Salinity stress and drought stress differentially affect biomass allocation, leaf orientation and performance of two sesame (Sesamum indicum L.) cultivars

Salinity stress and drought stress differentially affect biomass allocation, leaf orientation and performance of two sesame (Sesamum indicum L.) cultivars | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Salinity stress and drought stress differentially affect biomass allocation, leaf orientation and performance of two sesame (Sesamum indicum L.) cultivars Taha Mohamed El-Katony, Nemat Mohamed Hassan, Samia Helmy, Shaimaa Nassim Abdelfatah This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7362732/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Drought and salinity stresses are major threats to crop productivity in the arid lands. This work investigates the differential impact of salinity and drought on growth and performance of two sesame cultivars: Sohg and Shnd at water potentials of 0, -100, -250 and − 450 kPa with the same nutrient supply in a hydroponic sand culture. Under no stress, leaves of Sohg exhibited vertical orientation with higher pigment and soluble sugar contents but lower polysaccharide content relative to leaves of Shnd. Salinity and drought differentially affect leaf orientation only in Sohg, where drought induced horizontal leaf position but salinity led to the opposite effect. Whereas drought induced stronger reduction in foliage biomass than did salinity, with enhancement of root biomass, salinity reduced foliage and root biomass with favored allocation of plant biomass to root. Salinity reduced leaf pigment content, particularly in Sohg but drought either increased it or was without effect. Both drought and salinity, post a threshold of -100 kPa, reduced rates of photosynthesis and transpiration as well as stomatal conductance, with stronger impact of salinity than drought. The lowering in carbohydrate content under stress, being more aggressive under salinity than drought, was associated with increasing proportion of pectin at the expense of starch and variable changes in soluble sugar proportions. Drought. Salinity. Sesame genotypes. Gas exchange. Carbohydrate fractions. Leaf orientation Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 1 Introduction In natural habitats, plants may experience combined stresses simultaneously such as the combined stress of high temperature, water deficit and high irradiance in the hot dry summer of arid habitats [ 1 ]. Among the different abiotic stresses affecting plants, salinity stress and water stress are particularly serious and mostly interrelated. Mostly, saline soils are abundant in semiarid and arid regions where rainfall is insufficient for leaching of excessive soluble salts out of the rhizosphere [ 2 ]. In turn, irrigation with poor quality water, a practice common in the arid developing countries, can induce secondary salinization. Soil is considered saline when electrical conductivity of its saturation extract exceeds 4 dS m − 1 (equivalent to 40 mM NaCl) with an exchangeable sodium percentage lower than 15 [ 3 ]. Although the majority of crop plans are glycophytes, few species are salt-tolerant and can even benefit from mild salinity. Justification of crop salt response relies upon two criteria: the threshold salinity and the slope of the salinity-growth response line beyond the threshold. Differences in salt tolerance can occur among different species and cultivars as well as for the different plant developmental stages [ 4 , 5 ]. Similarly, drought stress represents a major challenge to plant productivity and land use, particularly in the arid regions. Even in the tropical rain forests, land plants can experience different degrees of transient water stress when water absorption lags behind evapotranspiration [ 6 ]. The witnessed global warming will further aggravate the threat of drought to plant productivity. Intrinsically, plants have low water use efficiency, with 90% of the absorbed mineral nutrients [ 1 ]. Water is translocated within the plant down a gradient of 1) water potential (Ψ w ) across membranes from the soil to the root and throughout the root, 2) hydrostatic pressure in the xylem vessels and 3) water vapor gradient from the leaf to air. Under the impact of salinity, plants suffer from several hazards including osmotic stress, ion toxicity, nutrient imbalance and oxidative stress; the relative effect of either factor depends on the level, duration and mode of exposure to salt stress as well as the plant developmental stage. For example, long-term salinity stress leads mainly to ion toxicity in the older leaves but to water deficit and shortage of carbohydrates in the younger leaves [ 7 ]. Salinity and drought cause morphological, physiological, biochemical and molecular changes in the plant. To cope with stress, the plant has to expend additional energy for ion exclusion or compartmentalization (salinity stress) as well as for osmotic adjustment and detoxification of reactive oxygen species (salinity and drought stresses), which will slow down plant growth [ 8 ]. As a rule, shoot growth is more depressed by salinity and drought than root growth, partially because of favored biomass allocation to roots at the expense of shoots [ 9 ]. In addition to the direct toxicity of high levels of Na + and Cl - to the cytoplasm, salt toxicity may also be attributed to interference with respiration, photosynthesis and protein synthesis. Salinity can cause insufficient cellular K + concentrations for enzyme activation, protein synthesis, photosynthesis and stomatal movement [ 10 ]. Hence, the ability of plants to protect enzymes from the adverse effects of Na + or Cl - can be attributed to the synthesis of compatible solutes such as sugars and amino acids (e.g. proline) that can stabilize the structure of membranes and macromolecules [ 11 ]. The role of compatible solutes in plant protection is also evident under drought stress, where plant cells adjust Ψ w by accumulating inorganic ions and organic acids in the vacuoles, with the synthesis of compatible solutes such as glycine betaine, sorbitol, and proline in the cytoplasm. The salinity-induced reduction in photosynthesis can be attributed to the reductions in leaf area (a capacity factor) and net CO 2 fixation per unit leaf area (an efficiency factor) in addition to the decreased demand of sink tissues for assimilates [ 12 ]. Further, the effect of salinity on photosynthetic efficiency includes both stomatal and nonstomatal limitations. Stomatal limitations are manifested as lowered intercellular CO 2 concentration while nonstomatal factors include the decrease in quantum efficiency and inhibited Ribusco activity [ 13 ]. Stomatal closure and the lowered stomatal conductance represent a way to prevent excessive dehydration of the leaves as a feedforward response to anticipate water deficit before the development of severe drought. Sesame ( Sesamum indicum L.), a Pedaliaceae member and one of the most valuable oil cash crops in Africa and Asia [ 14 ], is a robust crop with wide niche and marked drought tolerance. The high oil content of sesame seeds (up to 63%) is characterized with high proportion of unsaturated fatty acids, particularly omega 3 and γ-tocopherol [ 15 ]. Sesame seeds are also rich in human-compatible proteins, vitamins, dietary fibers and dietary minerals, particularly Ca 2+ [ 16 , 17 ]. Furthermore, sesame seeds contain bioactive components such as polyphenols, lignans, phytosterols, quinones and triterpenoids [ 18 ]. The present work investigates the differential the impact of NaCl salinity and water-withholding drought, at the same Ψ w , on growth and performance of two Egyptian sesame cultivars: Sohag 1 (Sohg) and Shandaweel 3 (Shnd). Partitioning of biomass, photosynthetic pigments and carbohydrate fractionations have been integrated with photosynthetic efficiency to achieve a plausible explanation of plant tolerance to abiotic stress. 2 Materials and Methods 2.1 Plant material Seeds of cultivars Sohag 1 (Sohg) and Shandaweel 3 (Shnd) of sesame ( Sesamum indicum L.) were obtained from the Agricultural Research Center (ARC), Egypt. The two experimental sesame cultivars are national cultivars, produced and released by Oil Crops Research Department, Field Crops Research Institute (FCRI), Agricultural Research Center (ARC), Ministry of Agricultural and Land Reclamation, Egypt according to The International Union for the Protection of New Varieties of Plants (UPOV) guideline (Egypt is a member of UPOV) and the Intellectual Property Rights (IPR) national low. The acquired names "Sohag 1" and "Shandaweel 3" are given by the breeder program of ARC and are related to national districts of Egypt. 2.2 Effect of salinity and drought on sesame growth and performance Seeds were selected for homogeneity, surface-sterilized in 10% (v/v) Clorox for 20 minutes, washed thoroughly with tap water and sown on plastic pots of 15 cm diameter and 20 cm height full of 1.1 kg of water-washed sand, five seeds per pot. Seeds were watered with 0.2 mM CaSO 4 for 7 days; thereafter, seedlings were successively thinned to one per pot and received the nutrient solution in a successive manner. First, seedlings were watered with a one-tenth strength nutrient solution for three days, followed by half-strength solution for six days and finally the full-strength solution for 13 days. The full-strength nutrient solution had the following composition: Macronutrients (mM): N (as NO 3 − ) 16, K 7.5, P 1.5, Ca 5, Mg 1.5 and S 1.5. Micronutrients (µM): Fe 50, Mn 5, Cu 0.5, Zn 0.5, B 25, Mo 0.25 and Na 50. After 13 days of irrigation with the full nutrient solution, plants of 31 days-old and eight true leaves were subjected to stress at four ψ w : 0, -100, -250 and − 450 kPa, induced either as NaCl stress or drought stress. The experimental water potentials corresponded to 0, 25, 61 and 110 mM NaCl, respectively for salinity stress and 100%, 23.6%, 14.1% and 10.2% field capacity, respectively or 20, 4.71, 2.83, 2.04 g water/100 g sand, respectively for drought stress. The water holding capacity of the sand used as growth medium was estimated as 20% (w/w). To achieve the required ψ w of salinity and drought stresses, the formula of [ 19 ] and the ψ w -water content curves of [ 20 ], respectively were consulted. To ensure homogenous supply of water and nutrients, irrigation was dispended over four evenly-spaced daily intervals, with the same dose of nutrients throughout. This implied proportional increase in strength of the nutrient solution as the supply of water decreased in drought stress. The nutrient solution was prepared as a stock solution so as 1 ml of which will provide the nutrient supply for one pot per day. The scheduled supply of nutrient solution, water (for drought stress) and NaCl (for salt stress) is shown in Table 1 . Plants were grown in the greenhouse at the Faculty of Science, Damietta University during the period from July to August 2023. Irradiance ranged from 1500 to 2000 µmol m − 2 s − 1 from natural sunlight, with a temperature of 38/25°C in a 14/10 h light/dark period and relative humidity of about 75% in average. Plants were harvested six days after application of stress, when they were 37 days old. Just before harvest, measurements of leaf angle, leaf dimensions, photosynthetic pigments and gas exchange were carried out in the third youngest leaf pair. Plants were thoroughly washed from sand, separated into leaves, stem and root, blotted gently and fresh weights were recorded. An aliquot of fresh leaves was kept frozen at -80°C for estimation of photosynthetic pigments and carbohydrate fractions. Dry weights were recorded after drying of the fresh plant material at 80°C for 48 h and were corrected for the leaf portions kept frozen. Table 1 Supply of nutrients, water and NaCl solution to the sand culture to obtain the experimental water potentials with the same dose of nutrients under the impact of salinity stress and drought stress kPa 0.00 -100 -250 -450 Salinity stress mM NaCl 0 25 61 110 ml water before addition of stocks 200 200 200 200 ml stock NS per pot per day 1.0 1.0 1.0 1.0 ml NaCl (2 M) per pot per day 0 2.75 6.71 12.1 Final volume (ml) 220 220 220 220 ml working solution per pot per 6 h 55 55 55 55 Drought stress Water content of sand (% DW) 20 4.71 2.83 2.04 Water content (% field capacity) 100 23.6 14.1 10.2 Stock N.S. (ml) per pot per day 1.0 1.0 1.0 1.0 Water (up to ml) per pot per day 220 51.92 31.02 22.44 ml working solution per pot per 6 h 55 12.98 7.76 5.61 2.3 Plant analysis 2.3.1 Estimation of photosynthetic pigments Photosynthetic pigments were determined according to the method described by [ 21 ]. Frozen leaf discs were macerated in 80% acetone using cold mortar and pestle in dim light with a pinch of magnesium carbonate to neutralize the plant acids. The slurry was centrifuged and the clear extract was brought up to volume with 80% acetone where the absorbance was read at 470, 646 and 663 nm using a UNICO 7200 series spectrophotometer. The concentrations of chlorophyll a (Chl a), chlorophyll b (Chl b) and carotenoids (Carot) were calculated (µg ml − 1 ) according to the following equations: Chl a = 12.21 E 663 – 2.81 E 646 Chl b = 20.13 E 646 – 5.03 E 663 Carot = \(\:\frac{(1000\:\text{E}470\: -\:3.27\:\text{C}\text{h}\text{l}\text{a}\: -\:104\:\text{C}\text{h}\text{l}\:\text{b})}{229}\) Concentrations were presented as µmol pigment g − 1 leaf FW. 2.3.2 Estimation of gas exchange parameters Photosynthesis rate (A), stomatal conductance to CO 2 (G s ), substomatal CO 2 concentration (C i ) and transpiration rate (E) of the third youngest leaf pair were measured at 10:00 a.m. using an LCA-4 portable gas exchange system (Analytical Development Company Ltd, England). Measurements were conducted with leaf area of 6.25 cm 2 , leaf chamber CO 2 concentration of 390 ppm at chamber temperature of 44°C and photosynthetic photon flux density of 1500 µmol m − 2 s − 1 . 2.3.3 Estimation of carbohydrate fractions Soluble sugars, starch, hemicellulose and pectic substances were sequentially assayed in the leaves. For extraction of soluble sugars, a known weight of the frozen leaf material was grinded to a fine powder in liquid nitrogen, then thoroughly extracted in 1.2 ml of 80% ethanol and kept at -4°C overnight [ 22 ]. The mixture was centrifuged at 8000 × g for 10 minutes; the supernatant was quantitatively transferred to clean Eppendorf tubes, evaporated to dryness at 70°C and re-dissolved in distilled water for determination of soluble sugars. An aliquot of the soluble carbohydrate extract was completed to 1 ml with distilled water, then carefully mixed with 3 ml of the anthrone reagent and heated at 80°C for 10 min in a water bath. The mixture was cooled for 30 min in an ice bath and absorbance was recorded at 623 nm against the reagent blank. The plant debris left after ethanolic extraction was used for determination of the insoluble carbohydrate fractions. For estimation of starch, the debris remained after extraction of soluble sugars was re-suspended in 1.5 ml of 3% HCl with boiling for 3 hours and the mixture was centrifuged at 8000 × g for 10 min [ 23 ]. The liberated glucose units were estimated by the anthrone method as mentioned above, and starch content was expressed as µg glucose equivalents (Gluc. eq.). For estimation of hemicellulose, the residue left after extraction of starch was re-suspended in 1.2 ml of 5% H 2 SO 4 , boiled for 2.5 hours and then centrifuged at 8000 × g for 10 min. The supernatant was quantitatively transferred to clean Eppendorf tubes, and the liberated glucose units were estimated by the anthrone method [ 24 ]. For estimation of pectic substances, the residue left after extraction of hemicellulose was re-suspended in 1ml of 72% H 2 SO 4 at 4°C for 24 hours, then centrifuged at 8000 × g for 10 min. The supernatant was quantitatively transferred to clean Eppendorf tubes, and the liberated glucose units were estimated by the anthrone method [ 24 ]. The anthrone reagent was prepared by dissolving 1.67 g anthrone in 1 l of 80% H 2 SO 4 (v/v) for 8.6 mM anthrone. The different carbohydrate fractions were estimated with reference to a glucose calibration curve in the range of 0–100 µg/ml. 2.4 Definitions and calculations The photosynthetic water use efficiency was defined as the photosynthesis rate (A)/transpiration rate (E) ratio [ 1 ]. RWR, StWR and LWR refer to the proportions of the plant dry matter allocated to root, stem and leaves respectively. Similarly, Chl a ratio, Chl b ratio and Carot ratio refer to the proportions of Chl a, Chl b and carotenoids, respectively in the total leaf pigment content. 2.5 Experimental design and statistical analysis The experiment was factorial with three factors and three replications in a completely randomized design. The main factors were: 1) sesame cultivar with two levels: Sohg and Shnd, 2) type of abiotic stress with two levels: salinity and drought and 3) severity of stress in terms of Ψ w of sand with four levels: 0, -100, -250 and − 450 kPa. Data were subjected to three-way ANOVA using SPSS version 22, and the effects of the main factors and their interactions were assessed. Mean separation was performed using the Duncan's multiple range test at P < 0.05. The correlations between plant growth and physiology were depicted in terms of the significant Pearson correlation coefficients. 3 Results 3.1 Plant growth The most affected growth attributes, justified by the magnitude of the F ratio, was the leaf angle, while the least affected one was leaf dimensions (Table 2 ). In absence of stress, Shnd exhibited horizontal leaf position relative to Sohg. The effect of stress on leaf angle was observed only in Sohg, which experienced 39% reduction by salt stress versus 52% increase by drought stress across the whole range of ψ w ; this in contrast to non-significant decrease and increase, respectively in Shnd (Fig. 1 a). Lowering ψ w of sand from 0 to -450 kPa led to progressive decrease in leaf length and width by 27% and 20%, respectively as an average for the two types of stress in Sohg but to non-significant reductions in Shnd (Fig. 1 b, c). Consequently, stress led to progressive mild increase in the leaf width/length ratio in Sohg but to a transient top at moderate stress, followed by a decrease at severe stress in Shnd (Fig. 1 d). Whereas the dry weights of root and leaves were comparable in the two cultivars irrespective of the stress regime, stem dry weight was higher in Sohg than Shnd in absence of stress but the reverse was true under stress. Lowering ψ w from 0 to -450 kPa reduced leaf dry weight of Sohg and Shnd by an average of 44% under salinity stress post a beneficial threshold salinity of -100 kPa, but drought stress led to a progressive average reduction of 42% for the two cultivars (Fig. 2 a). However, progression of salinity and drought stress from 0 to -450 kPa reduced stem dry weight by an average of 64% and 20% for Sohg and Shnd, respectively (Fig. 2 b). The lesser impact of salinity than drought on foliage biomass was manifested as the emergence of -100 kPa beneficial threshold salinity, along with less-aggressive reduction under moderate stress. Lowering ψ w from 0 to -450 kPa reduced root dry weight of Sohg and Shnd by an average of 65% post a threshold of -100 kPa under salinity stress but led to progressive increases of 38% and 75% in Sohg and Shnd, respectively under drought stress (Fig. 2 c). Shoot water content was non-significantly affected by genotype and stress except for the significant reduction only in Shnd under severe salinity. By contrast, the genotypic difference emerged in water content of root, with higher values in Sohg than Shnd particularly under stress, and stronger reduction under drought than salinity (Table 3 ). The differential impact of stress on dry weights of plant organs resulted in alteration of biomass partitioning among leaves, stem and root. Under salinity stress, Sohg increased allocation of biomass to leaves at the expense of stem and root; whereas Shnd increased allocation to stem at the expense of root with marginal increase in leaf proportion. By contrast, under drought stress both cultivars increased allocation of biomass to root at the expense of foliage (stem and leaves) (Fig. 3 ). Table 2 Three-way ANOVA showing the effect of the main factors: sesame cultivar (Cv), type of stress (Stress) and water potential of the medium (kPa) and their interactions on leaf dimensions, biomass and biomass partitioning and water content of plant organs Source of variation df Leaf angle Leaf length Leaf width Leaf W/L ratio Root water Shoot water F P F P F P F P F P F P Cv 1 83.55 0.000 0.611 0.440 0.361 0.552 3.121 0.087 7.651 0.009 0.378 0.543 Stress 1 10.16 0.003 0.126 0.725 0.361 0.552 0.269 0.608 39.24 0.000 0.499 0.485 kPa 3 0.709 0.554 11.16 0.000 3.252 0.034 3.003 0.045 11.18 0.000 4.894 0.007 Cv × Stress 1 5.491 0.025 1.823 0.186 2.373 0.133 0.488 0.490 1.574 0.219 1.775 0.192 Cv × kPa 3 0.919 0.443 1.715 0.184 1.053 0.383 1.291 0.294 2.576 0.071 3.063 0.042 Stress × kPa 3 5.133 0.005 0.423 0.738 0.065 0.978 0.430 0.733 20.02 0.000 1.536 0.224 Cv × Stress × kPa 3 2.150 0.113 0.880 0.462 0.627 0.603 0.375 0.772 1.297 0.293 2.771 0.058 Root dry weight Stem dry weight Leaf dry weight RWR StWR LWR F P F P F P F P F P F P Cv 1 0.498 0.486 0.827 0.370 0.691 0.412 1.360 0.252 4.987 0.033 1.043 0.315 Stress 1 96.52 0.000 21.73 0.000 48.02 0.000 272.2 0.000 25.20 0.000 69.17 0.000 kPa 3 1.368 0.270 21.99 0.000 30.42 0.000 10.45 0.000 4.264 0.012 0.683 0.569 Cv × Stress 1 0.778 0.384 0.282 0.599 0.036 0.851 0.002 0.963 0.643 0.428 0.428 0.518 Cv × kPa 3 1.610 0.206 8.508 0.000 1.172 0.336 1.358 0.273 5.591 0.003 1.950 0.141 Stress × kPa 3 41.62 0.000 7.303 0.001 14.22 0.000 48.01 0.000 4.181 0.013 13.78 0.000 Cv × Stress × kPa 3 0.167 0.918 0.442 0.724 0.371 0.775 0.016 0.997 1.086 0.369 0.945 0.431 Table 3 Effect of salt stress and drought stress on water content of shoot and root of cvs Sohg and Shnd of sesame. Each value is the mean of three replicates ± SE. Means with common letters are not significantly different at p ≤ 0.05 Stress and water potential (kPa) Shoot water (% FW) Root water (% FW) Sohg Salt stress 0 86.02 ± 0.10 a 91.17 ± 0.86 ab -100 87.22 ± 0.16 a 91.81 ± 0.87 ab -250 86.37 ± 0.39 a 92.03 ± 0.30 ab -450 86.63 ± 1.60 a 94.02 ± 1.26 a Drought stress 0 86.02 ± 0.10 a 91.17 ± 0.86 ab -100 87.40 ± 0.49 a 93.89 ± 0.09 a -250 86.02 ± 0.29 a 89.65 ± 1.30 bc -450 85.49 ± 0.27 a 84.79 ± 1.57 d Shnd Salt stress 0 86.69 ± 0.42 a 91.86 ± 0.56 ab -100 86.84 ± 0.44 a 92.40 ± 0.70 ab -250 88.17 ± 0.20 a 91.08 ± 0.89 abc -450 80.50 ± 3.44 b 91.35 ± 1.11 ab Drought stress 0 86.69 ± 0.42 a 91.86 ± 0.56 ab -100 87.43 ± 0.16 a 89.74 ± 0.31 bc -250 86.51 ± 0.64 a 88.02 ± 2.48 c -450 85.80 ± 1.08 a 81.62 ± 0.84 e 3.2 Photosynthetic pigments The contents of Chl a, Chl b and carotenoids were significantly higher in Sohg than Shnd; the genotypic difference was either most evident under mild stress (Chl a and Chl b) or was maitained under all levels of stress (carotenoids). The impact of salinity on photosynthetic pigments was more aggressive than that of drought, which exerted beneficial effect in Shnd. Increasing salinity from 0 to -450 kPa reduced the contents of Chl a, Chl b and carotenoids by 45%, 35% and 35%, respectively as averages for Sohg and Shnd; but whereas the reductions were progressive in Sohg, they occurred post a threshold of -100 kPa in Shnd (Fig. 4 ). Meanwhile, drought stress non-significantly affected pigment content of Sohg but led to 55% and 78% increases in Chla and Chl b contents, respectively in Shnd, without effect on carotenoid content (Fig. 4 ). Partitioning of photosynthetic pigments among Chl a, Chl b and carotenoids showed mild response to treatments, except for: 1) salinized Sohg–where carotenoid proportion was increased at the expense of Chl a, and 2) droughted Shnd–where chlorophyll proportion was increased at the expense of carotenoids (Fig. 5 ). 3.3 Gas exchange Table 4 reveals that the most affected gas exchange parameters were the rate of photosynthesis (A), followed by stomatal conductance (Gs) and transpiration rate (E) while the least affected parameter was substomatal CO 2 concentration (Ci). In contrast to the increased Ci by stress, E, Gs and A were lowered. The increases in Ci, across the whole range of ψ w , amounted to 54% and 39% in Sohg and Shnd, respectively (with − 100 kPa threshold under salinity stress) but were progressive with lower magnitude (33% and 14% in Sohg and Shnd, respectively) under drought stress (Fig. 6 a). By contrast, the reduction in E amounted to 77.5% under salinity stress and 47.5% under drought stress as an average for the two cultivars post a threshold of -100 kPa (Fig. 6 b). The genotypic gap in G s and A was marked, in favor of Sohg, in absence of stress but were narrowed under stress. Lowering ψ w from 0 to -450 kPa reduced Gs and A by an average of 85% under salinity stress and 57% under drought stress in the two cultivars post a threshold of -100 kPa (Fig. 6 c, d). The genotypic difference in water use efficiency (WUE) was marked, in absence of stress and under severe drought stress, in favor of Sohg. Increasing salinity from 0 to -450 kPa reduced WUE by an average of 47% for the two cultivars, either progressively in Sohg or post − 100 kPa threshold in Shnd. Meanwhile, the decrease in WUE across the whole range of ψ w under drought stress amounted to 3% in Sohg and 38% in Shnd post a threshold of -250 kPa (Fig. 6 e). 3.4 Sugar fractions ANOVA revealed that among the assayed sugar fractions, the most affected component was hemicelluloses followed by soluble sugars, whereas pectic substances were the least affected one (Tables 5 ). Soluble sugar (SS) content of leaves was markedly higher in Sohg than Shnd in absence of stress and under drought stress but turned in favor of Shnd under salinity stress. SS content of Sohg leaves experienced 58% and 38% reductions under salinity stress and drought stress, respectively upon lowering of ψ w from 0 to -450 kPa. Meanwhile, SS content of Shnd leaves experienced overall non-significant changes under stress, although approaching a minimum at -250 kPa (Fig. 7 a). In contrast to SS, the leaf content of polysaccharides (starch, hemicellulose and pectic substances) was higher in Shnd than Sohg in absence of stress; but the genotypic difference was either diminished or turned in favor of Sohg under stress. In Sohg, lowering ψ w post a threshold of -100 kPa down to -450 kPa reduced starch content of leaves by 51% as an average for salinity stress and drought stress; but in Shnd, the reduction was progressive across the whole range of ψ w , with stronger magnitude under salinity (72%) than under drought (56%) (Fig. 7 b). The low hemicellulose content of Sohg leaves was subjected to moderate changes under stress (either a reduction under salinity or an increase under drought). By contrast, the high hemicellulose content of Shnd leaves experienced 70% average reduction across the whole range of ψ w ; either progressively under drought or post a transient increase at -100 kPa under salinity (Fig. 7 c). Across the whole range of ψ w , salinity stress (on Sohg) as well as salinity and drought stress (on Shnd) led to an average 20% reduction in the leaf pectic substances; only drought stress led to non-significant increase in Sohg (Fig. 7 d). Starch constituted the greatest carbohydrate fraction of sesame leaves (> 50%), followed by soluble sugars and pectic substances (about 20% each), with minor hemicellulose component. In Sohg, mild salinity and drought stresses lowered SS ratio in favor of starch, but severe stresses led to the opposite effect; meanwhile the fraction of pectic substances or that of pectic substances and hemicellulose were progressively increase (Fig. 8 a, b). In Shnd, stress–particularly salinity–led to progressive increase in the ratios of SS and pectic substances at the expense of starch and hemicellulose (Fig. 8 c, d). Table 4 Three-way ANOVA showing the effect of the main factors: sesame cultivar (Cv), type of stress (Stress) and water potential of the medium (kPa) and their interactions on the content of photosynthetic pigments and gas exchange of sesame leaves Source of variation df Chl a Chl b Carot Chl a ratio Chl b ratio Carot ratio F P F P F P F P F P F P Cv 1 26.05 0.000 15.66 0.000 29.73 0.000 2.662 0.113 1.854 0.183 5.011 0.032 Stress 1 16.14 0.000 12.02 0.002 16.45 0.000 0.109 0.744 2.567 0.119 1.335 0.256 kPa 3 0.807 0.499 1.077 0.373 5.003 0.006 1.682 0.190 3.960 0.017 3.096 0.041 Cv × Stress 1 2.142 0.153 0.104 0.750 7.180 0.012 0.487 0.490 6.970 0.013 0.569 0.456 Cv × kPa 3 1.835 0.161 1.332 0.281 0.281 0.839 2.580 0.071 2.882 0.051 5.485 0.004 Stress × kPa 3 6.889 0.001 2.257 0.101 5.215 0.005 3.792 0.020 1.553 0.220 1.793 0.168 Cv × Stress × kPa 3 0.699 0.560 0.422 0.738 1.059 0.380 0.383 0.766 1.096 0.365 0.686 0.567 Substomatal CO 2 (Ci) Transpiration rate (E) Stomatal conductance (Gs) Photosyn-thesis rate (A) A/E ratio F P F P F P F P F P Cv 1 0.071 0.791 2.307 0.139 11.19 0.002 13.54 0.001 6.306 0.017 Stress 1 0.089 0.768 10.68 0.003 7.701 0.009 17.55 0.000 10.10 0.003 kPa 3 17.43 0.000 61.65 0.000 61.56 0.000 96.24 0.000 10.02 0.000 Cv × Stress 1 0.077 0.783 0.749 0.393 6.509 0.016 1.068 0.309 0.824 0.371 Cv × kPa 3 1.247 0.309 0.504 0.682 1.016 0.398 2.866 0.052 2.452 0.081 Stress × kPa 3 1.836 0.161 3.583 0.024 1.688 0.189 8.000 0.000 5.336 0.004 Cv × Stress × kPa 3 0.016 0.997 0.567 0.641 2.361 0.090 0.353 0.787 1.070 0.376 4 Discussion Understanding of the mechanisms, the plant can adopt to withstand drought and salinity stress is crucial for maintenance of crop productivity under unfavorable conditions. Stress effects on plant growth and performance are genotype-dependent but also depend on the severity and type of stress. Sesame, a moderately salt- and drought-tolerant crop [ 25 , 26 ] is often cultivated under arid and semi-arid conditions. Salinity and drought differentially affected shoot and root growth of sesame with more stress tolerance of cv. Shnd than Sohg. The less aggressive impact of salinity than drought on foliage growth of sesame can be viewed in light of the hypothesis that the accumulating salt ions can aid in lowering of water potential of plant tissues and hence allowing efficient water absorption under stress [ 27 ]. The beneficial effect of mild salinity to foliage growth and the lesser dehydrating effect of salinity Table 5 Three-way ANOVA showing the effect of the main factors: sesame cultivar (Cv), type of stress (Stress) and osmotic potential of the medium (kPa) and their interactions on the content of carbohydrate fractions of sesame leaves Source of variation df Soluble sugars Starch Hemicellulose Pectic substances F P F P F P F P Cv 1 31.48 0.000 0.243 0.625 181.9 0.000 2.811 0.103 Stress 1 0.642 0.429 0.513 0.479 4.795 0.036 6.783 0.014 kPa 3 27.92 0.000 79.90 0.000 58.64 0.000 11.74 0.000 Cv × Stress 1 4.540 0.041 4.050 0.053 64.88 0.000 1.715 0.200 Cv × kPa 3 13.18 0.000 13.63 0.000 64.70 0.000 7.530 0.001 Stress × kPa 3 0.216 0.885 3.138 0.039 4.750 0.008 3.307 0.032 Cv × Stress × kPa 3 2.298 0.096 1.709 0.185 8.668 0.000 0.699 0.559 Soluble sugars ratio Starch ratio Hemicellulose ratio Pectic substances ratio F P F P F P F P Cv 1 31.18 0.000 0.995 0.326 165.1 0.000 0.202 0.656 Stress 1 2.344 0.136 0.016 0.902 1.868 0.181 0.136 0.715 kPa 3 41.88 0.000 31.97 0.000 10.81 0.000 28.13 0.000 Cv × Stress 1 16.97 0.000 36.93 0.000 81.70 0.000 0.747 0.394 Cv × kPa 3 47.87 0.000 10.61 0.000 28.92 0.000 0.942 0.432 Stress × kPa 3 3.677 0.022 1.468 0.242 0.968 0.420 0.465 0.709 Cv × Stress × kPa 3 6.934 0.001 7.246 0.001 12.54 0.000 0.552 0.651 than drought on root is in line with this hypothesis. However, the stronger dehydrating effect of salinity than drought on shoot suggests different patterns of osmotic susceptibility between shoot and root. In contrast to its more aggressive effect on foliage biomass, drought improved root biomass of sesame. However, decreased root volume and root hydraulic conductivity of three wheat varieties by PEG-drought stress and NaCl-salinity stress has been reported [ 28 ]. The reduction in root volume along with increased root density by drought reported by [ 29 ] implies either less severe impact of drought on root biomass than on root volume or even enhanced root biomass production by drought. A functional equilibrium is always achieved by plants to balance their biomass allocation between root and shoot [ 1 ]. Accordingly, it is expected that plants invest more biomass in the root at the expense of foliage under water deficit to optimize acquisition of water and nutrients and to restrict transpirational water loss. The signaling processes regulating this balance may involve sugars, phytohormones and small RNAs [ 30 ]. The present work suggests that salinity favors allocation of plant biomass to the foliage at the expense of root while drought led to the opposite effect. However, this pattern was subjected to genotypic intervention; whereas the favored allocation of plant biomass to the shoot under salinity occurred in favor of either leaves (Sohg) or stem (Shnd), the favored biomass allocation to root under drought occurred at the expense of either stem and leaves (Sohg) or only leaves (Shnd). Drought has been claimed to increase root to shoot biomass ratio of plants [ 31 ] and specifically to favor allocation of plant biomass to root and stem at the expense of leaves [ 32 ]. By contrast, it has been claimed that root growth is less salt-sensitive than shoot growth, thus leading to increased root/shoot ratio of salinized plants, as has been reported by [ 33 ] and [ 34 ]. Hence, it seems that the effect of stress on biomass allocation is controversial and depends on the experimental conditions and genotype [ 35 ]. Leaf weight ratio has been reported to decrease in Medicago sativa [ 36 ] but to increase in Setaria viridis [ 37 ] by drought. In contrast to the inherent horizontal leaf orientation of Shnd with marginal effect of stress, the vertical leaf orientation of Sohg was differentially affected by salinity and drought; being more vertical under salinity but less vertical under drought. This fact lends support to the observation of preferred conservation of leaf biomass by Sohg and of stem biomass by Shand under stress. The positive correlations (under salinity stress) and the negative correlations (under drought stress) between leaf angle and the rates of photosynthesis and transpiration observed only in Sohg (Fig. 9 ) implies that, in the stress-sensitive Sohg, horizontal leaf orientation can improve gas exchange under salinity stress but vertical leaf orientation is appropriate under drought stress. Leaf orientation is an important factor in regulation of plant carbon–water–energy nexus to optimize photosynthesis and light interception under the impact of abiotic stress. However, it has been claimed that, in general, leaves are vertical in hot dry environments but more horizontal in mesic and light-limited environments [ 38 ]. Unfavorable environmental conditions can evoke leaf movements, such as reversible epinastic responses [ 39 ], which can aid in plant adaption to stress conditions via protecting the leaves from photo-inhibition by reducing the leaf area exposed to solar irradiation [ 40 ]. The differential effect of salt stress and drought stress on photosynthetic pigments of sesame leaves was manifested as stronger negative impact of salinity on Sohg than Shnd, versus a mild reduction (Sohg) or even improvement (Shnd) by drought. The increased photosynthetic pigments of Cynanchum acutum leaves under drought was attributed to drought-induced P deprivation [ 32 ]. Furthermore, abiotic stress can minimize the genotypic gap in chlorophyll content between sesame cultivars that is approximating the less green Shnd to the greener Sohg. The symptoms of salt stress on plants include chlorosis and scorching of leaves [ 41 ]. Chlorophyll content of leaves was decreased under both salinity and drought stress in Robinia pseudoacacia [ 42 ] and under drought in wheat [ 43 ], canola [ 44 ] and barley [ 45 ]. However, the effect of salinity on chlorophyll content of wheat leaves was genotype-dependent, being promotive in salt-tolerant genotypes but inhibitory in salt-sensitive genotypes [ 46 ]. The effect of stress on partitioning of photosynthetic pigments among Chl a, Chl b and carotenoids was particularly evident in Shnd, in which stress, can increase Chl a proportion at the expense of carotenoids. In accordance with the present findings, Chl a and Chl b contents were subjected to differential impact of salinity in rice leaves [ 47 ] and of drought in wheat [ 43 ]. The present findings suggest stronger impact of salinity than drought on gas exchange of sesame leaves, a pattern that contradicts the response of foliage growth. Photosynthesis rate (A), transpiration rate (E) and stomatal conductance (Gs) are positively related to water potential of the medium [ 48 ]. Lowered photosynthesis under stress is usually linked to stomatal closure but can also result from impaired biochemical and photochemical constraints [ 49 ]. The intimate association between the patterns of retardation in A, E and Gs under the impact of salinity and drought indicates that the reduced photosynthetic activity under stress can be attributed to the stress-induced stomatal closure and lowering of stomatal conductance (Fig. 9 ). The fact that A, E and Gs are positively correlated with each other but negatively correlated with Ci, along with the coincidence of minimal Ci with maximal A and Gs at mild salinity (Fig. 6 ) implies that stress can inhibit photosynthetic efficiency without effect on or even enhancement of respiration but mild salinity can enhance photosynthetic efficiency through increasing stomatal conductance. In agreement with the present findings, drought stress has been found to induce stomatal closure with consequent severe inhibition in photosynthesis along with mild inhibition or even promotion in respiration [ 50 , 32 ]. Stomatal movement under stress can control photosynthesis and transpiration in a way to increase water use efficiency. The present findings suggest that cv. Sohg with high photosynthesis potential exhibited marked drought robustness; in contrast to cv. Shnd of low photosynthesis potential, which although benefited from mild stress experienced more susceptibility to severe drought. Transpiration rate, in accordance with Gs, benefited from mild salinity in Shnd and mild drought in Sohg but was reduced under severe stress, with stronger impact of salinity than drought. According to the present finding, abiotic stress seems to reduce the water use efficiency (A/E ratio) of sesame with stronger impact of salinity on Sohg and of drought on Shnd. It has been reported that water-use efficiency was increased in Robinia pseudoacacia under salinity and drought stress ([ 42 ] and in Vicia faba [ 34 ] under salinity stress. The lowered water use efficiency of sesame under severe drought, observed in the present work, can be attributed either to greater sensitivity of the photosynthetic machinery than transpiration to water deficit or to progression of water loss despite of stomatal closure, probably via cuticular transpiration as proposed by [ 51 ]. Stomatal closure is a physiological adaptation to water deficit to avoid desiccation arising from transpirational water loss [ 52 ]; but the closed stomata will certainly inhibit photosynthesis by cutting down the influx of CO 2 to the mesophyll with the probability of continuation of water loss via alternative avenues such as the cuticular transpiration. It has been claimed that abiotic stress affects cell wall composition and production of compatible solutes [ 53 ]. The inhibited photosynthesis, concomitant with the unaffected or favored respiration under abiotic stress results in lowering of carbohydrate reserves of sesame leaves, which can be assigned to the decline in the major carbohydrate fraction (starch), followed by soluble sugars with marginal changes in pectic substances. However, the two sesame cultivars adopted different patterns of carbohydrate fractionation in response to stress. In Sohg, abiotic stress induced reductions in proportions soluble sugars and starch in favor of either pectic substances (salinity) or pectic substances and hemicellulose (drought). By contrast, in Shnd, abiotic stress increased the proportions of soluble sugars and pectic substances at the expense of starch and hemicellulose. Thus, the shared stress-induced response, irrespective of the genotype and type of stress, is the reduced proportion of starch in favor of pectic substances. The increased proportions of soluble sugars and pectic substances in Shnd under the impact of drought stress can aid in drought tolerance by contributing either to osmotic adjustment (soluble sugars) or by increasing cell wall elasticity (pectic substances). Osmotic and elastic adjustment are alternative strategies for a species to acclimate to water stress. Cells with highly elastic walls contain more water at full turgor; hence, their volume can decrease more before the turgor-loss point is reached. Elasticity of the cell wall depends on the chemical interactions between cell wall components [ 1 ]. It has been proposed that pectins, the major polyanionic polysaccharide in the walls of growing cells, play a critical role in cell wall expansion [ 54 ]. However, the role of soluble sugars in osmotic adjustment under stress is evident only in Shnd, despite the stronger impact of salinity and drought on leaf carbohydrate content of Shnd than Sohg. Conclusions The difference in stress-resistance was marked between the two sesame cultivars, with stronger aggressive impact of drought than salinity on foliage growth, which points to the role of salt ions in facilitating water absorption via lowering of water potential of foliage. Salinity shifted allocation of plant biomass to the foliage (specifically leaves of Sohg and stem of Shnd) at the expense of root; meanwhile, drought favors allocation of plant biomass to the root at the expense of stem of Sohg and leaf of Shnd. Thus, despite the contrasting effect of salinity and drought on rootage, Sohg seems to conserves leaves under stress but Shnd conserves stem. In a way to improve gas exchange, the sensitive Sohg adopts horizontal leaf orientation under salinity stress but vertical leaf orientation under drought stress. Abiotic stress narrowed the genotypic gap in chlorophyll content between sesame cultivars that is approximating the less green Shnd to the greener Sohg. In contrast to the influence on foliage growth, the impact of salinity on gas exchange of leaves was stronger than that of drought. The reduced photosynthetic activity under stress is related to stomatal closure and was accompanied with either no effect on or enhancement of respiration; however, mild salinity can enhance photosynthesis through increasing stomatal conductance. The photosynthetically less efficient genotype (Shnd) was more impacted by drought but at the same time, benefited from mild stress. Both E and Gs, benefited from mild salinity in Shnd and mild drought in Sohg but were subjected to stronger impact of salinity than drought. Stress reduced water use efficiency of sesame with stronger impact of salinity on Sohg and of drought on Shnd. The lowered water use efficiency can be attributed either to greater sensitivity of the photosynthetic machinery than transpiration to water deficit or to progression of water loss despite of stomatal closure, probably via cuticular transpiration. The stress-induced lowering in leaf carbohydrate reserves can be assigned to the decline in starch, either along with soluble sugars in favor of pectic substances (in Sohg) or of starch in favor of soluble sugars and pectic substances (in Shnd). The increased proportions of soluble sugars and pectic substances in Shnd under drought can aid in drought tolerance by contributing either to osmotic adjustment (soluble sugars) or to increased cell wall elasticity (pectic substances). Declarations Acknowledgments The authors acknowledge the Oil Crops Research Department, Field Crops Research Institute (FCRI), Agricultural Research Center at Giza, Egypt for providing seeds of the two sesame cultivars. Funding This work was supported by a research grant from Damietta University, Egypt as a part of M.Sc. program (Grant No. 1259/11/2021). Open access funding provided by The Science, Technology & Innovation Funding Authority (STDF) in cooperation with The Egyptian Knowledge Bank (EKB). Author contributions Conceptualization: TME; Methodology: SNA, SHA; Formal analysis and investigation: SNA, NMH; Writing - original draft preparation: SNA, TME; Writing - review and editing: TME, NMH; Funding acquisition: SHA; Resources: SNA; Supervision: TME, NMH. All authors read and approved the final manuscript. Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Data availability All data of this study are cited within the paper; raw data and calculations will be available on request. Permission to collect the seeds The manuscript is a purely eco-physiological study on performance of two sesame ( Sesamum indicum L.) cultivars and does not include any human participants, groups of humans or animal subjects. Seeds of cultivars Sohag 1 (Sohg) and Shandaweel 3 (Shnd) of sesame ( Sesamum indicum L.) were obtained from the Agricultural Research Center (ARC), Egypt. The two experimental sesame cultivars are national cultivars, produced and released by Oil Crops Research Department, Field Crops Research Institute (FCRI), Agricultural Research Center (ARC), Ministry of Agricultural and Land Reclamation, Egypt according to The International Union for the Protection of New Varieties of Plants (UPOV) guideline (Egypt is a member of UPOV) and the Intellectual Property Rights (IPR) national low. The acquired names "Sohag 1" and "Shandaweel 3" are given by the breeder program of ARC and are related to national districts of Egypt. Ethics declaration: not applicable . 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Each value is the mean of three replicates ± SE\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/9c1543d9ab4ed5d0d6aa8713.png"},{"id":98209160,"identity":"788113b1-b92e-4b97-b46e-dbfaa932f52e","added_by":"auto","created_at":"2025-12-15 09:13:23","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":36996,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of salt stress and drought stress on dry weight of leaves (a), stem (b) and root (c) of cvs Sohg and Shnd of sesame. Each value is the mean of three replicates ± SE\u003c/p\u003e\n\u003cp\u003eTable 3 Effect of salt stress and drought stress on water content of shoot and root of cvs Sohg and Shnd of sesame. Each value is the mean of three replicates ± SE. Means with common letters are not significantly different at p ≤ 0.05\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/322affae85f15d720281fc96.png"},{"id":98432154,"identity":"3383d16a-71bd-48b7-b562-0ec8276bf1b9","added_by":"auto","created_at":"2025-12-17 16:49:07","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":28246,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of salt stress and drought stress on biomass partitioning among leaves, stem and root of Sohg NaCl (a), Sohg drought (b), Shnd NaCl (c) and Shnd drought (d) of sesame. Each value is the mean of three replicates\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/0562f363d6323cbab6d51787.png"},{"id":98209162,"identity":"7a4b7409-4334-4310-82e5-24b36b1d0e93","added_by":"auto","created_at":"2025-12-15 09:13:23","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":35527,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of salt stress and drought stress on the content of Chl a (a), Chl b (b) and Carot (c) of cvs Sohg and Shnd of sesame leaves. Each value is the mean of three replicates ± SE\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/9403e53f508d87516b2c85fd.png"},{"id":98433323,"identity":"034ea1fd-3f4f-43bb-a034-19a3c14f33bd","added_by":"auto","created_at":"2025-12-17 16:50:37","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":22940,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of salt stress and drought stress on pigment partitioning among Chl a, Chl b and Carot of Sohg NaCl (a), Sohg drought (b), Shnd NaCl (c) and Shnd drought (d) of sesame. Each value is the mean of three replicates\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/2eab95237a8c0e496bec3c62.png"},{"id":98430849,"identity":"eac1cb55-8063-41b9-bf59-5e32a2519773","added_by":"auto","created_at":"2025-12-17 16:46:21","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":64186,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of salt stress and drought stress on substomatal [CO\u003csub\u003e2\u003c/sub\u003e] (a), transpiration rate (b), stomatal conductance (c), photosynthesis rate (d) and water use efficiency in terms of the A/E ratio (e) of cvs Sohg and Shnd of sesame. Each value is the mean of three replicates ± SE\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/0dca4b7f7441beca6d64c4df.png"},{"id":98209169,"identity":"736cddeb-625b-45ab-9aec-5f74b28c945c","added_by":"auto","created_at":"2025-12-15 09:13:23","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":51968,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of salt stress and drought stress on the content of soluble sugars (a), starch (b), hemicellulose (c) and pectic substances (d) in the leaves of cvs Sohg and Shnd of sesame. Each value is the mean of three replicates ± SE\u003c/p\u003e","description":"","filename":"7.png","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/585db9d3f90a9e9820877c75.png"},{"id":98209171,"identity":"32f2235c-850e-491c-9442-a69dab3f04a5","added_by":"auto","created_at":"2025-12-15 09:13:23","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":23227,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of salt stress and drought stress on carbohydrate partitioning among soluble sugars, starch, hemicellulose and pectic substances in the leaves of Sohg NaCl (A), Sohg drought (B), Shnd NaCl (C) and Shnd drought (D) of sesame. Each value is the mean of three replicates\u003c/p\u003e","description":"","filename":"8.png","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/8c602095bdeeb9f03ffb4236.png"},{"id":98431850,"identity":"f6e3add3-bfee-40c4-ae73-130ea39583ad","added_by":"auto","created_at":"2025-12-17 16:48:28","extension":"png","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":193446,"visible":true,"origin":"","legend":"\u003cp\u003eCorrelation analysis showing the relationships between gas exchange measurements, leaf angle (θ) and leaf biomass of cvs. Sohg and Shnd of sesame under salinity stress and drought stress. \u003cstrong\u003e+ + +\u003c/strong\u003e and \u003cstrong\u003e- - -\u003c/strong\u003e denote very highly significant (P\u0026lt;0.001) positive and negative correlation, respectively; \u003cstrong\u003e+ +\u003c/strong\u003e and \u003cstrong\u003e- -\u003c/strong\u003e highly significant (P\u0026lt;0.01) correlation, \u003cstrong\u003e+\u003c/strong\u003e and - just significant (P\u0026lt;0.05) correlation\u003c/p\u003e","description":"","filename":"9.png","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/70de481829739a951ebcc210.png"},{"id":104402078,"identity":"0e601e51-9432-428f-b150-9bd31bfc7c5b","added_by":"auto","created_at":"2026-03-11 12:14:13","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1853905,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7362732/v1/15321a8a-7139-4c2f-951c-72982cede0c9.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Salinity stress and drought stress differentially affect biomass allocation, leaf orientation and performance of two sesame (Sesamum indicum L.) cultivars","fulltext":[{"header":"1 Introduction","content":"\u003cp\u003eIn natural habitats, plants may experience combined stresses simultaneously such as the combined stress of high temperature, water deficit and high irradiance in the hot dry summer of arid habitats [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. Among the different abiotic stresses affecting plants, salinity stress and water stress are particularly serious and mostly interrelated. Mostly, saline soils are abundant in semiarid and arid regions where rainfall is insufficient for leaching of excessive soluble salts out of the rhizosphere [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. In turn, irrigation with poor quality water, a practice common in the arid developing countries, can induce secondary salinization. Soil is considered saline when electrical conductivity of its saturation extract exceeds 4 dS m\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (equivalent to 40 mM NaCl) with an exchangeable sodium percentage lower than 15 [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eAlthough the majority of crop plans are glycophytes, few species are salt-tolerant and can even benefit from mild salinity. Justification of crop salt response relies upon two criteria: the threshold salinity and the slope of the salinity-growth response line beyond the threshold. Differences in salt tolerance can occur among different species and cultivars as well as for the different plant developmental stages [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]. Similarly, drought stress represents a major challenge to plant productivity and land use, particularly in the arid regions. Even in the tropical rain forests, land plants can experience different degrees of transient water stress when water absorption lags behind evapotranspiration [\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e]. The witnessed global warming will further aggravate the threat of drought to plant productivity. Intrinsically, plants have low water use efficiency, with \u0026lt;\u0026thinsp;1% of the absorbed water being invested in the biomass versus \u0026gt;\u0026thinsp;90% of the absorbed mineral nutrients [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. Water is translocated within the plant down a gradient of 1) water potential (Ψ\u003csub\u003ew\u003c/sub\u003e) across membranes from the soil to the root and throughout the root, 2) hydrostatic pressure in the xylem vessels and 3) water vapor gradient from the leaf to air.\u003c/p\u003e\u003cp\u003eUnder the impact of salinity, plants suffer from several hazards including osmotic stress, ion toxicity, nutrient imbalance and oxidative stress; the relative effect of either factor depends on the level, duration and mode of exposure to salt stress as well as the plant developmental stage. For example, long-term salinity stress leads mainly to ion toxicity in the older leaves but to water deficit and shortage of carbohydrates in the younger leaves [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e]. Salinity and drought cause morphological, physiological, biochemical and molecular changes in the plant. To cope with stress, the plant has to expend additional energy for ion exclusion or compartmentalization (salinity stress) as well as for osmotic adjustment and detoxification of reactive oxygen species (salinity and drought stresses), which will slow down plant growth [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]. As a rule, shoot growth is more depressed by salinity and drought than root growth, partially because of favored biomass allocation to roots at the expense of shoots [\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eIn addition to the direct toxicity of high levels of Na\u003csup\u003e+\u003c/sup\u003e and Cl\u003csup\u003e-\u003c/sup\u003e to the cytoplasm, salt toxicity may also be attributed to interference with respiration, photosynthesis and protein synthesis. Salinity can cause insufficient cellular K\u003csup\u003e+\u003c/sup\u003e concentrations for enzyme activation, protein synthesis, photosynthesis and stomatal movement [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e]. Hence, the ability of plants to protect enzymes from the adverse effects of Na\u003csup\u003e+\u003c/sup\u003e or Cl\u003csup\u003e-\u003c/sup\u003e can be attributed to the synthesis of compatible solutes such as sugars and amino acids (e.g. proline) that can stabilize the structure of membranes and macromolecules [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. The role of compatible solutes in plant protection is also evident under drought stress, where plant cells adjust Ψ\u003csub\u003ew\u003c/sub\u003e by accumulating inorganic ions and organic acids in the vacuoles, with the synthesis of compatible solutes such as glycine betaine, sorbitol, and proline in the cytoplasm. The salinity-induced reduction in photosynthesis can be attributed to the reductions in leaf area (a capacity factor) and net CO\u003csub\u003e2\u003c/sub\u003e fixation per unit leaf area (an efficiency factor) in addition to the decreased demand of sink tissues for assimilates [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]. Further, the effect of salinity on photosynthetic efficiency includes both stomatal and nonstomatal limitations. Stomatal limitations are manifested as lowered intercellular CO\u003csub\u003e2\u003c/sub\u003e concentration while nonstomatal factors include the decrease in quantum efficiency and inhibited Ribusco activity [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]. Stomatal closure and the lowered stomatal conductance represent a way to prevent excessive dehydration of the leaves as a feedforward response to anticipate water deficit before the development of severe drought.\u003c/p\u003e\u003cp\u003eSesame (\u003cem\u003eSesamum indicum\u003c/em\u003e L.), a Pedaliaceae member and one of the most valuable oil cash crops in Africa and Asia [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e], is a robust crop with wide niche and marked drought tolerance. The high oil content of sesame seeds (up to 63%) is characterized with high proportion of unsaturated fatty acids, particularly omega 3 and γ-tocopherol [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e]. Sesame seeds are also rich in human-compatible proteins, vitamins, dietary fibers and dietary minerals, particularly Ca\u003csup\u003e2+\u003c/sup\u003e [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. Furthermore, sesame seeds contain bioactive components such as polyphenols, lignans, phytosterols, quinones and triterpenoids [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]. The present work investigates the differential the impact of NaCl salinity and water-withholding drought, at the same Ψ\u003csub\u003ew\u003c/sub\u003e, on growth and performance of two Egyptian sesame cultivars: Sohag 1 (Sohg) and Shandaweel 3 (Shnd). Partitioning of biomass, photosynthetic pigments and carbohydrate fractionations have been integrated with photosynthetic efficiency to achieve a plausible explanation of plant tolerance to abiotic stress.\u003c/p\u003e"},{"header":"2 Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003e2.1 Plant material\u003c/h2\u003e\u003cp\u003eSeeds of cultivars Sohag 1 (Sohg) and Shandaweel 3 (Shnd) of sesame (\u003cem\u003eSesamum indicum\u003c/em\u003e L.) were obtained from the Agricultural Research Center (ARC), Egypt. The two experimental sesame cultivars are national cultivars, produced and released by Oil Crops Research Department, Field Crops Research Institute (FCRI), Agricultural Research Center (ARC), Ministry of Agricultural and Land Reclamation, Egypt according to The International Union for the Protection of New Varieties of Plants (UPOV) guideline (Egypt is a member of UPOV) and the Intellectual Property Rights (IPR) national low. The acquired names \"Sohag 1\" and \"Shandaweel 3\" are given by the breeder program of ARC and are related to national districts of Egypt.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e\u003ch2\u003e2.2 Effect of salinity and drought on sesame growth and performance\u003c/h2\u003e\u003cp\u003eSeeds were selected for homogeneity, surface-sterilized in 10% (v/v) Clorox for 20 minutes, washed thoroughly with tap water and sown on plastic pots of 15 cm diameter and 20 cm height full of 1.1 kg of water-washed sand, five seeds per pot. Seeds were watered with 0.2 mM CaSO\u003csub\u003e4\u003c/sub\u003e for 7 days; thereafter, seedlings were successively thinned to one per pot and received the nutrient solution in a successive manner. First, seedlings were watered with a one-tenth strength nutrient solution for three days, followed by half-strength solution for six days and finally the full-strength solution for 13 days. The full-strength nutrient solution had the following composition:\u003c/p\u003e\u003cp\u003eMacronutrients (mM): N (as NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e) 16, K 7.5, P 1.5, Ca 5, Mg 1.5 and S 1.5.\u003c/p\u003e\u003cp\u003eMicronutrients (\u0026micro;M): Fe 50, Mn 5, Cu 0.5, Zn 0.5, B 25, Mo 0.25 and Na 50.\u003c/p\u003e\u003cp\u003eAfter 13 days of irrigation with the full nutrient solution, plants of 31 days-old and eight true leaves were subjected to stress at four ψ\u003csub\u003ew\u003c/sub\u003e: 0, -100, -250 and \u0026minus;\u0026thinsp;450 kPa, induced either as NaCl stress or drought stress. The experimental water potentials corresponded to 0, 25, 61 and 110 mM NaCl, respectively for salinity stress and 100%, 23.6%, 14.1% and 10.2% field capacity, respectively or 20, 4.71, 2.83, 2.04 g water/100 g sand, respectively for drought stress. The water holding capacity of the sand used as growth medium was estimated as 20% (w/w). To achieve the required ψ\u003csub\u003ew\u003c/sub\u003e of salinity and drought stresses, the formula of [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e] and the ψ\u003csub\u003ew\u003c/sub\u003e-water content curves of [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e], respectively were consulted.\u003c/p\u003e\u003cp\u003eTo ensure homogenous supply of water and nutrients, irrigation was dispended over four evenly-spaced daily intervals, with the same dose of nutrients throughout. This implied proportional increase in strength of the nutrient solution as the supply of water decreased in drought stress. The nutrient solution was prepared as a stock solution so as 1 ml of which will provide the nutrient supply for one pot per day. The scheduled supply of nutrient solution, water (for drought stress) and NaCl (for salt stress) is shown in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e\u003cp\u003ePlants were grown in the greenhouse at the Faculty of Science, Damietta University during the period from July to August 2023. Irradiance ranged from 1500 to 2000 \u0026micro;mol m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e from natural sunlight, with a temperature of 38/25\u0026deg;C in a 14/10 h light/dark period and relative humidity of about 75% in average. Plants were harvested six days after application of stress, when they were 37 days old. Just before harvest, measurements of leaf angle, leaf dimensions, photosynthetic pigments and gas exchange were carried out in the third youngest leaf pair. Plants were thoroughly washed from sand, separated into leaves, stem and root, blotted gently and fresh weights were recorded. An aliquot of fresh leaves was kept frozen at -80\u0026deg;C for estimation of photosynthetic pigments and carbohydrate fractions. Dry weights were recorded after drying of the fresh plant material at 80\u0026deg;C for 48 h and were corrected for the leaf portions kept frozen.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eSupply of nutrients, water and NaCl solution to the sand culture to obtain the experimental water potentials with the same dose of nutrients under the impact of salinity stress and drought stress\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"5\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c5\" namest=\"c2\"\u003e\u003cp\u003ekPa\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-100\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-250\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-450\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSalinity stress\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003emM NaCl\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e25\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e61\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e110\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eml water before addition of stocks\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e200\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e200\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e200\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e200\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eml stock NS per pot per day\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eml NaCl (2 M) per pot per day\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.75\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.71\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e12.1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eFinal volume (ml)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e220\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e220\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e220\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e220\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eml working solution per pot per 6 h\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e55\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eDrought stress\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eWater content of sand (% DW)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.71\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2.83\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.04\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eWater content (% field capacity)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e100\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e23.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e14.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10.2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStock N.S. (ml) per pot per day\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eWater (up to ml) per pot per day\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e220\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e51.92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e31.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e22.44\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eml working solution per pot per 6 h\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e12.98\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e7.76\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5.61\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec5\" class=\"Section2\"\u003e\u003ch2\u003e2.3 Plant analysis\u003c/h2\u003e\u003cdiv id=\"Sec6\" class=\"Section3\"\u003e\u003ch2\u003e2.3.1 Estimation of photosynthetic pigments\u003c/h2\u003e\u003cp\u003ePhotosynthetic pigments were determined according to the method described by [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]. Frozen leaf discs were macerated in 80% acetone using cold mortar and pestle in dim light with a pinch of magnesium carbonate to neutralize the plant acids. The slurry was centrifuged and the clear extract was brought up to volume with 80% acetone where the absorbance was read at 470, 646 and 663 nm using a UNICO 7200 series spectrophotometer. The concentrations of chlorophyll a (Chl a), chlorophyll b (Chl b) and carotenoids (Carot) were calculated (\u0026micro;g ml\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) according to the following equations:\u003c/p\u003e\u003cp\u003eChl a\u0026thinsp;=\u0026thinsp;12.21 E\u003csub\u003e663\u003c/sub\u003e \u0026ndash; 2.81 E\u003csub\u003e646\u003c/sub\u003e\u003c/p\u003e\u003cp\u003eChl b\u0026thinsp;=\u0026thinsp;20.13 E\u003csub\u003e646\u003c/sub\u003e \u0026ndash; 5.03 E\u003csub\u003e663\u003c/sub\u003e\u003c/p\u003e\u003cp\u003eCarot = \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{(1000\\:\\text{E}470\\: -\\:3.27\\:\\text{C}\\text{h}\\text{l}\\text{a}\\: -\\:104\\:\\text{C}\\text{h}\\text{l}\\:\\text{b})}{229}\\)\u003c/span\u003e\u003c/span\u003e\u003c/p\u003e\u003cp\u003eConcentrations were presented as \u0026micro;mol pigment g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e leaf FW.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec7\" class=\"Section3\"\u003e\u003ch2\u003e2.3.2 Estimation of gas exchange parameters\u003c/h2\u003e\u003cp\u003ePhotosynthesis rate (A), stomatal conductance to CO\u003csub\u003e2\u003c/sub\u003e (G\u003csub\u003es\u003c/sub\u003e), substomatal CO\u003csub\u003e2\u003c/sub\u003e concentration (C\u003csub\u003ei\u003c/sub\u003e) and transpiration rate (E) of the third youngest leaf pair were measured at 10:00 a.m. using an LCA-4 portable gas exchange system (Analytical Development Company Ltd, England). Measurements were conducted with leaf area of 6.25 cm\u003csup\u003e2\u003c/sup\u003e, leaf chamber CO\u003csub\u003e2\u003c/sub\u003e concentration of 390 ppm at chamber temperature of 44\u0026deg;C and photosynthetic photon flux density of 1500 \u0026micro;mol m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec8\" class=\"Section3\"\u003e\u003ch2\u003e2.3.3 Estimation of carbohydrate fractions\u003c/h2\u003e\u003cp\u003eSoluble sugars, starch, hemicellulose and pectic substances were sequentially assayed in the leaves. For extraction of soluble sugars, a known weight of the frozen leaf material was grinded to a fine powder in liquid nitrogen, then thoroughly extracted in 1.2 ml of 80% ethanol and kept at -4\u0026deg;C overnight [\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]. The mixture was centrifuged at 8000 \u003cem\u003e\u0026times; g\u003c/em\u003e for 10 minutes; the supernatant was quantitatively transferred to clean Eppendorf tubes, evaporated to dryness at 70\u0026deg;C and re-dissolved in distilled water for determination of soluble sugars. An aliquot of the soluble carbohydrate extract was completed to 1 ml with distilled water, then carefully mixed with 3 ml of the anthrone reagent and heated at 80\u0026deg;C for 10 min in a water bath. The mixture was cooled for 30 min in an ice bath and absorbance was recorded at 623 nm against the reagent blank.\u003c/p\u003e\u003cp\u003eThe plant debris left after ethanolic extraction was used for determination of the insoluble carbohydrate fractions. For estimation of starch, the debris remained after extraction of soluble sugars was re-suspended in 1.5 ml of 3% HCl with boiling for 3 hours and the mixture was centrifuged at 8000 \u003cem\u003e\u0026times; g\u003c/em\u003e for 10 min [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]. The liberated glucose units were estimated by the anthrone method as mentioned above, and starch content was expressed as \u0026micro;g glucose equivalents (Gluc. eq.). For estimation of hemicellulose, the residue left after extraction of starch was re-suspended in 1.2 ml of 5% H\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e, boiled for 2.5 hours and then centrifuged at 8000 \u003cem\u003e\u0026times; g\u003c/em\u003e for 10 min. The supernatant was quantitatively transferred to clean Eppendorf tubes, and the liberated glucose units were estimated by the anthrone method [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. For estimation of pectic substances, the residue left after extraction of hemicellulose was re-suspended in 1ml of 72% H\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e at 4\u0026deg;C for 24 hours, then centrifuged at 8000 \u003cem\u003e\u0026times; g\u003c/em\u003e for 10 min. The supernatant was quantitatively transferred to clean Eppendorf tubes, and the liberated glucose units were estimated by the anthrone method [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. The anthrone reagent was prepared by dissolving 1.67 g anthrone in 1 l of 80% H\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e (v/v) for 8.6 mM anthrone. The different carbohydrate fractions were estimated with reference to a glucose calibration curve in the range of 0\u0026ndash;100 \u0026micro;g/ml.\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\u003ch2\u003e2.4 Definitions and calculations\u003c/h2\u003e\u003cp\u003eThe photosynthetic water use efficiency was defined as the photosynthesis rate (A)/transpiration rate (E) ratio [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. RWR, StWR and LWR refer to the proportions of the plant dry matter allocated to root, stem and leaves respectively. Similarly, Chl a ratio, Chl b ratio and Carot ratio refer to the proportions of Chl a, Chl b and carotenoids, respectively in the total leaf pigment content.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e\u003ch2\u003e2.5 Experimental design and statistical analysis\u003c/h2\u003e\u003cp\u003eThe experiment was factorial with three factors and three replications in a completely randomized design. The main factors were: 1) sesame cultivar with two levels: Sohg and Shnd, 2) type of abiotic stress with two levels: salinity and drought and 3) severity of stress in terms of Ψ\u003csub\u003ew\u003c/sub\u003e of sand with four levels: 0, -100, -250 and \u0026minus;\u0026thinsp;450 kPa. Data were subjected to three-way ANOVA using SPSS version 22, and the effects of the main factors and their interactions were assessed. Mean separation was performed using the Duncan's multiple range test at P\u0026thinsp;\u0026lt;\u0026thinsp;0.05. The correlations between plant growth and physiology were depicted in terms of the significant Pearson correlation coefficients.\u003c/p\u003e\u003c/div\u003e"},{"header":"3 Results","content":"\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003e3.1 Plant growth\u003c/h2\u003e\u003cp\u003eThe most affected growth attributes, justified by the magnitude of the F ratio, was the leaf angle, while the least affected one was leaf dimensions (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In absence of stress, Shnd exhibited horizontal leaf position relative to Sohg. The effect of stress on leaf angle was observed only in Sohg, which experienced 39% reduction by salt stress versus 52% increase by drought stress across the whole range of ψ\u003csub\u003ew\u003c/sub\u003e; this in contrast to non-significant decrease and increase, respectively in Shnd (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea). Lowering ψ\u003csub\u003ew\u003c/sub\u003e of sand from 0 to -450 kPa led to progressive decrease in leaf length and width by 27% and 20%, respectively as an average for the two types of stress in Sohg but to non-significant reductions in Shnd (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb, c). Consequently, stress led to progressive mild increase in the leaf width/length ratio in Sohg but to a transient top at moderate stress, followed by a decrease at severe stress in Shnd (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ed).\u003c/p\u003e\u003cp\u003eWhereas the dry weights of root and leaves were comparable in the two cultivars irrespective of the stress regime, stem dry weight was higher in Sohg than Shnd in absence of stress but the reverse was true under stress. Lowering ψ\u003csub\u003ew\u003c/sub\u003e from 0 to -450 kPa reduced leaf dry weight of Sohg and Shnd by an average of 44% under salinity stress post a beneficial threshold salinity of -100 kPa, but drought stress led to a progressive average reduction of 42% for the two cultivars (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). However, progression of salinity and drought stress from 0 to -450 kPa reduced stem dry weight by an average of 64% and 20% for Sohg and Shnd, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb). The lesser impact of salinity than drought on foliage biomass was manifested as the emergence of -100 kPa beneficial threshold salinity, along with less-aggressive reduction under moderate stress. Lowering ψ\u003csub\u003ew\u003c/sub\u003e from 0 to -450 kPa reduced root dry weight of Sohg and Shnd by an average of 65% post a threshold of -100 kPa under salinity stress but led to progressive increases of 38% and 75% in Sohg and Shnd, respectively under drought stress (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec).\u003c/p\u003e\u003cp\u003eShoot water content was non-significantly affected by genotype and stress except for the significant reduction only in Shnd under severe salinity. By contrast, the genotypic difference emerged in water content of root, with higher values in Sohg than Shnd particularly under stress, and stronger reduction under drought than salinity (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe differential impact of stress on dry weights of plant organs resulted in alteration of biomass partitioning among leaves, stem and root. Under salinity stress, Sohg increased allocation of biomass to leaves at the expense of stem and root; whereas Shnd increased allocation to stem at the expense of root with marginal increase in leaf proportion. By contrast, under drought stress both cultivars increased allocation of biomass to root at the expense of foliage (stem and leaves) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eThree-way ANOVA showing the effect of the main factors: sesame cultivar (Cv), type of stress (Stress) and water potential of the medium (kPa) and their interactions on leaf dimensions, biomass and biomass partitioning and water content of plant organs\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"14\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c14\" colnum=\"14\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eSource of variation\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003edf\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e\u003cp\u003eLeaf angle\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003eLeaf length\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c8\" namest=\"c7\"\u003e\u003cp\u003eLeaf width\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003eLeaf W/L\u003c/p\u003e\u003cp\u003eratio\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eRoot water\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u003cp\u003eShoot water\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c12\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c13\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c14\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e83.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.611\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.440\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.361\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.552\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e3.121\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.087\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e7.651\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.009\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.378\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.543\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e10.16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.003\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.126\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.725\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.361\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.552\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.269\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.608\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e39.24\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.499\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.485\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ekPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.709\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.554\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e11.16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e3.252\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.034\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e3.003\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.045\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e11.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e4.894\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.007\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5.491\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.025\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.823\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.186\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e2.373\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.133\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.488\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.490\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e1.574\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.219\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e1.775\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.192\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.919\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.443\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.715\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.184\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.053\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.383\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e1.291\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.294\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e2.576\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.071\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e3.063\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.042\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5.133\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.005\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.423\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.738\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.065\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.978\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.430\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.733\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e20.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e1.536\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.224\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.150\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.113\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.880\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.462\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.627\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.603\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.375\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.772\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e1.297\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.293\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e2.771\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.058\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e\u003cp\u003eRoot dry\u003c/p\u003e\u003cp\u003eweight\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003eStem dry\u003c/p\u003e\u003cp\u003eweight\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c8\" namest=\"c7\"\u003e\u003cp\u003eLeaf dry\u003c/p\u003e\u003cp\u003eweight\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003eRWR\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eStWR\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u003cp\u003eLWR\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.498\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.486\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.827\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.370\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.691\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.412\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e1.360\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.252\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e4.987\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.033\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e1.043\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.315\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e96.52\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e21.73\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e48.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e272.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e25.20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e69.17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ekPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.368\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.270\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e21.99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e30.42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e10.45\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e4.264\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.012\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.683\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.569\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.778\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.384\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.282\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.599\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.036\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.851\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.002\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.963\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e0.643\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.428\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.428\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.518\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.610\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.206\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8.508\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.172\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.336\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e1.358\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.273\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e5.591\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.003\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e1.950\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.141\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e41.62\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e7.303\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e14.22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e48.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e4.181\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.013\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e13.78\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.167\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.918\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.442\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.724\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.371\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.775\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.016\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.997\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e1.086\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.369\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.945\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.431\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eEffect of salt stress and drought stress on water content of shoot and root of cvs Sohg and Shnd of sesame. Each value is the mean of three replicates\u0026thinsp;\u0026plusmn;\u0026thinsp;SE. Means with common letters are not significantly different at p\u0026thinsp;\u0026le;\u0026thinsp;0.05\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"3\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress and water\u003c/p\u003e\u003cp\u003epotential (kPa)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eShoot water\u003c/p\u003e\u003cp\u003e(% FW)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eRoot water\u003c/p\u003e\u003cp\u003e(% FW)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003eSohg\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSalt stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e86.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e91.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.86\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-100\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e87.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e91.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.87\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-250\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e86.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e92.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.30\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-450\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e86.63\u0026thinsp;\u0026plusmn;\u0026thinsp;1.60\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e94.02\u0026thinsp;\u0026plusmn;\u0026thinsp;1.26\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDrought stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e86.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e91.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.86\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-100\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e87.40\u0026thinsp;\u0026plusmn;\u0026thinsp;0.49\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e93.89\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-250\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e86.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e89.65\u0026thinsp;\u0026plusmn;\u0026thinsp;1.30\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-450\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e85.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.27\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e84.79\u0026thinsp;\u0026plusmn;\u0026thinsp;1.57\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003eShnd\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSalt stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e86.69\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e91.86\u0026thinsp;\u0026plusmn;\u0026thinsp;0.56\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-100\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e86.84\u0026thinsp;\u0026plusmn;\u0026thinsp;0.44\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e92.40\u0026thinsp;\u0026plusmn;\u0026thinsp;0.70\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-250\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e88.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e91.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.89\u003csup\u003eabc\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-450\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e80.50\u0026thinsp;\u0026plusmn;\u0026thinsp;3.44\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e91.35\u0026thinsp;\u0026plusmn;\u0026thinsp;1.11\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDrought stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e86.69\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e91.86\u0026thinsp;\u0026plusmn;\u0026thinsp;0.56\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-100\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e87.43\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e89.74\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-250\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e86.51\u0026thinsp;\u0026plusmn;\u0026thinsp;0.64\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e88.02\u0026thinsp;\u0026plusmn;\u0026thinsp;2.48\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e-450\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e85.80\u0026thinsp;\u0026plusmn;\u0026thinsp;1.08\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e81.62\u0026thinsp;\u0026plusmn;\u0026thinsp;0.84\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003e3.2 Photosynthetic pigments\u003c/h2\u003e\u003cp\u003eThe contents of Chl a, Chl b and carotenoids were significantly higher in Sohg than Shnd; the genotypic difference was either most evident under mild stress (Chl a and Chl b) or was maitained under all levels of stress (carotenoids). The impact of salinity on photosynthetic pigments was more aggressive than that of drought, which exerted beneficial effect in Shnd. Increasing salinity from 0 to -450 kPa reduced the contents of Chl a, Chl b and carotenoids by 45%, 35% and 35%, respectively as averages for Sohg and Shnd; but whereas the reductions were progressive in Sohg, they occurred post a threshold of -100 kPa in Shnd (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Meanwhile, drought stress non-significantly affected pigment content of Sohg but led to 55% and 78% increases in Chla and Chl b contents, respectively in Shnd, without effect on carotenoid content (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Partitioning of photosynthetic pigments among Chl a, Chl b and carotenoids showed mild response to treatments, except for: 1) salinized Sohg\u0026ndash;where carotenoid proportion was increased at the expense of Chl a, and 2) droughted Shnd\u0026ndash;where chlorophyll proportion was increased at the expense of carotenoids (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003e3.3 Gas exchange\u003c/h2\u003e\u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e reveals that the most affected gas exchange parameters were the rate of photosynthesis (A), followed by stomatal conductance (Gs) and transpiration rate (E) while the least affected parameter was substomatal CO\u003csub\u003e2\u003c/sub\u003e concentration (Ci). In contrast to the increased Ci by stress, E, Gs and A were lowered. The increases in Ci, across the whole range of ψ\u003csub\u003ew\u003c/sub\u003e, amounted to 54% and 39% in Sohg and Shnd, respectively (with \u0026minus;\u0026thinsp;100 kPa threshold under salinity stress) but were progressive with lower magnitude (33% and 14% in Sohg and Shnd, respectively) under drought stress (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea). By contrast, the reduction in E amounted to 77.5% under salinity stress and 47.5% under drought stress as an average for the two cultivars post a threshold of -100 kPa (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eb). The genotypic gap in G\u003csub\u003es\u003c/sub\u003e and A was marked, in favor of Sohg, in absence of stress but were narrowed under stress. Lowering ψ\u003csub\u003ew\u003c/sub\u003e from 0 to -450 kPa reduced Gs and A by an average of 85% under salinity stress and 57% under drought stress in the two cultivars post a threshold of -100 kPa (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ec, d).\u003c/p\u003e\u003cp\u003eThe genotypic difference in water use efficiency (WUE) was marked, in absence of stress and under severe drought stress, in favor of Sohg. Increasing salinity from 0 to -450 kPa reduced WUE by an average of 47% for the two cultivars, either progressively in Sohg or post \u0026minus;\u0026thinsp;100 kPa threshold in Shnd. Meanwhile, the decrease in WUE across the whole range of ψ\u003csub\u003ew\u003c/sub\u003e under drought stress amounted to 3% in Sohg and 38% in Shnd post a threshold of -250 kPa (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ee).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e\u003ch2\u003e3.4 Sugar fractions\u003c/h2\u003e\u003cp\u003eANOVA revealed that among the assayed sugar fractions, the most affected component was hemicelluloses followed by soluble sugars, whereas pectic substances were the least affected one (Tables\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Soluble sugar (SS) content of leaves was markedly higher in Sohg than Shnd in absence of stress and under drought stress but turned in favor of Shnd under salinity stress. SS content of Sohg leaves experienced 58% and 38% reductions under salinity stress and drought stress, respectively upon lowering of ψ\u003csub\u003ew\u003c/sub\u003e from 0 to -450 kPa. Meanwhile, SS content of Shnd leaves experienced overall non-significant changes under stress, although approaching a minimum at -250 kPa (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea).\u003c/p\u003e\u003cp\u003eIn contrast to SS, the leaf content of polysaccharides (starch, hemicellulose and pectic substances) was higher in Shnd than Sohg in absence of stress; but the genotypic difference was either diminished or turned in favor of Sohg under stress. In Sohg, lowering ψ\u003csub\u003ew\u003c/sub\u003e post a threshold of -100 kPa down to -450 kPa reduced starch content of leaves by 51% as an average for salinity stress and drought stress; but in Shnd, the reduction was progressive across the whole range of ψ\u003csub\u003ew\u003c/sub\u003e, with stronger magnitude under salinity (72%) than under drought (56%) (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eb). The low hemicellulose content of Sohg leaves was subjected to moderate changes under stress (either a reduction under salinity or an increase under drought). By contrast, the high hemicellulose content of Shnd leaves experienced 70% average reduction across the whole range of ψ\u003csub\u003ew\u003c/sub\u003e; either progressively under drought or post a transient increase at -100 kPa under salinity (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ec). Across the whole range of ψ\u003csub\u003ew\u003c/sub\u003e, salinity stress (on Sohg) as well as salinity and drought stress (on Shnd) led to an average 20% reduction in the leaf pectic substances; only drought stress led to non-significant increase in Sohg (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ed).\u003c/p\u003e\u003cp\u003eStarch constituted the greatest carbohydrate fraction of sesame leaves (\u0026gt;\u0026thinsp;50%), followed by soluble sugars and pectic substances (about 20% each), with minor hemicellulose component. In Sohg, mild salinity and drought stresses lowered SS ratio in favor of starch, but severe stresses led to the opposite effect; meanwhile the fraction of pectic substances or that of pectic substances and hemicellulose were progressively increase (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003ea, b). In Shnd, stress\u0026ndash;particularly salinity\u0026ndash;led to progressive increase in the ratios of SS and pectic substances at the expense of starch and hemicellulose (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003ec, d).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eThree-way ANOVA showing the effect of the main factors: sesame cultivar (Cv), type of stress (Stress) and water potential of the medium (kPa) and their interactions on the content of photosynthetic pigments and gas exchange of sesame leaves\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"14\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c14\" colnum=\"14\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eSource of variation\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003edf\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e\u003cp\u003eChl a\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003eChl b\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c8\" namest=\"c7\"\u003e\u003cp\u003eCarot\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003eChl a ratio\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eChl b ratio\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u003cp\u003eCarot ratio\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c12\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c13\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c14\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e26.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e15.66\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e29.73\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e2.662\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.113\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e1.854\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.183\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e5.011\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.032\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e16.14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e12.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.002\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e16.45\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.109\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.744\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e2.567\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.119\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e1.335\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.256\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ekPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.807\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.499\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.077\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.373\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e5.003\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e1.682\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.190\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e3.960\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.017\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e3.096\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.041\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.142\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.153\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.104\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.750\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e7.180\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.012\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.487\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.490\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e6.970\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.013\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.569\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.456\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.835\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.161\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.332\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.281\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.281\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.839\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e2.580\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.071\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e2.882\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.051\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e5.485\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.004\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e6.889\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.257\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.101\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e5.215\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.005\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e3.792\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.020\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e1.553\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.220\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e1.793\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.168\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.699\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.560\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.422\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.738\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.059\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.380\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.383\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.766\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e1.096\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.365\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u003cp\u003e0.686\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u003cp\u003e0.567\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e\u003cp\u003eSubstomatal CO\u003csub\u003e2\u003c/sub\u003e (Ci)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003eTranspiration\u003c/p\u003e\u003cp\u003erate (E)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c8\" namest=\"c7\"\u003e\u003cp\u003eStomatal conductance (Gs)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003ePhotosyn-thesis rate (A)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eA/E ratio\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.071\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.791\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.307\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.139\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e11.19\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.002\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e13.54\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e6.306\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.017\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.089\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.768\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10.68\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.003\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e7.701\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.009\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e17.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e10.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.003\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ekPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e17.43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e61.65\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e61.56\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e96.24\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e10.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.077\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.783\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.749\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.393\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e6.509\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.016\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e1.068\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.309\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e0.824\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.371\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.309\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.504\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.682\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.016\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.398\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e2.866\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.052\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e2.452\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.081\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.836\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.161\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3.583\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.024\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.688\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.189\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e8.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e5.336\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.004\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.016\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.997\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.567\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.641\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e2.361\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.090\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.353\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.787\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e1.070\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e0.376\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c14\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"4 Discussion","content":"\u003cp\u003eUnderstanding of the mechanisms, the plant can adopt to withstand drought and salinity stress is crucial for maintenance of crop productivity under unfavorable conditions. Stress effects on plant growth and performance are genotype-dependent but also depend on the severity and type of stress. Sesame, a moderately salt- and drought-tolerant crop [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e] is often cultivated under arid and semi-arid conditions.\u003c/p\u003e\u003cp\u003eSalinity and drought differentially affected shoot and root growth of sesame with more stress tolerance of cv. Shnd than Sohg. The less aggressive impact of salinity than drought on foliage growth of sesame can be viewed in light of the hypothesis that the accumulating salt ions can aid in lowering of water potential of plant tissues and hence allowing efficient water absorption under stress [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]. The beneficial effect of mild salinity to foliage growth and the lesser dehydrating effect of salinity\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eThree-way ANOVA showing the effect of the main factors: sesame cultivar (Cv), type of stress (Stress) and osmotic potential of the medium (kPa) and their interactions on the content of carbohydrate fractions of sesame leaves\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"10\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eSource of variation\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003edf\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e\u003cp\u003eSoluble sugars\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003eStarch\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c8\" namest=\"c7\"\u003e\u003cp\u003eHemicellulose\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003ePectic substances\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e31.48\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.243\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.625\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e181.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e2.811\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.103\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.642\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.429\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.513\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.479\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e4.795\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.036\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e6.783\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.014\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ekPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e27.92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e79.90\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e58.64\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e11.74\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.540\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.041\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4.050\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.053\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e64.88\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e1.715\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.200\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e13.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e13.63\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e64.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e7.530\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.216\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.885\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3.138\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.039\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e4.750\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.008\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e3.307\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.032\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.298\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.096\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.709\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.185\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e8.668\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.699\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.559\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e\u003cp\u003eSoluble sugars\u003c/p\u003e\u003cp\u003eratio\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003eStarch ratio\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c8\" namest=\"c7\"\u003e\u003cp\u003eHemicellulose\u003c/p\u003e\u003cp\u003eratio\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003ePectic substances ratio\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e31.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.995\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.326\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e165.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.202\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.656\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.344\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.136\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.016\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.902\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.868\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.181\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.136\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.715\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ekPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e41.88\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e31.97\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e10.81\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e28.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e16.97\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e36.93\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e81.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.747\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.394\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e47.87\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10.61\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e28.92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.942\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.432\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3.677\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.022\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.468\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.242\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.968\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.420\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.465\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.709\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCv \u0026times; Stress \u0026times; kPa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e6.934\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e7.246\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e12.54\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.552\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.651\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003ethan drought on root is in line with this hypothesis. However, the stronger dehydrating effect of salinity than drought on shoot suggests different patterns of osmotic susceptibility between shoot and root. In contrast to its more aggressive effect on foliage biomass, drought improved root biomass of sesame. However, decreased root volume and root hydraulic conductivity of three wheat varieties by PEG-drought stress and NaCl-salinity stress has been reported [\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e]. The reduction in root volume along with increased root density by drought reported by [\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e] implies either less severe impact of drought on root biomass than on root volume or even enhanced root biomass production by drought.\u003c/p\u003e\u003cp\u003eA functional equilibrium is always achieved by plants to balance their biomass allocation between root and shoot [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. Accordingly, it is expected that plants invest more biomass in the root at the expense of foliage under water deficit to optimize acquisition of water and nutrients and to restrict transpirational water loss. The signaling processes regulating this balance may involve sugars, phytohormones and small RNAs [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]. The present work suggests that salinity favors allocation of plant biomass to the foliage at the expense of root while drought led to the opposite effect. However, this pattern was subjected to genotypic intervention; whereas the favored allocation of plant biomass to the shoot under salinity occurred in favor of either leaves (Sohg) or stem (Shnd), the favored biomass allocation to root under drought occurred at the expense of either stem and leaves (Sohg) or only leaves (Shnd). Drought has been claimed to increase root to shoot biomass ratio of plants [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e] and specifically to favor allocation of plant biomass to root and stem at the expense of leaves [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]. By contrast, it has been claimed that root growth is less salt-sensitive than shoot growth, thus leading to increased root/shoot ratio of salinized plants, as has been reported by [\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e] and [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e]. Hence, it seems that the effect of stress on biomass allocation is controversial and depends on the experimental conditions and genotype [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]. Leaf weight ratio has been reported to decrease in \u003cem\u003eMedicago sativa\u003c/em\u003e [\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e] but to increase in \u003cem\u003eSetaria viridis\u003c/em\u003e [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e] by drought.\u003c/p\u003e\u003cp\u003eIn contrast to the inherent horizontal leaf orientation of Shnd with marginal effect of stress, the vertical leaf orientation of Sohg was differentially affected by salinity and drought; being more vertical under salinity but less vertical under drought. This fact lends support to the observation of preferred conservation of leaf biomass by Sohg and of stem biomass by Shand under stress. The positive correlations (under salinity stress) and the negative correlations (under drought stress) between leaf angle and the rates of photosynthesis and transpiration observed only in Sohg (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e) implies that, in the stress-sensitive Sohg, horizontal leaf orientation can improve gas exchange under salinity stress but vertical leaf orientation is appropriate under drought stress. Leaf orientation is an important factor in regulation of plant carbon\u0026ndash;water\u0026ndash;energy nexus to optimize photosynthesis and light interception under the impact of abiotic stress. However, it has been claimed that, in general, leaves are vertical in hot dry environments but more horizontal in mesic and light-limited environments [\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]. Unfavorable environmental conditions can evoke leaf movements, such as reversible epinastic responses [\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e], which can aid in plant adaption to stress conditions via protecting the leaves from photo-inhibition by reducing the leaf area exposed to solar irradiation [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eThe differential effect of salt stress and drought stress on photosynthetic pigments of sesame leaves was manifested as stronger negative impact of salinity on Sohg than Shnd, versus a mild reduction (Sohg) or even improvement (Shnd) by drought. The increased photosynthetic pigments of \u003cem\u003eCynanchum acutum\u003c/em\u003e leaves under drought was attributed to drought-induced P deprivation [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]. Furthermore, abiotic stress can minimize the genotypic gap in chlorophyll content between sesame cultivars that is approximating the less green Shnd to the greener Sohg. The symptoms of salt stress on plants include chlorosis and scorching of leaves [\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]. Chlorophyll content of leaves was decreased under both salinity and drought stress in \u003cem\u003eRobinia pseudoacacia\u003c/em\u003e [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e] and under drought in wheat [\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e], canola [\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e] and barley [\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e]. However, the effect of salinity on chlorophyll content of wheat leaves was genotype-dependent, being promotive in salt-tolerant genotypes but inhibitory in salt-sensitive genotypes [\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e]. The effect of stress on partitioning of photosynthetic pigments among Chl a, Chl b and carotenoids was particularly evident in Shnd, in which stress, can increase Chl a proportion at the expense of carotenoids. In accordance with the present findings, Chl a and Chl b contents were subjected to differential impact of salinity in rice leaves [\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e] and of drought in wheat [\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eThe present findings suggest stronger impact of salinity than drought on gas exchange of sesame leaves, a pattern that contradicts the response of foliage growth. Photosynthesis rate (A), transpiration rate (E) and stomatal conductance (Gs) are positively related to water potential of the medium [\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e]. Lowered photosynthesis under stress is usually linked to stomatal closure but can also result from impaired biochemical and photochemical constraints [\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e]. The intimate association between the patterns of retardation in A, E and Gs under the impact of salinity and drought indicates that the reduced photosynthetic activity under stress can be attributed to the stress-induced stomatal closure and lowering of stomatal conductance (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e). The fact that A, E and Gs are positively correlated with each other but negatively correlated with Ci, along with the coincidence of minimal Ci with maximal A and Gs at mild salinity (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e) implies that stress can inhibit photosynthetic efficiency\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003ewithout effect on or even enhancement of respiration but mild salinity can enhance photosynthetic efficiency through increasing stomatal conductance. In agreement with the present findings, drought stress has been found to induce stomatal closure with consequent severe inhibition in photosynthesis along with mild inhibition or even promotion in respiration [\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eStomatal movement under stress can control photosynthesis and transpiration in a way to increase water use efficiency. The present findings suggest that cv. Sohg with high photosynthesis potential exhibited marked drought robustness; in contrast to cv. Shnd of low photosynthesis potential, which although benefited from mild stress experienced more susceptibility to severe drought. Transpiration rate, in accordance with Gs, benefited from mild salinity in Shnd and mild drought in Sohg but was reduced under severe stress, with stronger impact of salinity than drought. According to the present finding, abiotic stress seems to reduce the water use efficiency (A/E ratio) of sesame with stronger impact of salinity on Sohg and of drought on Shnd. It has been reported that water-use efficiency was increased in \u003cem\u003eRobinia pseudoacacia\u003c/em\u003e under salinity and drought stress ([\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e] and in \u003cem\u003eVicia faba\u003c/em\u003e [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e] under salinity stress. The lowered water use efficiency of sesame under severe drought, observed in the present work, can be attributed either to greater sensitivity of the photosynthetic machinery than transpiration to water deficit or to progression of water loss despite of stomatal closure, probably via cuticular transpiration as proposed by [\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e]. Stomatal closure is a physiological adaptation to water deficit to avoid desiccation arising from transpirational water loss [\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e]; but the closed stomata will certainly inhibit photosynthesis by cutting down the influx of CO\u003csub\u003e2\u003c/sub\u003e to the mesophyll with the probability of continuation of water loss via alternative avenues such as the cuticular transpiration.\u003c/p\u003e\u003cp\u003eIt has been claimed that abiotic stress affects cell wall composition and production of compatible solutes [\u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e53\u003c/span\u003e]. The inhibited photosynthesis, concomitant with the unaffected or favored respiration under abiotic stress results in lowering of carbohydrate reserves of sesame leaves, which can be assigned to the decline in the major carbohydrate fraction (starch), followed by soluble sugars with marginal changes in pectic substances. However, the two sesame cultivars adopted different patterns of carbohydrate fractionation in response to stress. In Sohg, abiotic stress induced reductions in proportions soluble sugars and starch in favor of either pectic substances (salinity) or pectic substances and hemicellulose (drought). By contrast, in Shnd, abiotic stress increased the proportions of soluble sugars and pectic substances at the expense of starch and hemicellulose. Thus, the shared stress-induced response, irrespective of the genotype and type of stress, is the reduced proportion of starch in favor of pectic substances. The increased proportions of soluble sugars and pectic substances in Shnd under the impact of drought stress can aid in drought tolerance by contributing either to osmotic adjustment (soluble sugars) or by increasing cell wall elasticity (pectic substances). Osmotic and elastic adjustment are alternative strategies for a species to acclimate to water stress. Cells with highly elastic walls contain more water at full turgor; hence, their volume can decrease more before the turgor-loss point is reached. Elasticity of the cell wall depends on the chemical interactions between cell wall components [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. It has been proposed that pectins, the major polyanionic polysaccharide in the walls of growing cells, play a critical role in cell wall expansion [\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e]. However, the role of soluble sugars in osmotic adjustment under stress is evident only in Shnd, despite the stronger impact of salinity and drought on leaf carbohydrate content of Shnd than Sohg.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThe difference in stress-resistance was marked between the two sesame cultivars, with stronger aggressive impact of drought than salinity on foliage growth, which points to the role of salt ions in facilitating water absorption via lowering of water potential of foliage. Salinity shifted allocation of plant biomass to the foliage (specifically leaves of Sohg and stem of Shnd) at the expense of root; meanwhile, drought favors allocation of plant biomass to the root at the expense of stem of Sohg and leaf of Shnd. Thus, despite the contrasting effect of salinity and drought on rootage, Sohg seems to conserves leaves under stress but Shnd conserves stem. In a way to improve gas exchange, the sensitive Sohg adopts horizontal leaf orientation under salinity stress but vertical leaf orientation under drought stress. Abiotic stress narrowed the genotypic gap in chlorophyll content between sesame cultivars that is approximating the less green Shnd to the greener Sohg. In contrast to the influence on foliage growth, the impact of salinity on gas exchange of leaves was stronger than that of drought. The reduced photosynthetic activity under stress is related to stomatal closure and was accompanied with either no effect on or enhancement of respiration; however, mild salinity can enhance photosynthesis through increasing stomatal conductance. The photosynthetically less efficient genotype (Shnd) was more impacted by drought but at the same time, benefited from mild stress. Both E and Gs, benefited from mild salinity in Shnd and mild drought in Sohg but were subjected to stronger impact of salinity than drought. Stress reduced water use efficiency of sesame with stronger impact of salinity on Sohg and of drought on Shnd. The lowered water use efficiency can be attributed either to greater sensitivity of the photosynthetic machinery than transpiration to water deficit or to progression of water loss despite of stomatal closure, probably via cuticular transpiration. The stress-induced lowering in leaf carbohydrate reserves can be assigned to the decline in starch, either along with soluble sugars in favor of pectic substances (in Sohg) or of starch in favor of soluble sugars and pectic substances (in Shnd). The increased proportions of soluble sugars and pectic substances in Shnd under drought can aid in drought tolerance by contributing either to osmotic adjustment (soluble sugars) or to increased cell wall elasticity (pectic substances).\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors acknowledge the Oil Crops Research Department, Field Crops Research Institute (FCRI), Agricultural Research Center at Giza, Egypt for providing seeds of the two sesame cultivars.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by a research grant from Damietta University, Egypt as a part of M.Sc. program (Grant No. 1259/11/2021).\u003c/p\u003e\n\u003cp\u003eOpen access funding provided by The Science, Technology \u0026amp; Innovation Funding Authority (STDF) in cooperation with The Egyptian Knowledge Bank (EKB).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eConceptualization: TME; Methodology: SNA, SHA; Formal analysis and investigation: SNA, NMH; Writing - original draft preparation: SNA, TME; Writing - review and editing: TME, NMH; Funding acquisition: SHA; Resources: SNA; Supervision: TME, NMH. All authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of competing interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll data of this study are cited within the paper; raw data and calculations will be available on request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePermission to collect the seeds\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe manuscript is a purely eco-physiological study on performance of two sesame (\u003cem\u003eSesamum indicum\u003c/em\u003e L.) cultivars and does not include any human participants, groups of humans or animal subjects.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eSeeds of cultivars Sohag 1 (Sohg) and Shandaweel 3 (Shnd) of sesame (\u003cem\u003eSesamum indicum\u0026nbsp;\u003c/em\u003eL.) were obtained from the Agricultural Research Center (ARC), Egypt. The two experimental sesame cultivars are national cultivars, produced and released by Oil Crops Research Department, Field Crops Research Institute (FCRI), Agricultural Research Center (ARC), Ministry of Agricultural and Land Reclamation, Egypt according to The International Union for the Protection of New Varieties of Plants (UPOV) guideline (Egypt is a member of UPOV) and the Intellectual Property Rights (IPR) national low. The acquired names \"Sohag 1\" and \"Shandaweel 3\" are given by the breeder program of ARC and are related to national districts of Egypt. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics declaration:\u0026nbsp;\u003c/strong\u003enot applicable\u003cstrong\u003e.\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to publish declaration:\u0026nbsp;\u003c/strong\u003enot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to participate declaration:\u003c/strong\u003e not applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eClinical Trial Number\u003c/strong\u003e: not applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eLambers H, Oliveira RS. Plant water relations. In: Lambers H, Oliveira RS, editors. Plant Physiological Ecology. 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JACS Au. 2024;4:177\u0026ndash;88.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Drought. Salinity. Sesame genotypes. Gas exchange. Carbohydrate fractions. Leaf orientation","lastPublishedDoi":"10.21203/rs.3.rs-7362732/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7362732/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eDrought and salinity stresses are major threats to crop productivity in the arid lands. This work investigates the differential impact of salinity and drought on growth and performance of two sesame cultivars: Sohg and Shnd at water potentials of 0, -100, -250 and \u0026minus;\u0026thinsp;450 kPa with the same nutrient supply in a hydroponic sand culture. Under no stress, leaves of Sohg exhibited vertical orientation with higher pigment and soluble sugar contents but lower polysaccharide content relative to leaves of Shnd. Salinity and drought differentially affect leaf orientation only in Sohg, where drought induced horizontal leaf position but salinity led to the opposite effect. Whereas drought induced stronger reduction in foliage biomass than did salinity, with enhancement of root biomass, salinity reduced foliage and root biomass with favored allocation of plant biomass to root. Salinity reduced leaf pigment content, particularly in Sohg but drought either increased it or was without effect. Both drought and salinity, post a threshold of -100 kPa, reduced rates of photosynthesis and transpiration as well as stomatal conductance, with stronger impact of salinity than drought. The lowering in carbohydrate content under stress, being more aggressive under salinity than drought, was associated with increasing proportion of pectin at the expense of starch and variable changes in soluble sugar proportions.\u003c/p\u003e","manuscriptTitle":"Salinity stress and drought stress differentially affect biomass allocation, leaf orientation and performance of two sesame (Sesamum indicum L.) cultivars","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-12-15 09:13:18","doi":"10.21203/rs.3.rs-7362732/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"479dde6c-ce8d-4108-a0fd-28ccb1a4e0a6","owner":[],"postedDate":"December 15th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-04-21T23:38:52+00:00","versionOfRecord":[],"versionCreatedAt":"2025-12-15 09:13:18","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7362732","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7362732","identity":"rs-7362732","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}