Proline Accumulation, Photosynthetic Abilities and Growth Characters of Sugarcane (Saccharum officinarum L.) Plantlets in Response to Iso-Osmotic Salt and Water-Deficit Stress

Agricultural Sciences in China

January 2009

2009, 8(1): 51-58

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 isoosmotic salt and water-deficit stress. Disease-free sugarcane plantlets derived from meristem cuttings were photoautotrophically 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 waterdeficit 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 netphotosynthetic 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

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 detrimentally 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

Das 2005). In general, osmotic stresses caused by 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 generated from salt contaminated soil has negative effects on plant growth and development (Tester and Davenport 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: [email protected]

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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, leading to growth retardation and yield reduction. Plant biochemicals [ascorbate peroxidase (AOX), glutamine synthetase (GS), proline, glycine betaine, photosynthetic pigments, soluble proteins and mineral elements] and physiological changes [relative water content (RWC), stomatal conductance (Gs), water potential ( 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 (Chaum 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 breeding 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 maximum 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 propagated 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 without 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. Plantlets 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 supporting material for 7 days. The number of air-exchanges in the glass vessels was adjusted to 2.32 h-1 by punching a hole in the plastic cap (Ø 1 cm) and covering 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 biochemical analysis.

Data measurement Chlorophyll a (Chl a), chlorophyll b (Chl b), total chlorophyll, and total carotenoid (Cx+c) concentrations

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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 hundred 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 reagent (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 container in an ice bath. The reaction mixture was vigorously mixed with 2 mL toluene. After warming at 25°C, the chromophore was measured by Spectrophotometer 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 monitored with a Fluorescence Monitoring System in the pulse amplitude modulation mode, as previously described by Loggini et al. (1999). A leaf, adapted to dark conditions for 30 min using leaf-clips, was initially exposed to the modulated measuring beam of farred light (LED source with typical peak at wavelength 735 nm). Original (Fo) and maximum (Fm) fluorescence 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 fluorescence yield (Fv) was calculated by the equation of FmFo. 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 quenching (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 temperature 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 plantlets were dried at 110°C in a hot-air oven for 2 days, and then incubated in desiccators before the measurement of dry weight. The leaf area of sugarcane plantlets was measured using a leaf area meter DT-scan.

Experiment design The experiment was designed as completely randomized design (CRD) with ten replicates and four plantlets per replicate. The mean values obtained were compared 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 osmotic pressure in the culture media (r2 = 0.93) (Fig.1). Chl a, Chl b, TC and Cx+c contents in the -1.20 MPa

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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 relating to the osmotic stress, especially salt induced osmotic stress (Table 1). Proline was generally accumulated in osmotically-stressed sugarcane plantlets and played a key role in osmoregulation and antioxidant defense mechanisms. Moreover, the total chlorophyll degradation 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 significantly reduced when compared to those of plantlets of the control group (-0.23 MPa), while those parameters in plantlets grown in -1.20 MPa mannitol were unchanged (Table 2). On the other hand, non-photochemical quenching (NPQ) of osmotically stressed plantlets 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 waterdeficit 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) -0.23 (control) -0.67 mannitol -0.67 NaCl -1.20 mannitol -1.20 NaCl ANOVA

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)

371.46 a 271.42 b 163.73 c 238.77 b 129.68 c

154.63 a 88.02 b 64.00 bc 71.18 b 32.73 c

526.10 a 359.44 b 227.74 c 309.95 b 162.41 c

71.15 a 62.32 ab 37.92 bc 59.56 ab 21.94 c

318 d 518 d 770 c 1 027 b 1 412 a

**

**

**

**

**

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 and pigment degradation of sugarcane plantlets grown under isoosmotic drought (mannitol) and salt (NaCl) stress for 7 days.

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

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

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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) -0.23 (control) -0.67 mannitol -0.67 NaCl -1.20 mannitol -1.20 NaCl ANOVA

Fv/Fm

PSII

0.893 a 0.868 a 0.859 ab 0.855 ab 0.825 b

0.678 a 0.664 ab 0.655 ab 0.625 ab 0.615 b

*

NPQ

*

Different letters in each column show significant difference at P

Pn (μmol m-2 s-1)

0.133 b 0.138 b 0.185 ab 0.188 ab 0.253 a *

0.01 (**) or P

Gs (μmol H2O m-2 s-1)

7.14 a 2.71 b 1.61 c 1.58 c 0.47 d

7.82 a 4.93 b 4.21 b 1.28 a 0.31 a

**

**

E (mmol m-2 s-1) 0.106 a 0.045 b 0.024 bc 0.017 bc 0.006 c **

0.05 (*) by Duncan’s new multiple range test (DMRT).

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

Fig. 4 Relationship between net-photosynthetic rate (Pn) and dry weight of sugarcane plantlets grown under iso-osmotic drought (mannitol) and salt (NaCl) stress for 7 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 0.877 ** 0.915 ** -0.681 ** 0.516 * -0.360 0.784 ** 0.725 ** 0.818 **

Significant levels at P

0.05 and P

Chl b

Cx+c

PRO

Fv/Fm

NPQ

Pn

Gs

E

0.755 ** -0.811 ** 0.637 ** -0.542 * 0.903 ** 0.826 ** 0.895 **

-0.723 ** 0.464 ** -0.342 0.607 ** 0.759 ** 0.699 **

-

-

-

-

-

-

0.796 **

-

-0.667 ** 0.619 ** -0.805 ** -0.964 ** -0.790 **

-0.478 * 0.663 ** 0.703 ** 0.556 *

-0.504 * -0.548 * -0.406

0.812 ** 0.903 **

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) -0.67 mannitol -0.67 NaCl -1.20 mannitol -1.20 NaCl ANOVA

161.76 a 129.17 b 120.60 bc 111.27 bc 102.75 c

26.29 a 21.75 b 20.02 bc 18.71 cd 17.30 d

31.09 a 26.20 b 23.06 c 21.70 c 18.26 d

9.91 a 6.38 b 6.03 bc 5.52 cd 4.80 d

1 059 a 785 b 612 c 542 d 358 e

**

**

**

**

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 chlorophyll and carotenoid degradation in sugarcane grown under conditions of soil salinity and water defi-

cit have been reported (Wahid and Ghazanfar 2006; Silva et al. 2007). Those pigments are sharply reduced, depending on the levels of osmotic treatments, the num-

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Suriyan Cha-um et al.

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 RL LA

FW

DW

SH

RL

LA

0.858 ** 0.687 ** 0.779 ** 0.779 **

0.741 ** 0.824 ** 0.855 **

0.764 ** 0.869 **

0.872 **

-

Significant level at P coefficients.

0.01 is represented by

**

using Pearson’s correlation

ber of days after stress and the sugarcane genotypes (tolerant or susceptible). The pigment concentrations in tolerant cultivars [CP-4333 (salt tolerant), HOCP85845, TCP02-4548, TCP02-4620, and US01-40 (waterdeficit 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), Centaurea 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 plantlets grown under both NaCl salt stress and water-deficit stress was accumulated, relating to osmotic pressure 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 accumulation 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 waterdeficit stress (Errabii et al. 2006). In the case of isoosmotic 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

(cultivar CoC-671) callus culture peaked in conditions of 85.6 mM NaCl (Patade et al. 2008). The proline content in the leaf tissues of osmotic sensitive genotypes (salt susceptible CP-71-3002 and drought susceptible CP59-73) exposed to salt stress (Wahid 2004) or water-deficit stress (Errabii et al. 2006) increased to 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 pigment degradation caused by iso-osmotic stress in sugarcane 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 closure (low Gs) and low transpiration rate (E), leading to growth reduction. There are many documents reporting the photosynthetic responses in sugarcane to salinity (Wahid and Ghazanfar 2006) and water-deficit conditions (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, including leaf area, biomass and plant height (Smit and Singels 2006; Wahid and Ghazanfar 2006; Silva et al. 2007). In conclusion, chlorophyll pigments and the photosynthetic 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 investigation should be applied as salt and water-deficit tolerance indicators in sugarcane, as well as further assisting the sugarcane cultivation cultural practices in environments affected by soil salinity and water-deficit.

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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 supplying 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-BC4905).

References Akhtar S, Wahid A, Rasul E. 2003. Emergence, growth and nutrient composition of sugarcane sprouts under NaCl salinity. Biologia Plantarum, 46, 113-116. Ashraf M, Foolad M R. 2007. Role of glycine betaine and proline in improving plant abiotic stress resistance. Environmental and Experimental Botany, 59, 206-216. Ashraf M, Harris P J C. 2004. Potential biochemical indicators of salinity tolerance in plants. Plant Science, 166, 3-16. Bates L S, Waldren R P, Teare I D. 1973. Rapid determination of free proline for water-stress studies. Plant and Soil, 39, 205207. Castillo E G, Tuong T P, Ismail A M, Inubushi K. 2007. Response to salinity in rice: Comparative effects of osmotic and ionic stresses. Plant Production Science, 10, 159-170. Cha-um S, Supaibulwatana K, Kirdmanee C. 2006. Water relation, photosynthetic ability, and growth of Thai Jasmine rice (Oryza sativa L. ssp. indica cv. KDML105) to salt stress by application of exogenous glycinebetaine and choline. Journal of Agronomy and Crop Science, 192, 25-36. Cha-um S, Supaibulwatana K, Kirdmanee C. 2007. Glycinebetaine accumulation, physiological characterizations, and growth efficiency in salt tolerant and salt sensitive lines of indica rice (Oryza sativa L. spp. indica) response to salt stress. Journal of Agronomy and Crop Science, 193, 157-166. Davenport R, James R A, Zakrisson-Plogander A, Tester M, Munns R. 2005. Control of sodium transport in durum wheat. Plant Physiology, 137, 807-818. Errabii T, Gandonou C B, Essalmani H, Abrini J, Idomar M, Senhaji N S. 2006. Growth, proline and ion accumulation in sugarcane callus cultures under drought-induced osmotic stress and its subsequent relief. African Journal of Biotechnology, 5, 1148-1493. Errabii T, Gandonou CB, Essalmani H, Abrini J, Idaomar M, Senhaji N S. 2007. Effect of NaCl and mannitol induced stress on sugarcane (Saccharum sp.) callus cultures. Acta Physiologia Plantarum, 29, 95-102. Gandonou C B, Errabii T, Abrini J, Idaomar M, Senhaji N S. 2006. Selection of callus cultures of sugarcane (Saccharum

57

sp.) tolerant to NaCl and their responses to salt stress. Plant Cell, Tissue and Organ Culture, 87, 9-16. Gangopadhyay G, Basu S, Mukherjee B B, Gupta S. 1997. Effects of salt and osmotic shocks on unadapted and adapted callus lines of tobacco. Plant Cell, Tissue and Organ Culture, 49, 45-52. Hasegawa P M, Bressan R A, Zhu J K, Bohnert H J. 2000. Plant cellular and molecular responses to high salinity. Annual Review in Plant Physiology and Molecular Biology, 51, 463499. de Herralde F, Biel C, Savé R, Morales M A, Torrecillas A, Alarcón J J, Sánchez-Blanco M J. 1998. Effect of water and salt stresses on the growth, gas exchange and water relations in Argyranthemum coronopifolium plants. Plant Science, 139, 9-17. Hu Y, Burucs Z, von Tucher S, Schmidhalter U. 2007. Shortterm effects of drought and salinity on mineral nutrient distribution along growing leaves of maize seedlings. Environmental and Experimental Botany, 60, 268-275. Lichtenthaler H K. 1987. Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes. Methods in Enzymology, 148, 350-380. Loggini B, Scartazza A, Brugnoli E, Navari-Izzo F. 1999. Antioxidant defense system, pigment composition, and photosynthetic efficiency in two wheat cultivars subjected to drought. Plant Physiology, 119, 1091-1099. Luo Q, Yu B, Liu Y. 2005. Differential sensitivity to chloride and sodium ions in seedlings of Glycine max and G. soja under NaCl stress. Journal of Plant Physiology, 162, 1003-1012. Lutts S, Almansouri M, Kinet J M. 2004. Salinity and water stress have contrasting effects on the relationship between growth and cell viability during and after stress exposure in durum wheat callus. Plant Science, 167, 9-18. Maggio A, de Pascale S, Ruggiero C, Barbieri G. 2005. Physiological response of field-grown cabbage to salinity and drought stress. European Journal of Agronomy, 23, 5767. Mahajan S, Tuteja N. 2005. Cold, salinity and drought stresses: An overview. Archives in Biochemistry and Biophysics, 444, 139-158. Maricle B R, Cobos D R, Campbell C S. 2007. Biophysical and morphological leaf adaptations to drought and salinity in salt marsh grasses. Environmental and Experimental Botany, 60, 458-467. Maxwell K, Johnson G N. 2000. Chlorophyll fluorescence – A practical guide. Journal of Experimental Botany, 51, 659668. Munns R, James R A, Läuchli A. 2006. Approaches to increasing the salt tolerance of wheat and other cereals. Journal of Experimental Botany, 57, 1025-1043.

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

58

Suriyan Cha-um et al.

Nable R O, Robertson M J, Berthelsen S. 1999. Response of shoot growth and transpiration to soil drying in sugarcane.

physiological parameters as fast tools to screen for drought tolerance in sugarcane. Brazilian Journal of Plant Physiology,

Plant and Soil, 207, 59-65. Pagter M, Bragato C, Malagoli M, Brix H. 2009. Osmotic and

19, 193-201. Slama I, Ghnaya T, Hessini K, Messedi D, Savouré A, Abdelly

ionic effects of NaCl and Na2SO 4 salinity on Phragmites australis. Aquatic Botany, 90, 43-51.

C. 2007. Comparative study of the effects of mannitol and PEG osmotic stress on growth and solute accumulation in

Parida A K, Das A B. 2005. Salt tolerance and salinity effects on plants: a review. Ecotoxicology and Environmental Safety,

Sesuvium portulacastrum. Environmental and Experimental Botany, 61, 10-17.

60, 324-349. Patade V Y, Suprasanna P, Bapat V A. 2008. Effects of salt

Smit MA, Singels A. 2006. The response of sugarcane canopy development to water stress. Field Crops Research, 98, 91-

stress in relation to osmotic adjustment on sugarcane (Saccharum officinarum L.) callus cultures. Plant Growth

97. Teixeira J, Pereira S. 2007. High salinity and drought act on an

Regulation, 55, 169-173. Plaut Z, Meinzer FC, Federman E. 2000. Leaf development,

organ-dependent manner on potato glutamine synthetase expression and accumulation. Environmental and

transpiration and ion uptake and distribution in sugarcane cultivars grown under salinity. Plant and Soil, 218, 59-69.

Experimental Botany, 60, 121-126. Tester M, Davenport R. 2003. Na+ tolerance and Na+ transport

Pujol J A, Calvo J F, Ramírez-Díaz L. 2000. Recovery of germination from different osmotic conditions by four

in higher plants. Annals of Botany, 91, 503-527. Tonon G, Kevers C, Faivre-Rampant O, Graziani M, Gaspar T.

halophytes from southeastern Spain. Annals of Botany, 85, 279-286.

2004. Effect of NaCl and mannitol iso-osmotic stresses on proline and free polyamine levels in embryogenic Fraxinus

Radi S, Proliæ M, Pavlica M, Pevalek-Kozlina B. 2005. Cytogenetic effects of osmotic stress on the root meristem

angustifolia callus. Journal of Plant Physiology, 161, 701708.

cells of Centaurea ragusina L. Environmental and Experimental Botany, 54, 213-218.

Wahid A. 2004. Analysis of toxic and osmotic effects of sodium chloride on leaf growth and economic yield of sugarcane.

Radi S, Radiæ-Stojkoviæ M, Pevalek-Kozlina B. 2006. Influence of NaCl and mannitol on peroxidase activity and lipid

Botanical Bulletin of Academia Sinica, 45, 133-141. Wahid A, Ghazanfar A. 2006. Possible involvement of some

peroidation in Centaurea ragusina L. roots and shoots. Journal of Plant Physiology, 163, 1284-1292.

secondary metabolites in salt tolerance of sugarcane. Journal of Plant Physiology, 163, 723-730.

Reddy A R, Chitanya K V, Vivekanandan M. 2004. Droughtinduced responses of photosynthesis and antioxidant

Wahid A, Rao A, Rasul E. 1997. Identification of salt tolerance traits in sugarcane lines. Field Crops Research, 54, 9-17.

metabolism in higher plants. Journal of Plant Physiology, 161, 1189-1202.

Wang B, Li Z, Eneji A E, Tian X, Zhai Z, Li J, Duan L. 2008. Effects of coronatine on growth, gas exchange traits,

Sairam R K, Tyagi A. 2004. Physiology and molecular biology of salinity stress tolerance in plants. Current Science, 86, 407-

chlorophyll content, antioxidant enzymes and lipid peroxidation in maize (Zea mays L.) seedlings under simulated

421. Shabala S N, Shabala S I, Martynenko A I, Babourina O, Newman

drought stress. Plant Production Science, 11, 283-290. Wang W, Vinocur B, Altman A. 2003. Plant responses to drought,

I A. 1998. Salinity effect on bioelectric activity, growth, Na+ accumulation and chlorophyll fluorescence of maize leaves:

salinity and extreme temperatures: towards genetic engineering for stress tolerance. Planta, 218, 1-14.

a comparative survey and prospects for screening. Australia Journal of Plant Physiology, 25, 609-616. de Silva ALC, de Costa W A J M. 2004. Varietal variation in growth, physiology and yield of sugarcane under two contrasting water regimes. Tropical Agricultural Research, 16, 1-12. Silva M A, Jifon J L, da Silva J A G, Sharma V. 2007. Use of

Wang W, Vinocur B, Shoseyov O, Altman A. 2001. Biotechnology of plant osmotic stress tolerance: physiological and molecular considerations. Acta Horticulturae, 560, 285-292. Yupsanis T, Kefalas P S, Eleftheriou P, Kotinis K. 2001. RNase and DNase activities in the alfalfa and lentil grown in isoosmotic solutions of NaCl and mannitol. Journal of Plant Physiology, 158, 921-927. (Edited by ZHAO Qi)

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