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SEED PHYSIOLOGY, PRODUCTION & TECHNOLOGY

Osmotic and Salt Stress-Induced Alteration in Soluble Carbohydrate Content in Wheat Seedlings

^a Dep. of Analytical and Structural Chemistry, Janus Pannonius Univ., H-7624 Pécs, Hungary
^b Agricultural Research Institute of the Hungarian Academy of Sciences, H-2462 Martonvásár, Hungary

ilda{at}ttk.jpte.hu

	ABSTRACT

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The effect of drought and salt stresses on the water soluble carbohydrate content in wheat (Triticum aestivum L.) seedlings was examined to characterize the involvement of major sugar components in the adaptive processes. Hydroponically grown seedlings of four wheat varieties differing in drought and salt tolerance were exposed to consecutive water (polyethylene glycol, PEG) and salinity (NaCl) stresses. Total water-soluble carbohydrate (WSC), glucose, fructose, sucrose, and fructan content of stems (non-photosynthetic tissue) were determined. Tolerant genotypes accumulated more soluble carbohydrate than did sensitive ones. Both ionic and non-ionic stresses increased the concentration of reducing sugars, sucrose, and fructans. Drought tolerant varieties accumulated sucrose to a significantly greater level than did sensitive ones under non-ionic stress condition. Changes in fructan content of plants after transfer from PEG to NaCl containing solutions were genotype dependent, increasing in salt tolerant and decreasing in salt sensitive cultivars. These results indicate that WSC might be a useful marker for selecting genotypes that are more drought or salt tolerant. The type of sugar comprising the increase in WSC appears to be a less reliable marker since the initial response was an increase in monosaccharides and the delayed response was an increase in fructan.

Abbreviations: DW, dry weight • PEG, polyethylene glycol • WSC, water-soluble carbohydrate • RWC, relative water content • RWL, relative water loss

	INTRODUCTION

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WATER DEFICIT ASSOCIATED WITH SALINITY in irrigation water is the major limiting factor in the Mediterranean region where plants are subjected to extreme water deficit during the dry season. In this climatic condition, salts may accumulate in the soil because of high evaporative demand and insufficient leaching of ions because of low precipitation. Therefore, soil salinity is sometimes a key factor in determining the ecological distribution of drought-adapted species (Gucci et al., 1997).

A better understanding of physiological responses under these conditions may help in programs in which the objective is to improve the drought and/or salt tolerance of crop varieties. During the course of these stresses, active solute accumulation of compatible solutes such as amino acids, polyamines, and carbohydrates is claimed to be an effective stress tolerance mechanism (Martin et al., 1993; Galiba, 1994; McKersie and Leshem, 1994; Colmer et al., 1995; Rosa-Ibarra and Maiti, 1995).

Carbohydrate changes are of particular importance because of their direct relationship with such physiological processes as photosynthesis, translocation, and respiration. Among the soluble carbohydrates, sucrose and fructans have a potential role in adaptation to these stresses (Williams et al., 1992; Housley and Pollock, 1993; McKersie and Leshem, 1994).

Sucrose can act in water replacement to maintain membrane phospholipids in the liquid-crystalline phase and to prevent structural changes in soluble proteins. The role of reducing sugars (glucose and fructose) in the adaptive mechanism is more controversial, and even their accumulation can be detrimental from several points of view.

Glucose participates in crosslinking with protein by a complex glycosylation reaction between amino and carbonyl groups known as the Maillard reaction (Koster and Leopold, 1988). As respiratory substrates, monosaccharides promote respiration and mitochondrial electron transport which would seem to oppose the onset of quiescence and favor metabolism, energy production, and the formation of oxygen radicals (Leprince et al., 1993).

Fructan is not only a reserve carbohydrate, but it is considered to play a key role in stress-induced metabolic processes. There have been many reports on the effect of different environmental conditions on fructan accumulation in wheat (Pollock and Cairns, 1991; Tognetti et al., 1990; Virgona and Barlow, 1991; Albrecht et al., 1993; Bancal and Triboi, 1993; Galiba et al., 1997). Treatments which enhance photosynthetic carbon fixation lead to increased fructan accumulation. It has also been suggested that fructan metabolism was initiated by intermittent drought (Hendry and Wallace, 1993). Further, introduction of the bacterial levansucrase, which synthesizes fructan, in tobacco (Nicotiana tabacum L.) renders the transgenic plant more tolerant to drought (Pilon-Smits et al., 1995). The mechanism of how fructans can ameliorate stress is not known. Perhaps fructans can protect membranes or other cellular component from the adverse effects of drought, in a manner similar to other carbon compounds, or perhaps fructans influence growth process directly (Pilon-Smits et al., 1995).

In our earlier studies, distinguishing between non-ionic osmotic and salt stresses was reported on the basis of their differential effect on polyamine biosynthesis in wheat calli (Galiba et al., 1993). Furthermore, on the basis of the changes in free amino acid content under osmotic conditions, chromosomes involved in osmoregulation were identified (Galiba et al., 1992). Soluble sugar content proved to be a better marker for selecting improvement of drought tolerance in durum wheat (Triticum durum Desf.) than was proline content (Al Hakimi et al., 1995). In this earlier experiment, only the total sugar content was determined without the identification of specific sugar components.

In this study, we attempt to characterize the involvement of major sugar components in the adaptive processes of various wheat genotypes under consecutive drought and salt stresses. The experiments were undertaken to check whether differences in stress tolerance among genotypes also exist in the quantity of carbohydrate accumulation. More precisely, our purpose was to distinguish between the effects of non-ionic (PEG) and ionic (salinity) osmotic stress conditions on sugar metabolism and to describe the degree of salt and drought tolerance on the basis of changes in concentration of specific sugar components in wheat seedlings.

	Materials and methods

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Plant Materials
Seedlings of bread wheat varieties were grown in half-strength modified Hoagland nutrient solution (Nagy and Galiba, 1995) for 2 wk after germination prior to the stress period in a plant growth chamber (Conviron, Ontario, Canada). The nutrient solution was composed of the following: KNO₃, 5 mM; Ca(NO₃)₂ · 4H₂O, 5 mM; MgSO₄ · 7H₂O, 2 mM; KH₂PO₄, 1 mM; NaFe-EDTA, 0.1 mM. Micronutrient concentrations were as follows: H₃BO₃, 11.5 µM; MnCl₂ · 4H₂O, 4.6 µM; ZnSO₄ · 7H₂O, 0.2 µM; Na₂MO₄ · 2H₂O, 0.12 µM; and CuSO₄ · 5H₂O, 0.08 µM. The cultivars were chosen because of their known response to drought and salinity: Sakha-8 (S) from Egypt is regarded as drought and salt tolerant (Trivedi et al., 1991), Kobomugi (K) from Japan as drought tolerant but salt sensitive (Erdei et al., 1990; Nagy and Galiba, 1995), Chinese Spring (CS) from Hungary as moderately drought and salt tolerant (Galiba et al., 1989, 1993), and Regina (R) from Germany as drought and salt sensitive (Trivedi et al., 1991, Nagy and Galiba, 1995). Day/night (16/8 h) temperature and relative humidity were 15/10°C and 60/75%, respectively. Light intensity during the daytime was 300 µmol m^-2 s^-1 .

Osmotic stress was imposed at the beginning of the third week by application of PEG 4000 at the concentration of 180 g/kg in the nutrient solution for 7 d. After the PEG treatment, some plants were transferred either to equi-osmolal NaCl (200 mM) containing half-strength Hoagland solution for 4 d, or to the same nutrient solution without any supplements to study the recovery. An iso-osmotic transfer of adapted plants from the PEG medium to the corresponding salt supplemented medium allowed the experimental distinction from osmotic and salt-mediated growth effect. This approach was suggested by Harms and Oertli (1985) for cell cultures to study salt tolerance in vitro. We recently applied this method successfully to hydroponically grown seedlings of two bread wheat cultivars (Nagy and Galiba, 1995).

Control seedlings were grown in Hoagland solution throughout the experiment. Samples of leaf blades, stems (crown plus whorl) and roots were taken on 2, 7, and 11 d after treatment.

Determination of Growth and Water Relation Parameters
Shoot biomass production was determined on a dry weight basis after drying overnight at 80°C. Relative water content (RWC) and relative water loss (RWL) were calculated according to Ali Dib et al. (1990) and Yang et al. (1991), respectively.

Chemical Analysis
Water-soluble carbohydrate content was determined on lyophylized plant material. Samples of 200 mg dry weight were extracted according to Kerepesi et al. (1996). Oligosaccharides were hydrolyzed by boiling in 50 g/kg HCl for 60 min. The amount of free (analyzed before hydrolysis) and bound (analyzed after hydrolysis) oligosaccharides, glucose, fructose, and sucrose was measured with Boehringer Mannheim GmbH glucose/fructose/ sucrose, No. 716 260 Kits (Wagner et al., 1983). Total water soluble carbohydrate determination was based on the phenol-sulfuric-acid method (Dubois et al., 1956).

Statistical Analysis
The experimental design was a random complete block, with three replications. The data were analyzed by the STATGRAPHICS (Statistical Graphics Corporation, Princeton USA) statistical package by the t-test and ANOVA functions to assess significant differences among means.

	Results

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Application of PEG and NaCl to the nutrient solution increasingly stressed the plants as judged from the significant inhibition of dry weight production, RWC and RWL (not shown). These tendencies were observed in all cultivars, but to different degrees. Genotype related changes were found in dry weight and RWL. The PEG induced a drop in the total shoot biomass which was the greatest in Sakha (from 283–180 mg/g), while the decrease in Kobomugi biomass was greatest under salinity (from 224–87 mg/g). RWL was the lowest in Sakha (0.44) and the highest in Regina (0.69) under PEG treatment; there was no difference in RWL among the cultivars after salt stress.

The effects of osmotic and salt stress on water-soluble carbohydrate (WSC) content were studied in the roots, leaf blades, and stems of seedlings. As expected, differences were found among the organs. However, only the results from stems are presented here because this organ contained the highest sugar concentrations (one-and-a-half to two fold) and showed the most characteristic, genotype-related changes under stress conditions.

Control
Differences in stem WSC of 14-d-old control, unstressed seedlings were detected among genotypes (Table 1) , which were correlated with their drought sensitivity. Tolerant cultivars, S, K and CS, contained significantly (P < 0.05) more sugars than sensitive Cultivar R. Sucrose content ranged from 17.05 to 35.9 mg/g dw with Cultivars R and CS having significantly less sucrose than the other two genotypes, and with Cultivar S having significantly greater sucrose content than the other three genotypes (Fig. 1) . The greatest monosaccharide content (Fig. 1 and Table 2) was measured in Cultivar S (83.3 mg/g dw), followed by Cultivar K (73.8 mg/g dw), Cultivar CS (66.0 mg/g dw), and Cultivar R (43.5 mg/g dw). Fructan content (Fig. 2) in the unstressed seedlings were similar in all genotypes, ranging from 43.7 mg/g dw to 54.2 mg/g dw.

View larger version (25K): [in this window] [in a new window]   Fig. 1 Changes in glucose and sucrose level in stems of four wheat seedlings, grown in 180 g/kg PEG treated culture solution until Day 7 and then transferred to equi-osmolar 200 mM NaCl solution until Day 11. *, **, and *** indicate significance at P < 0.05, 0.01, and 0.001, respectively

View larger version (25K): [in this window] [in a new window]   Fig. 2 Fructan concentration in the stems of four wheat cultivars exposed to 180 g/kg PEG treated culture until day 7 and then transferred to equi-osmolar 200mMI NaCl solution until day 11. *, **, and *** indicate significance differences between control and stress treatments at P < 0.05, 0.01, and 0.001, respectively. Genotypes: S, Sakha; K, Kobomugi; CS, Chinese Spring; R, Regina

After the 2 d of PEG treatment, concentrations of all the main sugar components (glucose, fructose, and sucrose) increased in each genotype. The change in concentration of glucose (Fig. 1) was equivalent to fructose (Table 2). In drought tolerant cultivars, glucose and fructose increased two fold (P < 0.001) by the second day of treatment (Fig. 1a, c, and Table 2). Drought sensitive Cultivar R accumulated both monosaccharides and sucrose more slowly (Fig. 1g, h), significantly (monosaccharide P < 0.001, sucrose P < 0.05) surpassing the control concentrations only by Day 7 of the PEG treatment. In drought tolerant Cultivars S and K and moderately tolerant Cultivar CS, the content of monosaccharides decreased by the Day 7 (Fig. 1a, c, e), but were still above control levels. However, sucrose concentration increased until Day 7 in Cultivars S and K during PEG treatment (Fig. 1b, d). The changes in sucrose content for the moderately tolerant Cultivar CS had a maximum value on the Day 2, and decreased to about the control level at Day 7 (Fig. 1f).

The initial fructan level was similar in all cultivars. Fructan concentrations increased significantly (P < 0.001) after 2 d of treatment in Cultivars S and R but not Cultivars K or CS. Fructan concentrations increased further at Day 7 of PEG treatment but changed most dramatically in Cultivars K and CS (Fig. 2).

Transfer from PEG to NaCl
After 7 d of PEG treatment, the plants were transferred to an equi-osmolar medium containing NaCl. Sugars were analyzed after 4 d. The WSC content increased in Cultivar S decreased in Cultivar K but changed little in Cultivar CS and in R (Table 1).

After NaCl treatment, considerable differences in mono- and disaccharide content were measured in the cultivars. In Cultivar S, glucose and sucrose (Fig. 1a, b) content increased significantly (P < 0.001). Changes in sugar content in the two salt sensitive Cultivars K and R were different; both mono- and disaccharide increased considerably in Cultivar R (Table 2, Fig. 1g, h) while in Cultivar K a decrease in fructose (Table 2), glucose, and sucrose (Fig. 1 c, d) levels were observed. In Cultivar CS, monosaccharide content declined (Table 2, Fig. 2e) while sucrose accumulated (Fig. 2f).

The fructan content after salt treatment was positively correlated with the degree of salt tolerance (Fig. 2). Fructan in the salt resistant (S) and moderately tolerant (CS) cultivars, increased, while fructan content decreased in the sensitive cultivars (K, R) following the transfer from PEG to a NaCl medium.

Transfer from PEG to Hoagland
To study recovery after water stress, plants were transferred from PEG back to the original half-strength Hoagland medium and their carbohydrate content determined after 4 d. The accumulated WSC decreased in all cultivars to about the control level (Table 1). The component distribution in fructose and fructan was approximately equal to that of the control as well.

	Discussion

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In our experiment, WSC and fructan were sensitive markers for drought or salt tolerance and thus might prove useful in selecting genotypes for improvement. This work was based on the comparison of stems containing the crown and whorl tissues. Our results that the highest sugar content was observed in stems and the most characteristic, genotype-related changes were found in this organ also are consistent with the finding that the survival of wheat seedlings during stress periods depends on survival of the crown (Karen and McKersie, 1985). Further, because stem tissue is basically non-photosynthetic, the accumulation of carbohydrates and the composition of those carbohydrates reflected translocation and subsequent metabolism. The differences between cultivars in the ability to accumulate carbohydrates was evident in control and applied stress conditions. The tolerant Cultivar S accumulated the greatest WSC followed by Cultivars K, CS, and R. This order is in agreement with the apparent stress tolerance of these cultivars.

There have been contradictory results on the effect of water and salt stress on sugar accumulation in wheat. Some studies have reported that sugar content rose (Jones and Turner, 1980; Munns and Weir, 1981) while others have found sugar content decreased (Hanson and Hitz, 1982) or remained constant (Morgan, 1992) during stress conditions. We presented evidence that PEG-induced drought stress increased the WSC concentrations in all cultivars. This observation was also reported recently by Al Hakimi et al. (1995) and Kameli and Lösel (1993). The Cultivar K, which accumulate more WSC proved to be the more drought resistant than Cultivar S.

It is clear that salt stress also increased WSC compared with recovery in all cultivars. The rate of increase in WSC was the highest in most tolerant cultivar (S) followed by moderately tolerant Cultivar CS and sensitive Cultivars R and K. The conclusion that Cultivar K would be considered more sensitive to salt than Cultivar R is in agreement with the classification of Nagy and Galiba (1995), which was based on the measurements of photosynthesis and abscisic acid concentration under similar experimental conditions.

The amount and distribution of WSC components in the stem is an indirect indication of the translocation and net metabolic activity of sink tissues during stress treatments. In tolerant genotypes, sucrose translocated to the stem was initially metabolized to monosaccharides, which caused their content to rise (Day 2) and was followed by a decline (Day 7). These results support the finding that lower sucrose levels allow an increase in the quantities of reducing sugars (Farrant et al., 1993; Leprince et al., 1993). Moreover, the increase in monosaccharides is in agreement with Munns and Weir (1981) where initial changes in osmotic potential were largely due to changes in reducing sugars. In drought tolerant cultivars, as opposed to sensitive ones, the proportion of sugar content was more radically altered by PEG treatment. High fructose concentration in drought sensitive wheat was found by Martin et al. (1993). But glucose and fructose were similar in content in agreement with our results.

Drought-related fructan accumulation has been reported by numerous authors (Pollock and Cairns, 1991, Hendry and Wallace, 1993). Pilon-Smits et al. (1995) observed that transgenic tobacco plants that accumulate bacterial fructan (wild type does not synthesize fructan) performed significantly better under PEG induced water stress than wild-type plants. In our experiments, the lower concentration of WSC in the stem was likely the result of sucrose utilization to synthesize fructan. In all cultivars, the concentrations of fructans increased under osmotic stress but the accumulation rate was not necessarily correlated with drought tolerance. On the other hand, fructans proved to be good marker molecules of salt tolerance since considerably more accumulation was found in salt-tolerant genotypes.

Our data, while providing further evidence on relationship between carbohydrate accumulation and degree of salt and drought tolerance, indicate that total soluble carbohydrate content might be a useful trait to select drought and/or salt-tolerant wheat genotypes. Further, the initial response to drought stress appears to be an increase in monosaccharides, while the more delayed response was an increase in fructan. On the other hand, fructan content could be a useful indicator of degree of salt tolerance.

On the bases of our results, higher WSC levels in the stem (non-photosynthetic tissues) could indicate a greater degree of drought tolerance, and be a useful marker for selecting drought or salt tolerant genotypes.

	ACKNOWLEDGMENTS

 
The authors thank Dr. Thomas L. Housley (Agronomy Department, Purdue University, West Lafayette, IN) for discussion and correction of the manuscript. This research was supported by the Hungarian National Scientific Research funds No. 5474 and 021118.

Received for publication October 17, 1998.

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This Article Abstract Figures Only Full Text (PDF) Free Alert me when this article is cited Alert me if a correction is posted Services Similar articles in this journal Similar articles in ISI Web of Science Alert me to new issues of the journal Download to citation manager Citing Articles Citing Articles via HighWire Citing Articles via ISI Web of Science (17) Citing Articles via Google Scholar Google Scholar Articles by Kerepesi, I. Articles by Galiba, G. Search for Related Content PubMed Articles by Kerepesi, I. Articles by Galiba, G. Agricola Articles by Kerepesi, I. Articles by Galiba, G.