willd.
Digitaria insularis (L.) Mez ex Ekman
Eleusine indica (L.) Gaertn.
Emilia sonchifolia (L.) DC.
Euphorbia heterophylla L.
Asteraceae
Asteraceae
Asteraceae
Amaranthaceae
Amaranthaceae
Asteraceae
Poaceae
Poaceae
Commelinaceae
Euphorbiaceae
Poaceae
Poaceae
Poaceae
Asteraceae
Euphorbiaceae
Paraguayan burr (Carrapichinho)
Bristly starburr (Carrapicho de carneiro)
Billygoat-weed (Mentrasto/Ageratum)
Sanguinaria (Apaga-fogo)
Pigweed (Caruru)
Hairy beggarticks (Picão-preto)
Signalgrass (Capim marmelada)
Southern sandbur (Capim carrapicho)
Dayflower (Trapoeraba)
Lobed croton (Cróton)
Jamaican Crabgrass (Capim colchão)
Sourgrass (Capim-amargoso)
Indian goosegrass (Capim pé-de-galinha)
Lilac tasselflower (Falsa-serralha)
Mexican fireplant (Leiteiro)
Convolvulaceae
Portulacaceae
Rubiaceae
Poaceae
Asteraceae
Morning glories (Cordas-de-viola)
Little hogweed (Beldroega)
Tropical Mexican clover (Poaia branca)
Ichtgrass (Capim-camalote)
Common sowthistle (Serralha)
Poaceae
Poaceae
Poaceae
Cyperaceae
Poaceae
Malvaceae
Poaceae
Spreading liverssed grass (Capim-braquiária)
Para grass (Capim-braquiária)
Bermuda grass (Grama-seda)
Nut Grass (Tiririca)
Guinegrass (Capim-colonião)
Fanpetals (Guanxuma)
Johnson grass (Capim-massambará)
Ipomea spp.
Portulaca oleraceaL.
Richardia brasiliensisGomes
Rottboellia exaltata (L.) L.f.
Sonchus oleraceus L.
Perennial
Brachiaria decumbensStapf
Brachiaria mutica (Forssk.) Stapf
Cynodon dactylon (L.) Pers.
Cyperus rotundus (L.)
Panicum maximum Jacq.
Sida spp.
Sorghum halepense (L.) Pers.
78
Tropical Plant Biol. (2011) 4:62–89
Table 6 Registered products to control weeds in sugarcane fields in Brazil. Source: AGROFIT (2010)
Common name¹
Chemical group
Commercial name
Acetochlor
Alaclhor
Ametryn
chloroacetanilide
chloroacetanilide
triazine
Amicarbazone
Asulam
Atrazine
triazolinone
Sulphanilyl
carbamate
triazine
Fist EC, Surpass
Alaclor + Atrazina SC Nortox, Alaclor Nortox, Boxer, Laço EC
Agritin SC, Ametrex 500 SC, Ametrina Agripec, Ametrina Anator 50 SC, Ametron, Ametron
SC, Bimetron, Gesapax 500 Ciba-Geisy, Herbipak WG, Herbipak 500 BR, Krismat WG,
Metrimex, Metrimex 500 SC, Simetrex SC, Sinerge EC, Stopper 500 SC, Topeze SC
Dinamic
Asulox 400
Carfentrazone-ethyl
Clomazone
triazolone
isoxazolidinone
Paraquat dichloride
Diclosulam
Diuron
bipyridilium
triazolopyrimidine
sulfonanilide
urea
Ethoxysulfuron
Flazasulfuron
Glyphosate
sulfonylurea
sulfonylurea
substituted glycine
Glyphosate
isopropylamine
salt
Halosulfuron methyl
Hexazinone
substituted glycine
Imazapic
Imazapyr
Iodosulfuron-methyl
Imidazolinone
imidazolinone
sulfonylurea
Isoxaflutol
MCPA
Metribuzin
Metsulfuron-methyl
MSMA
isoxazol
aryloxyalkanoyl
triazinone
sulfonylurea
organoarsenic
Provence 750 WG
Agritin SC
Lexone SC, Sencor BR, Sencor WG, Sencor 480, Sencor 70 WG, Soccer SC
Ally, Wolf
Ancosar 720, Ansar 720, Daconate 480, Dessecan, Fortex SC, MSMA Sanachem 720 SL,
MSMA 720, MSMA 720 Volagro, Volcane
Oxadiazon
Oxifluorfen
Pendimethalin
Picloram
oxadiazolone
diphenyl ether
dinitroaniline
pyridinecarboxylic
acid
triazine
chloroacetanilide
Ronstar 250 BR
Galigan 240 EC, Goal BR
Herbadox, Herbadox 400 EC
Dontor
Simazine
S-metolachlor
sulfonylurea
triazinone
Alaclor + Atrazina SC Nortox, Atrazina Nortox 500 SC, Atraxinax 500, Boxer, Genius WG,
Gesaprim GrDa, Gesaprim 500 Ciba-Geisy, Herbitrin 500 BR, Proof, Siptran 500 SC, Siptram
800 WP, Sprint
Aurora, Aurora 400 EC, Quicksilver 400 EC
Clomanex 500 EC, Clomazone 500 EC FMC, Discover 500 WP, Escudo, Gamit, Gamit Star,
Gamit 360 CS, Magister, Ranger, Reator 360 CS, Sinerge SC
Gramocil, Gramoxone 200, Helmoxone, Paradox
Coact
Advance, Agritin SC, Ametron, Ametron SC, Bimate SA, Bimetron, Cention SC, Confidence,
Dihex, Direx 500 SC, Diurex Agricur 500 SC, Diurex Agricur 800 SC, Diurex WG,
Diuromex, Diuron Fersol 500 SC, Diuron Milenia WG, Diuron Nortox, Diuron Nortox 500
SC, Diuron 500 Agritec, Diuron 500 SC, Diuron 500 SC Milenia, Diuron 80 Volagro, Diuron
80 Volcano, Dizone, Fortex SC, Gramocil, Herburon WG, Herburon 500 BR, Hexaron,
Hexaron WG, Jump, Karmex, Karmex 800, Netun 500 SC, Netum 800 SC, Rancho, Scopus,
Soldier, Soligard, Velpar Max, Velpar-K, Velpar-K WG
Gladium
Katana
Direct, Fera, Gliato, Glifos, Glifos Concept, Glifos N, Glifos Plus, Glifosato Atanor, Glifosato
Atar 48, Glifosato Nortox, Glifosato Nortox WG, Glifosato Nufarm, Glifosato 480 Agripec,
Gliphogan 480, Glister, Gliz 480 SL, Glyox, Glyphotal, Icaro, Pilarsato, Polaris, Pretorian,
Radar, Rodeo, Ronat-A, Roundup Original, Roundup Transorb, Roundup WG, Rustler,
Samurai, Scuder, Stinger, Sumô, Trop
Glifosato Atanor 40, Gli-Up 480 SL, Gliz Plus, Glizmax, Sumô, Tupan
Sempra
Advance, Broker 750 WG, Confidence, Destaque, Dihex, Discover 500 WP, Dizone, Hexaron,
Hexaron WG, Hexazinona Nortox, Hexazinona Nortox 250 SL, Jump, Perform 240 SL,
Rancho, Ranger, Scopus, Soldier, Soligard, Style, Velpar Max, Velpar-K, Velpar-K WG
Plateau
Contain
Hussar
Simetrex SC, Topeze SC
Dual Gold
Tropical Plant Biol. (2011) 4:62–89
79
Table 6 (continued)
Common name¹
Chemical group
Commercial name
Sulfentrazone
Sulfometuronmethyl ²
Tebuthiuron
Thiazopyr
triazolone
sulfonylurea
Boral 500 SC, Explorer 500 SC
Curavial
urea
pyridinecarboxylic
acid
sulfonylurea
Aval, Aval 800, Bimate SA, Butiron, Combine 500 SC, Lava, Lava 800
Visor 240 EC
dinitroaniline
aryloxyalkanoyl
Novolate, Premerlin 600 EC, Trifuralina Nortox Gold
Aminamar, Aminol 806, Bratt, Brion, Capri, Dez, DMA 806 BR, Dontor, Grant, Herbi D-480,
Navajo, Tento 867 SL, U 46 BR, U 46 D-Fluid 2,4-D, Weedar 806, 2,4D Agritec, 2,4-D
Amina 72, 2,4-D Fersol
Trifloxysulfuron
sodium
Trifluralin
2,4-D
Envoke, Krismat WG
Potential Invasiveness
Weediness of Commercial Hybrids and Related Species
As a result of continuous breeding and selection for
agronomic traits of value, sugarcane has lost the competitiveness or invasiveness of the original species; modern cultivars
have largely lost the ability to persist in non-agricultural
habitats and only poorly perpetuate without human assistance
(Holm et al., 1997; OGTR, 2008). Sugarcane hybrid cultivars
do not possess true rhizomes or produce vigorous seedlings.
It is possible to find leftover stools in cultivated areas, but
there is no indication that these stools will perpetuate, and
there is even less evidence that they have any invasive
capacity. It is possible that if ratoons are not properly
eradicated at the end of a cultivation cycle, they can regrow
Table 7 List of common diseases occurring at Brazilian sugarcane fields. Source: Dinardo-Miranda (2008a); Dinardo-Miranda (2008b); Almeida
(2008)
Species
Type
Disease
English (Portuguese) name
Puccinia melanocephala Syd. & P. Syd.
Puccinia kuehnii (W. Krüger) E.J. Butler
Ustilago scitaminea Syd.
Fungal
Fungal
Fungal
Rust (Ferrugem)
Rust (Ferrugem)
Smut (Carvão)
Mycovellosiella koepkei (W. Kruger) Deighton
Bipolaris sacchari (E.J. Butler) Shoemaker
Cercospora longipes E. J. Butler
Glomerella tucumanensis (Speg.) Arx & E. Müll.
Anamorph: Colletotrichum falcatum Went
Gibberella fujikuroi (Sawada) Wollenw.
Anamorph: Fusarium moniliforme J. Sheld.
Gibberella subglutinans (E.T. Edwards) P.E. Nelson,
Toussoun & Marasas
Anamorph: Fusarium subglutinans (Wollenw. & Reinking)
P.E. Nelson, Toussoun & Marasas
Thielaviopsis paradoxa (De Seynes) V. Hohny
Anamorph: Ceratocystis paradoxa (Dade) C. Moreau
Leifsonia xyli supsp. xyli (Davis et al.) Evtushenko
Xanthomonas albilineans (Ashby) Dowson
Xanthomonas axonopodis pv. vasculorum (Cobb) Vauterin,
Hoste, Kersters & Swings
Acidovorax avenae subsp. avenae (Manns) Willens, Goor,
Thielemans, Gillis, Kersters e De Ley
Sugarcane mosaic virus (SCMV)
Sugarcane yellow leaf virus (SCYLV)
Sugarcane bacilliform virus (SCBV)
Fungal
Fungal
Fungal
Fungal
Yellow spot (Mancha amarela)
Eye spot (Mancha ocular)
Brown spot (Mancha parda)
Red rot (Podridão vermelha)
Fungal
Stem Rot (Podridão de Fusarium)
Fungal
Pokkah-boeng (Pokkah-boeng)
Fungal
Pineapple disease (Podridão abacaxi)
Bacterial
Bacterial
Bacterial
Ratoon stunting disease (Raquitismo da soqueira)
Leaf scald (Escaldadura das folhas)
Gumming disease (Gomose)
Bacterial
Red stripe (Estria vermelha)
Virus
Virus
Virus
Mosaic (Mosaico)
Yellow leaf (Amarelinho)
Sugarcane bacilliform virus
80
and become volunteer plants in the next crop. However, since
the ratoons do not possess the ability to spread, they remain as
isolated stools in the new crop.
Some species from which modern sugarcane hybrids are
derived are classified as weeds. S. spontaneum has an invasive
potential because it produces rhizomes that contribute to
natural vegetative propagation and enable the species to adapt
to a great range of environmental conditions. S. spontaneum
is classified as a noxious weed in the United States (USDA,
2008) and is considered a weed in many other countries
(GCW, 2009). It is important to emphasize that the
rhizomatous nature of S. spontaneum was not incorporated
into modern sugarcane hybrids. S. arundinaceum (Syn:
Erianthus arundinaceum) is also recorded as a weed in the
United States and in some Asian countries (GCW, 2009). The
other species of the Saccharum complex (S. robustum, S.
sinensis and S. edule) that may have been involved in the
evolution of sugarcane varieties have not been recorded
as exhibiting weed potential (GCW, 2009). Among other
genera of the Saccharum complex (Miscanthus, Narenga
and Sclerostachia), only Miscanthus and Narenga represent species classified as having weed potential (GCW,
2009). Narenga porphyrochoma (Syn=Saccharum narenga)
is cited as a weed in Vietnam (Koo et al., 2000 in GCW,
2009). The genus Miscanthus contains five species that
have been reported as having weed potential in some parts
of the world: M. floridulus (Syn =M. japonicus), M.
nepalensis, M. purpurascens (Syn=Miscanthus sinensis
Subsp. purpurascens), M. sacchariflorus and M. sinensis
(GCW, 2009).
There are other Saccharum species, which are not
involved in the origin of sugarcane hybrids, that have also
been recorded as having weed potential in some parts of the
world, including: S. angustifoulius, S. bengalense, S.
florindum, S. procerum, S. ravenae and S. villosum (Syn=
S. trinii) (GCW, 2009).
Weediness of Commercial Sugarcane Varieties
and Their Related Species in Brazil
After more than five centuries of sugarcane cultivation in
Brazil, there is no evidence that this crop presents any traits
that favor persistence and invasibility and there is no
evidence of dispersion outside of agricultural environments.
Commercial propagation is vegetative, and seedlings from
breeding programs lack vigor.
As stated previously, no species that were involved in
the origin of sugarcane hybrids are native to Brazil, but
some species of Saccharum did originate in this country.
Among those species, only S. angustifolium and S
villosum are recorded as having some weed potential. S.
angustifolium is listed as a weed in pastures in southern
Brazil, where it originated. The plant is not consumed by
Tropical Plant Biol. (2011) 4:62–89
cattle, thus allowing it to occupy areas that might
otherwise be occupied by more desirable species. In
addition, S. angustifolium is able to spread into abandoned land and roadsides (Kissmann 1997; Lorenzi,
2000). Unlike S. angustifolium, which seems to be
restricted to southern Brazil, Saccharum villosum (Syn=
Saccharum trinii) occurs throughout the country, but it is
not recorded as having weed potential. However, this
latter species has been recorded as an alien naturalized
species in Mexico (Villaseñor and Espinosa-Garcia, 2004
in GCW, 2009).
There are no reports of the presence of Narenga species
in Brazil, but there have been reports of the introduction of
Miscanthus species into the country for their ornamental
potential (Lorenzi and Sousa 2001; Bastos, 2008). Nothing
is known about how these introduced species will behave in
the Brazilian environment. Although some of them are
already recorded as weeds in other countries, there have
been no reports of their occurrence as weeds in Brazil
(GCW, 2009; Lorenzi, 2000).
Persistence of Commercial Varieties in Agricultural
Systems
Unlike most crops, modern sugarcane varieties are propagated vegetatively, and seeds are not disseminated in
agricultural areas. As a consequence, the sexual reproduction of sugarcane has been inadequately studied; most
information related to this field has come from breeders.
There have been virtually no studies on the fertility and
longevity of seeds produced in commercial fields or on
their germination responses to environmental variables.
However, sugarcane breeders invest great effort to obtain
and preserve germinability of seed produced in their
breeding programs. Preliminary studies have shown that
the optimal temperature for seed germination is 36°C. Seed
could also germinate well after 8 weeks of storage at 24°C,
indicating a possible dormancy characteristic (OlivaresVillegas et al., 2008).
In Brazil, sugarcane occasionally blooms in commercial fields and it produces seeds in the northern/
northeastern regions much more often than in midwestern/
southern regions. If these seeds fall onto the ground and
encounter high humidity conditions, they may germinate
to produce new plants. However, due to heavy competition with the existing sugarcane, weeds, the actions of
pathogenic agents and predators in the non- cultivated
areas, or herbicides and weeding in the cultivated areas,
these volunteer plants do not survive for long periods of
time. If ratoons are not properly eradicated, they can
regrow in the next crop, behaving as a source of
volunteer plants, but there has been no reported case
of volunteers spreading throughout the field. However,
Tropical Plant Biol. (2011) 4:62–89
to avoid the presence of volunteer plants, the application
of herbicides during the eradication phase is highly
recommended.
Impacts on Human and Animal Health
Sugarcane has a long history of safe use as a food for
humans and animal feed. It is commercially cultivated for
use as a source of sucrose. Its byproducts are commonly
used as components of ruminant feeds: bagasse as a fiber
source and molasses as an energy source. The syrup is also
used as a sugar substitute in food.
The large-scale use of sugarcane as a source of fuel
ethanol began with the implementation of the Pró-Álcool
program in the 1970s. Industrial processing of sugarcane to
obtain sugar and ethanol involve several phases (heating,
flocculation, filtration, fermentation, distillation), which
produces crystallized sugar and/or ethanol. These products
are practically free of any contamination by other organic
molecules (Leme Junior and Borges, 1965).
Sucrose is a molecule with an extensive history of
human consumption. It is consumed as a sweetener and
energy source and is classified as non-toxic to humans, with
an LD50 in rats of 29.7 g/kg of body weight (SigmaAldrich, 2007a). Although consumption of standard doses
of sucrose has always been considered safe, excessive oral
consumption of sucrose may cause gastrointestinal problems. While there is no evidence of a direct correlation
between sucrose consumption and toxicity, many studies
suggest that average consumption should be reduced due to
a possible association with health problems such as
cardiovascular diseases, type II diabetes, obesity and
hypertension (Howard and Wylie-Rosett, 2002). In addition, the relationship between sucrose consumption and an
increased risk of developing dental cavities has been
established (Rugg-Gunn and Murray, 1983; Sreebny, 1982).
The consumption of ethanol in alcoholic beverages may
also be harmful to human health. The LD50 in rats is 7 g/kg
of body weight (Sigma-Aldrich, 2007b). Ethanol is considered toxic to humans if it is consumed in high doses and
inhalation for a long period of time may provoke coughing,
respiratory insufficiency, dizziness and intoxication. Eye
contact may cause severe irritation. The excessive consumption of alcoholic beverages causes damage to practically all organs, particularly the liver, kidneys and central
nervous system. The acute effects of ethanol ingestion
range from dizziness and intoxication to alcoholic coma
and death. Excessive consumption of alcoholic beverages
during pregnancy is associated with the induction of fetal
alcohol syndrome in the offspring and the occurrence of
low weight and asphyxia at birth, among other problems
(Sigma-Aldrich, 2007b).
81
Sugarcane pollen, like that of many other plants, has
allergenic potential and may cause immunological hypersensitivity in humans who come into contact with it through
the respiratory tract. In an allergy skin test conducted in
India by Chakraborty et al. (2001), land workers having
respiratory disorders showed enhanced reactivity to the
pollen of plants of different botanical families, including
sugarcane and rice.
Industrial Processing (Sugar, Ethanol, Vinasse,
Filtercake and Biomass)
The objective of industrial sugarcane processing is to
obtain highly purified sugar and ethanol. The process
involves pressing of the sugarcane to obtain juice, which
goes through several phases of purification and concentration, followed by crystallization (in the case of sugar
production) or fermentation and distillation (in the case
of ethanol production). Sucrose and ethanol, which are
pure and chemically defined substances, are obtained at
the conclusion of both processes. The byproducts are
vinasse (also called vinhoto) and bagasse (biomass).
Figure 8 illustrates the phases involved in the industrial
production of sugarcane products and byproducts.
Ethanol Ethyl alcohol, or ethanol, is a flammable liquid
substance that is obtained through the distillation of
fermented sugars. The main substrate for ethanol production from sugarcane is the sucrose contained in the
juice. Hydrated ethanol, the final product of the process,
is a binary mixture of ethanol and water, with an ethanol
content of approximately 96° GL (96° Gay-Lussac, 96%
ethanol + 4% water). This product may be used directly
as transportation fuel or may be dehydrated, generating
anhydrous ethanol. Anhydrous ethanol (99.5° GL) is
used in Brazil as a gasoline additive. According to
Brazilian Law No. 10696/2003, the volume of anhydrous
ethanol added to gasoline may vary from 20 to 25%
(Copersucar, 2008).
Sugar The raw sugar obtained directly from sugarcane
processing consists of 99.8% sucrose and 0.2% impurities
(0.04% humidity; 0.07% minerals; 0.07% inverted sugar;
0.02% insoluble material). Refined white sugar is obtained
by dissolving raw sugar and removing the insoluble
material and natural colorants through physical processes
(Quast, 1986). After this additional purification step, the
sucrose content of refined white sugar reaches 99.93%
(Clarke, 1988).
In some countries of the European Union, Australia,
Mexico, Canada, the United States and Japan, sugar
produced from glyphosate and gluphosinate resistant,
82
Tropical Plant Biol. (2011) 4:62–89
Fig. 8 Sugarcane industrial processing
genetically modified sugarbeets has already been approved
for human consumption. In those cases, the composition
analysis of the sugarbeet roots detected negligible amount
of total protein and the analysis of refined sugar were not
able to detect heterologous protein at the final product
(CERA 2008).
Vinasse Vinasse is a residue of industrial sugarcane
processing that consists of suspended solids and organic
and mineral substances, mainly potassium (Almeida, 1952).
Orlando Filho (1983) presented options for the use of
vinasse that include: protein production through anaerobic
fermentation, methane gas production, use in the formulation of animal feed (following treatment to bring the
concentration to 60° Brix), and as fertilizer on fields.
Despite the potential diversity of uses, vinasse is almost
solely used in Brazil as a fertilizer in fields surrounding
ethanol-producing mills.
Filter cake Filter cake is a byproduct of industrial sugarcane processing that is obtained from the rotation filters
after residual sucrose is extracted from the sugar production
leftover (sludge). Filter cake composition is variable, but in
general, the residue is rich in minerals (nitrogen, phosphorus, potassium, calcium, magnesium and sulfur) and
organic matter, mainly proteins and lipids. This residue is
commonly used as a fertilizer or in animal feed (Nardin,
2007; Diaz et al., 1998).
Biomass The bagasse obtained after sugarcane pressing
consists of lignocellulosic biomass. The volume of bagasse
obtained ranges between 240 kg and 280 kg per ton of
sugarcane. In the mills, this byproduct of sugarcane
processing is burned to generate energy (Copersucar,
2008). Currently, this process is so efficient that mills
generate excess electric energy that is added to the grid,
providing electrical energy to nearby cities, especially
during the dry season, when hydroelectric plants have
difficulty in operating at full capacity due to the low water
levels in rivers. Bagasse may also be used as a raw material
for ethanol production through acid or enzymatic hydrolysis, where cellulose and hemicellulose fractions can be
converted into hexoses and pentoses. After a purification
process, the mixture can be fermented to produce ethanol.
However, this technology is still under development, and its
economic feasibility has yet to be proven.
Tropical Plant Biol. (2011) 4:62–89
Cachaça (Sugarcane Spirit)
In Brazil, cachaça (or aguardente) production started
during the colonial period, shortly after sugarcane was
introduced into the Capitania de São Vicente in the 16th
century and the first sugar mill was installed in this region
(Lima, 1992). Cachaça production units have different
names depending on the scale of production and the
region in which they are produced; industrial cachaças
are produced in distilleries; artisanal (or boutique)
cachaças that are created in northeastern Brazil are
produced in engenhos, a holdover from the colonial
period; and artisanal cachaças that are created in southern
and southeastern Brazil are produced in alambiques,
which is the name of the equipment where distillation is
conducted (SEBRAE, 2005).
Cachaça results from the distillation of fermented
sugarcane juice; it has ethanol content between 38% and
54% by volume at 20°C. According to a survey
conducted by Martinelli et al. (2000), cachaça, with an
annual consumption of seven liters per capta, is the most
popular alcoholic beverage in the country after beer
(ABRABE, 2008). Cachaça production is estimated at
1.3 billion liters, which essentially represents the internal
Fig. 9 Flow chart of artisanal
sugarcane processing
83
market because exports represent less than 1% of total
production. It is estimated that there are 30,000 cachaça
production units throughout the country. The activity
generates annual revenues of US$ 500 million and
approximately 400,000 direct and indirect jobs (SEBRAE,
2005).
Artisanal Processing (Rapadura, Muscovado Sugar
and Sugarcane Syrup)
Rapadura, muscovado sugar, and sugarcane syrup are the
main products of the artisanal sugarcane production
system. These speciality products are produced on small
farms that are characterized by their low technology
levels and intensive use of labor. Since there is little
boutique market integration, these products are sold in
local markets, and their processing is simplified, as
shown in Fig. 9.
Rapadura (Jaggery) Rapadura is the Portuguese word for
jaggery, a concentrated product of sugarcane juice
without the separation of molasses from the crystals
whose color can vary from golden to dark brown. It is a
84
whole, unrefined sweetener that can be used in the same
way as sugar with the additional flavor of molasses. The
consumption of this product is concentrated in the rural
areas of Brazil, mostly in the northeastern region where
it is considered part of the cultural identity of the
northeastern population (Coutinho, 2003). Data collected
by FAO (Borray, 1997) showed that jaggery is produced in
approximately 30 countries. India is the largest producer,
as it is responsible for 67% of the world production;
Colombia is the second largest producer, with the highest
consumption per capita (32 kg/per capita/year). Brazil
ranks seventh among the world’s major jaggery producers
and has a per capita consumption of 1.4 kg/year
(SEBRAE, 2005).
Muscovado Sugar Muscovado sugar production is similar to
that of jaggery but with a process to achieve higher
concentrations of soluble solids (Fig. 9) (Cesar et al., 2003).
Industrial production of white sugar in combination with
consumers’ rejection of its dark color has caused muscovado
sugar to nearly disappear. Thus, the muscovado sugar market
has shrunk with a threat to the continuity of its production.
Recently, however, muscovado sugar has been rediscovered
by consumers seeking more “natural” products.
In poorer Brazilian regions, muscovado sugar and jaggery
play important roles in children’s diets because they provide
excellent sources of low-cost energy in addition they contain
an impressive level of minerals and proteins (calculated in mg
per 100 g of product): potassium (60–400), calcium (50–350),
magnesium (30–80), phosphorus (30–100), sodium (30–80),
iron (2–10), manganese (1–5), zinc (1–4), and proteins (280).
The vitamin content is not significant, snce vitamins are
destroyed by heat (Chaves et al., 2003).
Sugarcane Syrup Sugarcane syrup, known in the northeast
as “mel de engenho,” is syrup produced through the
concentration of sugarcane juice. It is also called “liquid
rapadura” due to the similarity of these substances (Fig. 9).
In general, rapadura and sugarcane syrup are processed in
the same production unit (SEBRAE, 2005). It is an
excellent source of energy and minerals, and because of
its high iron level, it is considered an anti-anemic product.
It has many uses in human diets depending on the region of
Brazil in which it is being used (Chaves et al., 2003).
Tropical Plant Biol. (2011) 4:62–89
very popular in Brazil where it is consumed by people of all
ages and social strata, especially during the summer. The
juice is extracted in electric or manual presses, sieved and
served with ice, either pure or mixed with fruit juice (Lubatti,
1999; Soccol et al., 1990; Oliveira, 2007). Because this is an
almost entirely informal activity, there is virtually no
literature or reliable statistics on the sugarcane juice market
(Oliveira, 2007). Studies conducted on samples collected
from the product sold in cities of São Paulo State have
shown poor hygienic conditions (Oliveira, 2007).
Sugarcane bagasse is the fibrous residue of juice
extraction, consisting of water (49%), fibers (48.7%) and
a small quantity of soluble solids (2.3%) (Paturau, 1969).
Bagasse fiber is a water-insoluble mixture that is mainly
composed of cellulose, hemicellulose, pentosanes and
lignin (Valsechi and Oliveira, 1964; Paturau, 1969; Delgado
and Delgado, 1999; ICIDCA-GEPLACEA-PNUD 1990).
Due to its high fiber content, sugarcane bagasse has been
used as part of ruminant feed. Although its digestibility is
low (approx. 25%), digestibility can be increased to 65%
through chemical, biological or thermomechanical treatments (Allen et al., 1997; Pate, 1982; de Medeiros and
Machado, 1993; de la Cruz, 1990). In natura sugarcane is
an economically feasible alternative to forage especially
during the drought season, when pastures are not sufficient
to feed herds (Embrapa Gado de Leite, 2008).
Chemical Industry
Biopolymers are polymeric-based materials that are structurally classified as polysaccharides, polyesters and polyamides. The basic raw material for production is a source of
renewable carbon. The most important biopolymers are
polylactate (PLA), polyhydroxyalkanoate (PHA), starch
polymers (PA) and xantane (Xan). According to Pradella
(2006), of all raw materials that are available for biopolymer production, sugarcane has one of the best profiles as a
carbon source. The lignocellulosic fiber content of bagasse
give sugarcane a unique competitive advantage compared
to other carbon sources since bagasse may also be used to
generate the energy used in biopolymer production.
Biotechnological Development
Use of Sugarcane in Natura
Unprocessed sugarcane is used as human food and animal
feed. As a food item, sugarcane may be consumed in natura
or as juice (garapa). In natura consumption is common in
Brazilian rural areas, but juice consumption is much more
common. Sugarcane juice is a nutritious energy drink that is
The sugarcane genome is among the most complex of
cultivated crops. This complexity has hindered our understanding of sugarcane genetics and our ability to improve
the crop using biotechnology tools (D’Hont et al., 2008). In
the late 1990s, in situ hybridization studies helped clarify
how the sugarcane genome is organized. A series of studies
described part of its complexity and clearly established the
Tropical Plant Biol. (2011) 4:62–89
ploidy level of the species, while revealing the coexistence
of two distinct genome organization modes in modern
varieties (D’Hont et al., 1996; D’Hont et al., 1998; Ha et
al., 1999). Tomkins et al. (1999) built a bacterial artificial
chromosome (BAC) library for the genome with over
100,000 clones, which facillitated beginning stages of
physical mapping of sugarcane chromosomes and comparisons with other grasses. In the same period, several
research groups started EST (expressed sequence tags)
sequencing projects in South Africa (Carson and Botha,
2000), Australia (Casu et al., 2001) and Brazil (Vettore et
al., 2001). Up to present, Brazilian sequencing project,
called SUCEST, was the project with greater sugarcane
EST sequence contribution, responsible for sequencing
238,000 ESTs from 26 different cDNA libraries.
Methods for the genetic transformation of sugarcane were
reported decades ago. The first experiments in sugarcane
genetic transformation were conducted in the late 1980 s
(Chen et al., 1987) when a kanamycin-resistance gene was
introduced into protoplasts through electroporation and
polyethylene glycol treatment. Transformed plants were
produced through particle bombardment (biolistics) of cell
suspensions and embryogenic calli (Bower and Birch, 1992).
Sun et al. (1993) obtained plants that were resistant to the
herbicide amonium-gluphosinate through biolistics using the
bar gene, which encodes phosphinothricin acetyltransferase.
Gambley et al. (1994) bombarded meristematic tissues with
microprojectiles and regenerated plants that expressed the
luciferase gene. Later, Arencibia et al. (1995) reported a
procedure for the stable transformation of meristematic
tissues using electroporation. Gallo-Meagher and Irvine
(1996) described the development of plants of the commercial cultivar NCo 310 containing the bar gene, which was
transformed through biolistics.
Currently, many studies have demonstrated Agrobacterium–mediated and particle bombardment transformation of
sugarcane with sufficient efficiency to produce commercial
varieties (Bower and Birch, 1992; Birch and Maretzki 1993;
Bower et al., 1996; Birch, 1997; Irvine and Mirkov, 1997;
Joyce et al. 1998a, b; Arencibia et al., 1998; EnriquezObregon et al., 1998; Elliott et al. 1998; Moore, 1999; Nutt
et al., 1999; Elliott et al. 1999; Manickavasagan et al., 2004).
Additionaly, several genes of commercial interest have been
introduced into sugarcane by genetic transformation, conferring herbicide tolerance, resistance to diseases and pests,
tolerance to drought, increased sucrose content, and
improvements in sugar quality and color (Falco et al.,
2000; Braga et al., 2001; Braga et al., 2003; Zhang et al.,
2006; Molinari et al., 2007). Sugarcane has also been
transformed with genes with the intention of using the plant
as a biofactory, producing high-value-added products such as
bioplastics (Lakshmanan et al., 2005) and high-value
isomers of sucrose (Wu and Birch, 2007).
85
In Brazil, the National Biosafety Commission (CTNBio)
has approved more than 40 applications to conduct field trials
of genetically modified sugarcane that contain genes conferring higher sucrose content, herbicide tolerance, insect
resistance and drought tolerance. Field trials have also been
conducted or are ongoing in South Africa, Australia and the
United States. These trials will almost certainly result in
biotechnology products. At present, however, transgenic
sugarcane cultivars have not been commercially released in
Brazil, or elsewhere in the world.
Open Access This article is distributed under the terms of the
Creative Commons Attribution Noncommercial License which permits any noncommercial use, distribution, and reproduction in any
medium, provided the original author(s) and source are credited.
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