Rapid and tissue-specific accumulation of solutes in the growth zone of barley leaves in response to salinity

The aim of the present study was to test whether rapid accumulation of solutes in response to salinity in leaf tissues of barley (Hordeum vulgare L.) contributes to recovery and maintenance of residual elongation growth. Addition of 100 mM NaCl to the root medium caused an immediate reduction close to zero in elongation velocity of the growing leaf 3. After 20–30 min, elongation velocity recovered suddenly, to 40–50% of the pre-stress level. Bulk osmolality increased first, after 60 min, significantly in the proximal half of the elongation zone. Over the following 3 days, osmolality increases became significant in the distal half of the elongation zone, the adjacent, enclosed non-elongation zone and finally in the emerged portion of the blade. The developmental gradient and time course in osmolality increase along the growing leaf was reflected in the pattern of solute (Cl, Na and K) accumulation in bulk tissue and epidermal cells. The partitioning of newly accumulated solutes between epidermis and bulk tissue changed with time. Even though solute accumulation does not contribute to the sudden and partial growth recovery 20–30 min after exposure to salt, it does facilitate residual growth from 1 h onwards. This is due to a high sink strength for solutes of the proximal part of the growth zone and its ability to accumulate solutes rapidly and at high rates.Abbreviations EDX analysis Energy-dispersive X-ray analysis - LEV Leaf elongation velocity - LVDT Linear variable differential transformer - REGR Relative elemental growth rate     

Biophysical limitation of leaf cell elongation in source-reduced barley

The biophysical basis of reduced leaf elongation rate in source-reduced barley ( Hordeum vulgare L. cv Golf) was studied. Reduction in source strength was achieved by removing the blade of leaves 1 and 2 at the time leaf 3 had emerged 3.0-6.7 cm from the encircling sheath. Third leaves of source-reduced plants elongated at 10-36% lower velocities than those of control plants. Removal of source leaves had no significant effect on maximum relative elemental growth rates (REGRs) and the length of the elongation zone (42-46 mm) but caused a shift of high REGR towards the basal portion of the elongation zone. Cell turgor was similar between treatments in the zone of maximal REGR (16-24 mm from base), but was significantly lower in source-reduced plants in the distal part of the elongation zone, where REGR was also lower. Throughout the elongation zone, osmolality and growth-associated water potential gradients were significantly smaller in source-reduced plants; bulk concentrations of sugars (hexoses, sucrose) were also lower. However, even in control plants, sugars contributed little to bulk osmotic pressure (6-11%). The most likely biophysical limitation to leaf (cell) elongation in source-reduced barley was a reduction in turgor in the distal half of the elongation zone. It is proposed that in the proximal half, increase in average tissue hydraulic conductance enabled source-reduced plants to maintain turgor and REGR at control level, while spending less energy on solute transport.     

The short-term growth response to salt of the developing barley leaf

Recent results concerning the short-term growth response to salinity of the developing barley leaf are reviewed. Plants were grown hydroponically and the growth response of leaf 3 was studied between 10 min and 5 d following addition of 100 mM NaCl to the root medium. The aim of the experiments was to relate changes in variables that are likely to affect cell elongation to changes in leaf growth. Changes in hormone content (ABA, cytokinins), water and solute relationships (osmolality, turgor, water potential, solute concentrations), gene expression (water channel), cuticle deposition, membrane potential, and transpiration were followed, while leaf elongation velocity was monitored. Leaf elongation decreased close to zero within seconds following addition of NaCl. Between 20 and 30 min after exposure to salt, elongation velocity recovered rather abruptly, to about 46% of the pre-stress level, and remained at the reduced rate for the following 5 d, when it reached about 70% of the level in non-stressed plants. Biophysical and physiological analyses led to three major conclusions. (i) The immediate reduction and sudden recovery in elongation velocity is due to changes in the water potential gradient between leaf xylem and peripheral elongating cells. Changes in transpiration, ABA and cytokinin content, water channel expression, and plasma membrane potential are involved in this response. (ii) Significant solute accumulation, which aids growth recovery, is detectable from 1 h onwards; growing and non-growing leaf regions and mesophyll and epidermis differ in their solute response. (iii) Cuticular wax density is not affected by short-term exposure to salt; transpirational changes are due to stomatal control.     

Control of leaf cell elongation in barley. Generation rates of osmotic pressure and turgor, and growth-associated water potential gradients

In a previous study on the effects of N-supply on leaf cell elongation, the spatial distribution of relative cell elongation rates (RCER), epidermal cell turgor, osmotic pressure (OP) and water potential (Ψ) along the elongation zone of the third leaf of barley was determined (W. Fricke et al. 1997, Planta 202: 522–530). The results suggested that in plants receiving N at fixed relative addition rates (N-supply limitation of growth), cell elongation was rate-limited by the rate of solute provision, whereas in plants growing on complete nutrient solution containing excessive amounts of N (N-demand limitation), cell elongation was rate-limited by the rate of water supply or wall yielding. In the present paper, these suggestions were tested further. The generation rates of cell OP, turgor and Ψ along the elongation zone were calculated by applying the continuity equation of fluid dynamics to the previous data. To allow a more conclusive interpretation of results, anatomical data were collected and bulk solute concentrations determined. The rate of OP generation generally exceeded the rate of turgor generation. As a result, negative values of cell Ψ were created, particularly in demand-limited plants. These plants showed highest RCER along the elongation zone and a Ψ gradient of at least −0.15 MPa between water source (xylem) and expanding epidermal cells. The latter was similar to a theoretically predicted value (−0.18 MPa). Highest rates of OP generation were observed in demand-limited plants, with a maximum rate of 0.112 MPa · h⁻¹ at 16–20 mm from the leaf base. This was almost twice the rate in N-supply-limited plants and implied that the cells in the leaf elongation zone were capable of importing (or synthesising) every minute almost 1 mM of osmolytes. Potassium, Cl⁻ and NO₃ ⁻ were the main inorganic osmolytes (only determined for demand-limited plants). Their concentrations suggest that, unlike the situation in fully expanded epidermal cells, sugars are used to generate OP and turgor. Anatomical data revealed that the zone of lateral cell expansion extended distally beyond the zone of cell elongation. It is concluded that leaf cell expansion in barley relies on high rates of water and solute supply, rates that may not be sustainable during periods of sufficient N-supply (limitation by water supply: Ψ gradients) or limiting N-supply (limitation by solute provision: reduced OP-generation rates). To minimise the possibility of growth limitation by water and osmolyte provision, longitudinal and lateral cell expansion peak at different locations along the growth zone. Received: 15 October 1997 / Accepted: 12 March 1998     

Stress-induced osmotic adjustment in growing regions of barley leaves

Young barley seedlings were stressed using nutrient solutions containing NaCl or polyethylene glycol and measurements were made of leaf growth, water potential, osmotic potential and turgor values of both growing (basal) and nongrowing (blade) tissues. Rapid growth responses similar to those noted for corn (Plant Physiology 48: 631-636) were obtained using either NaCl or polyethylene glycol treatments by which exposure of seedlings to solutions with water potential values of −3 to −11 bars effected an immediate cessation of leaf elongation with growth resumption after several minutes or hours. Latent periods were increased and growth resumption rates were decreased as water potential values of nutrient solutions were lowered. In unstressed transpiring seedlings, water potential and osmotic potential values of leaf basal tissues were usually −6 to −8 bars, and −12 to −14 bars, respectively. These tissues began to adjust osmotically when exposed to any of the osmotic solutions, and hourly reductions of 1 to 2 bars in both water potential and osmotic potential values usually occurred for the first 2 to 4 hours, but reduction rates thereafter were lower. When seedlings were exposed to solutions with water potential values lower than those of the leaf basal tissues, growth resumed about the time water potential values of those tissues fell to that of the nutrient solution. After 1 to 3 days of seedling exposure to solutions with different water potential values, cumulative leaf elongation was reduced as the water potential values of the root medium were lowered. Reductions in water potential and osmotic potential values of tissues in leaf basal regions paralleled growth reductions, but turgor value was largely unaffected by stress. In contrast, water potential, osmotic potential, and turgor values of leaf blades were usually changed slightly regardless of the degree and duration of stress, and blade water potential values were always higher than water potential values of the basally located cells. It is hypothesized that blades have high water potential values and are generally unresponsive to stress because water in most of the mesophyll cells in this area does not exchange readily with water present in the transpiration stream.     

Spatial distribution of growth rates and of epidermal cell lengths in the elongation zone during leaf development in Lolium perenne L.

Relative elemental growth rates (REGR) and lengths of epidermal cells along the elongation zone of Lolium perenne L. leaves were determined at four developmental stages ranging from shortly after emergence of the leaf tip to shortly before cessation of leaf growth. Plants were grown at constant light and temperature. At all developmental stages the length of epidermal cells in the elongation zone of both the blade and sheath increased from 12 m at the leaf base to about 550 m at the distal end of the elongation zone, whereas the length of epidermal cells within the joint region only increased from 12 to 40 m. Throughout the developmental stages elongation was confined to the basal 20 to 30 mm of the leaf with maximum REGR occurring near the center of the elongation zone. Leaf elongation rate (LER) and the spatial distributions of REGR and epidermal cell lengths were steady to a first approximation between emergence of the leaf tip and transition from blade to sheath growth. Elongation of epidermal cells in the sheath started immediately after the onset of elongation of the most proximal blade epidermal cells. During transition from blade to sheath growth the length of the blade and sheath portion of the elongation zone decreased and increased, respectively, with the total length of the elongation zone and the spatial distribution of REGR staying near constant, with exception of the joint region which elongated little during displacement through the elongation zone. Leaf elongation rate decreased rapidly during the phase when only the sheath was growing. This was associated with decreasing REGR and only a small decrease in the length of the elongation zone. Data on the spatial distributions of growth rates and of epidermal cell lengths during blade elongation were used to derive the temporal pattern of epidermal cell elongation. These data demonstrate that the elongation rate of an epidermal cell increased for days and that cessation of epidermal cell elongation was an abrupt event with cell elongation rate declining from maximum to zero within less than 10 h.Abbreviations LER leaf elongation rate - REGR relative elemental growth rates     

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.     

Short- and Long-Term Effects of Salinity on Leaf Growth in Rice (Oryza sativa L.)

Addition of 50 mM NaCl to Oryza sativa L. had little effectupon the time of leaf initiation, but leaf mortality prior tothe normal phase of senescence was increased and the onset ofsenescence was advanced. There was no significant effect uponthe day-to-day pattern of growth, nor upon the ultimate length,of leaves that were developing at the time of, or shortly after,salinization with 50 mM NaCI. Leaves that developed after prolongedexposure of the plants to salinity were shorter. Addition ofNaCl, KC₁ or mannitol to the root medium brought about a cessationof leaf elongation within one minute. Growth at a reduced raterestarted abruptly after a lag period that depended upon theexternal concentration. Elongation rate recovered its originalvalue within 24 h after exposure to 50 mM NaCl, though not athigher concentrations. Addition of NaCl at concentrations upto 100 mM elicited no short-term effects upon photosyntheticgas exchange. Na uptake contributed to osmotic adjustment ofthe growing zone. When plants were rapidly exposed to 50 mMNaCl, no change in turgor pressure was detectable in the growingzone with the resolution of the miniature pressure probe used(about 70 kPa). It is concluded that the initial growth reductionin rice caused by salinization is due to a limitation of watersupply. A clear distinction is made between the initial effectsof low salinity which are recoverable and the long-term effectswhich result from the accumulation of salt within expanded leaves. Key words: Leaf elongation, gas exchange, photosynthesis, water relations     

Leaf Elongation in Relation to Leaf Water Potential in Soybean

Leaf water potential, turgor pressure, and leaf elongation ratewere measured in soybeans growing in controlled environmentchambers, greenhouses, and outdoors. Plants in chambers hadthe highest water potentials and turgor pressures, and plantsoutdoors the lowest. In all three environments there was a linearrelationship between elongation rate and turgor pressure. Leavesof plants in drier environments required less turgor for elongation,and showed a greater increase in elongation rate per unit increasein turgor. Elongation rates over a 72 h period were equal inthe three environments. Leaves reached the largest final sizein the greenhouse (intermediate in water potential). Epidermalcells were larger in chamber- and greenhouse-grown leaves thanin leaves of plants grown outdoors. The number of epidermalcells per leaf was greater in the greenhouse and outdoors thanin the chamber. Leaf elongation characteristics of greenhouseplants were duplicated by mildly stressing chamber plants, andleaf elongation characteristics of field plants were duplicatedby more severely stressing chamber plants. Leaves of mildlystressed chamber plants also reached a larger final size thanleaves of more severely stressed chamber plants, or leaves ofcontrol plants in the chamber. Water stress in the chamber increasedthe number of epidermal cells per leaf. More severe water stressin the chamber reduced epidermal cell size. Based on the waterstress experiments it is concluded that the differences in plantwater status in the chamber, greenhouse, and field caused differencesin elongation characteristics, and were responsible for thedifferences in leaf size.     

The intercellular distribution of vacuolar solutes in the epidermis and mesophyll of barley leaves changes in response to NaCl

Concentrations of inorganic and organic solutes were measuredin sap extracted from individual mesophyll and epidermal cellsof the third leaf of barley. During the development of the thirdleaf plants were grown in various salt solutions (NaCl; 2, 50,100, and 150 mM, KCI; 100 mM or KNO₃; 100 mM). Leaves were analysed2–4 d after full expansion. Cell-sap was extracted usinga modified pressure probe and analysed for its osmolality, concentrationsof P, Na⁺ K⁺ Ca²⁺, and Cl^– and, in some cases, of nitrate,hexoses and total amino acids. Salt treatment caused differentialchanges in the concentrations of solutes in mesophyll and epidermalcells, but did not affect the basic pattern of solute compartmentationbetween these tissues. Calcium was found at osmotically significantconcentrations only in the epidermis, whereas P and organicsolutes were almost exclusively found in the mesophyll. Chlorideand Na⁺ accumulated preferentially in the epidermis, althoughmesophyll concentrations also increased considerably. At 150mM external NaCl, mesophyll cells contained 302 mM Na and 167mM Cl^–, compared to 29 mM Na⁺ and 16 mM Cl^– in thecontrol. Mesophyll Cl^– levels were even higher in the100 mM KCl treatment (216 mM) where mesophyll and epidermalK⁺ accumulated to 424 and 491 mM, respectively. These huge increasesin mesophyll Na⁺ Cl^– and K⁺ were not associated with abreakdown in leaf performance since net rates of photosynthesisdecreased only by less than 20%. Under control (2 mM NaCl) conditions,solutes followed patterned gradients between the various epidermalcell types. The extent of these gradients changed with leafage. During 50 mM NaCl treatment, gradients in Cl^–, nitrateand malate concentrations progressively disappeared, with malateconcentrations approaching zero. Potassium and Na⁺ exhibitedaltered distribution profiles, whereas Ca²⁺ distribution wasunaffected. NaCl-dependent increases in osmolalities differedbetween cells. Exposure of plants to 150 mM NaCl caused qualitativelysimilar changes in both epidermal solute and osmolality profiles,although absolute values differed from those at 50 mM NaCl.In particular, epidermal Cl^– and Na⁺ increased to about500 mM and K⁺ disappeared (<<5 mM) from the vacuole ofcertain epidermal cell types completely. Key words: Barley leaf epidermis, mesophyll, salt stress, single-cell analysis, vacuolar solutes     

Do pH changes in the leaf apoplast contribute to rapid inhibition of leaf elongation rate by water stress? Comparison of stress responses induced by polyethylene glycol and down‐regulation of root hydraulic conductivity

We have dissected the influences of apoplastic pH and cell turgor on short-term responses of leaf growth to plant water status, by using a combination of a double-barrelled pH-selective microelectrodes and a cell pressure probe. These techniques were used, together with continuous measurements of leaf elongation rate (LER), in the (hidden) elongating zone of the leaves of intact maize plants while exposing roots to various treatments. Polyethylene glycol (PEG) reduced water availability to roots, while acid load and anoxia decreased root hydraulic conductivity. During the first 30 min, acid load and anoxia induced moderate reductions in leaf growth and turgor, with no effect on leaf apoplastic pH. PEG stopped leaf growth, while turgor was only partially reduced. Rapid alkalinization of the apoplast, from pH 4.9 ± 0.3 to pH 5.8 ± 0.2 within 30 min, may have participated to this rapid growth reduction. After 60 min, leaf growth inhibition correlated well with turgor reduction across all treatments, supporting a growth limitation by hydraulics. We conclude that apoplastic alkalinization may transiently impair the control of leaf growth by cell turgor upon abrupt water stress, whereas direct hydraulic control of growth predominates under moderate conditions and after a 30-60 min delay following imposition of water stress.     

Molecular Cloning of emip,a Member of the Major Intrinsic Protein (MIP) Gene Family,Preferentially Expressed in Epidermal Cells of Barley Leaves

Cloning and characterization of a barley gene named emip is reported that encodes a member of the major intrinsic protein family. A λ-unizap cDNA library synthesized from poly A⁺?mRNA of leaf epidermis was screened differentially with epidermis-versus mesophyll-derived probes. One of the clones epi 3-2 was sequenced and further analyzed. The open reading frame of the full length clone codes for a polypeptide of 288 amino acids with a molecular mass of 30,634 Da exhibiting a high degree of homology with members of the major intrinsic protein family. Hydropathy analysis predicts six potential membrane-spanning helices. mRNA levels were high in the growing zone of barley leaves and declined towards the tip of the fully expanded leaf blade. Expression was high in epidermal strips, lower in roots and very low in the leaf mesophyll. In order of decreasing response, wilting, salt shock and heat shock resulted in stimulated expression. mRNA levels remained low during slow salting up experiments. The expressional pattern suggests a role of EMIP in turgor regulation, particularly under stress.     

Effects of short- and long-term salinity on leaf water relations, gas exchange, and growth in Ipomoea pes-caprae

Ipomoea pes-caprae is widespread in pantropical coastal areas along the beach. The aim of this study was to investigate the salinity tolerance level and physiological mechanisms that allow I. pes-caprae to endure abrupt increases in salinity under brief or prolonged exposure to salinity variations. Xylem sap osmolality (X_osm), leaf water relations, gas exchange, and number of produced and dead leaves were measured at short- (1-7 d) and long- (22-46 d) term after a sudden increase in soil salinity of 0, 85, 170, and 255 mM NaCl. In the short-term, X_osm was not affected by salinity, but in the long-term there was a significant increase in plants grown in presence of salt compared with control plants. After salt addition, the plants showed osmotic stress with temporal cell turgor loss. However, the water potential gradient for water uptake was re-established at 4, 7 and 22 d after salt addition, at 85, 170 and 255 mM NaCl, respectively. In the short-term I. pes-caprae was able to tolerate salinities of up to 255 mM NaCl without significant reduction in carbon assimilation or growth. With the duration of stress, leaf ion concentration continued to increase and reached toxic levels at high salinity with a progressive decrease in photosynthetic rate, reduced leaf formation and accelerated senescence. Then, if high levels of soil salts from tidal inundation occur for short periods, the survival of I. pes-caprae is possible, but prolonged exposure to salinity may induce metabolic damage and reduce drastically the plant growth.     

Decreased reactive oxygen species concentration in the elongation zone contributes to the reduction in maize leaf growth under salinity

Reactive oxygen species (ROS) in the apoplast of cells in the growing zone of grass leaves are required for elongation growth. This work evaluates whether salinity-induced reductions in leaf elongation are related to altered ROS production. Studies were performed in actively growing segments (SEZ) obtained from leaf three of 14-d-old maize (Zea mays L.) seedlings gradually salinized to 150 mM NaCl. Salinity reduced elongation rates and the length of the leaf growth zone. When SEZ obtained from the elongation zone of salinized plants (SEZs) were incubated in 100 mM NaCl, the concentration where growth inhibition was approximately 50%, O2*- production, measured as NBT formazan staining, was lower in these than in similar segments obtained from control plants. The NaCl effect was salt-specific, and not osmotic, as incubation in 200 mM sorbitol did not reduce formazan staining intensity. SEZs elongation rates were higher in 200 mM sorbitol than in 100 mM NaCl, but the difference could be cancelled by scavenging or inhibiting O2*- production with 10 mM MgCl2 or 200 microM diphenylene iodonium, respectively. The actual ROS believed to stimulate growth is *OH, a product of O2*- metabolism in the apoplast. SEZ(s) elongation in 100 mM NaCl was stimulated by a *OH-generating medium. Fusicoccin, an ATPase stimulant, and acetate buffer pH 4, could also enhance elongation in these segments, although both failed to increase ROS activity. These results show that decreased ROS production contributes to the salinity-associated reduction in grass leaf elongation, acting through a mechanism not associated with pH changes.     

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.     

Growth, proline accumulation and water relations of NaCl-selected and non-selected callus lines of Dactylis glomerata L

Sodium chloride-tolerant calli were selected from leaf-derived embryogenic calli of Dactylis glomerata L. on agar solidified medium supplemented with 200 mM NaCl, a concentration lethal to non-selected calli. Growth characteristics, water relations and proline accumulation pattern were compared in selected and non-selected lines. The objective was to gain an understanding of the mechanism(s) of tolerance in the NaCl-tolerant line. Growth in the selected line, as expressed in terms of tolerance index (ratio of fresh wt. on NaCl medium:fresh wt. on NaCl free medium x 100), was greater than that of the non-selected line at all levels of NaCl between 50 and 300 mM. There was no significant difference in proline accumulation in the selected and non-selected lines. Maintenance of turgor by osmotic adjustment was observed in the non-selected line despite decreased growth. In contrast, the selected line lost either the need or the ability to adjust osmotically. There was little or no increase in symplastic osmolality in the selected line when exposed to NaCl. Presumably, selection was made for a salt-excluding tissue that has lost the ability to accumulate solutes and adjust turgor with NaCl stress.     

Participation of Plant Hormones in Growth Resumption of Wheat Shoots Following Short-Term NaCl Treatment

The effects of sodium-chloride salinity on the leaf elongation rate, transpiration rate, cell sap osmolality, and phytohormone content in 7-day-old shoots of durum wheat (Triticum durum L.) were studied. Leaf growth was suppressed under the salinity stress and resumed 1 h after NaCl removal. The resumption of leaf growth coincided with a decrease in the transpiration rate due to the rapid ABA accumulation in the differentiation leaf zone. The increased IAA concentration in the growing leaf zone promoted the formation of the attraction signal. The authors concluded that the changes in phytohormonal status in wheat plants occurred already following short-term (up to 1 h) salinity and were directed to the maintenance of plant growth under these conditions.     

Measurement of a growth-induced water potential gradient in tall fescue leaves

Spatial distribution of cell turgor pressure, cell osmotic pressure and relative elemental growth rate were measured in growing tall fescue leaves ( Festuca arundinacea ). Cell turgor pressure (measured with a pressure probe) was c . 0.55 MPa in expanding cells but increased steeply (+0.3 MPa) in cells where elongation had stopped. However, cell osmotic pressure (measured with a picolitre osmometer) was almost constant at 0.85 MPa throughout the leaf. The water potential difference between the growth zone and the mature zone (0.3 MPa) was interpreted as a growth-induced water potential gradient. This and further implications for the mechanism of growth control are discussed.     

Maintenance of low cl concentrations in mesophyll cells of leaf blades of barley seedlings exposed to salt stress

The concentrations of vacuolar Na⁺ and Cl⁻ in the epidermal and mesophyll cells of the leaf blade and sheath of Hordeum vulgare seedlings (cv California Mariout and Clipper) were measured by means of quantitative electron probe x-ray microanalysis. A preferential accumulation of Cl⁻ in vacuoles of epidermal cells in both blade and sheath and a low level in mesophyll cells of the blade were evident in plants grown in full strength Johnson solution. The concentration of Cl⁻ in the mesophyll cells of the blade remained at a low level after exposure to 50 or 100 millimolar NaCl for 1 day or to 50 millimolar for 4 days, while at the same time the concentration of Cl⁻ in the epidermis and mesophyll of the sheath showed a dramatic increase. Clipper generally contained more Cl⁻ in the mesophyll cells of the blade than California Mariout. A greater accumulation of Na⁺ in the mesophyll of the sheath relative to that of the blade was only apparent after treatment with 100 millimolar NaCl for 1 day or 50 millimolar for 4 days. These results confirm the suggestion that sheath tissue is capable of accumulating excess Cl⁻ (and to a lesser extent Na⁺) and suggest that the site of regulation of Cl⁻ concentration in the barley leaf is located in the mesophyll cells of the blade.     

Growth maintenance of the maize primary root at low water potentials involves increases in cell-wall extension properties, expansin activity, and wall susceptibility to expansins

Previous work on the growth biophysics of maize (Zea mays L.) primary roots suggested that cell walls in the apical 5 mm of the elongation zone increased their yielding ability as an adaptive response to low turgor and water potential (psi w). To test this hypothesis more directly, we measured the acid-induced extension of isolated walls from roots grown at high (-0.03 MPa) or low (-1.6 MPa) psi w using an extensometer. Acid-induced extension was greatly increased in the apical 5 mm and was largely eliminated in the 5- to 10-mm region of roots grown at low psi w. This pattern is consistent with the maintenance of elongation toward the apex and the shortening of the elongation zone in these roots. Wall proteins extracted from the elongation zone possessed expansin activity, which increased substantially in roots grown at low psi w. Western blots likewise indicated higher expansin abundance in the roots at low psi w. Additionally, the susceptibility of walls to expansin action was higher in the apical 5 mm of roots at low psi w than in roots at high psi w. The basal region of the elongation zone (5-10 mm) did not extend in response to expansins, indicating that loss of susceptibility to expansins was associated with growth cessation in this region. Our results indicate that both the increase in expansin activity and the increase in cell-wall susceptibility to expansins play a role in enhancing cell-wall yielding and, therefore, in maintaining elongation in the apical region of maize primary roots at low psi w.