Gradual Drought Research Articles

Effects of gradual soil drought stress on the growth, biomass partitioning, and chlorophyll fluorescence of Prunus mongolica seedlings

The present studyaimed to determine the effects of gradual drought stress on Prunus mongolica seedlings. The growth and chlorophyll fluorescence of the seedlings were investigated under drought and control conditions. The short-term (21 day) decline in soil water content from 30.46% to 15.55% did not significantly influence the growth of P. mongolica seedlings but increased the height, basal diameter, crown, leaf number, stem mass, leaf mass, and root mass of the seedlings. The growth of the drought group reached the maximum, but remained lower than that of the control group, within 21 days. Compared with the control group, the drought group showed 1.46%, 9.65%, 9.44%, 5.19%, 29.09%, 19.20%, and 0.03% lower height, basal diameter, crown, leaf number, stem mass, leaf mass, and root mass, respectively. With the gradual manifestation of soil drought through the decline in soil water content from 15.55% to 11.38% from 21 days to 42 days, growth of the drought group became significantly lower than that of the control group (P < 0.05). The minimal fluorescence decreased and then increased, whereas the nonphotochemical quenching initially increased and then decreased. Thus, P. mongolica can protect the PSII reaction center from damage at the early stage of drought stress. The maximal fluorescence,the maximum quantum yield of PSII, and the photochemical quenching slightly decreased during the initial period of drought stress. These results suggest that drought can slightly influence the openness of PSII reaction centers. This influence was further proven by the decreased quantum yield of PSII. This study provided insights into the adaptive mechanism of P. mongolica seedlings under drought and may serve as a theoretical basis to protect the P. mongolica population. Comparing these responses is crucial for elucidating the drought-tolerance mechanisms in P. mongolica cultivars.

Sodium replacement of potassium in physiological processes of olive trees (var. Barnea) as affected by drought.

Potassium (K) is a macro-nutrient understood to play a role in the physiological performance of plants under drought. In some plant species, sodium (Na) can partially substitute K. Although a beneficial role of Na is well established, information regarding its nutritional role in trees is scant and its function under conditions of drought is not fully understood. The objective of the present study was to evaluate the role of K and its possible replacement by Na in olive's (Olea europaea L.) response to drought. Young and bearing olive trees were grown in soilless culture and exposed to gradual drought. In the presence of Na, trees were tolerant of extremely low K concentrations. Depletion of K and Na resulted in ∼50% reduction in CO2 assimilation rate when compared with sufficiently fertilized control plants. Sodium was able to replace K and recover the assimilation rate to nearly optimum level. The inhibitory effect of K deficiency on photosynthesis was more pronounced under high stomatal conductance. Potassium was not found to facilitate drought tolerance mechanisms in olives. Moreover, stomatal control machinery was not significantly impaired by K deficiency, regardless of water availability. Under drought, leaf water potential was affected by K and Na. High environmental K and Na increased leaf starch content and affected the soluble carbohydrate profile in a similar manner. These results identify olive as a species capable of partly replacing K by Na. The nutritional effect of K and Na was shown to be independent of plant water status. The beneficial effect of Na on photosynthesis and carbohydrates under insufficient K indicates a positive role of Na in metabolism and photosynthetic reactions.

Enzymatic changes in phenylalanine ammonia-lyase, cinnamic-4-hydroxylase, capsaicin synthase, and peroxidase activities in capsicum under drought stress.

Penylalanine ammonia-lyase (PAL), cinnamic-4-hydroxylase (C4H), capsaicin synthase (CS), and peroxidase (POD) are involved in the capsaicinoid biosynthesis pathway and may be altered in cultivars with different pungency levels. This study clarified the action of these enzymes under drought stress for hot Capsicum cultivars with low, medium,and high pungency levels. At the flowering stage, control plants were watered at field capacity, whereas drought-induced plants were subjected to gradual drought stress. Under drought stress, PAL, C4H, CS, and POD enzyme activities increased as compared to the non-drought-stressed plants. A novel discovery was that PAL was the critical enzyme in capsaicinoid biosynthesis under drought stress because its activities and capsaicinoid increased across the different pungency levels of hot pepper cultivars examined.

Interactive Effects of Sudden and Gradual Drought Stress and Foliar‐applied Glycinebetaine on Growth, Water Relations, Osmolyte Accumulation and Antioxidant Defence System in Two Maize Cultivars Differing in Drought Tolerance

AbstractInfluence of sudden and gradual drought stress (DS) and foliar‐applied glycinebetaine (GB) on growth, water relations, osmolyte accumulation and antioxidant defence system were investigated in the plants of two maize (Zea mays L.) cultivars, that is, drought‐tolerant Shaandan 9 (S9) and drought‐sensitive Shaandan 911 (S911). Sudden DS caused less accumulation of GB and free proline, but a more accumulation of malondialdehyde (MDA), which resulted in a greater reduction in leaf relative water content (RWC) and dry matter (DM) in both cultivars compared with the gradual DS. Exogenous GB application caused a rise in DM, RWC, contents of GB and free proline as well as the activities of SOD, CAT and POD along with a decline in MDA content to various extent in both cultivars under both types of DS. A more pronounced effectiveness of GB application was observed in S911 than that in S9 under the same type of DS. It seemed that the more serious damage of DS was on maize plants, and the better positive role of GB was observed in terms of mitigating the adverse effects of DS. From this study, it was possible to propose that hardening for drought resistance by gradual DS treatment and GB application are effective to make plants robust to thrive under water‐deficit conditions.

Impact of Drought Stress on the Accumulation of Capsaicinoids in Capsicum Cultivars with Different Initial Capsaicinoid Levels

Capsaicinoids are the alkaloids in hot pepper that cause the sensation of heat when eaten and are affected by a genetic and environment interaction. Drought stress is well recognized as an environmental condition that influences capsaicinoid accumulation. This investigation identified the responses of capsaicinoid accumulation in hot pepper cultivars under drought stress condition. A total of nine cultivars with a different initial pungency level, i.e., low, medium, and high, was subjected to gradual drought stress during the flowering stage. Plants in this drought stress group were supplied with reduced water applications of 25%, 50%, and 75% by volume at 10, 20, and 30 days after flowering (DAF), respectively. Leaf water potential and relative water content were recorded to measure the level of drought stress. The results indicated that all cultivars were subjected to drought stress because of their decrease in leaf water potential and changes in physiological characteristics, e.g., growth and yield performance. In addition, leaf area and shoot-to-root ratio were good criteria for identifying hot pepper cultivars under drought stress because their responses were correlated with the stress level and yield components. Yield performances of the high pungency group did not decrease under drought stress, whereas those of the low pungency group did decrease. In conclusion, capsaicinoid levels increased for all cultivars studied when subjected to drought stress, except for the cultivars in the high pungency group. A yield response under drought stress for the medium pungency group varied and was not found to be associated with drought stress.

Polyamine metabolic canalization in response to drought stress in Arabidopsis and the resurrection plant Craterostigma plantagineum

In this work, we have studied the transcriptional profiles of polyamine biosynthetic genes and analyzed polyamine metabolic fluxes during a gradual drought acclimation response in Arabidopsis thaliana and the resurrection plant Craterostigma plantagineum. The analysis of free putrescine, spermidine and spermine titers in Arabidopsis arginine decarboxylase (adc1-3, adc2-3), spermidine synthase (spds1-2, spds2-3) and spermine synthase (spms-2) mutants during drought stress, combined with the quantitative expression of the entire polyamine biosynthetic pathway in the wild-type, has revealed a strong metabolic canalization of putrescine to spermine induced by drought. Such canalization requires spermidine synthase 1 (SPDS1) and spermine synthase (SPMS) activities and, intriguingly, does not lead to spermine accumulation but to a progressive reduction in spermidine and spermine pools in the wild-type. Our results suggest the participation of the polyamine back-conversion pathway during the drought stress response rather than the terminal catabolism of spermine. The putrescine to spermine canalization coupled to the spermine to putrescine back-conversion confers an effective polyamine recycling-loop during drought acclimation. Putrescine to spermine canalization has also been revealed in the desiccation tolerant plant C. plantagineum, which conversely to Arabidopsis, accumulates high spermine levels which associate with drought tolerance. Our results provide a new insight to the polyamine homeostasis mechanisms during drought stress acclimation in Arabidopsis and resurrection plants.

Organic matter availability during pre- and post-drought periods in a Mediterranean stream

Mediterranean streams are characterized by water flow changes caused by floods and droughts. When intermittency occurs in river ecosystems, hydrologic connectivity is interrupted and this affects benthic, hyporheic and flowing water compartments. Organic matter use and transport can be particularly affected during the transition from wet to dry and dry to wet conditions. In order to characterize the changes in benthic organic matter quantity and quality throughout a drying and rewetting process, organic matter, and enzyme activities were analyzed in the benthic accumulated material (biofilms growing on rocks and cobbles, leaves, and sand) and in flowing water (dissolved and particulate fractions). The total polysaccharide, amino acid, and lipid content in the benthic organic matter were on average higher in the drying period than in the rewetting period. However, during the drying period, peptide availability decreased, as indicated by decreases in leucine aminopeptidase activity, as well as amino acid content in the water and benthic material, except leaves; while polysaccharides were actively used, as indicated by an increase in β-glucosidase activity in the benthic substrata and an increase in polysaccharide content of the particulate water fraction and in leaf material. During this process, microbial heterotrophs were constrained to use the organic matter source of the lowest quality (polysaccharides, providing only C), since peptides (providing N and C) were no longer available. During the flow recovery phase, the microbial community rapidly recovered, suggesting the use of refuges and/or adaptation to desiccation during the previous drought period. The scouring during rewetting was responsible for the mobilization of the streambed and loss of benthic material, and the increase in high quality organic matter in transport (at that moment, polysaccharides and amino acids accounted for 30% of the total DOC). The dynamics of progressive and gradual drought effects, as well as the fast recovery after rewetting, might be affected by the interaction of the individual dynamics of each benthic substratum: sand sediments and leaves providing refuge for microorganisms and organic matter storage, while on cobbles, an active bacterial community is developed in the rewetting. Since global climate change may favor a higher intensity and frequency of droughts in streams, understanding the effects of these disturbances on the materials and biota could contribute to reliable resource management. The maintenance of benthic substrata heterogeneity within the stream may be important for stream recovery after droughts.

Comparison of drought-induced polypeptides and ion leakage in three tomato cultivars

An attempt has been made to determine if drought-induced proteins could be used as a selection marker to differentiate between tolerant and sensitive cultivars. Three Indian tomato (Lycopersicon esculentum Mill.) cultivars (Pusa Ruby, Arka Vikas and Pusa Early Dwarf) were subjected to drought stress in vivo as well as in vitro and the pattern of polypeptide expression was determined using one-dimensional SDS-PAGE. In all the three cultivars, a new 29 kDa polypeptide accumulated in leaves, in response to gradual drought stress and its accumulation was fastest in Pusa Ruby. Drought stress also resulted in an increase in ion leakage from leaf discs of all the three cultivars but the rate was lower in Pusa Ruby than in other two. Therefore, it was concluded that Pusa Ruby is most tolerant to drought stress among the three tomato cultivars investigated.

Physiological responses to gradual drought stress in the diploid hybrid Pinus densata and its two parental species

Pinus densata is a homoploid hybrid species, originating from P. tabuliformis × P. yunnanensis. The physiological fitness of this natural hybrid compared to its two parental species remains unknown. In this study, we investigated physiological responses of the three species by exposing artificially breed seedlings of each to drought stress lasting 28 days. Our results suggest that, in all three species, drought affected the contents of the plants’ chlorophyll, stomatal conductance, TBARS, hydrogen peroxide, and free proline and increased the activities of antioxidant enzymes, including superoxide dismutase (SOD; EC 1.15.1.1), catalase (CAT; EC 1.11.1.6), and peroxidase (POD; EC 1.11.1.7). The drought stress also induced significant changes in the activity of ascorbate peroxidase (APX; EC 1.11.1.11), monodehydroascorbate reductase (MDHAR; EC 1.6.5.4), dehydroascorbate reductase (DHAR; EC 1.8.5.1), glutathione reductase (GR; EC 1.6.4.2), and levels of ascorbate and glutathione in the ascorbate–glutathione cycle. The hybrid species P. densata appeared to achieve greater drought tolerance and exhibit hybrid superiority in antioxidant processes and other related physiological traits compared to the two parental species, although a few of the hybrid’s measured variables were similar to those of P. tabuliformis. However, P. yunnanensis was more sensitive to drought and appeared to have the lowest resistance to such stress. These physiological differences are largely consistent with the species’ habitat preferences, which may reflect their early genetic divergences and niche differentiation. These findings provide important information for management and forest restoration efforts of these species in the future.

Dehydrin gene expression provides an indicator of low temperature and drought stress: transcriptome-based analysis of Barley (Hordeum vulgare L.)

Low temperature and drought have major influences on plant growth and productivity. To identify barley genes involved in responses to these stresses and to specifically test the hypothesis that the dehydrin (Dhn) multigene family can serve as an indicator of the entire transcriptome response, we investigated the response of barley cv. Morex to: (1) gradual drought over 21 days and (2) low temperature including chilling, freeze-thaw cycles, and deacclimation over 33 days. We found 4,153 genes that responded to at least one component of these two stress regimes, about one fourth of all genes called "present" under any condition. About 44% (1,822 of 4,153) responded specifically to drought, whereas only 3.8% (158 of 4,153) were chilling specific and 2.8% (119 of 4,153) freeze-thaw specific, with 34.1% responsive to freeze-thaw and drought. The intersection between chilling and drought (31.9%) was somewhat smaller than the intersection between freeze-thaw and drought, implying an element of osmotic stress response to freeze-thaw. About 82.4% of the responsive genes were similar to Arabidopsis genes. The expression of 13 barley Dhn genes mirrored the global clustering of all transcripts, with specific combinations of Dhn genes providing an excellent indicator of each stress response. Data from these studies provide a robust reference data set for abiotic stress.

Differential photosynthetic and survival responses to soil drought in two evergreen Nothofagus species

We asked if differences in distribution between Nothofagus nitida and N. dombeyi were associated with differences in drought tolerance. Survival, gas exchange and chlorophyll fluorescence were measured on seedlings subjected to a gradual drought. At a predawn leaf water potential (Ψm) of −2.7 MPa, survival of N. nitida was 50%, compared to 100% in N. dombeyi. Under well-watered conditions, the two species displayed similar stomatal conductance (g w ) and transpiration (E), but net photosynthesis (A) and instantaneous water-use efficiency (WUE i ) were slightly higher in N. nitida. A, E and g w declined in N. nitida along the gradual drought but increased in N. dombeyi at a Ψm between −1.5 and −2.5 MPa, and declined then drastically at a Ψm below < −2.5 MPa. As N. dombeyi was able to maintain A at higher levels despite declining g w , this species displayed significantly increased WUE i at Ψm below −2.5 MPa. Photochemical efficiency of PSII in the light (ΔF/Fmr) and photochemical quenching (qP) were always lower in N. nitida and along with the photochemical efficiency in the dark (Fv/Fm) they declined in both species. Non-photochemical quenching (NPQ) increased slowly in N. dombeyi along with the gradual drought, whilst it decreased in N. nitida. These results show that differences in drought tolerance are in agreement with sorting of Nothofagus species along moisture gradients in south-central Chile.

Whole Gene Family Expression and Drought Stress Regulation of Aquaporins

Since many aquaporins (AQPs) act as water channels, they are thought to play an important role in plant water relations. It is therefore of interest to study the expression patterns of AQP isoforms in order to further elucidate their involvement in plant water transport. We have monitored the expression patterns of all 35 Arabidopsis AQPs in leaves, roots and flowers by cDNA microarrays, specially designed for AQPs, and by quantitative real-time reverse transcriptase PCR (Q-RT-PCR). This showed that many AQPs are pre-dominantly expressed in either root or flower organs, whereas no AQP isoform seem to be leaf specific. Looking at the AQP subfamilies, most plasma membrane intrinsic proteins (PIPs) and some tonoplast intrinsic proteins (TIPs) have a high level of expression, while NOD26-like proteins (NIPs) are present at a much lower level. In addition, we show that PIP transcripts are generally down-regulated upon gradual drought stress in leaves, with the exception of AtPIP1;4 and AtPIP2;5, which are up-regulated. AtPIP2;6 and AtSIP1;1 are constitutively expressed and not significantly affected by the drought stress. The transcriptional down-regulation of PIP genes upon drought stress could also be observed on the protein level.

Cavitation vulnerability in roots and shoots: does Populus euphratica Oliv., a poplar from arid areas of Central Asia, differ from other poplar species?

Populus euphratica is a poplar species growing in arid regions of Central Asia, where its distribution remains nevertheless restricted to river-banks or to areas with an access to deep water tables. To test whether the hydraulic architecture of this species differs from that of other poplars with respect to this ecological distribution, the vulnerability to cavitation of P. euphratica was compared with that of P. alba and of P. trichocarpa x koreana. The occurrence of a potential hydraulic segmentation through cavitation was also investigated by assessing the vulnerability of roots, stems, and leaf mid-rib veins. Cryo-scanning electron microscopy (cryo-SEM) was used to assess the level of embolism in fine roots and leaf mid-ribs and a low pressure flowmeter (LPFM) was used for stems and main roots. The cryo-SEM technique was validated against LPFM measurements on paired samples. In P. alba and P. trichocarpa x koreana, leaf mid-ribs were more vulnerable to cavitation than stems and roots. In P. euphratica, leaf mid-ribs and stems were equally vulnerable and, contrary to what has been observed in other species, roots were significantly less vulnerable than shoots. P. euphratica was by far the most vulnerable. The water potential inducing 50% loss of conductivity in stems was close to -0.7 MPa, against approximately -1.45 MPa for the two others species. Such a large vulnerability was confirmed by recording losses of conductivity during a gradual drought. Moreover, significant stem embolism was recorded before stomatal closure, indicating the lack of an efficient safety margin for hydraulic functions in this species. Embolism was not reversed by rewatering. These observations are discussed with respect to the ecology of P. euphratica.

Gradual Drought Under Field Conditions Influences the Glutathione Metabolism, Redox Balance and Energy Supply in Spring Wheat

Glutathione (GSH) metabolism, redox balance and energy supply in spring wheat (Triticum aestivum L.) during gradual drought stress under field conditions were investigated. Although levels of total and reduced GSH were decreased, the ratio of GSH/GSSG (glutathione disulfide) was markedly increased by drought. Levels of GSH biosynthetic precursors, cysteine (Cys) and c-glutamylcysteine (c-GC), and the activities of their biosynthetic enzymes, c-glutamylcysteine synthetase (c-GCS) and glutathione synthetase (GSHS) were also significantly increased in stressed plants. Glutathione reductase (GR) activity, which is responsible for the conversion of GSSG to GSH, was also increased under this field stress. However, two other important enzymes in GSH metabolism, glutathione peroxidase (GP) and glutathione S-transferase (GST), showed decreased activity in the droughted plants. These results suggest that the higher ratio of GSH/GSSG, the rate of GSH biosynthesis and the capacity of its redox cycling rather than GSH accumulation might be essential for drought resistance of plants. Activities of the two key Calvin-cycle enzymes possessing exposed sulfhydryl groups, NADP + -dependent glyceraldehydes-3-phosphate dehydrogenase (G3PD) and fructose-1,6-bisphosphatase (FBPase) were not affected by drought stress, whereas, activity of the key enzyme in the pentose-phosphate pathway (PPP), 6phosphogluconate dehydrogenase (6-PGD), increased in the droughted plants. The ratios of NADPH/NADP + , NADH/NAD + and ATP/ADP increased in the droughted plants, indicating that an up-regulation of the reduced redox state and the energy supply in the plant cells might be an important physiological strategy for plants responding to drought stress. A simple correlation between the high ratio of GSH/GSSG, the rate of GSH biosynthesis and the redox cycle and the high reduction states of redox status in the plant cells was also observed under field drought.

Effects of Gradual Drought on the Fatty Acid Composition of Polar Lipids, H+-ATPase and 5&amp;prime;-AMPase Activities in the Plasma Membranes of Two Spring Wheat Leaves

研究了田间缓慢干旱胁迫下,抗旱性不同的两个小麦(Triticum aestivum)品种的生长状况、质膜极性脂脂肪酸组成以及质膜关键酶活力的变化.在小麦生长发育的早期,干旱胁迫使其叶片质膜极性脂脂肪酸不饱和度下降、质膜微囊消耗O2的速率升高、膜蛋白含量降低、H+-ATPase (EC 3.6.1.35)活力下降、5′-AMPase (EC 3.1.3.5)活力大幅度升高;在小麦发育的后期,随着干旱的持续,小麦叶片质膜的极性脂脂肪酸不饱和度不变或升高、质膜微囊消耗O2的速率降低、膜蛋白含量与H+-ATPase活力升高、5′-AMPase活力下降.以上结果表明,小麦在发育的早期阶段对干旱较敏感,其细胞质膜流动性降低、细胞中能荷贮备降低;而在后期,则又表现出对干旱的适应.这些结果将有助于阐明自然干旱条件下植物的抗旱机制.

Physiological and morphological adaptations of the fruit tree Ziziphus rotundifolia in response to progressive drought stress.

The physiological basis of drought resistance in Ziziphus rotundifolia Lamk., which is an important, multipurpose fruit tree of the northwest Indian arid zone, was investigated in a greenhouse experiment. Three irrigation regimes were imposed over a 34-day period: an irrigation treatment, a gradual drought stress treatment (50% of water supplied in the irrigation treatment) and a rapid drought stress treatment (no irrigation). Changes in gas exchange, water relations, carbon isotope composition and solute concentrations of leaves, stems and roots were determined. The differential rate of stress development in the two drought treatments did not result in markedly different physiological responses, but merely affected the time at which they were expressed. The initial response to decreasing soil water content was reduced stomatal conductance, effectively maintaining predawn leaf water potential (Psi(leaf)), controlling water loss and increasing intrinsic water-use efficiency, while optimizing carbon gain during drought. Carbon isotope composition (delta13C) of leaf tissue sap provided a more sensitive indicator of changes in short-term water-use efficiency than delta13C of bulk leaf tissue. As drought developed, osmotic potential at full turgor decreased and total solute concentrations increased in leaves, indicating osmotic adjustment. Decreases in leaf starch concentrations and concomitant increases in hexose sugars and sucrose suggested a shift in carbon partitioning in favor of soluble carbohydrates. In severely drought-stressed leaves, high leaf nitrate reductase activities were paralleled by increases in proline concentration, suggesting an osmoprotective role for proline. As water deficit increased, carbon was remobilized from leaves and preferentially redistributed to stems and roots, and leaves were shed, resulting in reduced whole-plant transpiration and enforced dormancy. Thus, Z. rotundifolia showed a range of responses to different drought intensities indicating a high degree of plasticity in response to water deficits.

Poplar nutrient resorption in fall or drought: influence of nutrient status and clone

Nutrient mobilization in hybrid poplars was studied in two greenhouse experiments. Half the clones in each experiment were drought resistant and half were drought sensitive. In experiment 1, four clones were grown at three levels of N, two levels of P, and with four drought treatments. In experiment 2, six clones received two levels of N and three levels of K. Well-watered trees, trees subjected to drought conditions once, and hardened drought-stressed trees were harvested in late summer, and other well-watered trees were harvested in fall. Mobilization of nutrients differed with speed and intensity of drought. During gradual drought in the first experiment, N and P were resorbed but not K or Cu. In experiment 2, drought was too rapid for resorption. Instead, nutrients became more concentrated: increased N facilitated hardening and osmotic adjustment, while K moved out of the leaves on rewatering. In fall, N, P, Cu, and K were resorbed. Increased K supply increased fall resorption efficiency of Cu, N, and P but not K. Increased N supply improved fall resorption proficiency of K at low levels of supply. However, clonal productivity on dry sites was more related to leaf retention in drought and in fall than to resorption efficiency.

Interactive effects of elevated [CO2] and drought on cherry (Prunus avium) seedlings I. Growth, whole‐plant water use efficiency and water loss

Seeds of cherry (Prunus avium) were germinated and grown for two growing seasons in ambient (∼350 μmol mol−1) or elevated (ambient+∼350 μmol mol−1) CO2 mole fractions in six open‐top chambers. The seedlings were fertilized once a week, following Ingestad principles in order to supply mineral nutrients at free‐access rates. In the first growing season gradual drought was imposed on rapidly growing cherry seedlings by withholding water for a 6‐wk drying cycle. In the second growing season, the rapid onset of drought was imposed at the height of the growing season on the seedlings which had already experienced drought in the first growing season. Elevated [CO2] significantly increased total dry‐mass production in both water regimes, but did not ameliorate the growth response to drought of the cherry seedlings subjected to two sequential drying cycles. Water loss did not differ in either well watered or droughted seedlings between elevated and ambient [CO2]; consequently whole‐plant water‐ use efficiency (the ratio of total dry mass produced to total water consumption) was significantly increased. Similar patterns of carbon allocation between shoot and root were found in elevated and ambient [CO2] when the seedlings were the same size. Thus, elevated [CO2] did not improve drought tolerance, but it accelerated ontogenetic development irrespective of water status.

The role of solute accumulation, osmotic adjustment and changes in cell wall elasticity in drought tolerance in Ziziphus mauritiana (Lamk.)

bulk tissue elastic modulus (wall rigidity) which resulted in turgor loss at the same h in both stressed Ber (Ziziphus mauritiana Lamk.) is a major fruit tree and unstressed leaves. The possible ecological signicrop of the north-west Indian arid zone. In a study of ficance of maintenance of turgor potential and cell the physiological basis of drought tolerance in this volume at low water potentials for drought tolerance species, two glasshouse experiments were conducted in Ziziphus is discussed. in which trees were droughted during single stresscycles. In the first experiment, during a 13 d drying Key words: Ziziphus mauritiana, drought, solute accumulacycle, pre-dawn leaf water (Y leaf ) and osmotic (Y p ) tion, osmotic adjustment, proline. potentials in droughted trees declined from ’0.5 and ’1.4 MPa to ’1.7 and ’2.2 MPa, respectively, for a decrease in relative water content (h) of 14%. During Introduction drought stress, changes in sugar metabolism were associated with significant increases in concentra- Drought is probably the most important factor limiting tions of hexose sugars (3.8-fold), cyclitol (scyllo- crop productivity world-wide (Jones and Corlett, 1992). inositol; 1.5-fold), and proline (35-fold; expressed per In semi-arid regions, where the amounts and patterns of unit dry weight), suggesting that altered solute parti- seasonal rainfall are often erratic and unpredictable, deeptioning may be an important factor in drought toler- rooting perennial species may exploit deep soil water ance of Ziziphus. On rewatering, pre-dawn Y leaf and h reserves. This facilitates production of yields in situations recovered fully, but Y p remained depressed by 0.4 MPa of high light coupled with drought, where shallow-rooted relative to control values, indicating that solute con- annual crops would normally fail. The use of perennials centration per unit water content had changed during may help buVer farmers’ production against year-to-year the drought cycle. fluctuations in yields from annual crop species. Under Evidence for osmotic adjustment was provided from such conditions, an ability to maintain cell water status a second study during which a gradual drought (e.g. through osmotic adjustment and/or leaf area adjustwas imposed. Pressure‐volume analysis revealed a ment) and cell integrity (by protecting cells against photo0.7 MPa reduction in osmotic potential at full turgor, and chemical oxidation) may be an advantage. There are with Y leaf at turgor loss depressed by ~1 MPa in many cases where plants growing in hostile environments drought-stressed leaves. Coupled with osmotic adjust- exhibit increased oxy-stress enzyme activities to combat lipid peroxidation by free radicals, hydrogen peroxide

The role of sucrose synthase in the response of soybean nodules to drought

Experiments were carried out to investigate the effects of drought stress on enzymatic activities related to carbon and nitrogen metabolism in soybean nodules. Gradual drought stress, imposed by withholding water nutrients, resulted in declines in the water potential of leaves and nodules consistent with a significant decline in N2 fixation. However, the amounts of nitrogenase components 1 and 2 were virtually unaffected by drought stress. Similarly, no significant changes could be detected in aspartate aminotransferase, phosphoenolpyruvate carboxylase, glutamine synthetase or alkaline invertase activities throughout the experiment. In contrast, sucrose synthase (SS), one of the enzymes involved in sucrose metabolism in legume nodules, declined dramatically in activity and in content within a few days of withholding water. Coincident with this decline in SS activity were significant increases in the nodule contents of sucrose, total free amino acids and ureides. The amounts of proline, however, did not increase until some days later. It is suggested that SS may play a key role in the regulation of nodule carbon metabolism and, therefore, of nitrogen fixation under drought stress conditions.