Tropical Plant Biol.

(2011) 4:22–30
DOI 10.1007/s12042-011-9076-3

Sugarcane Underground Organs: Going Deep

for Sustainable Production
Sizuo Matsuoka & Antonio Augusto Franco Garcia

Received: 2 February 2011 / Accepted: 14 March 2011 / Published online: 30 March 2011
# Springer Science+Business Media, LLC 2011

Abstract Sugarcane breeding has greatly advanced in
recent decades, but many aspects of sugarcane physiology
are still poorly understood, including the root-shoot
relationships that ultimately affect yield. Traditional methods
for studying root systems are imprecise due to methodological
difficulties of in situ assessment and sampling; this seems
especially true for the sugarcane root system. Studies on
sugarcane roots lag well behind those on other crops, in part
due to the large plant stature and long crop cycle.
Commercial sugarcane cultivars are hybrids from crosses
mostly between Saccharum officinarum and S. spontaneum
made by breeders at the beginning of the last century. These
hybrids have a genomic structure composed of 80% S.
officinarum, 10% S. spontaneum and 10% recombinants of
these two species. S. spontaneum is included in large part for
the robustness of its underground organs (root and rhizome).
The S. spontaneum genes controlling these characteristics
may be lost during recurrent backcrosses with S. officinarum
to increase sugar content and yield. Thus, ratooning ability is
one of the most desired traits. Ratooning ability comes
mainly from the rhizomatousness of S. spontaneum, but this
trait has been diluted during the selection process so that the
stubble of hybrids does not have rhizomes sensu stricto. In
this review, we revisit some basic aspects of the sugarcane
root system, mainly from an ecophysiological view, and
point out considerations for breeders to consider in designing
the architecture of a new sugarcane cultivar that can meet the
need for sustainable agricultural production.
Keywords Saccharum officinarum . Saccharum
spontaneum . Rhizomatousness . Sugarcane breeding .
Energy cane
Abbreviations
QTLs Quantitative Trait Loci
GHG Greenhouse Gas
DMA Dry Matter Accumulation
SOC Soil Organic Carbon

Communicated by: Paulo Arruda

S. Matsuoka
Universidade Federal de São Carlos,
Campus de Araras-SP,
Introduction
Araras, Brazil
The first interspecific crosses to produce the noble canes
S. Matsuoka (*) were made as early as 1893 in Java, but success with this
Rua Ciro Lagazzi,
approach did not appear until the 1920s with the production
01. 13603–027, Araras, SP, Brazil
e-mail: sizuo.matsuoka@gmail.com of world famous hybrids POJ 2725 and POJ 2878, which
are progenitors of most cane cultivars worldwide. These
A. A. F. Garcia successes stimulated the establishment of sugarcane breeding
Departamento de Genética, Escola Superior de Agricultura Luiz
and research centers in the main sugar-producing regions of
de Queiroz (ESALQ), Universidade de São Paulo (USP),
CP 83, CEP 13400–970, Piracicaba, SP, Brazil the world, with further advances in knowledge supporting the
e-mail: aafgarci@esalq.usp.br expansion of the sugarcane industry (Tew 1987; Matsuoka et
Tropical Plant Biol. (2011) 4:22–30
al. 2009). In spite of the technological advancements made
during the last century in techniques for sugarcane farming
throughout the world, researchers have reported that practical
yields still lag behind the theoretical physiological limit
(Garside et al. 1997; Moore 2005).
Plant-ecosystem interactions are governed by many
complex factors so that complete control over all of them
is not possible in a variable and large agrosystem like a
sugarcane plantation, even when all of the available
advanced technologies are adopted. Moreover, the climatic
conditions over the large area and long crop cycle will be
less than optimal for maximum performance of the crop.
One important and often neglected issue is the functioning
of the root system and the root-shoot interactions, which
ultimately affect productivity and the agro-environment
(Lynch 2007). In this context, the comprehensive review on
the “growth and function of the sugarcane root system” by
Smith et al. (2005) is of particular value. However, in
addition to affecting agricultural production, the root
system has a major influence on carbon and mineral
nutrient cycling in the ecosystem (Gill and Jackson 2000;
Kumar et al. 2006) and these issues are particularly relevant
in managing the shift from traditional farming to a
sustainable agriculture system. Thus, the aim of this paper
is to revisit the sugarcane underground organs with a
holistic view, discussing issues that have received little
attention to date, but which we recognize as extremely
important in developing a new paradigm for sugarcane.
The Importance of Underground Organs
Plants have evolved to exhibit a characteristic balance
between the mass contained in the shoot and root proportions
in a given ecophsiological stage, and this ratio is characteristic
for each species or genotype (Marschner 1995). The
ecophysiological stage is defined by specific environmental
conditions dictated by light, temperature, wind, water,
nutrients, noxious substances, physical constraints and
biological interactions (diseases, pests, plant competition,
parasitism and symbiosis) that affect the morphology and
physiology of both the aboveground and belowground
organs (Eissenstat and Yanai 2002). Despite the importance
of roots in plant ecophysiology, relatively little is known
about the complex root-soil-shoot-atmosphere interactions.
This lack of knowledge can be attributed to the inherent
difficulties involved in comprehensive studies of root
systems and their relationships with the environment
(Marschner 1995). The anatomy and morphology of sugar-
cane roots have been reviewed elsewhere (Moore 1987a;
Smith et al. 2005) and is largely omitted from the present
review, which concentrates more on the genetic basis of the
root system functioning.
23
Most studies of the sugarcane root system have
disregarded the substantial influence of the genetic consti-
tution of the cultivars under study. Modern cultivars are
highly heterozygous interspecific hybrids derived mainly
from the introgression of a small genomic proportion of
accharum spontaneum into a predominantly S. officinarum
genome. Due to complex cytogenetic process the hybrids
have about 80% of the S. officinarum genome and 10% of
the S. spontaneum genome, with the remaining 10% of the
hybrid genome as recombinant chromosomes (D’Hont et al.
1996; Grivet and Arruda 2001; Cuadrado et al. 2004). S.
officinarum transmites high sucrose content and good
milling characteristics, while S. spontaneum contributes
physiological characters that allow the hybrids to be grown
in diverse, frequently unfavorable, environments (Tew
1987). The ability to adapt to harsh environments is
possible because S. spontaneum is more diverse, both
within the species and a single genome, than S. officinarum,
and contributes characteristics such as adaptation to biotic
and abiotic stresses, high tillering, and increased ratooning
ability (Jackson 1994; Legendre and Burner 1995). One
example of greater adaptability to distinct environmental
conditions as the result of a higher fraction of S.
spontaneum in the hybrid can be seen in the performance
of two cultivars, R570 and NCo376. R570 is a modern
cultivar with a chromosome constitution of 80:10:10 of S.
officinarum, S. spontaneum, and recombinants between the
two, respectively, whereas the old cultivar NCo376 has a
chromosome constitution of 70:20:10 (Ming et al. 2006).
NCo376, released in 1955, is one of the cultivars with the
longest commercial life in the last century (Wilson 1974),
while the cultivation R570 is restricted to specific, more
ideal areas. One can hypothesize that the success of
NCo376 could be attributed to its adaptation to the stressful
climate-soil-management conditions of South Africa and
other southern and central African countries (Meyer 2007).
The roots of NCo376 have been shown to extend deeper
and more profusely than those of other cultivars under
unfavorable conditions (Smit and Singels 2006). Other pair-
wise comparisons like this are strongly needed to foster the
hypothesis.
Molecular studies have suggested that the minor chromosome
complement inherited from S. spontaneum confers diversity and
adaptability to modern cultivars (Lu et al. 1994; Jannoo et al.
1999; D’Hont et al. 2008), and that plant selections for
cultivation in subtropical conditions generally give rise to a
greater fraction of S. spontaneum chromosomes than selections
for cultivation under tropical conditions (Jannoo et al. 1999;
Selvi et al. 2005). Thus, the underground part of S. spontaneum
can be imputed as largely responsible for the success of the
hybrid cultivars throughout the world and the potential for
increased use of S. spontaneum for adverse environments
merits more attention.
24
The Rhizome
Like many other grass species, S. spontaneum forms
rhizomes, but S. officinarum does not (Paterson 2009).
Thus, the rhizomatous characteristics of commercial sugarcane
come from S. spontaneum (Hu et al. 2003). The F1 generation
derived from pioneer crosses between S. spontaneum and S.
officinarum gave rise to rhizomatous populations. However,
due to the successive backcrosses and selection to augment
the traits of S. officinarum, rhizomatousness of the progeny,
together with other traits of S. spontaneum, gradually declined
(Burner and Legendre 1995). Nevertheless, a set of rhizome-
specific genes have probably been maintained in some
individuals, in a process similar to what happened during
the domestication of cereal crops (Hu et al. 2003).
Rhizomes develop from axillary buds at the lowermost
nodes of the erect shoot. Rhizome growth is oriented
perpendicular to the force of gravity (diageotropic), and
they retain the ability to produce geotropic shoots that
become independent ramets. Rhizomes are key to the
ability of many species to survive in harsh environments
(Hu et al. 2003; Paterson 2009). For example, in Louisiana,
USA, sugarcane planting is done in the fall early enough to
allow development of underground organs as a way to
avoid the annual freezing temperatures to which sugarcane
shoots are very sensitive. Freezing winter temperatures kill
the young shoots in late December, but the stubble stays
protected underground during the coldest temperatures;
when spring comes, new shoots are formed from the
overwintering belowground buds. The developed cane is
then harvested in the next fall, before the frost, and the
underground buds overwinter and sprout new shoots in the
spring to produce the ratoon crop. In studies with grasses,
QTLs for rhizomatousness and ratooning have been
identified and have been shown to be associated with one
another as a result of an ancient duplication, suggesting
long-term functional conservation of the underlying genes
(Jang et al. 2006).
Good ratoonability in sugarcane is undoubtedly linked to
high tillering ability that is in turn linked to high
rhizomatousness. Others have noted that genes associated
with tillering in the Poaceae may have some relationship to
rhizomes (Jang et al. 2006). It is possible that rhizome-
specific genes may serve multiple functions for plant
growth and development as well as serving regulatory
functions (Jang et al. 2009). The temperate region sugar-
cane cultivars showing better production performance are
generationally less distance removed from the ancestral S.
spontaneum (Tew and Cobill 2008). Pan et al. (2004) found
more than 60% homology between US56-15-8, a S.
spontaneum clone, and the highly successful cultivar in
Louisiana, LCP85-384, which is a BC4 derivative of that
ancestor. The maintenance of rhizomatousness to permit
Tropical Plant Biol. (2011) 4:22–30
overwintering and produce ratoon canes may explain this
fact, as the success of LCP85-384 has been attributed to its
capacity to produce additional ratoons when compared to
other contemporary cultivars. Therefore, rhizomatousness
should be considered a key trait to be incorporated in
sugarcane hybrids to encourage improved ratooning ability.
In S. officinarum, the primary shoot grows vertically, and
subsequent generations of shoots arise from buds at the
small bend at the previous shoot base to reach an upright
position. In S. spontaneum, the primary shoot first grows
horizontally, to variable extents, before emerging from the
ground; thus the primary shoot may be more properly
called a rhizome (Moore 1987a). The buried section of the
stalk and the rhizome are essential for tillering and for the
regrowth of the stool after the harvest of the stalks
(ratooning ability). As each stalk forms its own root
system, this tillering ability influences the final root system
of the stubble. Given the high cost of establishing a new
sugarcane crop, the greater the number of harvested ratoon
crops, the greater the profitability. The rhizomatous trait
from S. spontaneum has been diluted during the selection
process so that the stubble of modern hybrid cultivars does
not have rhizomes sensu stricto. Therefore, increasing the
rhizomatousness of cultivars is not only of direct agronomic
value but could also represent a great contribution to a
sustainable agriculture (Glover et al. 2007; Glover et al.
2010). A successful case was demonstrated when an early-
generation hybrid with a highly prolific stalk population
and deep root system exhibited outstanding ratoon regrowth
without a decrease in cane yield through eight ratoons,
benefiting from its rhizomatous growth habit (Giamalva
et al. 1984).
Studies on sugarcane roots usually do not consider the
rhizome or the underground stubble. Thus there is a paucity
of information on the total dry mass of the belowground
parts of any Saccharum species or hybrid cultivars. The
limited data that exists is generally only fresh weight,
which is of little use for a dependable comparative analysis.
The new demand for biomass production to help
mitigate the greenhouse gas (GHG) effect on the planet
has driven plant scientists to seek the development of crop
plants having a high efficiency in fixing atmospheric CO2
while being as environmentally friendly as possible in its
management. These lines of investigation frequently focus
on the potential use of C4 grasses, as opposed to C3 plants
because of their more efficient conversion of solar energy;
greater capacity to produce in suboptimal environments
where they are exposed to stresses such as drought, salinity,
sodicity, low temperature, low nutrient availability, etc.;
higher resistance to diseases and pests; perennially to
require less tillage and reduced inputs of fertilizer and
herbicide, thus causing less soil erosion and creating fewer off-
site consequences (Glover et al. 2007; Jonhson et al. 2007;
Tropical Plant Biol. (2011) 4:22–30
Yuan et al. 2008; Gonzalez-Herandez et al. 2009; Jakob et al.
2009). C4 plants have a high capacity to sequester C, both
aboveground and belowground. Grasses producing rhizomes
fit well with these characteristics (Paterson 2009) and,
thus, increasing the effect of rhizomatousness in sugarcane
cultivars is not only of direct agricultural value but should
also contribute to a more sustainable agriculture (Glover
et al. 2010).
A recent trend in sugarcane development is a shift to the
so-called “energy cane” (e.g., Mislevy et al. 1995; Bischoff
et al. 2008; Tew and Cobill 2008; Matsuoka et al. 2010),
which is a cultivar developed for energy production, rather
than conventional sugar production. Energy cane will add
considerable efficiency to the alternative biomass energy
model once the technology of second-generation biofuels
becomes more prominent (Sticklen 2008; Sainz 2009;
Kalluri and Keller 2010). This efficiency will result because
the new energy cane will maintain many of the characteristics
of its wild ancestors, including rhizomatousness. Paterson
(2009) stated that “growth and development of rhizomes is an
area of plant biology that remains unexplored” adding that
much remains to be learned on complex traits like rhizoma-
tousness and regrowth. Although rhizomatousness may be an
undesirable trait in weeds, it is advantageous in crops to be
used as biomass-bioenergy feedstock in a more sustainable
way (Hu et al. 2003; Jang et al. 2006; Paterson 2009). The
genus Saccharum is one of the best candidates for use in
breeding for that purpose because of its high energy-
conversion efficiency (Ming et al. 2006; Tew and Cobill
2008) and strong ratooning ability as well as other favorable
traits. The most likely candidate species is S. spontaneum
(Ming et al. 2006; Wang et al. 2008).
The Root
There are two groups of roots in sugarcane. The first
consists of the roots that grow from the root primordia of
the root band of the stalk nodes. These have been
designated as the sett roots (sett is a word used to denote
a section of stalk or culm containing one or more nodes
with an axillary bud for asexual propagation). The growth
rate of sett roots is higher than that of the roots originating
at the base of the shoot (called shoot roots) possibly to
quickly supply enough water and nutrients to the highly
demanding young shoot. However, not all of the root
primordia on the sett form roots. Some primordia remain
dormant and germinate roots only if the roots that are
already growing are severely damaged. Thus, the spare root
primordia provide a survival mechanism to supply water
and nutrients for the young shoot in its early phase; later, as
the shoot grows and develops its own shoot roots, the sett
roots gradually die off (Moore 1987a; Smith et al. 2005).
25
The higher number of sett roots produced by S. officinarum
as compared to S. spontaneum (Moore 1987a) is probably
also a survival strategy of the former, since, in the tropical
conditions where it evolved, vigorous shoot growth is
essential to competing and surviving in a vigorous plant
community. S. spontaneum, on the other hand, is generally
growing in a harsh environment where there are not many
vigorous competitors; the survival strategy in this case is to
develop strong underground tissues (rhizome) to withstand
environmental stresses instead of developing a large
number of shoot roots. This is probably why F1 hybrids
of S. spontaneum and Miscanthus usually have slow growth
during the initial months, and the aerial mass is usually
lower in the plant crop cycle than in the immediately
subsequent ratoons (Bischoff et al. 2008).
As indicated above, the second group of sugarcane roots
is those coming from the base of the shoots. These originate
from the base of the primary shoot as well as from the bases
of the young tillers (Smith et al. 2005). In this group of
roots, there are three types: “superficial roots”, “buttress
roots” and “rope systems” (Moore 1987a; Smith et al.
2005). Superficial roots are thin and abundant, spreading
horizontally in the upper 6 to 10 cm of topsoil and forming
an entangled mat. They extend from the tiller nodes near
the surface and undergo abundant branching. They are
covered by root hairs and are the principal absorbing roots
in sugarcane. The buttress roots grow from the basal nodes
of the young shoot. They are white, succulent and thicker
than the superficial roots. Growing at 45 to 60º angles, they
are not much branched, and their main function is to anchor
the plant. Rope systems are those roots that penetrate the soil
profile vertically and are characteristic of S. spontaneum
(Moore 1987a).
Characteristics of sugarcane root system depend on the
proportion of S. officinarum and S. spontaneum genomes in
the cultivar, and usually, the fewer generations removed
from the original ancestor, the higher the contribution of S.
spontaneum. Roots consist of various functional zones from
the root cap to the base, with the root hair zone being the
most permeable and thus playing a role in water uptake
(Segal et al. 2008). As root hairs may vary in length,
number and positioning along the root according to the
genotype (Marschner 1995; Segal et al. 2008; Zhu et al.
2010), their physiological activity should vary accordingly,
and this may play a role in the agronomic value of a
cultivar. In sugarcane, 80 to 90% of the root mass is
concentrated in the top 60 cm of soil, and more than 50% is
in the top 200 mm (Ball-Coelho et al. 1992; Vasconcelos
et al. 2003; Otto et al. 2009). It has been shown that
nitrogen fertilization via the application of vinasse, a
nitrogen rich aqueous residue from alcohol distillation,
favors intense root growth in the surface layer of soil (Otto
et al. 2009). Continuous irrigation also promotes root
26
development mostly superficially whereas water starvation
promotes roots to grow more evenly and deeply (Smith et
al. 2005). These observations indicate that providing plants
with a medium environment quality would stimulate
growth of roots to cover a larger volume of soil, thus
reducing root damage from episodes of environmental
stress. Roots may also develop more in a superficial patch
as a result of soil strength of the sub-soil layer (Monteith
and Banath 1965; Marschner 1995), restriction of essential
nutrients in the lower soil layers (Dias et al. 1999), sub-soil
acidity and the presence of toxic chemicals ions such as
aluminum (Matsumoto 2002). Root also grows deeper in
advanced ratoons as compared to plant cane (Vasconcelos
et al. 2003). It is well established among sugarcane farmers
that ratoon crops have a faster rate of canopy development
than do plant crops. The persistence of roots from a previous
cycle could be a significant factor in more rapid development
of ratoon crops since part of the driving force for the
production of new shoots would be the input of water and
nutrients provided by these persistent roots, especially during
a dry period. Differences in root mass between crop cycles
may also result from climatic conditions prevailing during the
crop cycle or in the immediate period anteceding the
sampling time and field-management procedures (mainly
harvesting method, if manual or mechanical), in interaction
with the cultivar (Vasconcelos et al. 2003).
Root Shoot Ratio
Root and shoot dry matter accumulation (DMA) is used to
calculate a root/shoot ratio of a given plant in a crop field
environment. It must bear in mind that in sugarcane crop
shoot growth is much influenced by competition among
plants that results from the spacing of the rows (Kanwar
and Sharma 1974; Irvine and Benda 1980). Although most
root studies to date have been done in field conditions, the
root/shoot ratio has been mostly ignored which makes
effect of treatment comparisons difficult and weakens the
interpretations.
The harvestable portion of the stalk fluctuates roughly
between 60 and 85% of the total aerial biomass on a fresh
weight bases, depending on the cultivar, age, ripening stage
and environment (Muchow et al. 1994; Singels et al. 2005).
After planting, sugarcane starts its development by sprouting
and emergence of a primary shoot with its corresponding
shoot roots. The plant then proceeds through a stage of intense
tillering and shoot formation and development referred to as
the grand growth phase (Robertson et al. 1996; Singels and
Smit 2002). Later, many of the shoots die as a result of plant
competition, leaving only a fraction of the initial shoots at
harvest (Kanwar and Sharma 1974; Inman-Bamber 1994;
Robertson et al. 1996; Muchow et al. 1997; Zhou 2005).
Tropical Plant Biol. (2011) 4:22–30
Evensen et al. (1997) found a decrease in the proportion
of the belowground biomass from 17% of the total biomass
at 6 months to 11% at 12 months. Studies have shown that,
during the phase of high tiller mortality, there is no
impairment of the growth of living shoots, and shoot
growth may even be favored in the remaining shoots (Otto
et al. 2009). In field conditions, the constraints of
competition for limited resources occur intermittently; since
roots and shoots are intimately connected, root dynamics
depend on shoot dynamics and vice versa (Reich 2002).
Root growth responds strongly to the soil environment,
creating plasticity in the form and size of the root system
(Glover 1967). The most efficient cultivar in using limited
resources is one that loses few tillers, and thus fewer roots,
throughout its life span so that it maintains an adequate
number of roots even during stressful conditions. However,
a compromise is needed for satisfactory stubble regrowth,
which is usually linked to the capacity for bearing a high
number of tillers (Hu et al. 2003), which as emphasized
earlier is a trait inherited from S. spontaneum (Dunckelman
1974; Jackson 1994).
The higher the root mass, the greater the environmental
benefits, mainly via an increase in soil organic matter (with
all of its beneficial consequences in terms of fertility),
improved soil structure and, as result of these improve-
ments, higher erosion control (Kumar et al. 2006). As
discussed by Smith et al. (2005), studies determining
“living root biomass seriously underestimate C allocation
to roots”. The C sequestered in the root debris in the soil
profile is an important contribution by plants to alleviating
the GHG, but it represents only a small fraction of the total
soil organic carbon (SOC). A significant proportion of the
SOC results from rhizodeposition, i.e., the C released by
roots as exudates in the form of a wide array of organic
compounds (Kumar et al. 2006), together with the cells of
the root cap that is continuously sloughed off as the root
grows. On average, 30% to 60% of the C photosynthesized
by the plant is allocated to the root system, and of this, 40%
to 70% may be released to the rhizosphere (Lynch and
Whipps 1990). The rich substrate in the rhizosphere is a
medium for intense activity of a wealth of microorganisms
and soil fauna that may be beneficial, neutral or harmful to
the root (Jones and Hinsinger 2008).
Drought Stress Tolerance
Drought stress tolerance is a genetic trait in which breeders
of most crops are extremely interested. When analyzing
data for modern sugarcane cultivars, we must keep in mind
that we are dealing with hybrids having the genomic
constitutions discussed above. Drought-resistant S. spontaneum
clones have been shown to possess deeply growing rope root
Tropical Plant Biol. (2011) 4:22–30
systems that are not found in drought-susceptible cultivars.
Vigorous root systems correlated with drought tolerance
(Moore 1987b). It is possible that the fine horizontal roots
growing in the topsoil have fundamental functions in
absorbing water and nutrients under less stressful conditions,
however, as they are more sensitive to water deficit and can die
easily as it progresses, deeper and more robust roots will
gradually take their place to supply the plant’s necessities. A
remarkable characteristic of the sugarcane root system that
allows it to thrive satisfactorily under the intermittent water of
rain-fed conditions is the capacity of rope roots to grow down
to 400 to 600 cm, a depth to which other crop plant roots
rarely reach (Canadell et al. 1996). Quantitative cause-effect
relationships for characteristics such as drought stress tolerance
are not always simple to determine, since many interacting
factors may be acting simultaneously (Whitmore and Whalley
2009; Passioura 2010). In each case, the combination of factors
causing drought stress is unique. In one field, the main factor
could be soil compaction resulting from improper use of
vehicles (Yang 1977), in another, the primary factor could be a
chemical impediment (soil acidity or low levels of an essential
nutrients), while in another field, a root disease or a pest attack,
individually or in conjunction, may be the main cause
(Whitmore and Whalley 2009). Moreover, the complex and
active biota of the rhizosphere is another significant factor
influencing root dynamics (Jones and Hinsinger 2008).
Breeding of sugarcane for resistance to abiotic factors is
usually made indirectly via the effects they have on plant
development. As sugarcane is a very complex plant and
almost all of the agronomic characters are of a quantitative
nature, the biometric breeding and selection approach is
most utilized. In early breeding stages, selection is made for
general morphological appearance and for the introgression
of specific desired agronomic traits such as sugar content
and resistance to diseases. With later stage advancement of
selection and the corresponding reduction in the number of
selected individuals, multi-location yield trials are established
to evaluate performance of promising clones. Breeding and
selection programs never seriously consider features such as
root structure, even in those programs focused on drought
stress tolerance. Root structure selection is only derived
indirectly by selecting for better performance under drought
conditions.
The rhizome is an important organ for drought stress
tolerance. In some clones of S. spontaneum, the rhizome
extends far from the base of the plant, and tillering is very
intense. This characteristic, when transferred to new
cultivars, can lead to longer persistence of ratoons even
under conditions of intense mechanization or in poor soils
and marginal areas. Rhizomatous is a breeding and
selection trait worthy of consideration: the longer the crop
stays in the field without plowing, the less costs for
machinery, management operations, fertilizers, fuels, etc.
27
and, most importantly, the better is the soil conservation.
Re-growth of the stool after harvest is also an important
issue to consider in dealing with drought stress tolerance in
sugarcane. Water deprivation not only reduces normal plant
growth, thereby diminishing productivity, it also reduces
survival of the stool after harvest. Throughout the long
duration of the harvest season for the sugarcane crop (from
3 to 11 months, depending on the region), the occurrence of
a dry period in some fields is inevitable, and depending on
the intensity of the drought interacting with the temperature
(low in some regions and high in others), the stool may die.
The resulting number of gaps without plants in the field
may make it necessary to plow out the field precociously to
establish a new crop. This is a costly operation in sugarcane
farming, so that an ability to increase the number of
successive ratoons between plantings is a desire of all
farmers. Rhizomatous growth, as mentioned above, is
critically important for good ratooning. In hot regions,
such as in Australia and parts of Brazil where there is little,
or variable, rainfall, summer dormancy is an important trait
for improving survival rates of pasture grasses (Nie and
Norton 2009). Fully investigating the summer dormancy
trait could be an important contribution for the future
advancement of sugarcane cultivar improvement for hot
and dry regions.
Root diseases and pests represent an additional group of
factors that exert an indirect negative influence on drought
stress tolerance. Sugarcane roots are prone to being attacked
by several soil microorganisms. Such microorganisms can
cause root rot and other physiological disorders, as well as
inject harmful toxins capable of killing the roots and even the
young shoots (ex: pineapple disease and leaf scald). Like pests
that feed on the roots, or on the base of the shoots, or suck the
photoassimilates, they can exacerbate the drought sensitivity
of the plants. However, the degree of influence of these factors
depends on the growing conditions of the plant. A plant's
“spare capacity” can effectively buffer it against root loss or
sub-optimal soil environment conditions. However, this
spare capacity is effective only for limited periods of time,
since most times the agroecosystem is rather poor in several
factors. The role played by root lysis to increase water
penetration in the soil profile through the microchannels left
by this process is also an important area for investigation; the
consequences of root lysis may be important in alleviating
the level of water stress during dry spells as the resulting
micro-channels could favor water penetration in the soil.
It is important to note that drought conditions bring
about several side effects that might interfere with the
physiology of plants. For a given soil, mechanical impedance
increases as the soil dries out, and this is a stress factor
along with low soil water potential (Marschner 1995).
The effect of mechanical impedance on root growth is well
known (Passioura 1991; Marschner 1995), but other
28
factors also come into play, including oxygen deficiency,
elevated concentrations of ethylene and differential
uptake of nutrients, to mention a few (Marschner 1995;
Passioura 2010).
Concluding Remarks
Over the last 50 years, agriculture advancements have been
driven by the philosophy of the “Green Revolution”, that is,
agricultural research has focused on improving productivity.
Currently, a new era has begun that emphasizes sustainability in
addition to productivity. Sustainable agricultural production
will necessitate a compromise between maximum productivity
and sound management practices to maintain or restore the
health of agroecosystems for the long term. In other words,
actions taken to develop an environmentally benign agriculture
will help preserve world resources to benefit human popula-
tions, ultimately preserving the life of our planet. The only way
to adequately address the paramount requirement for food and
energy is to increase our focus on agriculture marginal areas,
both in terms of soil and climate, and to exploit these areas with
a minimum resource input. Lynch (2007) considered this shift
towards sustainability as a central role for the “second green
revolution”. Perennial grasses, either for food or for energy,
can greatly contribute to the development of this new
paradigm (Cox et al. 2002; Glover et al. 2007, 2010; Yuan
et al. 2008). Roots play a key role in the dynamics of the
planet’s ecosystem and are crucial in agroecosystems to
sustain productivity, contribute to the maintenance and/or
improvement of healthy systems and help with the balance of
carbon in the atmosphere by C-sequestration (Norby and
Jackson 2000; Kumar et al. 2006; Glover et al. 2007; Pierret
2008). On first inspection, plant roots and their functioning
seem to be simple, but science is now discovering that hidden
in the soil is a very complex interplay among roots, soil, biota,
water and air. What is already known gives us a some clues of
what to look for, but coming to terms with the complex issues
of climate change and environmentally friendly exploitation
of the resources of our planet provides a new impetus for
research aimed at a holistic understanding of the functioning
of root systems (Skaggs and Shouse 2008). Root studies are
notoriously difficult, presenting methodological obstacles in
addition to being time-consuming and costly. New methods
have been developed to evaluate roots in the soil profile or in
artificial conditions (Polomski and Kuhn 2002; Waisel 2002).
For sugarcane, most studies have been done in field
conditions, but a few have also been conducted in rhizotrons
and in pots.
Soil is a very complex substrate with variations occurring
at the millimeter scale. As pointed out by Pierret (2008) “to
date, the majority of studies on roots have been conducted
based on the false premises of homogenous soil conditions”
Tropical Plant Biol. (2011) 4:22–30
which do not reflect conditions in nature (Lynch 1995).
Realizing this problem, Zobel (1992) stated that the evidence
suggests that the roots routinely studied in laboratory
experiments are normally either nonfunctional in the field,
or have different functional characteristics than the majority
of the active roots of field grown plants”. This present
review shows that the sugarcane root system requires more
comprehensive studies. Any research project on roots should
be very carefully designed and executed without a restricted
disciplinary approach, but rather holistically as is being
increasingly done in the biological sciences (Kalluri and
Keller 2010) by an interdisciplinary team, for exploring the
biological, genetic, agronomical, chemical and physical
aspects of rooting. In this way, the results and the
interpretations will be robust and foster the development of
reliable knowledge. Cutting-edge technology for the study of
roots is being developed, including high throughput imagery
equipment formerly developed for industrial and medical
research and use, together with other modern instruments
and new procedures and protocols (e.g., Pierret 2008; Segal
et al. 2008; Skaggs and Shouse 2008; Tulinson et al. 2008),
with the added support of advanced modeling programs run
by high capacity computers (Pierret 2008; Skaggs and
Shouse 2008). The application of these tools and techniques
should speed the development of our understanding of
sugarcane underground systems.
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