Agricultural Sciences in China

2009, 8(1): 51-58 January 2009

Proline Accumulation, Photosynthetic Abilities and Growth Characters of

Sugarcane (Saccharum officinarum L.) Plantlets in Response to Iso-Osmotic
Salt and Water-Deficit Stress

Suriyan Cha-um and Chalermpol Kirdmanee

National Center for Genetic Engineering and Biotechnology (BIOTEC), National Science and Technology Development Agency (NSTDA),
Pathumthani 12120, Thailand

Abstract
The aim of this study was to investigate the biochemical, physiological and morphological responses of sugarcane to iso-
osmotic salt and water-deficit stress. Disease-free sugarcane plantlets derived from meristem cuttings were photo-
autotrophically grown in MS media and subsequently exposed to -0.23 (control), -0.67 or -1.20 MPa iso-osmotic NaCl (salt
stress) or mannitol (water-deficit stress). Chlorophyll a (Chl a), chlorophyll b (Chl b), total carotenoids (Cx+c), maximum
quantum yield of PSII (Fv/Fm), photon yield of PSII ( PSII), stomatal conductance (Gs) and transpiration rate (E) in the
stressed plantlets were significantly reduced when compared to those of plantlets of the control group (without mannitol
or NaCl), leading to net-photosynthetic rate (Pn) and growth reduction with positive correlation. In addition, physiological
changes and growth parameters of plantlets in the salt stress conditions were more sharply reduced than those in water-
deficit stress conditions. On the other hand, the proline content and non-photochemical quenching (NPQ) in the leaves
of stressed plantlets increased significantly, especially in response to iso-osmotic salt stress. The chlorophyll pigments
in iso-osmotic stressed leaves were significantly degraded (r2 = 0.93), related to low water oxidation (r2 = 0.87), low net-
photosynthetic rate (r2 = 0.81), and growth reduction (r2 = 0.97). The multivariate biochemical, physiological and growth
parameters in the present study should be further used to develop salt, or drought, tolerance indices in sugarcane
breeding programs.

Key words: growth performances, net-photosynthetic rate, pigment degradation, proline, water oxidation

Das 2005). In general, osmotic stresses caused by

INTRODUCTION
Abiotic stresses, especially water-deficit and soil salinity,
are major problems, reducing crop productivity by more
than 50% on agricultural land world-wide (Mahajan and
Tuteja 2005). Both water-deficit and salt-stresses det-
rimentally affect plant growth and developmental
processes, which have been reported in terms of
biochemical, physiological and morphological changes
(Hasegawa et al. 2000; Wang et al. 2001; Parida and
both soil salinity and water-deficit are well established
in halophyte (Pujol et al. 2000; Slama et al. 2007; Pagter
et al. 2009) and glycophyte species (Lutts et al. 2004;
Wahid 2004; Luo et al. 2005). Also, ionic toxicity gen-
erated from salt contaminated soil has negative effects
on plant growth and development (Tester and Daven-
port 2003; Davenport et al. 2005; Munns et al. 2006).
However, there are many defense mechanisms in plants
which are tolerant to water-deficit and salt stresses,
such as osmoregulation, ion homeostasis, antioxidant

Received 24 August, 2008 Accepted 16 December, 2008

Correspondence Suriyan Cha-um, Ph D, Tel: +66-2-564 6700, Fax: +66-2-564 6707, E-mail: suriyanc@biotec.or.th

© 2009 CAAS. All rights reserved. Published by Elsevier Ltd.

52
and hormonal systems (Hasegawa et al. 2000; Wang
et al. 2003; Reddy et al. 2004; Sairam and Tyagi 2004;
Mahajan and Tuteja 2005), helping plants to survive
and grow under severe environmental conditions prior
to their reproductive stages. In contrast, the defense
mechanisms in sensitive plant species are weaker, lead-
ing to growth retardation and yield reduction.
Plant biochemicals [ascorbate peroxidase (AOX),
glutamine synthetase (GS), proline, glycine betaine, pho-
tosynthetic pigments, soluble proteins and mineral
elements] and physiological changes [relative water
content (RWC), stomatal conductance (Gs), water po-
tential ( w), osmotic potential ( s), chlorophyll a
fluorescence, and net-photosynthetic rate (Pn)] in plants
growing under salt or water-deficit conditions have been
investigated in many plant species such as rice (Cha-
um et al. 2007; Castillo et al. 2007), cabbages (Maggio
et al. 2005), salt marsh grasses (Maricle et al. 2007),
maize (Hu et al. 2007; Wang et al. 2008), potatoes
(Teixeira and Pereira 2007), and Argyranthemum
coronopifolium (de Herralde et al. 1998). Biochemical
and physiological parameters in higher plants cultivated
in salt or water-deficit conditions have been developed
as effective indices for tolerant screening in plant breed-
ing programs (Ashraf and Harris 2004; Parida and Das
2005; Ashraf and Foolad 2007). Polyethylene glycol
(PEG), mannitol and sorbitol sugar alcohols are the major
chemical formulae which are added to the media or
nutrient solutions in order to control osmotic potential
and replicate water-deficit conditions. For salt stress,
NaCl, Na2SO4, MgSO4, and MgCl2 salts are generally
used. In the present study, mannitol and NaCl were
selected to induce water deficit and saline stress,
respectively, to adjust the iso-osmotic pressures in the
culture media to -0.67 and -1.20 MPa.
Sugarcane (Saccharum officinarum L.), belonging
to the Poaceae family, is a sugar producing plant species,
which grows well in tropical and subtropical regions.
Sugarcane is a high biomass produce, consuming large
amounts of water and nutrients from the soil for maxi-
mum productivity. Water irrigation management is an
important factor in sugarcane cultivation, especially in
arid and semi-arid zones. Moreover, sugarcane is a
glycophyte species, reported to be salt susceptible,
which is demonstrated by toxicity symptoms, low sprout
emergence, nutritional imbalance and overall growth
Suriyan Cha-um et al.
reduction, leading to low biomass (Wahid et al. 1997;
Plaut et al. 2000; Akhtar et al. 2003). The objective of
this investigation was to identify the physiological
changes and the growth parameters of sugarcane in
response to iso-osmotic salt, or water-deficit stresses.
MATERIALS AND METHODS
Plant materials
Disease-free sugarcane shoots (S. officinarum L. cv.
K84-200) derived from meristem cuttings were propa-
gated in MS media containing 8.88 μM benzyl adenine
(BA), 3% sucrose and 0.25% Phytagel® for 6 weeks.
Multiple shoots were elongated in the MS media with-
out plant growth regulators for 4 weeks, then, single
shoots were excised and roots induced in MS media
supplemented with 2.46 μM indole butyric acid (IBA),
3% sucrose and 0.25% Phytagel® for 2 weeks. Plant-
lets were cultured in vitro under conditions of (25 ± 2)°C
ambient temperature, (60 ± 5)% relative humidity (RH)
and (60 ± 5) μmol m-2 s-1 photosynthetic proton flux
density (PPFD), provided by fluorescent lamps with a
16 h d -1 photoperiod. Then, the sugarcane plantlets
were transferred to MS sugar-free liquid media
(photoautotrophic conditions) using vermiculite as sup-
porting material for 7 days. The number of air-ex-
changes in the glass vessels was adjusted to 2.32 h-1 by
punching a hole in the plastic cap (Ø 1 cm) and cover-
ing the hole with a micro-porous filter. The plantlets
were subsequently cultured in a plant growth incubator
with the same conditions as previously mentioned and
CO2 enrichment at (1 000 ± 100) μmol mol-1. Sodium
chloride (salt stress) and mannitol (water-deficit stress)
in the culture media were adjusted to -0.23 (control),
-0.67 or -1.20 MPa iso-osmotic pressures for 7 days.
Photosynthetic pigments, proline contents, chlorophyll
a fluorescence, net-photosynthetic rate (Pn) and growth
characters were measured for physiological and bio-
chemical analysis.
Data measurement
Chlorophyll a (Chl a), chlorophyll b (Chl b), total
chlorophyll, and total carotenoid (Cx+c) concentrations
© 2009, CAAS. All rights reserved. Published by Elsevier Ltd.
Proline Accumulation, Photosynthetic Abilities and Growth Characters of Sugarcane (Saccharum officinarum L.)
were analyzed following the methods of Shabala et al.
(1998) and Lichtenthaler (1987), respectively. One hun-
dred milligrams of leaf material were collected from
the second and third nodes of the shoot tip. The leaf
samples were placed in a 25 mL glass vial, along with
10 mL of 95.5% acetone, and blended using a
homogenizer. The glass vials were sealed with parafilm
to prevent evaporation and then stored at 4°C for 48 h.
The Chl a and Chl b concentrations were measured
using a UV-visible spectrophotometer at 662 nm and
644 nm wavelengths. The Cx+c concentration was also
measured by spectrophotometer at 470 nm. A solution
of 95.5% acetone was used as a blank.
The proline content of the leaves was extracted and
analyzed according to the method of Bates et al. (1973).
Fifty milligrams of fresh weight material were ground
with liquid nitrogen in a mortar. The homogenate powder
was mixed with 1 mL aqueous sulfosalicylic acid (3%
w/v) and filtered through filter paper (Whatman #1,
England). The extracted solution was reacted with an
equal volume of glacial acetic acid and ninhydrin re-
agent (1.25 mg ninhydrin in 30 mL of glacial acetic
acid and 20 mL 6 M H3PO4) and incubated at 95°C for
1 h. The reaction was terminated by placing the con-
tainer in an ice bath. The reaction mixture was vigor-
ously mixed with 2 mL toluene. After warming at 25°C,
the chromophore was measured by Spectrophotom-
eter DR/4000 at 520 nm using L-proline as a standard.
Chlorophyll a fluorescence emission from the adaxial
surface on the third leaf from the shoot tip was moni-
tored with a Fluorescence Monitoring System in the
pulse amplitude modulation mode, as previously de-
scribed by Loggini et al. (1999). A leaf, adapted to
dark conditions for 30 min using leaf-clips, was ini-
tially exposed to the modulated measuring beam of far-
red light (LED source with typical peak at wavelength
735 nm). Original (Fo) and maximum (Fm) fluores-
cence yields were measured under weak modulated red
light (< 0.5 μmol m-2 s-1) with 1.6 s pulses of saturating
light (> 6.8 μmol m-2 s-1 PAR) and autocalculated using
FMS software for Windows®. The variable fluores-
cence yield (Fv) was calculated by the equation of Fm-
Fo. The ratio of variable to maximum fluorescence
(Fv/Fm) was calculated as maximum quantum yield
of PSII photochemistry. The photon yield of PSII
( PSII) in the light was calculated by PSII = (Fm´ -
53
F)/Fm´ after 45 s of illumination, when steady state
was achieved. In addition, non-photochemical quench-
ing (NPQ) was calculated as described by Maxwell and
Johnson (2000).
The net-photosynthetic rate (Pn), transpiration rate
(E; mmol m-2 s-1) and stomata conductance (Gs; mmol
H2O m-2 s-1) of sugarcane plantlets were measured on
the leaf using an infra-red gas analyser. The E and Gs
were measured continuously by monitoring H2O of the
air entering, and existing in, the IRGA headspace
chamber. The flow-rate of air in the sample line was
adjusted to 500 μmol s -1. The micro-chamber tem-
perature was set at 25°C. The light intensity was fluxed
by 6400-02B red-blue LED light source at 1 000 μmol
m-2 s-1 PPFD (Cha-um et al. 2007).
Fresh weight, dry weight, shoot height, root length
and leaf area of sugarcane plantlets were measured as
described by Cha-um et al. (2006). Sugarcane plant-
lets were dried at 110°C in a hot-air oven for 2 days,
and then incubated in desiccators before the measure-
ment of dry weight. The leaf area of sugarcane plant-
lets was measured using a leaf area meter DT-scan.
Experiment design
The experiment was designed as completely random-
ized design (CRD) with ten replicates and four plantlets
per replicate. The mean values obtained were com-
pared by Duncan’s new multiple range test (DMRT)
and analyzed using SPSS software. The correlations
between physiological and biochemical parameters were
evaluated with Pearson’s correlation coefficients.
RESULTS
Photosynthetic pigments, including chlorophyll a (Chl a),
chlorophyll b (Chl b), total chlorophyll (TC) and total
carotenoid (Cx+c) in the osmotically stressed leaves of
sugarcane plantlets were sharply reduced, related to
the decrease in osmotic pressure in the culture media
(Table 1). Pigment degradation in the leaf tissues of
stressed plantlets was a rapid indicator of plant responses
to osmotic stress and was inversely related to the os-
motic pressure in the culture media (r2 = 0.93) (Fig.1).
Chl a, Chl b, TC and Cx+c contents in the -1.20 MPa
© 2009, CAAS. All rights reserved. Published by Elsevier Ltd.
54 Suriyan Cha-um et al.

salt-stressed plantlets were significantly reduced by
1.84, 2.17, 1.91, and 2.37 times, respectively, when
compared to those of -1.20 MPa water-deficit stressed
plantlets (Table 1). In contrast, proline content in the
osmotic stressed-leaves was increased, positively re-
lating to the osmotic stress, especially salt induced os-
motic stress (Table 1). Proline was generally accumu-
lated in osmotically-stressed sugarcane plantlets and
played a key role in osmoregulation and antioxidant de-
fense mechanisms. Moreover, the total chlorophyll deg-
radation due to osmotic stresses was inversely related
to maximum quantum yield of PSII (Fv/Fm) (r2 = 0.87)
(Fig.2). The chlorophyll a fluorescence parameters,
Fv/Fm and photon yield of PSII ( PSII) in sugarcane
plantlets grown under -1.20 MPa salt-stress were sig-
nificantly reduced when compared to those of plantlets
of the control group (-0.23 MPa), while those param-
eters in plantlets grown in -1.20 MPa mannitol were
unchanged (Table 2). On the other hand, non-photo-
chemical quenching (NPQ) of osmotically stressed plant-
lets increased, especially in response to -1.20 MPa salt
stress. The reduction of Fv/Fm in response to osmotic
stress was positively correlated with net-photosynthetic
rate (Pn) (r2 = 0.81) (Fig.3). The Pn, stomatal conduc-
tance (Gs), and transpiration rate (E) in osmotically
stressed plantlets were sharply reduced when exposed
to both salt stress and water-deficit stress (Table 2).
The biochemical and physiological data were subjected
to analysis using SPSS software to determine the
Pearson’s correlation coefficients, which are shown
in Table 3. The Pn reduction in osmotically stressed
plantlets was positively related to biomass production,
which was represented by dry weight (DW) (r2 = 0.97)
(Fig.4). Fresh weight (FW), shoot height (SH), root
length (RL) and leaf area (LA) in osmotically stressed
plantlets were significantly reduced, relating to osmotic
pressure in the culture media and salt stress (Table 4).
Also, there was a positive correlation between the
growth parameters (Table 5). In a recent study, the
iso-osmotic salt stress strongly inhibited growth and
development in sugarcane when compared to water-
deficit stress.
DISCUSSION
Photosynthetic pigments in sugarcane plantlets exposed
to osmotic stress using NaCl salt and mannitol iso-os-

Table 1 Chlorophyll a (Chl a), chlorophyll b (Chl b), total chlorophyll (TC), total carotenoids (Cx+c) and proline contents of sugarcane
plantlets grown under iso-osmotic drought (mannitol) and salt (NaCl) stress for 7 days
Osmotic potential (MPa) Chl a (μg g-1 FW) Chl b (μg g-1 FW) TC (μg g-1 FW) Cx+c (μg g-1 FW) Proline (μg g -1 FW)
-0.23 (control) 371.46 a 154.63 a 526.10 a 71.15 a 318 d
-0.67 mannitol 271.42 b 88.02 b 359.44 b 62.32 ab 518 d
-0.67 NaCl 163.73 c 64.00 bc 227.74 c 37.92 bc 770 c
-1.20 mannitol 238.77 b 71.18 b 309.95 b 59.56 ab 1 027 b
-1.20 NaCl 129.68 c 32.73 c 162.41 c 21.94 c 1 412 a
ANOVA ** ** ** ** **

Different letters in each column show significant difference at P 0.01 ( ) by Duncan’s new multiple range test (DMRT).
**

Fig. 1 Relationship between osmotic potential in the culture media Fig. 2 Relationship between pigment degradation and maximum
and pigment degradation of sugarcane plantlets grown under iso- quantum yield of PSII (Fv/Fm) of sugarcane plantlets grown under
osmotic drought (mannitol) and salt (NaCl) stress for 7 days. iso-osmotic drought (mannitol) and salt (NaCl) stress for 7 days.

© 2009, CAAS. All rights reserved. Published by Elsevier Ltd.

Proline Accumulation, Photosynthetic Abilities and Growth Characters of Sugarcane (Saccharum officinarum L.) 55

Table 2 Maximum quantum yield of PSII (Fv/Fm), photon yield of PSII ( PSII), non-photochemical quenching (NPQ), net-photosynthetic
rate (Pn), stomatal conductance (Gs) and transpiration rate (E) of sugarcane plantlets grown under iso-osmotic drought (mannitol) and salt
(NaCl) stress for 7 days
Osmotic potential (MPa) Fv/Fm PSII NPQ Pn (μmol m-2 s-1) Gs (μmol H2O m-2 s-1) E (mmol m-2 s-1)
-0.23 (control) 0.893 a 0.678 a 0.133 b 7.14 a 7.82 a 0.106 a
-0.67 mannitol 0.868 a 0.664 ab 0.138 b 2.71 b 4.93 b 0.045 b
-0.67 NaCl 0.859 ab 0.655 ab 0.185 ab 1.61 c 4.21 b 0.024 bc
-1.20 mannitol 0.855 ab 0.625 ab 0.188 ab 1.58 c 1.28 a 0.017 bc
-1.20 NaCl 0.825 b 0.615 b 0.253 a 0.47 d 0.31 a 0.006 c
ANOVA * * * ** ** **

Different letters in each column show significant difference at P 0.01 (**) or P 0.05 (*) by Duncan’s new multiple range test (DMRT).

Fig. 3 Relationship between maximum quantum yield of PSII (Fv/ Fig. 4 Relationship between net-photosynthetic rate (Pn) and dry
Fm) and net-photosynthetic rate (Pn) of sugarcane plantlets grown weight of sugarcane plantlets grown under iso-osmotic drought
under iso-osmotic drought (mannitol) and salt (NaCl) stress for 7 (mannitol) and salt (NaCl) stress for 7 days.
days.

Table 3 Relationship between physiological and biochemical parameters of sugarcane plantlets grown under iso-osmotic drought (mannitol)
and salt (NaCl) stress for 7 days
Parameters Chl a Chl b Cx+c PRO Fv/Fm NPQ Pn Gs E
Chl a - - - - - - - - -
Chl b 0.877 ** - - - - - - - -
Cx+c 0.915 ** 0.755 ** - - - - - - -
PRO -0.681 ** -0.811 ** -0.723 ** - - - - - -
Fv/Fm 0.516 * 0.637 ** 0.464 ** -0.667 ** - - - - -
NPQ -0.360 -0.542 * -0.342 0.619 ** -0.478 * - - - -
Pn 0.784 ** 0.903 ** 0.607 ** -0.805 ** 0.663 ** -0.504 * - - -
Gs 0.725 ** 0.826 ** 0.759 ** -0.964 ** 0.703 ** -0.548 * 0.812 ** - -
E 0.818 ** 0.895 ** 0.699 ** -0.790 ** 0.556 * -0.406 0.903 ** 0.796 ** -
Significant levels at P 0.05 and P 0.01 are represented by * and **, respectively using Pearson’s correlation coefficients.

Table 4 Growth characters, fresh weight (FW), dry weight (DW), shoot height (SH), root length (RL) and leaf area (LA) of sugarcane
plantlets grown under iso-osmotic drought (mannitol) and salt (NaCl) stress for 7 days
Osmotic pressure (MPa) FW (mg) DW (mg) SH (cm) RL (cm) LA (mm2)
-0.23 (control) 161.76 a 26.29 a 31.09 a 9.91 a 1 059 a
-0.67 mannitol 129.17 b 21.75 b 26.20 b 6.38 b 785 b
-0.67 NaCl 120.60 bc 20.02 bc 23.06 c 6.03 bc 612 c
-1.20 mannitol 111.27 bc 18.71 cd 21.70 c 5.52 cd 542 d
-1.20 NaCl 102.75 c 17.30 d 18.26 d 4.80 d 358 e
ANOVA ** ** ** ** **

Different letters in each column show significant difference at P 0.01 ( ) by Duncan’s new multiple range test (DMRT).
**

motic adjustments were significantly degraded. Total cit have been reported (Wahid and Ghazanfar 2006;
chlorophyll and carotenoid degradation in sugarcane Silva et al. 2007). Those pigments are sharply reduced,
grown under conditions of soil salinity and water defi- depending on the levels of osmotic treatments, the num-

© 2009, CAAS. All rights reserved. Published by Elsevier Ltd.

56
Table 5 Relationship between growth characters of sugarcane
plantlets grown under iso-osmotic drought (mannitol) and salt
(NaCl) stress for 7 days
Parameters FW DW SH
FW - - - -
DW 0.858 ** - -
SH 0.687 ** 0.741 ** - -
RL 0.779 ** 0.824 ** 0.764 **
LA 0.779 ** 0.855 ** 0.869 **
Significant level at P 0.01 is represented by **
using Pearson’s correlation
coefficients.
ber of days after stress and the sugarcane genotypes
(tolerant or susceptible). The pigment concentrations
in tolerant cultivars [CP-4333 (salt tolerant), HOCP85-
845, TCP02-4548, TCP02-4620, and US01-40 (water-
deficit tolerant)] treated with salt or water-deficit stress
are maintained better than those in sensitive cultivars
[HSF-240 (salt susceptible), CP72-1210, CP92-675,
H99-295, and TCP02-4624 (water-deficit susceptible)]
(Wahid and Ghazanfar 2006; Silva et al. 2007). In
general, the ionic toxicity of salt stress treatment causes
more damage to plant cells than that in mannitol drought
stress conditions, and plays a major role in membrane
injury, organelle damage and pigment degradation prior
to cell death, which is well documented in many plant
species such as sugarcane (Errabii et al. 2007), Cen-
taurea rugusina (Radi et al. 2005, 2006), Fraxinus
angustifolia (Tonon et al. 2004), durum wheat (Lutts
et al. 2004), lentils (Yupsanis et al. 2001), and tobacco
(Gangopadhyay et al. 1997). Proline in sugarcane plant-
lets grown under both NaCl salt stress and water-defi-
cit stress was accumulated, relating to osmotic pres-
sure in the culture media and type of stressors. Proline
accumulation is the first response of plants exposed to
salt stress and water-deficit stress in order to reduce
injury to cells (Ashraf and Foolad 2007). In sugarcane,
there are many reports from studies into proline accu-
mulation in the callus, plantlets and whole plants in field
trials when exposed to salt stress(Wahid 2004;
Gandonou et al. 2006; Patade et al. 2008) and water-
deficit stress (Errabii et al. 2006). In the case of iso-
osmotic salt stress or water-deficit stress, the proline
content in plants exposed to salt treatments reaches a
higher level than that in plants exposed to water-deficit
treatment (Errabii et al. 2007). In the present study,
proline accumulation was dependent on the type of
stress (NaCl salt stress or mannitol water-deficit stress).
In a previous report, proline content in sugarcane
Suriyan Cha-um et al.
(cultivar CoC-671) callus culture peaked in conditions
of 85.6 mM NaCl (Patade et al. 2008). The proline
RL LA content in the leaf tissues of osmotic sensitive geno-
- types (salt susceptible CP-71-3002 and drought sus-
- -
-
ceptible CP59-73) exposed to salt stress (Wahid 2004)
- - or water-deficit stress (Errabii et al. 2006) increased to
0.872 ** -
a greater degree than that in tolerant genotypes (salt
tolerant CP-4333 and drought tolerant R570). In
addition, other osmolytes, glycine, betaine and soluble
sugars are reported as playing major roles in osmotic
adjustment in sugarcane, as defense for coping with
salt stress and water-deficit stress (Wahid 2004;
Gandonou et al. 2006; Patade et al. 2008). The pig-
ment degradation caused by iso-osmotic stress in sug-
arcane plantlets was a major factor in the limitation of
photosynthetic activities, light reaction (Fv/Fm and
PSII) and dark reaction (Pn) as well as stomatal clo-
sure (low Gs) and low transpiration rate (E), leading to
growth reduction. There are many documents report-
ing the photosynthetic responses in sugarcane to salin-
ity (Wahid and Ghazanfar 2006) and water-deficit con-
ditions (Silva et al. 2007) which can be used to develop
effective indices for salt tolerance (Wahid et al. 1997;
Plaut et al. 2000; Akhtar et al. 2003) or water-deficit
tolerance screening (Nable et al. 1999; de Silva and de
Costa 2004; Smit and Singels 2006). The Fv/Fm and
Pn in salt or drought-tolerant genotypes of sugarcane
are maintained better than those in sensitive genotypes
(Wahid and Ghazanfar 2006; Silva et al. 2007). In
addition, there is a high correlation coefficient between
physiological characters and growth performance, in-
cluding leaf area, biomass and plant height (Smit and
Singels 2006; Wahid and Ghazanfar 2006; Silva et al.
2007).
In conclusion, chlorophyll pigments and the photo-
synthetic abilities of sugarcane plantlets grown under
iso-osmotic salt stress declined to a greater degree than
those of plants grown under iso-osmotic water-deficit
stress, leading to a greater reduction in growth rate.
The physiological and growth characters of sugarcane
plantlets were more sensitive to soil salinity than water
deficit. The basic knowledge gained by this investiga-
tion should be applied as salt and water-deficit toler-
ance indicators in sugarcane, as well as further assist-
ing the sugarcane cultivation cultural practices in envi-
ronments affected by soil salinity and water-deficit.
© 2009, CAAS. All rights reserved. Published by Elsevier Ltd.
Proline Accumulation, Photosynthetic Abilities and Growth Characters of Sugarcane (Saccharum officinarum L.)
Acknowledgements
The authors are grateful to the Mitr Phol Sugarcane
Research Center, Mitr Phol Group Co. Ltd. for supply-
ing sugarcane seed stock. This experiment was funded
by the Mitr Phol Sugarcane Research Center, Thailand
(BT-B-03-PT-BC-4930) and partially supported by the
National Center for Genetic Engineering and
Biotechnology, Thailand (BIOTEC) (BT-B-02-RG-BC-
4905).
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(Edited by ZHAO Qi)
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