Differences in responses of moderately salt-tolerant and salt-sensitive tree species to heterogeneous salinity

Growth responses of the moderately salt-tolerant velvet ash (Fraxinus velutina) and salt-sensitive poplar (Populus × euramericana) were investigated under heterogeneous root zone salinity. The salinity treatments imposed on the two root zones (lower-higher) were 137-137 (uniform), 103-171, 68-205, 34-239, and 0-273 mM NaCl for velvet ash, and 51-51 (uniform), 34-68, 17-85, and 0-103 mM NaCl for poplar. The leaf gas exchange of the plants was measured one month after these treatments were implemented, and the plants were sampled 75 d after treatment to measure other physiological parameters. Net photosynthetic rate, transpiration rate, total biomass, and fine root compensatory growth increased as the difference in salinity between the two root zones (i.e., salinity heterogeneity) increased in velvet ash. These parameters showed no significant difference among the treatments in poplar. The leaf Na⁺ content was lower under heterogeneous salinity than under uniform salinity in both tested species. The leaf proline content in velvet ash decreased under heterogeneous salinity compared to that under uniform salinity, whereas that of poplar increased. The soluble sugar content of velvet ash leaves increased under heterogeneous salinity, whereas no changes were observed in poplar. The increased fine root biomass in the lower salinity zone promoted velvet ash growth by decreasing the leaf Na⁺ and Cl^- content under heterogeneous salinity. The poplar’s undifferentiated root distribution and gas exchange in response to the heterogeneous salinity were attributed to its salt sensitivity.     

Responses of leaf growth and gas exchanges to salt stress during reproductive stage in wild wheat relative Aegilops geniculata Roth. and wheat (Triticum durum Desf.)

In order to investigate the effect of salinity on the growth and photosynthesis of the wild wheat and wheat, three accessions of Aegilops geniculata from Ain Zana, Zaghouan and Sbitla and one variety of durum wheat (Triticum durum) were grown in the INRAT greenhouse and treated with different salinity levels. The growth of leaves, water status and gas exchange parameters have been measured at the reproductive stage. The flag leaf length, total leaf dry weight, water status, CO₂ assimilation rate, stomatal conductance, intercellular CO₂ and transpiration for the three Ae. geniculata accessions and wheat variety significantly decreased with increasing salt. The decline in photosynthesis measured in response to salt stress was proportionally greater than the declines in transpiration, resulting in a reduction of water-use efficiency, at both the leaf and whole-plant levels. Among the factors inhibiting photosynthetic activity, those of a stomatal nature had a greater effect. This study has shown a high degree of variation of these characters mainly related to geographical origin. It was observed also that Sbitla accession was less affected by the imposed salt stress than all the others while Ain Zana was the most affected one.     

Separate and combined effects of silicon and selenium on salt tolerance of wheat plants

Soil salinity is the leading global abiotic stress which limits agricultural production with an annual increment of 10%. Therefore; a pot experiment was conducted with the aim to alleviate the salinity effects on wheat seedlings through exogenous application of silicon (Si) and selenium (Se). Treatments included in the study were viz. (Ck) control (no NaCl nor Si and Se added), only salinity (50 mM NaCl), salinity + Si (50 mM NaCl with 40 mM Si), salinity + Se (50 mM NaCl with 40 mM Se) and salinity + Si + Se (50 mM NaCl + 40 mM Si + 40 mM Si). The salt stress impaired the growth (root and shoot dry weight, root: shoot ratio, seedlings biomass), water relations, photosynthetic attributes, transpiration rate and chlorophyll contents of wheat seedlings. Nonetheless, the foliar application of Si and Se alone and in combination improved the growth, water relations, photosynthetic attributes, transpiration rate and chlorophyll contents of wheat seedlings under stressed conditions. Moreover, an increase in antioxidant enzyme activity and accumulation of osmo-protectants (proline, soluble protein and soluble sugar) was noted under stressed conditions, which was more pronounced in wheat seedling which experienced combined application of Si and Se. To conclude that, foliar application of Si alone mitigated the adverse effect of salinity, while the combined application of Si and Se was proved to be even more effective in alleviating the toxic effects of salinity stress on wheat seedlings.     

The Role of Hormones in Fast Growth Responses of Wheat Plants to Osmotic and Cold Shocks

The effects of nutrient-solution cooling and PEG addition to the nutrient solution on the phytohormone content, the rate of leaf growth, leaf extensibility under the influence of external mechanical action, osmotic potential, and transpiration were studied in seven-day-old wheat plants. Leaf growth rapidly ceased, and the transpiration rate was reduced in both treatments. Growth cessation induced by PEG was transient, and growth resumption was preceded by an increase in the leaf extensibility. The functional role of auxin accumulation in plant shoots in the control of extensibility as well as the relationship between the ABA accumulation and a decrease in the cytokinin content, on the one hand, and reduced transpiration, on the other hand, under stress conditions are discussed.     

Soil salinity effects on transpiration and net photosynthetic rates,stomatal conductance and Na+ and Cl− contents in durum wheat

Leaf gas exchange, plant growth and leaf ion content were measured in wheat (Triticum durum L. cv. HD 4502) exposed to steady- state salinities (1.6, 12.0 and 16.0 dS nr⁻¹) for 8 weeks. Salinity reduced leaf area and number of tillers, and increased Na⁺ and Cl⁻ concentrations in leaves. Leaf- to- leaf gradients of these ions were observed. The oldest leaf contained 6 to 8 times more Na⁺ and Cl⁻ than the flag leaf. Net photosynthetic rate (P_N), transpiration rate (E) and stomatal conductance (g_S) were the highest in flag leaf, declined in the middle and fully expanded leaves, and were minimum in the oldest leaves. These processes were reduced by salinity with similar leaf- to- leaf gradients. Intercellular CO₂ concentrations in the older leaves were higher than in the flag leaf in non-saline plants, and increased similarly with salinity. Leaf age was the major factor in reducing P_N, and senescence processes were promoted by salinity.     

Does night-time transpiration provide any benefit to wheat (Triticum aestivum L.) plants which are exposed to salt stress?

The study aimed to test whether night-time transpiration provides any potential benefit to wheat plants which are subjected to salt stress. Hydroponically grown wheat plants were grown at four levels of salt stress (50, 100, 150, and 200 mM NaCl) for 5–8 days prior to harvest (day 14–18). Salt stress caused large decreases in transpiration and leaf elongation rates during day and night. The quantitative relation between the diurnal use of water for transpiration and leaf growth was comparatively little affected by salt. Night-time transpirational water loss occurred predominantly through stomata in support of respiration. Diurnal gas exchange and leaf growth were functionally linked to each other through the provision of resources (carbon, energy) and an increase in leaf surface area. Diurnal rates of water use associated with leaf cell expansive growth were highly correlated with the water potential of the xylem, which was dominated by the tension component. The tissue-specific expression level of nine candidate aquaporin genes in elongating and mature leaf tissue was little affected by salt stress or day/night changes. Growing plants under conditions of reduced night-time transpirational water loss by increasing the relative humidity (RH) during the night to 95% had little effect on the growth response to salt stress, nor was the accumulation of Na⁺ and Cl⁻ in shoot tissue altered. We conclude that night-time gas exchange supports the growth in leaf area over a 24 h day/night period. Night-time transpirational water loss neither decreases nor increases the tolerance to salt stress in wheat.     

Salinity and drought interaction in wheat (Triticum aestivum L.) is affected by the genotype and plant growth stage

Salinity and drought are important agro-environmental problems occurring separately as well as together with the combined occurrence increasing with time due to climate change. Screening of bread wheat genotypes against salinity or drought alone is common; however, little information is available on the response of wheat genotypes to a combination of these stresses. This study investigates the response of a salt-resistant (SARC-1) and a salt-sensitive (7-Cerros) wheat genotype to drought at different growth stages under non-saline (ECe 2.1 dS m^?1) and saline soil (ECe 15 dS m^?1) conditions. Drought was applied by withholding water for 21 days at a particular growth stage viz. tillering, booting, and grain filling stages. At booting stage measurements regarding water relations, leaf ionic composition and photosynthetic attributes were made. At maturity grain yield and different yield, components were recorded. Salinity and drought significantly decreased grain yield and different yield components with a higher decrease in the case of combined stress of salinity × drought. The complete drought treatment (drought at tillering + booting + grain filling stages) was most harmful for wheat followed by drought at booting stage and grain filling–tillering stages, respectively. The salt-resistant wheat genotype SARC-1 performed better than the salt-sensitive genotype 7-Cerros in different stress treatments. A decrease in the water and turgor potentials, photosynthetic and transpiration rates, stomatal conductance, leaf K⁺, and increased leaf Na⁺ were the apparent causes of growth and yield reduction of bread wheat due to salinity, drought, and salinity × drought.     

Effect of salinity on tissue architecture in expanding wheat leaves

Salinity greatly reduces the leaf cross-sectional area of wheat (Triticum aestivum L.) during its development, which may lead to variation in the architectural properties of growing leaves that would result in a change in leaf physiological functions. Our objective was to characterize the effect of salinity on the spatial distribution of the cross-sectional area and the anatomy of large and small veins of a growing wheat leaf. Spring wheat was grown in a growth chamber in soils with or without 120 mM NaCl. Leaf 4 in both treatments was harvested 2–3 days after its emergence and then cut into five transverse segments. Examination of the transverse sections revealed that salinity significantly reduced the cross-sectional area, width, and radii of both epidermal and mesophyll cells along the leaf axis. Reduction in the cross-sectional area and width occurred mainly at the leaf base, indicating that these reductions occur during the period of leaf initiation. The reduction in cross-sectional area was attributed to a decrease in the size of the vein segments and a reduced number of medium and small veins. The thickness of the leaf was also reduced under the 120 mM NaCl treatment. A greater intercellular air space in the large vein segments under saline conditions was also found. The approximately 35% reduction observed in the number of veins under saline conditions (mainly in the number of small veins) may suggest that salinity reduces the capacity for re-translocation of mineral nutrients and assimilates. The reduced area of protoxylem and metaxylem in midrib and large vein segments in growing tissues may be responsible for lower water deposition into the growth zone under saline conditions.     

Root cooling strongly affects diel leaf growth dynamics,water and carbohydrate relations in Ricinus communis

In laboratory and greenhouse experiments with potted plants, shoots and roots are exposed to temperature regimes throughout a 24 h (diel) cycle that can differ strongly from the regime under which these plants have evolved. In the field, roots are often exposed to lower temperatures than shoots. When the root‐zone temperature in Ricinus communis was decreased below a threshold value, leaf growth occurred preferentially at night and was strongly inhibited during the day. Overall, leaf expansion, shoot biomass growth, root elongation and ramification decreased rapidly, carbon fluxes from shoot to root were diminished and carbohydrate contents of both root and shoot increased. Further, transpiration rate was not affected, yet hydrostatic tensions in shoot xylem increased. When root temperature was increased again, xylem tension reduced, leaf growth recovered rapidly, carbon fluxes from shoot to root increased, and carbohydrate pools were depleted. We hypothesize that the decreased uptake of water in cool roots diminishes the growth potential of the entire plant – especially diurnally, when the growing leaf loses water via transpiration. As a consequence, leaf growth and metabolite concentrations can vary enormously, depending on root‐zone temperature and its heterogeneity inside pots.     

The Effect of Rapid Temperature Increase on the Growth Rate of Wheat Leaves

The growth rate of the first leaf of eight-day-old wheat plants was measured using a DLT-2 highly sensitive linear displacement transducer. Leaf extensibility was evaluated from the growth rate under the increase in the pulling force by 2 g. An increase in the air temperature resulted in the doubling of the transpiration rate and immediate slowing of the leaf growth followed by the leaf shrinkage. However, growth was later resumed almost completely. Heat treatment did not induce any changes in the leaf extensibility, indicating that cell-wall mechanical properties were not changed. Growth retardation was supposed to result from a decrease in the water content in the leaf tissues because the balance between water influx from roots and its loss through transpiration was shifted toward the water loss. An initial drop in the relative water content (RWC) indicates such a misbalance. Subsequent growth resumption coincided with a decreased water deficiency. Since the rate of transpiration was not reduced, RWC and growth rate restoring evidently occurred due to the activated water uptake by roots, which can be explained by the increased hydraulic permeability detected in our experiments.     

Growth enhancement and antitranspirant activity following seed treatment with a derivative of 5-hydroxybenzimidazole (Ambiol) in four drought-stressed agricultural species

The biostimulating action of seed treatment with the synthetic antioxidant, Ambiol (2-methyl-4-Edimethylaminomethyl-5-hydroxybenzimidazole dihydrochloride) on subsequent growth and transpiration of seedlings was studied. To study growth and transpiration responses, seeds of four agricultural species, soybean ( Glycine max L.), rapeseed ( Brassica napus L.), winter wheat ( Trilicum aestivum L.) and corn ( Zea mays L.), were soaked in Ambiol for 24 h, using the following concentrations: 0, 0.01, 0.1, 1, 10 and 100 mg 1¹. The subsequent seedlings were subject to simulated soil drought, using computer-automated root misting chambers. The influence of Ambiot on transpiration rate under simulated air drought was studied by growing plants under low humidity in a controlled humidity chamber. Response to Ambiol varied, depending on its concentration, the species used and the environment. Compared to untreated plants, 10 mg 1-⁻¹ Ambiol reduced the mid-day transpiration rate and total daily water usage of soybean by approximately 25%. Under simulated soil drought in the root misting chamber, 10 and 100 mg 1-⁻¹ Ambiol increased growth of rapeseed and soybean by 25–45%, relative to the 0 mg 1-⁻¹ treatment, yielding plants comparable in size to the fully-irrigated controls. However, Ambiol failed to promote growth of two drought-stressed monocotyledons (corn and winter wheat). At 100 mg P. Ambiol inhibited growth of both well-watered wheat and rapeseed, although this inhibition was mitigated by drought.     

Leaf development,transpiration and ion uptake and distribution in sugarcane cultivars grown under salinity

The effects of salinity on leaf growth, initiation and senescence, on transpiration rates, on leaf water potential and on uptake and distribution of several ions were studied in two sugarcane cultivars differing in salinity sensitivity. Plants, growing in a growing mixture in pots, were exposed to salinized irrigation water for 68 days, starting 60 days after planting. EC values of the irrigation water were 1.0, 2.0, 4.0, 8.0 and 12 dS/m, obtained by using a mixture of NaCl and CaCl2. Plants were also grown in nutrient solution and were at a similar age when exposed to a salinity level of 3 dS/m for 30 days followed by 6.0 dS/m for an additional 30 days. Two Na:Ca ratios of 18:1 and 1:2 were used for salinization of the nutrient solution. Both leaf dry weight and area decreased with increasing salinity, but in the more salinity tolerant cultivar H69-8235, the decrease was moderate. Salinity hardly reduced average area per leaf in H69-8235, while the number of leaves declined sharply. This decline was caused by enhanced senescence of mature leaves and not by a decreased rate of leaf initiation. In the more sensitive cultivar, H65-7052, leaf area and initiation of new leaves were sharply reduced by salinity while leaf senescence was less affected. Leaf water potential decreased during the early stages of salinity exposure, and the reduction in water potential was larger in H69-8235. Salinity also decreased the rate of transpiration rate but to a lesser extent than leaf development and growth. The accumulation of Cl and Na in the TVD (top visible dewlap) leaf of the tolerant cultivar H69-8235 was greater than in the sensitive cultivar H65-7052. The concentration of Cl in the TVD leaf was more than 10 times that of Na in both cultivars. The concentration of both ions, but not of K, increased during the early stages of salinity exposure and then remained constant. A gradient in concentration of Cl and Na over the plant was found in both cultivars at all salinity levels, and was steepest between the TVD and younger leaves. No specific Na effect on leaf growth or transpiration could be detected. The accumulation of Cl and Na but not of K occurred primarily in the roots rather than in the leaves and stalks. This revised version was published online in August 2006 with corrections to the Cover Date.     

Spatial and temporal quantitative analysis of cell division and elongation rate in growing wheat leaves under saline conditions

Leaf growth in grasses is determined by the cell division and elongation rates, with the duration of cell elongation being one of the processes that is the most sensitive to salinity. Our objective was to investigate the distribution profiles of cell production, cell length and the duration of cell elongation in the growing zone of the wheat leaf during the steady growth phase. Plants were grown in loamy soil with or without 120 mmol/L NaCl in a growth chamber, and harvested at day 3 after leaf 4 emerged. Results show that the elongation rate of leaf 4 was reduced by 120 mmol/L NaCl during the steady growth phase. The distribution profile of the lengths of abaxial epidermal cells of leaf 4 during the steady growth stage shows a sigmoidal pattern along the leaf axis for both treatments. Although salinity did not affect or even increased the length of the epidermal cells in some locations in the growth zone compared to the control treatment, the final length of the epidermal cells was reduced by 14% at 120 mmol/L NaCl. Thus, we concluded that the observed reduction in the leaf elongation rate derived in part from the reduced cell division rate and either the shortened cell elongation zone or shortened duration of cell elongation. This suggests that more attention should be paid to the effects of salinity on those properties of cell production and the period of cell maturation that are related to the properties of cell wall.     

Effect of partial root excision on shoot water relations

Removing 4 out of 5 serminal roots from 7-day-old wheat seedlings arrested leaf elongation for 1.5 h. This effect can be explained by an initial decrease in foliar water content resulting from the smaller root surface area available for water uptake. Subsequently, leaf hydration increased with time and came to equal that of intact plants within 2 h. The rehydration was seemingly effected by an increasing conductivity of the one remaining root axis, since transpiration of the partially de-rooted plants did not fall below that of controls. With time, leaf elongation resumed, but at a slower rate than in intact plants. This slower growth may be attributed to a decrease in leaf extensibility since this was found to be reduced when measured by a counterweight technique involving linear displacement transducers. Loss of extensibility was associated with decreased IAA concentration in the leaf elongation zone.     

Effects of heterogeneous salinity on growth,water uptake,and tissue ion concentrations of alfalfa

Aims

Soil salinity varies greatly in the plant rhizosphere. The effect of nonuniform salinity on the growth and physiology response of alfalfa plants was determined to improve understanding of salt stress tolerance mechanisms of alfalfa.

Methods

Plant growth, predawn leaf water potential, water uptake, and tissue ionic content were studied in alfalfa plants grown hydroponically for 9 days using a split-root system, with uniform salinity or horizontally nonuniform salinity treatments (0/S, 75/S, and 150/S corresponding to 0, 75, and 150 mM NaCl on the low salt side, respectively).

Results

Compared with uniform high salinity, 0/S and 75/S treatments significantly increased the alfalfa shoot dry mass and stem extension rate. Compensatory water uptake by low salt roots of 0/S and 75/S treatments was observed. However, decreased leaf Na⁺ concentration, increased leaf K⁺/Na⁺, and compensatory growth of roots on the low salt side were observed only following the 0/S treatment.

Conclusions

Nonuniform salinity dose not enhance plant growth once a threshold NaCl concentration in low salinity growth medium has been reached. Compensation of water uptake from the low-salt root zone and regulation of K⁺/Na⁺ homeostasis in low salt root play more important role than regulation of leaf ions in enhancing alfalfa growth under nonuniform salinity.

     

Wide vessels sustain marginal transpiration flux and do not optimize inefficient gas exchange activity under impaired hydraulic control and salinity

Plants optimize water use and carbon assimilation via transient regulation of stomata resistance and by limiting hydraulic conductivity in a long-term response of xylem anatomy. We postulated that without effective hydraulic regulation plants would permanently restrain water loss and photosynthetic productivity under salt stress conditions. We compared wild-type tomatoes to a transgenic type (TT) with impaired stomatal control. Gas exchange activity, biomass, starch content, leaf area and root traits, mineral composition and main stems xylem anatomy and hydraulic conductivity were analyzed in plants exposed to salinities of 1 and 4 dS m⁻¹ over 60 days. As the xylem cannot easily readjust to different environmental conditions, shifts in its anatomy and the permanent effect on plant hydraulic conductivity kept transpiration at lower levels under unstressed conditions and maintained it under salt-stress, while sustaining higher but inefficient assimilation rates, leading to starch accumulation and decreased plant biomass, leaf and root area and root length. Narrow conduits in unstressed TT plants were related to permanent restrain of hydraulic conductivity and plant transpiration. Under salinity, TT plants followed the atmospheric water demand, sustained similar transpiration rate from unstressed to salt-stressed conditions and possibly maintained hydraulic integrity, due to likely impaired hydraulic regulation, wider conduits and higher hydraulic conductivity. The accumulation of salts and starch in the TT plants was a strong evidence of salinity tolerance via osmotic regulation, also thought to help to maintain the assimilation rates and transpiration flux under salinity, although it was not translated into higher growth.     

Silicon increases salt tolerance by influencing the two-phase growth response to salinity in wheat (Triticum aestivum L.)

Salt usually stresses plants in two ways, osmotic stress and ion toxicity. Plant responds to salinity in two distinct phases through time. It is known that silicon (Si) could alleviate salt stress by decreasing the Na⁺ accumulated in the leaf. In order to determine the function of Si in the two-phase growth response (osmotic and ion toxicity) to salinity, we selected the wheat cultivar “Changwu 134” out of 10 wheat cultivars, and confirmed that it responds to salinity in two distinct phases through time. The fresh weight, leaf area, and leaf Na⁺ concentration were measured during 31 days of 120 mM NaCl supplemented with 1 mM Si treatment. The results revealed that the growth of plants under salinity conditions both with and without Si application were in accordance with the two-phase growth model. Si alleviated the salt stress in the both two-phase growth, but the alleviative effects were more pronounced in the osmotic stress phase than ion toxicity phase. These results clearly showed that Si can enhance plant salt tolerance by alleviating the salt-induced osmotic stress.     

根茎在羊草响应短期NaCl胁迫过程中的作用

根茎是根状茎型克隆植物的特有结构,在养分储存、运输和分蘖茎的形成等方面起关键性作用。然而有关根茎生理学方面的研究却十分匮乏。为了探讨根茎在植物感应环境胁迫中的作用, 本文以羊草(Leymus chinensis)为实验材料,研究和比较了短期NaCl胁迫根、根茎、根和根茎3种处理方式下羊草对盐胁迫的响应。试验结果表明:200 mmol·L^-1 NaCl处理羊草根、根茎、根和根茎24 h,显著(p<0.05)降低羊草叶片净光合速率和蒸腾速率,增加叶片渗透浓度与脯氨酸含量;其中同时处理根和根茎叶片,蒸腾速率和净光合速率的降低程度显著高于分别处理根和根茎。在分别处理根与根茎的情况下,叶片含水量、脯氨酸含量、净光合速率、蒸腾速率均无显著性差异。不论单独胁迫根、根茎还是同时胁迫根和根茎,羊草根、根茎和叶片内Na⁺含量都显著高于对照,而羊草根、根茎和叶片内K⁺含量都显著低于对照。这些结果显示:1)根茎在羊草响应盐胁迫的生理过程中与根系具有类似的功能;2)羊草根茎在盐胁迫条件下能够有效地吸收Na⁺;3)鉴于根茎的生物量和表面积都明显地低于根系,在盐胁迫下羊草根茎吸收Na⁺的效率高于根系。     

Rapid and tissue-specific changes in ABA and in growth rate in response to salinity in barley leaves

The addition of 100 mM NaCl to the root medium of barley plantscaused the rapid cessation of elongation of the growing leafthree, followed by a sudden resumption of growth during thefollowing hour. The idea that resumption of growth is precededand mediated by rapid and tissue-specific changes in ABA concentrationand by changes in transpiration was tested. Leaf elongationvelocity was recorded continuously using linear variable displacementtransducers (LVDT), ABA was determined by immunoassay, and transpirationand stomatal conductivity were measured gravimetrically andby porometry, respectively. Within 10 min following additionof salt, ABA increased 6-fold in the distal portion of the leafelongation zone; in the proximal portion, ABA accumulated witha delay. In the portion of the growing blade that had emergedABA increased 3-fold and remained elevated during the following20 min. This preceded a decrease in transpiration and stomatalconductivity, which, in turn, coincided with growth resumption.Twenty hours following the addition of salt, the ABA concentrationshad returned to the level before stress. Leaf elongation velocitywas still reduced. It is concluded that NaCl causes a rapidincrease in ABA in the transpiring portion of the growing leaf.This leads to a decrease in transpiration. As a result, xylemwater potential is expected to rise. The moment that the waterpotential gradient between the xylem and the peripheral cellsin the growth zone favours water uptake again into the latter,leaf elongation resumes. The results suggest that ABA causesdifferent responses in different leaf regions, all aimed atpromoting the resumption of leaf growth. Key words: Abscisic acid, cell elongation, Hordeum vulgare, leaf growth, salinity, water relations.