OSMOTIC ADJUSTMENT OF PLANTS TO SALINE MEDIA. II. DYNAMIC PHASE

Bernstein , Leon . (U. S. Salinity Lab., Riverside, Calif.) Osmotic adjustment of plants to saline media. II. Dynamic phase. Amer. Jour. Bot. 50(4): 360–370. Illus. 1963.—The time-course of osmotic adjustment in bean and pepper plants to increased salinity of the medium was determined by periodic sampling of plants following salt additions to the medium. Bean plants adjusted to increases of 1 atm OP within a day, the adjustment in roots occurring primarily at night following salt addition at 6 pm , whereas leaves and stems made most of their adjustment in the daytime. Pepper plants did not adjust completely to 1.5 atm NaCl additions in 48 hr, but OP increased by about the same amount in both species (0.5—1.0 atm per day). Diurnal fluctuations in OP of leaves and stems of both species and in roots of pepper were matched by parallel fluctuations in K concentrations. Added NaCl caused increased concentrations of K in leaves and stems which were more or less replaced by more slowly absorbed ions, Ca and Mg in bean leaves and Na in bean stems. Other salts produced comparable immediate effects on K level, but K was replaced more rapidly if the cation added was readily accumulated by the bean (Ca). In roots, Na uptake predominated if Na salts were added but K uptake was important on the CaCl₂ treatment. The K effects suggest a passive distribution of K between the cell and the medium.     

Water relations and osmotic adjustment in Lycopersicon esculentum and L. pennellii during short-term salt exposure and recovery

Cultivated tomato Lycopersicon esculentum (L.) Mill. cv. P-73 and its wild salt-tolerant relative L. pennellii (Correll) D'Arcy accession PE-47 growing on silica sand in a growth chamber were exposed to 0, 70, 140 and 210 m M NaCl nutrient solutions 35 days after sowing. The saline treatments were imposed for 4 days, after which the plants were rinsed with distilled water. Salinity in L. esculentum reduced leaf area and leaf and shoot dry weights. The reductions were more pronounced when sodium chloride was removed from the root medium. Reduction in leaf area and weight in L. pennellii was only observed after the recovery period. In both genotypes salinity induced a progressive reduction in leaf water potential and leaf conductance. During the recovery period leaf water potential (ψ₁) and leaf conductance (g₁) reached levels similar to those of control plants in wild and cultivated species, respectively. Leaf osmotic potential at full turgor (ψ_os) decreased in the salt treated plants of both genotypes, whereas the bulk modulus of elasticity was not affected by salinity. Leaf water potential at turgor loss point (ψ_tlp) and relative water content at turgor loss point (RWC_tlp) appeared to be controlled by leaf osmotic potential at full turgor (ψ_os) and by bulk modulus of elasticity, respectively. At lowest salinity, the wild species carried out the osmotic adjustment based almost exclusively on Cl⁻ and Na⁺, with a marked energy savings. Under highest salinity, this species accommodate the stress through a higher expenditure of energy due to the contribution of organic solutes to the osmotic adjustment. The domesticated species carried out the osmotic adjustment based always on an important contribution of organic solutes.     

Leaf age effects on solute accumulation in water-stressed grapevines

The effects of leaf age on water relations, organic solute, and total ion accumulation were studied in mature and immature leaves of two-year-old grapevines (Vitis vinifera L., cv. Savatiano) grown under water stress conditions. Osmotic potential at full turgor decreased significantly in leaves of stressed plants, irrespective of leaf age, indicating the occurrence of an active osmotic adjustment. The apoplastic water fraction (A) increased during leaf ontogeny in both control and stressed plants. However, the values of A were lower in stressed plants. Starch concentration decreased significantly in both mature and immature leaves during the drought cycle, while the relative proportion of monosaccharides and sucrose was markedly different in immature leaves compared to mature. The accumulation of total inorganic ions, induced by drought, was also age dependent, increasing significantly with leaf age, while there were no significant differences in total amino acids content. Inorganic ions and carbohydrates seem to be the major component of osmotic adjustment in mature and immature grapevine leaves, respectively.     

Influence of Xylem Water Potential on Leaf Elongation and Osmotic Adjustment of Wheat and Lupin

The leaf elongation rate and osmotic pressure at full turgorof wheat (Triticum aestivum L.) and lupin (Lupinus cosentiniiGuss.) were measured in well watered plants, in plants thatwere allowed to dry the soil slowly over 7 d, and in plantsin which the water potential of the leaf xylem was maintainedhigh by applying pressure to the roots during the drying cycle.Maintenance of high xylem water potentials failed to preventa reduction in the rate of leaf elongation as the soil dried,while the osmotic pressure at full turgor and the degree ofosmotic adjustment increased as the soil water content decreased.The rate of leaf elongation was reduced more and the degreeof osmotic adjustment was higher in leaves with high xylem waterpotentials than in those in which leaf xylem potentials wereallowed to decrease as soil water content decreased. Osmoticadjustment was linearly correlated with the reduction in leafelongation rate in both wheat and lupin. Key words: Osmotic adjustment, leaf elongation, turgor regulation     

Leaf age and salinity influence water relations of pepper leaves

Plant growth is reduced under saline conditions even when turgor in mature leaves is maintained by osmotic adjustment. The objective of this study was to determine if young leaves from salt-affected plants were also osmotically adjusted. Pepper plants (Capsicum annuum L. cv. California Wonder) were grown in several levels of solution osmotic potential and various components of the plants' water relations were measured to determine if young, rapidly growing leaves could accumulate solutes rapidly enough to maintain turgor for normal cell enlargement. Psychrometric measurements indicated that osmotic adjustment is similar for both young and mature leaves although osmotic potential is slightly lower for young leaves. Total water potential is also lower for young leaves, particularly at dawn for the saline treatments. The result is reduced turgor under saline conditions at dawn for young but not mature leaves. This reduced turgor at dawn, and presumably low night value, is possibly a cause of reduced growth under saline conditions. No differences in leaf turgor occur at midday. Porometer measurements indicated that young leaves at a given salinity level have a higher stomatal conductance than mature leaves, regardless of the time of day. The result of stomatal closure is a linear reduction of transpiration.     

Osmoregulation,solute distribution,and growth in soybean seedlings having low water potentials

Soybean (Glycine max (L.) Merr.) seedlings osmoregulate when the supply of water is limited around the roots. The osmoregulation involves solute accumulation (osmotic adjustment) by the elongating region of the hypocotyls. We investigated the relationship between growth, solute accumulation, and the partitioning of solutes during osmoregulation. Darkgrown seedlings were transplanted to vermiculite containing 1/8 (0.13 x) the water of the controls. Within 12–15 h, the osmotic potential of the elongating region had decreased to-12 bar, but it was-7 bar in the controls. This osmoregulation involved a true solute accumulation by the hypocotyls, since cell volume and turgor were virtually the same regardless of the water regime. The hypocotyls having low water potentials elongated slowly but, when deprived of their cotyledons, did not elongate or accumulate solute. This result indicated a cotyledonary origin for the solutes and a dependence of slow growth on osmotic adjustment. The translocation of nonrespired dry matter from the cotyledons to the seedling axis was unaffected by the availability of water, but partitioning was altered. In the first 12 h, dry matter accumulated in the elongating region of the 0.13 x hypocotyls, and osmotic adjustment occurred. The solutes involved were mostly free amino acids, glucose, fructose, and sucrose, and these accounted for most of the increased dry weight. After osmotic adjustment was complete, dry matter ceased to accumulate in the hypocotyls and bypassed them to accumulate in the roots, which grew faster than the control roots. The proliferation of the roots resulted in an increased root/shoot ratio, a common response of plants to dry conditions.Osmotic adjustment occurred in the elongating region of the hypocotyls because solute utilization for growth decreased while solute uptake continued. Adjustment was completed when solute uptake subsequently decreased, and uptake then balanced utilization. The control of osmotic adjustment was therefore the rate of solute utilization and, secondarily, the rate of solute uptake. Elongation was inhibited by unknown factors(s) despite the turgor and substrates associated with osmotic adjustment. The remaining slow elongation depended on osmotic adjustment and represented some optimum between the necessary inhibition for solute accumulation and the necessary growth for seedling establishment.     

Tissue water relations and leaf growth of tropical corn cultivars under water deficits

Abstract This study reports on the effect of water deficit on the tissue water relations and leaf growth of six corn cultivars, growing in glasshouse conditions, in order to understand growth responses to drought of tropical corn. A mild water-stress treatment was imposed slowly; plants reached a minimum pre-dawn leaf water potential of about –1.5 MPa by day 12 after watering was withheld. Analysis of the water relation characteristics of growing leaves using the pressure–volume technique demonstrated that under water deficits all the cultivars changed their moisture-release curves compared with irrigated plants. Osmotic potential at full turgor was lowered in water-stressed plants of all the genotypes and the degree of such change was between 0.34 MPa and 0.58 MPa. Thus, turgor pressure was lost at a lower water potential in water-stressed plants than in irrigated plants of all the varieties. Volumetric elastic moduli were also increased under water deficits and the increase ranged between 10% and 141% among the cultivars. In all the genotypes, the stress imposed led to a reduction of leaf area and dry matter accumulation. Leaf expansion was very sensitive to low turgor pressure and it ceased when turgor reached 0.2 MPa. Thus, varieties able to maintain a higher degree of turgor pressure (i.e. by osmotic adjustment) under water deficits may be able to prolong leaf growth.     

Genotypic variation in tissue water relations of leaves and roots of black walnut (Juglans nigra) seedlings

Genetic variation in the drought response of leaf and root tissue water relations of seedlings of eight sources of black walnut ( Juglans nigra L.) was investigated using the pressure-volume technique. Tissue water relations were characterized at three stages of a drying cycle during which well-watered plants were allowed to desiccate and then were reirrigated.
Sources varied both in the capacity for and degree of leaf and root osmotic adjustment, and in the mechanism by which it was achieved. A decrease in osmotic potential at the turgor loss point (ψ_πp) of 0.4 MPa was attributable to increased leaf tissue elasticity in seedlings of four sources, while seedlings of an Ontario source exhibited a 0.7–0.8 MPa decline in ψ_πp as a result of both increased solute content and increased leaf tissue elasticity. Seedlings of a New York source showed no detectable osmotic adjustment.
In roots, decreased ψ_πp (osmotic potential at full hydration) and ψ_πp were observed under drought. Sources that exhibited significant leaf osmotic adjustment also generally showed a similar response in roots. Tissue elasticity and ψ_πp of roots were higher than those of shoots, whereas ψ_πp of the two organs was similar for most sources. Because of greater elasticity, roots exhibited a more gradual decline in turgor and total water potential than did leaves as tissue relative water content decreased.     

Abscisic acid accumulation in the roots of nutrient-limited plants: its impact on the differential growth of roots and shoots

We describe the involvement of abscisic acid (ABA) in the control of differential growth of roots and shoots of nutrient limited durum wheat plants. A ten-fold dilution of the optimal concentration of nutrient solution inhibited shoot growth, while root growth remained unchanged, resulting in a decreased shoot/root ratio. Addition of fluridone (inhibitor of ABA synthesis) prevented growth allocation in favour of the roots. This suggests the involvement of ABA in the redirecting of growth in favour of roots under limited nutrient supply. The ABA content was greater in shoots and growing apical root parts of starved plants than in nutrient sufficient plants. Accumulation of ABA in shoots of nutrient deficient plants was linked to a decrease in leaf turgor. Increased flow of ABA in the phloem apparently contributed to the accumulation of ABA in the apical part of the roots. Thus, partitioning of growth between roots and shoots of wheat plants limited in mineral nutrients appears to be modulated by accumulation of ABA in roots. This ABA may originate in the shoots, where its synthesis is stimulated by the loss of leaf turgor.     

Changes in the Water Balance and Photosynthesis of Onion, Bean and Cotton Plants under Saline Conditions

The osmotic concentration (osmotic potential) of onion leaf sap did not adjust to chloride salinity, and consequently water potential, turgor, stomatal aperture and transpiration were reduced. Although osmotic concentration of bean and cotton leaf sap did adjust to a saline root medium and turgor was no less in the salinized plants than in the controls, stomata of the salinized plants remained only partly open and transpiration was reduced. Net photosynthesis of onion plants was reduced by salinity (this effect being much enhanced in a hot dry atmosphere) but it could be rapidly raised to the level of the controls by inducing elevated leaf turgor. Stomatal closure was initially responsible for most of the ～30 % reduction in photosynthesis of salinized beans. This was due to interference with CO₂ diffusion and could be overcome by raising the CO₂ concentration in the air. At a later stage of growth, salinity affected the light reaction of bean photosynthesis, and elevation of the air CO₂ had little effect. Closure of stomata of salinized cotton plants had only a relatively small effect on net photosynthesis. Light intensity and CO₂ concentration experiments showed that salinity was reducing the photosynthesis of cotton leaves mainly by affecting the light reaction of photosynthesis. It is concluded that chloride salinity does affect the water balance and rate of photosynthesis of plants and that the nature and degree of the effects will depend upon climatic conditions and may be very different between plant species and in the same species at different periods of growth.     

Exogenous proline effects on water relations and ions contents in leaves and roots of young olive

The ability of exogenous compatible solutes, such as proline, to counteract salt inhibitory effects was investigated in 2-year-old olive trees (Olea europaea L. cv. Chemlali) subjected to different saline water irrigation levels supplied or not with exogenous proline. Leaf water relations [relative water content (RWC), water potential], photosynthetic activity, leaf chlorophyll content, and starch contents were measured in young and old leaves. Salt ions (Na⁺, K⁺, and Ca²⁺), proline and soluble sugars contents were determined in leaf and root tissues. Supplementary proline significantly mitigated the adverse effects of salinity via the improvement of photosynthetic activity (Pn), RWC, chlorophyll and carotenoid, and starch contents. Pn of young leaves in the presence of 25 mM proline was at 1.18 and 1.38 times higher than the values recorded under moderate (SS1) and high salinity (SS2) treatments, respectively. Further, the proline supply seems to have a more important relaxing effect on the photosynthetic chain in young than in old leaves of salt-stressed olive plants. The differential pattern of proline content between young and old leaves suggests that there would be a difference between these tissues in distinguishing between the proline taken from the growing media and that produced as a result of salinity stress. Besides, the large reduction in Na⁺ accumulation in leaves and roots in the presence of proline could be due to its interference in osmotic adjustment process and/or its dilution by proline supply. Moreover, the lower accumulation of Na⁺ in proline-treated plants, compared to their corresponding salinity treatment, displayed the improved effect of proline on the ability of roots to exclude the salt ions from the xylem sap flowing to the shoot, and thus better growth rates.     

Leaf Water Relations, Osmotic Adjustment, Cell Membrane Stability, Epicuticular Wax Load and Growth as Affected by Increasing Water Deficits in Sorghum

A field experiment was conducted with a non-irrigated waterstress treatment and an irrigated control using four sorghum(Sorghum bicolor L. Moench) cultivars. We investigated the effectsof water deficits on leaf water relations, osmotic adjustment,stomatal conductance, cuticular conductance, cell membrane stability(CMS) measured by the polyethylene glycol (PEG) test, epicuticularwax load (EWL), cytoplasmic lipid content, solute concentrationin cell sap, and growth. Osmotic adjustment was observed under water deficit conditions.Lower osmotic potential enabled plants to maintain turgor anddecreased the sensitivity of turgor-dependent processes. Sugarand K were identified as the major solutes contributing to osmoticpotential in sorghum. Sugar and K concentrations in cell sapincreased by 37·4% and 27%, respectively, under waterdeficit conditions in favour of decreasing osmotic potential.Stomatal conductance and cuticular conductance were lower inthe non-irrigated plants. A wide range in CMS among four cultivarswas observed. CMS increased with increasing water deficits.EWL increased on leaves of water deficient plants and was positivelycorrelated with cuticular conductance and CMS. Membrane phospholipidcontent increased in water-stressed plants. CMS as measured by the PEG test, was influenced by EWL, cuticularthickness, and osmotic concentration of leaf tissues. The cultivarswhich maintained higher CMS, higher EWL, lower cuticular conductance,higher turgor and higher osmotic adjustment under water deficitconditions were identified as drought tolerant. Key words: Sorghum bicolor, cell membrane stability, leaf water relationsosmotic adjustment, water stress     

Salinity effects on photosynthesis in isolated mesophyll cells of cowpea leaves

Mesophyll cells from leaves of cowpea (Vigna unquiculata [L.] Walp.) plants grown under saline conditions were isolated and used for the determination of photosynthetic CO₂ fixation. Maximal CO₂ fixation rate was obtained when the osmotic potential of both cell isolation and CO₂ fixation assay media were close to leaf osmotic potential, yielding a zero turgor pressure. Hypotonic and hypertonic media decreased the rate of photosynthesis regardless of the salinity level during plant growth. No decrease in photosynthesis was obtained for NaCl concentrations up to 87 moles per cubic meter in the plant growing media and only a 30% decrease was found at 130 moles per cubic meter when the osmotic potential of cell isolation and CO₂ fixation media were optimal. The inhibition was reversible when stress was relieved. At 173 moles per cubic meter NaCl, photosynthesis was severely and irreversibly inhibited. This inhibition was attributed to toxic effects caused by high Cl⁻ and Na⁺ accumulation in the leaves. Uptake of sorbitol by intact cells was insignificant, and therefore not associated with cell volume changes. The light response curve of cells from low salinity grown plants was similar to the controls. Cells from plants grown at 173 moles per cubic meter NaCl were light saturated at a lower radiant flux density than were cells from lower salinity levels.     

Water relations of the tos1 tomato mutant at contrasting evaporative demand

The tos1 (tomato osmotically sensitive) mutant, isolated from an in vitro screen of root growth during osmotic stress, was less sensitive to exogenous ABA, but accumulated more ABA under osmotic stress than WT plants. We assessed growth and water relations characteristics of hydroponically grown tos1 seedlings (in the absence of osmotic stress) at low and high evaporative demands. Growth of tos1 was severely inhibited at both high and low evaporative demands. Twenty DAS, WT and tos1 genotypes had a similar leaf water and turgor potential, but mature tos1 plants (45 day old) showed a significant diurnal loss of leaf turgor, with recovery overnight. Increased evaporative demand increased turgor loss of tos1 plants. High evaporative demand at the beginning of the day decreased stomatal conductance of tos1, without diurnal recovery, thus whole plant transpiration was decreased. De-topped tos1 seedlings showed decreased root hydraulic conductance and had a 1.4-fold increase in root ABA concentration. Impaired root function of tos1 plants failed to meet transpirational water demand and resulted in shoot turgor loss, stomatal closure and growth inhibition.     

Cell Membrane Stability and Leaf Water Relations as Affected by Potassium Nutrition of Water-Stressed Maize

A field experiment was conducted to investigate the effect ofK nutrition under water stress conditions on cell membrane stabilitymeasured by the polyethylene glycol test, plant growth, internalplant water relations and solute and mineral concentrationsin maize (Zea mays L.). Water-stressed plants showed greateradaptation to water deficits at higher K levels. Cell membranestability increased, leaf water potential and osmotic potentialdecreased, turgor potential increased and stomatal resistancedecreased with increasing K nutrition. Osmotic adjustment wasevident and it may have been influenced by increased K⁺ concentrationsin leaf tissues with increasing K nutrition. Higher leaf thicknessand higher leaf water content were observed at higher K levels.Results suggested that higher supplies of K nutrition may increaseplant production during periods of water stress. Key words: Zea mays L., cell membrane stability, leaf water potential, osmotic adjustment, osmotic potential, potassium nutrition, water stress     

Salt tolerance of Beta macrocarpa is associated with efficient osmotic adjustment and increased apoplastic water content

The chenopod Beta macrocarpa Guss (wild Swiss chard) is known for its salt tolerance, but the mechanisms involved are still debated. In order to elucidate the processes involved, we grew wild Swiss chard exposed to three salinity levels (0, 100 and 200 mm NaCl) for 45 days, and determined several physiological parameters at the end of this time. All plants survived despite reductions in growth, photosynthesis and stomatal conductance in plants exposed to salinity (100 and 200 mm NaCl). As expected, the negative effects of salinity were more pronounced at 200 mm than at 100 mm NaCl: (i) leaf apoplastic water content was maintained or increased despite a significant reduction in leaf water potential, revealing the halophytic character of B. macrocarpa; (ii) osmotic adjustment occurred, which presumably enhanced the driving force for water extraction from soil, and avoided toxic build up of Na⁺ and Cl^– in the mesophyll apoplast of leaves. Osmotic adjustment mainly occurred through accumulation of inorganic ions and to a lesser extent soluble sugars; proline was not implicated in osmotic adjustment. Overall, two important mechanisms of salt tolerance in B. macrocarpa were identified: osmotic and apoplastic water adjustment.     

Drought resistance of mycorrhizal pepper plants independent of leaf P concentration - response in gas exchange and water relations

Pepper ( Capsicum annuum L.) plants with and without the VA-mycorrhizal fungus Ghmus deserticola Trappe. Bloss and Menge (VAM and NVAM. respectively), were drought acclimated by four drought cycles (DA) or kept well watered (NDA). All plants were then subjected to an additional drought followed by a 3-day irrigation recovery period. Measurements of water relations, gas exchange and carbohydrates were made at selected intervals throughout the drought cycles and recovery. To equalize growth and avoid higher P in VAM plants. NVAM plants received higher P fertilization. Consequently, similar transpirational surface and shoot mass were achieved in all treatments, but NVAM had a higher tissue P concentration than VAM plants. Plants that were either VAM or DA, but especially the VAM-DA plants, tended to be high in net photosynthetic flux (A), A per unit of tissue P concentration (A/P), stomatal conductance (g) or leaf turgor (Ψ_p) during high environmental stress or recovery from stress. During this time, NVAM-NDA plants had low A. A/P and leaf chlorophyll, but high soluble carbohydrate concentrations in their leaves. All VAM and DA plants had some osmotic adjustment compared to the NVAM-NDA plants, but VAM-DA plants had the most. Osmotic adjustment was not due to accumulation of soluble carbohydrate. The high turgor, A and g in the VAM-DA plants during and following environmental stress indicated superior drought resistance of these plants; however, osmotic adjustment was only apparent during recovery and cannot account for the observed drought resistance during environmental stress. Drought resistance of VAM-DA plants was not attributable to high leaf P concentration or confounded by differences in plant transpirational surface.     

Proline accumulation and the adaptation of cultured plant cells to water stress

The transfer of cultured tomato cells (Lycopersicon esculentum cv VFNT-Cherry) to a low water potential environment resulted in an increased dry weight to fresh weight ratio accompanied by a rapid accumulation of proline. Proline content continued to increase as osmotic adjustment and growth occurred. The initial increase in proline concentration was accompanied by a drop in turgor. However, proline levels continued to increase with a gain in turgor during osmotic adjustment. Thus, the accumulation of proline depended not only on cell water potential, or on the initial loss of turgor but more closely on cell osmotic potential. The ultimate level of proline depended on the level of adaptation. Proline levels remained high after more than 100 cell generations in low water potential media, but declined rapidly after transfer to media with a less negative water potential. Addition of exogenous proline to the medium during water stress and during osmotic downshock alleviated the normally resulting inhibition of growth. The results suggest a positive role for proline accumulation in adaptation of cells to changing external water potentials.     

Effect of high salinity on Atriplex portulacoides: Growth,leaf water relations and solute accumulation in relation with osmotic adjustment

Atriplex (Halimione) portulacoides is a halophyte with potential interest for saline soil reclamation and phytoremediation. Here, we assess the impact of salinity reaching up to two-fold seawater concentration (0–1000 mM NaCl) on the plant growth, leaf water status and ion uptake and we evaluate the contribution of inorganic and organic solutes to the osmotic adjustment process. A. portulacoides growth was optimal at 200 mM NaCl but higher salinities (especially 800 and 1000 mM NaCl) significantly reduced plant growth. Na⁺ and Cl⁻ contents increased upon salt exposure especially in the leaves compared to the roots. Interestingly, no salt-induced toxicity symptoms were observed and leaf water content was maintained even at the highest salinity level. Furthermore, leaf succulence and high instantaneous water use efficiency (WUE_i) under high salinity significantly contributed to maintain leaf water status of this species. Leaf pressure–volume curves showed that salt-challenged plants adjusted osmotically by lowering osmotic potential at full turgor (Ψ_π¹⁰⁰) along with a decrease in leaf cell elasticity (values of volumetric modulus elasticity (ε) increased). As a whole, our findings indicate that A. portulacoides is characterized by a high plasticity in terms of salt-response. Preserving leaf hydration and efficiently using Na⁺ for the osmotic adjustment especially at high salinities (800–1000 mM NaCl), likely through its compartmentalization in leaf vacuoles, are key determinants of such a performance. The selective absorption of K⁺ over Na⁺ in concomitance with an increase in the K⁺ use efficiency also accounted for the overall plant salt tolerance.     

Leaf expansion of four sunflower (Helianthus annuus L.) cultivars in relation to water deficits. II. Diurnal patterns during stress and recovery

Abstract. Leaf expansion of four sunflower cultivars ( Helianthus annuus L. cvs. Hysun 31, Havasupai, Hopi and Seneca) was monitored continuously in a growth cabinet through the final stages of a drying cycle and then throughout the first 2 days after rewatering in order to study the responses of leaf expansion to water deficits. Comparable plants were also measured throughout a diurnal cycle in a glasshouse.
In the cabinet, leaf extension was faster in the dark than in the light, but an extended dark period suppressed leaf extension. At similar leaf water potentials, the rate of leaf extension was greater in the light than in the dark, but as the osmotic potential was lower in the light than in the dark, the relationship between turgor pressure and leaf extension rate was similar in both environments. Throughout the drying and recovery cycles turgor and leaf extension rate was positively correlated: no significant differences among cultivars were observed.
In the plants grown and measured in the glasshouse, leaf expansion occurred at lower leaf water potentials in stressed than in unstressed plants, but the relationship between leaf expansion and turgor was similar in both stressed and unstressed plants as a result of a lowering of the osmotic potential in the former. Diurnal turgor maintenance resulting from osmotic adjustment was almost half that occurring during a complete drying cycle. During the day, the leaf expansion rate increased linearly with turgor pressure in all cultivars: the expansion rate per unit turgor pressure was greater in the glasshouse than in the growth cabinet. Nocturnal leaf expansion in the stressed and unstressed plants was not, however, correlated with turgor pressure.