Field Crops Research 134 (2012) 95104

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Field Crops Research

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Sugarcane for water-limited environments: Theoretical assessment of suitable

traits
N.G. Inman-Bamber a, , P. Lakshmanan b , S. Park c
a

CSIRO Plant Industry, Building 145 - ATSIP, James Cook Drive, James Cook University Douglas Campus, Townsville, QLD 4811, Australia
David North Plant Research Centre, BSES Limited, 50 Meiers Road, Indooroopilly, QLD 4068, Australia
c
CSIRO Ecosystem Sciences, Clunies Ross Street, Black Mountain, Canberra, ACT 2601, Australia
b

a r t i c l e

i n f o

Article history:
Received 13 July 2011
Received in revised form 27 April 2012
Accepted 13 May 2012
Keywords:
Rooting depth
Transpiration efciency
Crop simulation
APSIM
GE

a b s t r a c t
In Australia water stress is estimated to cost the sugar industry an average of $260 million (AUD) per
annum in lost production. With the predicted increased frequency of drought events the industry is now
considering breeding for drought adaptation after water stress inicted yield losses of more than $400
million in the years 20022004, in one region alone. Dening drought adaptation broadly, including both
short and long periods of water stress, we took the rst step in improving sugarcane for such conditions
by assessing the potential benets of a number of traits in a simulation study. The APSIM-Sugarcane
model was used to simulate the biomass yield response to traits that may confer adaptation to drought
in a range of climates, some extremely dry at times, and in a shallow and a deep soil. Among the traits
studied, increased rooting depth resulted in 021% increase in mean dry biomass yield depending on
the climate and soil type. This trait was more benecial in the shallow than the deep soil which had a
smaller fraction of additional stored water to offer the more vigorous root system. The simulations showed
that breeding for reduced stomatal or root conductance (conductance) would increase biomass yield by
about 5% only in the driest climates and better soils. Other traits which conserved water such as leaf and
stalk senescence were generally unsuccessful in conferring adaptation to the water-limited production
environments considered. Simulations indicated that increased transpiration efciency (TE) at the leaf
level would nearly always help to improve sugarcane biomass yields in water-limited environments
if the increased TE arose from up-regulation of intrinsic water use efciency. However if increased TE
was increased through reduced conductance, which effectively reduces VPD during transpiration, yields
could be reduced in high rainfall climates and shallow soils and they could increase in moderate rainfall
climates and deeper soils. Thus increased rooting depth, increased intrinsic water use efciency and to
a lesser extent, reduced conductance leading to increased TE, are suggested as the best traits to consider
for selection of sugarcane clones in water-limited environments in the tropics and sub-tropics.
Crown Copyright 2012 Published by Elsevier B.V. All rights reserved.

1. Introduction
Water is the single most limiting factor worldwide to the
productivity of rainfed crops and those with supplementary irrigation (Juenger et al., 2005). It is likely to further constrain crop
production where seasonal rainfall is predicted to be more variable, and/or decline (Campos et al., 2004). In Australia, the 2002
drought reduced farm output by nearly 30% (Horridge et al., 2005).
In the sugarcane industry alone, water stress is estimated to
cost an average of $260 million per annum in lost production
(Inman-Bamber, 2007). These losses occur despite nearly 60% of

Corresponding author. Tel.: +61 7 47538587; mobile: +61 0424759841.

E-mail addresses: geoff.inmanbamber@gmail.com,
geoff.inman-bamber@csiro.au (N.G. Inman-Bamber), plakshmanan@bses.com.au
(P. Lakshmanan), sarah.park@csiro.au (S. Park).

the industry receiving supplementary irrigation (Inman-Bamber

and McGlinchey, 2003). All sugarcane-growing regions in Australia
experience extreme seasonal and annual variability in temper
atures and rainfall largely due to the inuence of the El Nino
Southern Oscillation (ENSO) (Nicholls and Kariko, 1993). It is therefore questioned whether developing drought tolerant and water
use-efcient sugarcane varieties may reduce these productivity
losses.
Whilst there appears to be clear benets from breeding cultivars
with improved drought-tolerance and water use efciency (WUE)
traits, the list of potential traits associated with drought tolerance
and WUE is extensive (Ludlow and Muchow, 1990). Further, the
traits expressed are known to depend greatly on the environment
in which the crop is grown, the specic climate experienced during crop growth and the management strategy imposed on the
crop (Chapman et al., 2000). The large genotype environment
(G E) interactions for crop yield mean that breeding programs

0378-4290/$ see front matter. Crown Copyright 2012 Published by Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.fcr.2012.05.004

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N.G. Inman-Bamber et al. / Field Crops Research 134 (2012) 95104

must select for the specic secondary physiological traits associated with particular drought tolerance and WUE strategies deemed
optimal for a dened water-stressed environment (Tardieu, 2003).
Some degree of drought adaptation has been reported for a
number of sugarcane cultivars. These showed drought avoidance
traits such as leaf rolling, early stomatal closure, leaf shedding
and reduced leaf area, rather than traits that confer drought tolerance (Inman-Bamber and Smith, 2005). Leaf rolling and early
stomatal closure are highly reversible. In a cultivar with these
two traits, CO2 assimilation and transpiration would be reduced
during a dry period, but would then resume quickly when water
stress was relieved either through increased soil water content or
reduced atmospheric demand for water. Cultivars vary considerably in regard to stomatal response to soil water decits (Saliendra
and Meinzer, 1989; Inman-Bamber and Smith, 2005). Two cultivars
subjected to water stress reduced their projected leaf width similarly through leaf rolling (Inman-Bamber and de Jager, 1986) but
Rutherford (1989) suggested that leaf rolling may vary considerably
amongst sugarcane genotypes.
Cultivar differences in leaf shedding in response to water stress
have been observed (Inman-Bamber and de Jager, 1986). This is a
more drastic response to water stress than leaf rolling or stomatal
closure and recovery would be relatively slow. Stalk senescence is
another option for the crop to reduce water-use during extended
dry periods. There are few studies on the nature of water stress
and stalk senescence but experience shows that crops lose stalks
and stools during severe drought, eventually requiring a replanting program. Stalk population in two cultivars remained unaffected
during a period without irrigation long enough to reduce biomass
accumulation by 70% (Inman-Bamber, 2004), suggesting the need
for severe stress for stalk loss to occur.
Clearly a mechanistic approach to conventional crop improvement programs requires knowledge of how plants manage
competing requirements to assimilate carbon and conserve water
throughout their growth and development. By capturing current scientic understanding of the physiological determinants of
crop growth and development, mechanistic and eco-physiological
crop models provide a useful tool for integrating physiological
understanding into empirical plant breeding procedures. Such an
approach can enable a quantitative assessment of the impact of specic traits on crop yield (Tardieu, 2005), aid the design of optimal
ideotypes for target environments (Shorter et al., 1991; Chapman
et al., 2002), facilitate an assessment of the variation in the G E
interaction for quantitative traits such as yield, and determine the
representativeness of the short-term climate and associated yields
obtained in plant breeding trials, to more long-term historic climate
and crop performance (Chapman et al., 2002).
In this paper we conduct a quantitative assessment of the
impact of specic traits such as conductance, rooting depth, leaf
senescence and transpiration efciency (TE) on biomass yield of
sugarcane to support a largely eld-based breeding program in
search of drought tolerance mechanisms and related traits in different production conditions. To the best of our knowledge this is
the rst study of its kind for sugarcane.

2. Materials and methods

In this simulation study, the APSIM-Sugarcane module (v 7.2)
was used in conjunction with the Agricultural Production Systems
sIMulator (APSIM) crop modelling system (McCown et al., 1996;
Keating et al., 2003). The model is designed to simulate cane yield,
sucrose yield, crop biomass, water use, crop nitrogen uptake and
partitioning of carbon and nitrogen for a uniform eld of sugarcane
(Lisson et al., 2005). The APSIM-Sugarcane module was developed from detailed growth analysis experiments, 14 of which were

conducted along the east coast of Australia (18.429.5 S), four on

the east coast of South Africa (26.229.5 S) and one in Hawaii
(21.5 N). The experiments included ve varieties and various levels
of N and water regimes (Keating et al., 1999). The Australian experiments were done largely with the commercial variety Q117 and
the South African experiments with NCo376 and N12. Coefcients
of determination for model predictions compared to observed
data (n > 150) were 0.79 for leaf area index (LAI), 0.93 for dry
biomass yield and 0.83 for sucrose yield (Keating et al., 1999).
Q117 is therefore one of the varieties that has been characterised
comprehensively in terms of its physiology and crop growth and
development and was used at the default variety (clone) for the
long-term simulations in this study.
APSIM-Sugarcane was used to simulate reduced water use as
would occur with early stomatal closure, rapid leaf shedding and
early stalk senescence in response to increased water stress. Two
additional traits for which there is some evidence of variation in
sugarcane genotypes are transpiration efciency (TE) (Saliendra
et al., 1996) and rooting depth distribution (Smith et al., 2005).
These were also varied within realistic limits in the APSIM simulations.
Two published eld experiments (Inman-Bamber, 2004) were
used to verify simulations of rapid leaf shedding responses to
water stress. We reported only biomass yields in this study because
in some situations, water stress was so severe that simulated
crops never advanced enough to accumulate signicant amounts
of sucrose. In addition, biomass is increasingly becoming the
starting point for integrated sugarcane-based industries that may
deliver sugar, ethanol, electricity and high-value products in future
(Waclawovsky et al., 2010).
2.1. Field experiments
Full details of the two experiments used for verication of the
leaf senescence trait were provided by Inman-Bamber (2004) and
only essential details are presented here.
The two experiments were planted simultaneously on 15 June
1998 at Kalamia Estate, Ayr, Queensland (147 25 E, 19 32 S). Both
experiments were of a randomized split plot design with cultivars
Q96 and Q124 as the sub-plot treatment and irrigation (wet) or
suspended irrigation (dry) as the whole plot treatment. Sub-plot
size was 45 m 10 rows, 1.5 m apart. There were ve replications
and hence 20 sub-plots in each experiment.
Experiment 1 was ratooned on 28 April 1999 in preparation for
imposition of a stress treatment in September to December and
experiment 2 was ratooned on 15 November 1999 in preparation
for imposition of a stress treatment after the annual wet season
(usually December to April). Both crops were maintained following standard crop management practices, other than irrigation.
Lodging was minimal in these trials. Standard industry irrigation
management was applied to all plots until it was time to impose
water stress to half of the plots in each experiment. For the rst
experiment the dry plots were irrigated on 21 September 1999 for
the last time and for the second experiment, dry plots were not
irrigated at all after the wet season. Biomass, LAI and stalk population were monitored at 23-week intervals through destructive
sampling as described by Inman-Bamber (2004).
The experiments were simulated using standard (default) settings in APSIM and canopy characteristics for one of the varieties
(Q96) used in the experiment. The other variety was Q124 but it
succumbed to orange rust in the second experiment and was therefore excluded (Inman-Bamber, 2004). Settings for the maximum
area of successive leaves of Q96 recognised the reduction in leaf
area in older leaves even when plants were well irrigated (InmanBamber, 2004). Leaf numbers 1, 14, 30 and 40 were assigned areas
of 50, 550, 550 and 350 cm2 and served as inection points for the

N.G. Inman-Bamber et al. / Field Crops Research 134 (2012) 95104

extrapolation of the area of other leaves. Climate data was obtained

from an automatic weather station installed within 2 km from the
site.
2.2. Reduced conductance
APSIM-Sugarcane does not simulate stomatal conductance as
such so the impact of reduced stomatal conductance was simulated
by modifying experimentally determined root water extraction
coefcients (KL ) for sugarcane (Inman-Bamber et al., 2001). There
is some evidence that root resistance and/or root signals, regulate
resistance to water ow though the soilplantatmosphere continuum (Saliendra and Meinzer, 1989) so the variation of KL to
simulate additional reduction in transpiration and CO2 assimilation
during water stress, has some foundation. Bulk canopy resistance
varied from 34 to 57 s m1 in studies on sugarcane by Grantz and
Meinzer (1991) and McGlinchey and Inman-Bamber (1996) thus
a 50% reduction in conductance (1/resistance) seemed plausible
in our simulation study. Standard and reduced values for KL were
used to compare standard and high resistance to water vapour and
CO2 ux. A 50% reduction in KL (Table 1) reduced transpiration by
about 20%. Photosynthesis was reduced in proportion to transpiration when atmospheric demand for water exceeded root water
supply (Keating et al., 1999).
2.3. Simulation of deep root distribution
The observed differences in rooting depth between modern cultivars are not large and some at least can extract water to 3 m and
possibly more (Smith et al., 2005). The water balance module (Soilwat) in APSIM divides the soil into layers (Table 1) to simulate water
ux in one dimension only but in two directions, with gravity and
capillarity involved (Keating et al., 1999). Roots were allowed to
explore one additional layer in each of the two soils (Yellow chromosol and Red dermosol) simulated in this study. Deeper roots
were also made more efcient by increasing the lower limit (LL)
to which water could be extracted from each soil layer beneath the
surface layer (Table 1). The LL for the surface layer was not altered
to avoid confounding the root depth and vigour traits with emergence rate which in the model depends on the amount of water
available in the top layer. Hypothetical clones with shallow and
less vigorous roots had access to 10% less water stored in the soil
layers where roots were present and 22% less water over all layers
for the shallow soil (60 mm and 76 mm for shallow and deep roots,

97

respectively) and 10% less water for the deep soil (258 mm and
287 mm for shallow and deep roots, respectively). The default
settings in APSIM-Sugarcane allow the rooting front to advance
10 mm d1 until the stalk elongation phase starts and then at
15 mm d1 thereafter (Keating et al., 1999). The model assumes that
rooting depth penetration is reduced when soil water content in a
particular layer falls below 25% but we removed this limitation in
our study.
2.4. Simulation of rapid leaf senescence
In APSIM sugarcane, water stress reduces photosynthesis in proportion to the extent to which root water supply fails to meet
atmospheric demand for water. The ratio of root water supply to
water demand is called the stress factor for photosynthesis (SP or
swdef photo in APSIM code). When SP is less than 1.5, green leaf
area is reduced each day by the fraction 0.01(1.0 SP ) to simulate
standard leaf senescence. To simulate rapid leaf senescence the die
back fraction was increased to 0.03(1 SP ) per day based on the leaf
area loss observed under water stress in the second experiment
of Inman-Bamber (2004). Initial rate of leaf senescence during the
onset of water stress was more than three times greater in one cultivar of soybean than another (Lawn and Likoswe, 2008) and it is
seems from the eld experiment (Inman-Bamber, 2004) that leaf
senescence could vary this much in sugarcane. Inman-Bamber and
de Jager (1986) concluded that cultivar N11 was better adapted
to water stress than another cultivar, NCo376, because leaf area
declined more rapidly during stress and increased more rapidly
after stress was relieved than was the case for NCo376. However
Smit and Singels (2006) found no difference in the rate of decline in
leaf area between N22 and NCo376 during a period of water stress.
2.5. Early stalk senescence
Stalk senescence is known to occur in sugarcane but this process
was omitted in the sugarcane module of APSIM because the exact
causes were not known at the time (Keating et al., 1999). Lodging was later found to be one of the causes of stalk senescence
(Singh et al., 2002) and a procedure for reducing stalk population was introduced to the sugarcane module. A senescence rate
up to 0.1 stalks m2 d1 was reported by Park and Attard (2005)
and Park et al. (2005) when attempting to explain reduced growth
in ageing sugarcane and a mid-range value of 0.05 stalks m2 d1
due to water stress is conceivable. Assuming a typical nal

Table 1
Volume fraction for lower limit of water availability, drained upper limit, saturation and saturated conductivity coefcient and standard, reduced and deep values for root
water extraction coefcient (KL ) in simulated layers for two soils (see Inman-Bamber et al., 2000). All units are fractions except for soil depth.
Yellow chromosol (poor soil)
Layer
Soil depth from surface (mm)
Lower limit shallow roots
Lower limit deep roots*
Drained upper limita
Saturationa
Saturated conductivity coefcient
KL standarda
KL reduced
Red dermosol (excellent soil)
Soil depth from surface (mm)
Lower limit shallow roots
Lower limit deep rootsa
Drained upper limita
Saturationa
Saturated conductivity coefcient
KL standard
KL reduceda
a

1
200
0.082
0.082
0.162
0.289
0.8
0.10
0.05

2
400
0.083
0.077
0.136
0.247
0.8
0.10
0.05

3
500
0.099
0.094
0.141
0.25
0.8
0.08
0.04

4
600
0.115
0.111
0.151
0.236
0.8
0.07
0.035

5
800
0.115
0.111
0.151
0.236
0.8
0.06
0.03

6
1000
0.115
0.111
0.151
0.236
0.8
0.05
0.025

7
1300
0.115
0.111
0.151
0.236
0.8
0.05
0.025

8
1600
0.151
0.111
0.151
0.236
0.8
0.05
0.025

200
0.171
0.171
0.282
0.358
0.5
0.10
0.05

400
0.235
0.225
0.325
0.379
0.5
0.10
0.05

500
0.280
0.27
0.371
0.402
0.5
0.08
0.04

600
0.327
0.317
0.42
0.427
0.5
0.07
0.035

800
0.329
0.319
0.416
0.425
0.5
0.06
0.03

1500
0.333
0.323
0.412
0.423
0.5
0.05
0.025

2300
0.333
0.323
0.412
0.423
0.5
0.05
0.025

3100
0.412
0.323
0.412
0.423
0.5
0.05
0.025

Based on measurements by Inman-Bamber et al. (2000, 2001).

98

N.G. Inman-Bamber et al. / Field Crops Research 134 (2012) 95104

Table 2
Traits and treatments with up to ve variations or levels used in 640 factorial combinations in the simulation.
Trait or treatment

Level 1

Level 2

Rooting depth and vigour

Root water extraction coefcient (KL )
Leaf senescence rate (m2 d1 )
Stalk senescence (% stalks m2 d1 when
leaf area index falls below 0.5)
Intrinsic water use efciency (g kPa kg1 )
Soil

Standard
Standard
0.01(1-Sp)
0.0

Reduced
Reduced
0.03 (1-Sp)
0.5

8.7
Yellow chromosol
(shallow)
Bundaberg
0.75

9.6
Red dermosol
(deep)
Gargett
0.67

Climate
Fraction of maximumminimum
saturation vapour for VPD estimation

population density of 10 stalks m2 without water stress (Bell and

Garside, 2005), early stalk senescence was invoked in half of the
simulations by reducing the stalk population by 0.5% per day when
leaf area index (LAI) declined to a low value of 0.5 after having
reached a value of 2.0. The loss of stalks then further reduced leaf
area and the accumulated amount of biomass but not radiation use
efciency.
2.6. Transpiration efciency (TE)
In the APSIM model, daily transpiration (T) is proportional to
daily biomass accumulation (W) multiplied by mean daily vapour
pressure decit (VPD).
T = W

VPD
k

(1)

where k is an empirical constant equal to transpiration efciency

(TE) at a VPD of 1 kPa (Keating et al., 1999; Sinclair et al., 2005).
Sinclair et al. (2005) called k the intrinsic water use efciency coefcient, a phrase we adopted here. Keating et al. (1999) reported
a k of 8.0 g kPa kg1 when simulating a number of growth analysis experiments mentioned above, mainly with Q117. However
Inman-Bamber and McGlinchey (2003) obtained 8.7 g kPa kg1 for
k from Bowen ratio energy balance and biomass accumulation work
on one sugarcane cultivar (Q127). It is reasonable to assume that a
10% variation in k could be found in the sugarcane germplasm similar to the variation implied for sorghum by Hammer et al. (1997).
In our study TE values of 8.7 and 9.6 g kPa kg1 were compared
(Table 2). This is a smaller range than the one used by Hammer
et al. (2005) (8 to 10 g kPa kg1 ) for a similar simulation exercise
with sorghum.
TE dened as W/T depends on the diurnal variation in VPD
and T particularly in drying soil conditions when partial stomatal
closure limits T during periods of high VPD. Meinzer and Grantz
(1990) presented evidence indicating that T in sugarcane is limited
by coordinated root and stomatal conductance which reached a
maximum of about 1.4 mmol s1 per plant with 0.8 m2 leaf area.
Thus maximum transpiration rates may be limited to 0.6 mm h1
using the maximum conductance of Meinzer and Grantz (1990) and
a leaf area index of 5 m2 m2 (Muchow et al., 1994; Inman-Bamber,
1994). Under water limited conditions or under high VPD (or both)
stomata start to close further limiting T and W (CO2 assimilation)
during the time of day when evaporative demand and VPD are high
(Inman-Bamber et al., 1986). Mean daily TE as dened will increase
as relatively more transpiration occurs towards the morning and
evening when VPD is low (Sinclair et al., 2005).
The effect on effective VPD by limiting T during the day was
analysed using hourly records of relative humidity and radiation
obtained from an automatic weather station at BSES. Eq. (1) was
used to derive T on an hourly basis assuming 1.8 g MJ1 for radiation use efciency (Keating et al., 1999) and 8.7 g kPa kg1 for k. The
long-term mean VPD was obtained as the sum of hourly T VPD

Level 3

Level 4

Level 5

Charters Towers

Pongola

Deep

Farleigh

divided by total daily T with T either not limited at all or limited

to 0.5 mm h1 (Sinclair et al., 2005). In APSIM-Sugarcane, VPD is
derived as a fraction of the difference in saturation vapour pressure at minimum and maximum daily temperatures. The default
fraction is 0.75 and this was reduced by 11% (to 0.67), which is the
% reduction in long term VPD when T was limited to 0.5 mm h1 .
It was assumed that reduced conductance could lead to reduced
effective VPD which would increase TE.
Thus TE was varied in two ways, rstly by a 10% increase in the
default intrinsic water use efciency (k) and secondly by combining
reduced conductance and reduced VPD in the APSIM model.
2.7. Climate
Three distinct climatic regions of the sugarcane industry in
Australia (southern, central and dry tropics) were selected from
the range of conditions where drought resistance traits could be of
benet. The SILO database operated by Queensland Climate Change
Centre of Excellence (http://www.longpaddock.qld.gov.au/silo/)
provided daily radiation, rainfall, class A-pan evaporation, and
maximum and minimum temperatures from 1960 to 2009 for the
BSES Ltd experimental farms near Bundaberg (southern region),
Farleigh Cooperative Sugar Mill and the Gargett Post Ofce in the
Mackay region (central region), and for Charters Towers (Towers)
in the dry tropics. The dry region of Pongola in South Africa was
also considered in order to broaden the study. Climate data for the
Pongola region was obtained from the South African Sugarcane
Research Institute through their web-enabled weather network
(http://portal.sasa.org.za/weatherweb).
Although mean annual rainfall (MAR) for Mackay is reasonably
high (1647 mm), drought has inicted severe yield losses of more
than $400 million AUD in the years 20022004 (Inman-Bamber,
2007). The Bundaberg/Maryborough region has one of the lowest
annual rainfalls (1022 mm for BSES Bundaberg) in the Australian
sugarcane belt. Despite the availability of modest amounts of irrigation water, the region suffers more from drought than any other
production area in Australia (Inman-Bamber, 2007). Sugarcane is
not grown commercially at Charters Towers with only 655 mm
MAR and no irrigation, but this region may represent the type of
climate that might be experienced with climate change with a possible 40% reduction in rainfall by 2070 (Park and Attard, 2005) or
the type of climate where sugarcane may be grown for biofuel to
help reduce CO2 emissions in future (Waclawovsky et al., 2010).
Pongola in South Africa has a similar MAR (648 mm) but irrigation
is available to support commercial sugarcane cultivation.
2.8. Other model settings
A typical crop cycle was simulated using the APSIM sugarcane
module. This consisted of a crop planted in April and harvested
in July followed by four 13-month-old ratoon crops. The nal stalk
population was 10, 10, 9, 8 and 7 stalks m2 for each successive crop

N.G. Inman-Bamber et al. / Field Crops Research 134 (2012) 95104

Fig. 1. Measured leaf area index for wet ( ) and dry ( ) treatments (bars denote
2 SEM). Simulated leaf area index for wet () and dry treatments (- - - -) using
),
standard senescence settings and using a high leaf senescence rate for wet (
).
and dry treatments (

in the planting and ratooning cycle to recognise, at least to some

extent, the loss of vigour in later ratoons. The soil was assumed to be
fallowed between harvesting the fourth ratoon crop in November
and replanting in April. This cycle was repeated from 1960 to 2009
for the Australian sites and between 1967 and 2009 for the South
African site. Crop residue was left on the soil surface to simulate
a green cane trash blanket which is a common practice in the
Mackay and Bundaberg regions. The residue, water balance and
nitrogen balances were continued (not reset) for the entire simulation period. An irrigation of 40 mm was applied to help with crop
germination and to acknowledge the likely and logical choice of
farmers to plant into moist soil. The time limit for germination was
removed but not the thermal time or the cumulative stress limits
which can cause the crop to fail. If these limits were exceeded, the
crop was harvested and the ground then lay fallow till the next
opportunity for planting in April. APSIM crop modules restrict root
penetration when the soil is dry but we removed this restriction
based on our unpublished observations of root penetration into
extremely dry soils and saprolite. A total of 640 combinations of
the various traits and treatments (Table 2) and a total of 9382 crops
were simulated.
3. Results
3.1. Simulation of eld experiments
The simulation of LAI (Fig. 1) and biomass (Fig. 2) illustrate the
effect of varying levels of sensitivity to water stress in relation
to leaf senescence. Using standard APSIM settings, simulated LAI
towards the end of the crop was low for experiment 1 and high
for experiment 2 compared to observed LAI but the simulated difference between the wet and dry treatments was realistic for both
experiments. Measured LAI of the dry treatment in experiment 2
was bounded by simulations with standard settings and with the

99

Fig. 3. Average total monthly rainfall (bars), mean monthly temperature (lines) and
line,
symbol), Farleigh
radiation (symbols) for Bundaberg (solid bars,
, ), Gargett (checked bars,
, ), Pongola (striped
(diagonal bars,
bars,
, ) and Charters Towers (open bars,
, ) from 1960 to 2009.

higher level of leaf senescence, indicating that the simulated variation in leaf senescence was realistic. In the simulation, conditions
for stalk senescence causing loss of leaf area did not occur even
with higher leaf senescence trait. In the eld experiments the water
stress treatments imposed were not sufciently severe to result in
stalk death (Inman-Bamber, 2004).
Simulated biomass was generally too high for experiment 1 and
too low for experiment 2, although nal estimates of biomass were
close to measured biomass in the case of the wet treatment in both
experiments (Fig. 2). High sensitivity of leaf senescence to water
stress had only a small negative effect on biomass accumulation
in both experiments because variations in LAI above a value of 2
have a diminishing effect on radiation interception which is 53, 68
and 78% with LAI values of 2, 3 and 4, respectively (based on an
extinction coefcient of 0.38; Keating et al., 1999).
Comparing measured and simulated responses to water stress
for the two experiments, indicated that the drought avoidance
mechanism of rapid leaf senescence is potentially available while
the trait for early loss of stalks may not be immediately available in
the local germplasm if it turns out to be a useful mechanism. However, Smit and Singels (2006) found that one South African variety
(N22) was considerably more susceptible to water stress in regard
to stalk senescence than another (NCo376).
3.2. Climate of target sites
Mean monthly rainfall totals for JanuaryApril were 1.52.5
times higher at Farleigh and Gargett in the Mackay region than
at the other sites (Fig. 3) and exceeded monthly evaporation totals
(data not shown). In all other cases, rainfall totals were lower than
evaporation totals for all months and sites. Water stress is likely
to be severe in spring particularly in Charters Towers where radiation, temperature and evaporation are both high and rainfall is
low. Spring temperatures were considerably lower at Pongola than
at the other sites and rainfall was relatively high so water stress at
this time of year could be less severe at Pongola than at the other
sites.
3.3. Biomass yields

Fig. 2. Measured biomass for wet () and dry () treatments (bars denote 2 SEM).
Simulated biomass for wet () and dry treatments (- - - -) using a standard leaf senes), and dry treatments
cence rate and using a high leaf senescence for wet (
).
(

The range in biomass yields was large for all sites (Fig. 4). Simulated dry biomass yields could be less than 10 t ha1 in the most
favourable site, Farleigh, and greater than 30 t ha1 in the least
favourable site, Towers. Although MAR was similar for Towers and
Pongola, yields were greater at Pongola because of the better distribution of rain through the year and the lower radiation and
temperature resulting in less water stress. Mean biomass yield for
Towers, Pongola, Bundaberg, Gargett and Farleigh was 11, 13, 30,
34 and 44 t ha1 respectively.

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N.G. Inman-Bamber et al. / Field Crops Research 134 (2012) 95104

15.2 0.3, 19.2 0.2, 19.1 0.2 and 18.9 0.2%, respectively and
for the deep soil the mean benet was 5.4 0.6, 4.2 0.3, 9.2 0.4,
13.7 0.4 and 13.7 0.4% for these sites, respectively.

1.0
0.9

Fraction of Data

0.8

3.5. Increased leaf senescence rate

0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
0

30

60

Biomass yield (t

90

3.6. Early stalk senescence

ha-1)

Fig. 4. Cumulative frequency distribution of biomass yield for all treatments for
), Pongola (
), Bundaberg (
),
ve climates Charters Towers (
) and Farleigh (
).
Gargett (

3.4. Increased rooting depth

Increased rooting depth was benecial to biomass yield in nearly
all situations (all years and all treatments other than climates and
soils) simulated with properties of a poor soil type (Yellow chromosol) (Fig. 5a). A deep and more vigorous rooting trait was not
as benecial in the good soil because the increase in soil water
exploited was not as great (10% compared to 22% for the poor soil).
More vigorous roots could also be a disadvantage in some circumstances where they encourage greater exploitation of water stored
in the soil leaving little or none available for the subsequent ratoon
crop. Deeper roots in the Red dermosol were more benecial for
the wetter sites Gargett and Farleigh because there was more additional water to exploit. For this soil at Gargett and Farleigh, deeper
roots increased biomass yield in 80% and 90% of all crops simulated respectively while in the other sites only 50% of the crops
benetted from this trait (Fig. 5b). The average yield increase from
a deeper and more vigorous root system in the shallow soil for
Towers, Pongola, Bundaberg, Gargett and Farleigh was 18.0 0.5,

The biomass yield response to increased leaf senescence rate

was mostly negative with yield losses of up to 40% for both the poor
and good quality soil (Fig. 6). Rapid leaf senescence was of benet
in less than 10% of situations. The negative affects were greater in
the shallow than in the deeper soil because of the limited storage
capacity of the shallow soil and the more frequent loss of leaf area.
In the deeper soil the negative effects were less serious in the wetter
climates than in the drier climates because leaf senescence was less
prevalent there. Generally this strategy failed to save enough water
to offset yield loss through reduced radiation capture except in a
very limited number of situations.

Stalk senescence initiated in about one third of all simulations

for both soils but in many cases this had little effect on biomass
yield because stalk senescence occurred shortly before harvesting,
or it occurred in situations where there was little recovery in LAI
regardless whether stalks senesced or not. The loss of stalks when
LAI declined to 0.5 through water stress was generally unsuccessful
in helping crops get through dry conditions (Fig. 7) despite the stalk
population being fully renewed at the start of each crop. A small
proportion (<15%) of crops in the Towers and Pongola climates in
the Red dermomsol beneted from stalk senescence because of the
reduced water use by a reduced crop canopy allowing water to be
available during a more critical period of growth.
Farmers could reduce sugarcane stands in dry regions by planting rows further apart but these simulations indicate that reducing
stalk population through genetic or management manipulation will
not improve average biomass yields even in very dry climates.
3.7. Reduced conductance
The mean response to reduced conductance was positive only
for the Red dermosol in the driest sites, Towers (+7.3 1.0%) and
Pongola (+3.4 0.5%) where the yield in some crops was greatly
improved by having water conserved during dry periods (Fig. 8).
In other situations yields were reduced by this trait because the

Yellow chromosol

Red dermosol

1.0

Fraction of Data

0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
0

50

100

150

200 0

50

100

150

200

Biomass yield response to deeper roots (%)

Fig. 5. Cumulative frequency distribution of biomass yield response to increased root depth for two soils and ve climates, Towers (
), Gargett (
) and Farleigh (
).
(

), Pongola (

), Bundaberg

N.G. Inman-Bamber et al. / Field Crops Research 134 (2012) 95104

Yellow chromosol

101

Red dermosol

1.0

Fraction of Data

0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
60

70

80

90

100

110 120 60

70

80

90

100 110

120

Biomass yield response to rapid leaf senescence (%)

), Pongola (

Fig. 6. Cumulative frequency distribution of biomass yield response to rapid leaf senescence for two soils and ve climates, Towers (
), Gargett (
) and Farleigh (
).
(

Yellow chromosol

), Bundaberg

Red dermosol

1.0

Fraction of Data

0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
50

70

90

110

130

150 50

70

90

110

130

150

Biomass yield response to early stalk senescence (%)

Fig. 7. Cumulative frequency distribution of biomass yield response to early stalk senescence for two soils and ve climates, Towers (
), Gargett (
) and Farleigh (
).
Bundaberg (

Yellow chromosol

), Pongola (

),

), Pongola (

),

Red dermosol

1.0

Fraction of Data

0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
50

100

150

200

250 50

100

150

200

250

Biomass yield response to reduced conductance (%)

Fig. 8. Cumulative frequency distribution of biomass yield response to reduced conductance for two soils and ve climates, Towers (
), Gargett (
) and Farleigh (
).
Bundaberg (

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N.G. Inman-Bamber et al. / Field Crops Research 134 (2012) 95104

Yellow chromosol

Red dermosol

1.0
0.9

Fraction of Data

0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
60

80

100

120

140

160 60

80

100

120

140

160

Biomass yield response to increased intrinsic water use efficiency (k) (%)
Fig. 9. Cumulative frequency distribution of biomass yield response to increased transpiration efciency combined with reduced conductance for two soils and ve climates,
), Pongola (
), Bundaberg (
), Gargett (
) and Farleigh (
).
Towers (

conservation of water within the life of the crop and for the next
crop was mostly insufcient to offset the loss of yield through
reduced CO2 assimilation. For the wetter climates of Bundaberg,
Gargett and Farleigh reduced conductance was less detrimental
(1.9 0.2%) in the deep than the shallow soil (14.7 0.2%). This
was because of the greater amount of water that could be conserved
in the deeper soil for later use, either within the crops duration or
for the next ratoon crop.
3.8. Increased TE through increased intrinsic water use efciency
(k)
Increased k was benecial in more than 95% of all cases in
all climates and both soils (Fig. 9). The yield increase was more
than 17% for 1 out of 10 simulated crops at Bundaberg and Pongola (Fig. 9). This is greater than the 10% increase in k because
of ow on effects of reduced water stress and more rapid canopy
development. Mean biomass yield increased most in the Pongola
climate (11.1 0.15% for the shallow soil and 13.7 0.16% for the
deep soil) and least in the wetter Farleigh climate (9.2 0.15% for
the shallow soil and 8.4 0.15% for the deep soil) (Fig. 9). The
response to increased k was intermediate in the dry Towers climate. Increased k is of no benet when water is either non-limiting
for photosynthesis and transpiration or it is totally limiting. These

extremes were more prevalent at Towers than at Pongola. In a very

limited number of cases increased k was detrimental for biomass
yield because preceding crops used more water with up-regulated
k, leaving subsequent crops with less water to start with. Mean
whole crop TE (biomass yield/total transpiration) was 6.18 g kg1
for k = 8.7 g kPa kg1 and 6.82 g kg1 (10% greater) for up-regulated
k = 9.6 g kPa kg1 . Variation around a 10% mean response in biomass
yield to a 10% up-regulation in TE was due to periods when water
was not limiting or when leaf area development was enhanced or
senescence was reduced by more favourable water relations with
up-regulated k.
3.9. Increased TE through decreased VPD combined with reduced
conductance
Mean whole crop TE was 6.13 g kg1 with the default estimate of VPD and 6.87 g kg1 (12% greater) when VPD was reduced
effectively by reduced conductance. Although whole crop TE was
increased more by reduced effective VPD than by up-regulated k,
the yield benets were equivocal (Fig. 10). The benet was greater
for the deep than for the shallow soil, due to the greater storage
capacity of the deeper soil where reduced conductance could be of
more benet than in the shallow soil in conserving water for use in
subsequent dry periods as explained above.

Fig. 10. Cumulative frequency distribution of biomass yield response to reduced VPD (hence increased transpiration efciency) combined with reduced conductance for two
), Pongola (
), Bundaberg (
), Gargett (
) and Farleigh (
).
soils and ve climates, Towers (

N.G. Inman-Bamber et al. / Field Crops Research 134 (2012) 95104

Yield benets were substantially greater in the drier climates

(Pongola, Towers) than the wetter climates (Farleigh and Gargett)
because of the combined effects of benets from reduced conductance at Pongola and Towers (Fig. 8) and increased TE at Pongola
and to a lesser extent at Towers (Fig. 9). Crops in the Pongola climate and on deep soils would nearly always benet from reduced
conductance associated with reduced effective VPD while 9 out of
10 crops at Farleigh in shallow soils would be worse off with this
trait.
The mean response in biomass yield to increased TE and reduced
conductance in the deep soil for Towers, Pongola, Bundaberg, Gargett and Farleigh climates was 13.6 0.7, 16.7 0.4, 13.7 0.4,
9.3 0.4 and 4.8 0.3%, respectively and the mean response for
the shallow soil in these climates was 8.0 0.6, 3.8 0.5, 0.8 0.5,
5.2 0.4 and 8.9 0.4%, respectively.

4. Discussion
In a review of a range of plant traits associated with droughttolerance, Ludlow and Muchow (1990) recommend eight traits that
are desirable for intermittent stress conditions in modern agriculture. The top three of these traits were plant phenology, osmotic
adjustment, and rooting depth. The reproductive phase is generally avoided in sugarcane production through crop management
and altering plant phenology to achieve better conditions for this
phase was not of interest in this study. Instead the traits identied
as potentially useful for improving drought tolerance in sugarcane
included rooting depth, rate of leaf senescence, trigger for stalk
senescence, hydraulic conductance and transpiration efciency.
At two of the climatic regions of this study (Towers and Pongola)
irrigation is mandatory for a viable sugarcane industry. Biomass
yields simulated for these regions illustrate this point with means
yields of less than 15 t ha1 at Towers and Pongola and 30 t ha1
or more at the other sites (Fig. 4). Irrigation was introduced relatively recently (in the past 20 years) in the Bundaberg and Mackay
regions but now growers believe their enterprises would not be
viable without irrigation. In this study the only irrigation allowed
was 40 mm at the time of planting the crop at the start of a seven
year cropping cycle. Simulated biomass yields at Bundaberg and
Gargett were less than 20 t ha1 (60 t cane ha1 ) for one out of
three crops (Fig. 4) thus supporting the need for irrigation in these
regions.
A root system capable of exploring an additional 10% of water
available in the root zone as well as extending the rootzone by
300800 mm (one additional soil layer) would be of considerable
benet (021% mean and 200% maximum yield increase in our
study) particularly for poor soils and dry climates. Of course this
benet can only be realised if there is sufcient rain periodically to
wet the soil to the additional depth. Rainfall during summer (Fig. 3)
is normally adequate for this purpose and the APSIM model used
in our study accounted for seasons when soil at depth was not fully
replenished with water. Smith et al. (2005) provided evidence for
root water extraction at a depth of 2.8 m in Australia. Battie Laclau
and Laclau (2009) found surprisingly little difference in the rate of
advance and the nal rooting depth between irrigated and rainfed
sugarcane in Brazil. The root front advanced about 5 mm d1 for the
rst four months and then about 18 mm d1 thereafter. Roots were
found at a depth of 4.7 m for the rainfed crop and at 4.2 m for the irrigated crop suggesting that rooting depth is not greatly inuenced
by water regime. However the rainfall during the 10 month study
period was not particularly low (940 mm) and only 1140 mm water
was required for the irrigated crop. Baran et al. (1974) also found
that roots were more deeply distributed when irrigation frequency
was reduced in sugarcane even though the rooting depth was similar between treatments. APSIM-Sugarcane makes no allowance for

103

an increase in root proliferation under moderately dry conditions

such as in the Brazilian experiment where withholding irrigation
resulted in 49% more intersects between roots in the vertical grid
used in the study of Battie Laclau and Laclau (2009). If root depth
penetration is constrained in dry soil then the benets of a more
vigorous root system as well as the benets of a deep soil would be
substantially diminished. However sugarcane is a perennial crop
and only part of the root system dies back after harvesting and
deep roots from the harvested crop can be critical for the survival
of subsequent ratoon crops (Smith et al., 2005).
The strategy underlying an increase in leaf senescence rate in
our study was to reduce water use early in response to drought in
order to conserve water for periods of water stress later on. The simulations suggest that regardless of soil type, climate or ratoon date
there were very few years when this strategy benetted biomass
accumulation and it is questionable whether reduced leaf area is
of any benet to sugarcane production at all, given the extremely
dry climates used in this study. Rather, scarce resources should be
targeted at strategies to build and maintain leaf area.
In the majority of years biomass yield did not respond to a trait
for stalk senescence triggered when LAI declined below 0.5. We
have little evidence as to what kind of water stress triggers stalk
death. Stress imposed at different stages of crop growth reduced
biomass accumulation by as much as 70% and this was not sufcient to reduce the stalk population in the cultivars Q96 and Q117
(Inman-Bamber, 2004).
Breeding for reduced stomatal or root conductance is unlikely
to provide benets in terms of increased crop yield in shallow soils.
However, in deep soils, where there is a greater potential for storing
water through reduced conductance this trait will be benecial but
could be applied only to a limited number of climates and soils in
the sugarcane industry.
The observations and interpretations on root depth, leaf and
stalk senescence and reduced conductance (without increased TE)
align with the analysis by Blum (2009) which suggests that traits
which conserve water and increase water use efciency are generally unsuccessful for improving economic yield in production
environments which may be water-limited but are generally more
favourable for plant survival compared to environments were survival rather than production is the key issue.
Our simulations indicated that increased TE at the leaf level
would nearly always help to improve sugarcane biomass yields
in water-limited environments if the increased TE arose from upregulation of intrinsic water use efciency (k). However if increased
TE was increased through reduced conductance, which effectively
reduces VPD during transpiration, the yield benets are not at all
certain. A large water storage (deep soil) would be required to take
advantage of water savings from reduced conductance. A moderate
climate such as the Pongola climate would also be required to make
most of the increased TE which is most benecial when root water
supply is limiting but not close to zero and not excessive.
TE or rather its surrogate, delta, which is derived from the ratio
of 13 C to 12 C captured during assimilation of CO2 by C3 plants,
is not always associated with yield (Blum, 2009). 13 C discrimination for measuring TE may not be suitable for sugarcane with C4
photosynthesis because of the small contribution of rubsico to the
assimilation process (Ranjith et al., 1995). However the potential
benets of improved TE needs to be explored more fully using
a model capable of integrating diurnal variation in VPD, stomatal conductance, transpiration and photosynthesis as in the work
of Sinclair et al. (2005). The APSIM model chosen for our study
was useful for considering a number of traits that may be useful in water limited environments however the daily time step of
APSIM allowed only an indirect link between reduced conductance
and increased TE. TE may also be increased through an increase
in intrinsic water efciency (k) and the APSIM modelling was

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N.G. Inman-Bamber et al. / Field Crops Research 134 (2012) 95104

adequate for this possibility. The 12% increase in whole crop TE due
to reduced conductance was less than the increase in TE derived
theoretically by Sinclair et al. (2005) who limited transpiration
at midday assuming partial stomatal closure when demand for
water was high. Transpiration and photosynthesis measurements
on soybean by Rawson et al. (1978) indicated that TE was highest in the morning and evening and lowest between 12 and 14 h
because of partial stomatal closure, increased leaf temperature and
VPD. Water stress further reduced TE during the middle of the day
because of further increases in leaf temperature and VPD. Diurnal
interactions between transpiration, photosynthesis, VPD and water
stress have not been studied in sugarcane and our study highlights
the need to do this in order to better understand and model the
proposed benets of increased TE.
Acknowledgements
This research was funded by the Australian Federal Government
and Sugar Industry through the Sugar Research and Development
Cooperation. We thank the Queensland Department of Natural
Resources and Water and the South African Sugarcane Research
Institute for providing the climate data used in this study.
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