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     

Expression of XET-related genes and its relation to elongation in leaves of barley (Hordeum vulgare L.)

Five cDNA clones were isolated from barley (Hordeum vulgare L.) that encoded mRNAs related to xyloglucan endotransglycosylase (XET). One of the clones encoded a protein with XET activity in vitro. Sequence comparisons revealed five families of XET-related sequences, one of which (containing two of the barley genes) was novel. Hybridization studies using clone-specific probes indicated that the corresponding genes were represented once, or possibly twice, in the barley genome. Treatment of dwarf mutants with gibberellic acid (GA₃), or homozygosity at the ‘slender’ (sln1) locus, resulted in a 2.5-fold (approximately) stimulation of blade elongation rate. Three of the five clones detected mRNAs that were maximally expressed towards the base of the blade, and present in greater quantities in GA₃-treated or slender seedlings. The remaining two clones detected mRNAs that were maximally expressed in the middle of the blade. Relative elemental growth rate (REGR) profiles of leaves growing with or without GA₃ treatment revealed similar maximal REGR values despite a 2.5-fold difference in leaf elongation rate. Segments of GA₃-treated leaves attained their maximal REGR values more rapidly, this being associated with enhanced expression of the three ‘basal’ XET-related mRNAs. Highest XET activities were detected in the base of the elongation zone, and in GA₃-treated seedlings a second activity peak was observed near the distal end of the elongation zone. We conclude that there are likely to be several XET isoenzymes with different expression patterns, and identify those XET-related proteins potentially involved in leaf elongation.     

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     

Changes in osmotic and turgor pressure in response to sugar accumulation in barley source leaves

Pressure-probe measurements and single-cell sampling and analysis techniques were used to determine the effect of photosynthetic production and accumulation of sugars on osmotic and turgor pressures of individual cells of barley ( Hordeum vulgare L.) source leaves. In control plants, the changes in osmotic pressure in individual cells during the photoperiod were different for mesophyll (increase of 276 mOsmol/kg), parenchymatous bundle sheath (PBS; increase of 100 mOsmol/kg) and epidermis (remains constant). There was also an increase in osmotic pressure at the tissue level. Cooling of roots and the shoot apical meristem restricted the export of sugars from leaves, and the resulting changes in osmotic and turgor pressure were monitored. In contrast to the control leaves, mesophyll, PBS, and epidermal cells showed a similar increase in osmotic pressure (up to 500 mOsmol/kg). Cooling also increased the turgor pressure in epidermal and (to a greater extent) PBS cells. The difference in turgor pressure between epidermal and PBS cells is consistent with the presence of a water potential gradient within the leaf, from the vascular bundles towards the leaf surface.     

The biophysics of leaf growth in salt-stressed barley. A study at the cell level

Biophysical parameters potentially involved in growth regulation were studied at the single-cell level in the third leaf of barley (Hordeum vulgare) after exposure to various degrees of NaCl stress for 3 to 5 d. Gradients of elongation growth were measured, and turgor pressure, osmolality, and water potentials (psi) were determined (pressure probe and picoliter osmometry) in epidermal cells of the elongation zone and the mature blade. Cells in the elongation zone adjusted to decreasing external psi through increases in cell osmolality that were accomplished by increased solute loads and reduced water contents. Cell turgor changed only slightly. In contrast, decreases in turgor also contributed significantly to psi adjustment in the mature blade. Solute deposition rates in the elongation zone increased at moderate stress levels as compared with control conditions, but decreased again at more severe NaCl exposure. Growth-associated psi gradients between expanding epidermal cells and the xylem were significant under control and moderate stress conditions (75 mM NaCl) but seemed negligible at severe stress (120 mM NaCl). We conclude that leaf cell elongation in NaCl-treated barley is probably limited by the rate at which solutes can be taken up to generate turgor, particularly at high NaCl levels.     

The decrease in growth of phosphorus-deficient maize leaves is related to a lower cell production

The spatial distribution of leaf elongation and adaxial epidermal cell production in leaf 6 of maize (Zea mays L. cv. Cecilia) plants grown in a growth chamber under two contrasting availabilities of P in the soil was investigated. Lower displacement velocities from 32.5 mm from leaf base and a shorter growth zone were found in low P (LP) leaves compared with control leaves. P deficiency significantly diminished maximum relative elemental growth rate and shifted its location closer to the leaf base. Cells were significantly longer in LP than in control leaves for all positions from the leaf base except at the end of the growth zone. For both treatments it took a similar time for a cell situated at the leaf base to reach the limit of the growth zone. The average length of the cell division zone was decreased by 21% in LP leaves. Significant differences were found in cell production and cell division rates from 12.5 mm from the leaf base although maximum values were similar between P treatments. A shorter zone of cell division with lower cell production rates along most of its length was the regulatory event that decreased cell production, and ultimately leaf elongation rates, in P‐deficient maize plants.     

The effect of gibberellic acid on the response of leaf extension to low temperature

The effect of cooling on leaf extension rate (LER) and on relative elemental growth rate (REGR) was measured in both gibberellic acid (GA)-responsive dwarf barley and in the same barley variety treated with GA. Seedlings were maintained at 20 degrees C while their leaf extension zone (LEZ) temperature was reduced either in steps to -6 degrees C in short-term cooling experiments, or to 10 degrees C for 48 h in long-term cooling experiments. Short-term cooling resulted in a biphasic response in LER, with a clear inflection point identified. Below this point, the activation energy for leaf extension becomes higher. The short-term response of LER to cooling was altered by the application of GA, which resulted in a lower base temperature (Tb), inflection point temperature and activation energy for leaf extension. Both GA-treated and untreated seedlings were less sensitive to cooling maintained for a prolonged period, with LER making a partial recover over the initial 5 h. Although long-term cooling reduced maximum REGR, it resulted in a longer LEZ and an increase in the length of mature interstomatal cells in GA-treated and untreated seedlings. These changes in overall physiology appear to enhance the ability of the leaves to continue expansion at suboptimal temperatures. In both GA-treated and cold-acclimated tissue, the occurrence of a longer LEZ was associated with a lower temperature sensitivity in LER.     

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.     

An analysis of relative elemental growth rate, epidermal cell size and xyloglucan endotransglycosylase activity through the growing zone of ageing maize leaves

The effect of development on leaf elongation rate (LER) andthe distribution of relative elemental growth rate (REGR), epidermalcell length, and xyloglucan endotransglycosylase (XET) activitythrough the growing zone of the third leaf of maize was investigated.As the leaf aged and leaf elongation slowed, the length of thegrowing zone (initially 35 mm) and the maximal REGR (initially0.09 mm mm^–1 h^–1) declined. The decline in REGRwas not uniform through the growth profile. Leaf ageing sawa maintenance of REGR towards the base of the leaf. Epidermalcell size was not constant at a given position in the growingzone, but was seen to increase as the leaf aged. There was apeak of XET activity close to the base of the growing zone.The peak of XET activity preceded the zone of maximum REGR.XET activity declined as leaves aged and their elongation rateslowed. When leaf elongation was complete a distinct peak ofXET activity remained close to the base of the leaf. Key words: Leaf elongation rate (LER), relative elemental growth rate (REGR), xyloglucan endotransglycosylase (XET)     

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     

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.     

Leaf growth and turgor in growing cells of maize (Zea mays L.) respond to evaporative demand under moderate irrigation but not in water-saturated soil

To test whether the inhibition of leaf expansion by high evaporative demand is a result of hydraulic processes, we have followed both leaf elongation rate (LER) and cell turgor in leaves of maize plants either normally watered or in water-saturated soil in which hydraulic resistance at the soil-root interface was abolished. Cell turgor was measured in situ with a pressure probe in the elongating zone of the first and sixth leaves, and LERs of the same leaves were measured continuously with transducers or by following displacements of marks along the growing leaves. Both variables displayed spatial variations along the leaf and positively correlated within the elongating zone. Values peaked at mid-distance of this zone, where the response of turgor to evaporative demand was further dissected. High evaporative demand decreased both LER and turgor for at least 5 h, with dose-effect linear relations. This was observed in five genotypes with appreciable differences in turgor maintenance among genotypes. In contrast, the depressing effects of evaporative demand on both turgor and LER disappeared when the soil was saturated, thereby opposing a negligible resistance to water flow at the soil-root interface. These results suggest that the response of LER to evaporative demand has a hydraulic origin, enhanced by the resistance to water flux at the soil-root interface. They also suggest that turgor is not completely maintained under high evaporative demand, and may therefore contribute to the reductions in LER observed in non-saturated soils.     

Why do leaves and leaf cells of N-limited barley elongate at reduced rates?

The objective of the present study was to assess whether, in barley, nitrogen supply limits the rate of leaf elongation through a reduction in (relative) cell elongation rate and whether this is attributable to a reduced turgor, a reduced availability of osmolytes or, by implication, changed wall properties. Plants were grown on full-strength Hoagland solution (“Hoagland”-plants), or on N-deficient Hoagland solution while receiving N at a relative addition rate of 16 or 8% N · plant-N⁻¹ · d⁻¹ (“16%-” and “8%-plants”). Hoagland-plants were demand-limited, whereas 16%- and 8%-plants were supply-limited in N. Third leaves were analysed for leaf elongation rate and final epidermal cell length, and, within the basal growing region, for the spatial distribution of relative segmental elongation rates (RSER, pin-pricking method), epidermal cell turgor (cell-pressure probe), osmotic pressure (OP, picolitre osmometry) and water potential (Ψ). During the development of the third leaf, plants grew at relative growth rates (relative increase in fresh weight ) of 18.2, 15.6 and 8.1% · d⁻¹ (Hoagland-, 16%- and 8%-plants, respectively). Final leaf length and leaf elongation rate were highest in Hoagland plants (ca. 34.1 cm and 2.33–2.60 mm · h⁻¹, respectively), intermediate in 16%- plants (31.0 cm and 1.89–1.96 mm · h⁻¹) and lowest in 8%-plants (29.4 cm and 1.41–1.58 mm · h⁻¹). These differences were accompanied by only small differences in final cell length, but large differences in cell-flux rates (146, 187 and 201 cells · cell-file⁻¹ · d⁻¹ in 8%-, 16%- and Hoagland-plants, respectively). The length of the growth zone (32–38 mm) was not much affected by N-levels (and nutrient technique). A decrease in RSER in the growth zone distal to 10 mm produced the significant effect of N-levels on leaf elongation rate. In all treatments, cell turgor was almost constant throughout the growing region, as were cell OP and Ψ in 16%- and 8%-plants. In Hoagland-plants, however, cell OP increased by ca. 0.1 MPa within the zone of highest elongation rates and, as a consequence, cell Ψ decreased simultaneously by 0.1 MPa. Cell Ψ increased considerably where elongation ceased. Within the zone where differences in RSERs were highest between treatments (10–34 mm from base) average turgor was lowest, OP highest and Ψ most negative in Hoagland- compared to 8%- and 16%-plants (P < 0.001), but not significantly different between 8%- and 16%-plants. Received: 9 January 1997 / Accepted: 6 March 1997     

Leaf illumination and root cooling inhibit bean leaf expansion by decreasing turgor pressure

Phaseolus vulgaris plants with expanding primary leaves weresubjected to dark-light or light-dark transition at a root temperatureof 25 °C, or to root cooling to 10 °C. Illuminationor darkening caused rapid changes in water flux through theplants and in epidermal turgor pressure when analysed by pressureprobe. However, these were not concurrent with variations inbulk leaf water potential and turgor pressure as determinedby the pressure chamber method. In addition, the turgor pressureof epidermis measured with the pressure probe was invariably0.05 to 0.15 MPa lower than that measured in bulk tissue withthe pressure chamber. Cooling roots to 10°C induced waterstress and wilting. Both techniques indicated a decrease ofturgor pressure, but a 20-30 min lag was observed with the pressurechamber. Due to stomatal closure and decreased transpiration,root-cooled plants regained cell turgor after 5-7 h of cooling,but bulk tissue and epidermal turgor (as well as leaf growthrate) remained significantly lower than control levels. Thesefindings indicate that changes in turgor pressure as the resultof hydraulic signalling are sufficient to explain the rapidchanges in growth rate following illumination or cooling reportedin earlier work (Sattin et al 1990). They also indicate thatdata obtained by use of the pressure chamber must be treatedwith caution. Key words: Phaseolus vulgaris, expansion growth, water relations, hydraulic signalling, pressure probe, pressure chamber     

Effects of Water Stress and Hardening on the Internal Water Relations and Osmotic Constituents of Cotton Leaves

Experiments were designed to test the hypothesis that the internal water relations of leaves are altered when cotton plants (Gossypium hirsutum L.‘Acala SJ-2′) are conditioned by several cycles of water stress. Preliminary experiments suggested that plants so conditioned are less sensitive to water deficits and that the change might be partly explained by an accumulation of solutes or by structural alterations attendant on development under conditions of water stress. Leaves of preconditioned plants maintained turgor to lower values of water potential than did leaves of well-watered plants. Accompanying this change was a lower osmotic potential at any given leaf water content in preconditioned plants. Tissue analysis of several osmotically active solutes indicated that soluble sugars and malate accumulate to about the same levels (dry-weight basis) in both conditioned and unconditioned plants exposed to stress. These accumulations could not account for the turgor change. Analysis of the data on relative water content indicated that the leaves of conditioned plants had less water per unit dry weight than did leaves of controls. This change accounts for a substantial fraction of the difference between the osmotic potential of conditioned and control plants. The results of a simple model suggest that structural changes may play a significant role in explaining differences in the responses of conditioned and control plants to water stress.     

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.     

Influence of arbuscular mycorrhizae and Rhizobium on nutrient content and water relations in drought stressed alfalfa

The objective of this research was to study the effect of drought on nutrient content and leaf water status in alfalfa (Medicago sativa L. cv Aragón) plants inoculated with a mycorrhizal fungus and/or Rhizobium compared with noninoculated ones. The four treatments were: a) plants inoculated with Glomus fasciculatum and Rhizobium meliloti 102 F51 strain, (MR); b) plants inoculated with R. meliloti only (R); c) plants with G. fasciculatum only (M); and d) noninoculated plants (N). Nonmycorrhizal plants were supplemented with phosphorus and nonnodulated ones with nitrogen to achieve similar size and nutrient content in all treatments. Plants were drought stressed using two cycles of moisture stress and recovery. The components of total leaf water potential (osmotic and pressure potentials at full turgor), percentage of apoplastic water volume and the bulk modulus of elasticity of leaf tissue were determined. Macronutrient (N, P, K, Ca, S and Mg) and micronutrient (Co, Mo, Zn, Mn, Cu, Na, Fe and B) content per plant were also measured. Leaves of N and R plants had decreased osmotic potentials and increased pressure potentials at full turgor, with no changes either in the bulk modulus of elasticity or the percentage of apoplastic water upon drought conditions. By contrast, M and MR leaves did not vary in osmotic and turgor potentials under drought stress but had increased apoplastic water volume and cell elasticity (lowering bulk modulus). Drought stress decreased nutrient content of leaves and roots of noninoculated plants. R plants showed a decrease in nutrient content of leaves but maintained some micronutrients in roots. Leaves of M plants were similar in content of nutrients to N plants. However, roots of M and MR plants had significantly lower nutrient content. Results indicate an enhancement of nutrient content in mycorrhizal alfalfa plants during drought that affected leaf water relations during drought stress.     

Does shoot water status limit leaf expansion of nitrogen-deprived barley?

The role of shoot water status in mediating the decline in leaf elongation rate of nitrogen (N)-deprived barley plants was assessed. Plants were grown at two levels of N supply, with or without the application of pneumatic pressure to the roots. Applying enough pressure (balancing pressure) to keep xylem sap continuously bleeding from the cut surface of a leaf allowed the plants to remain at full turgor throughout the experiments. Plants from which N was withheld required a greater balancing pressure during both day and night. This difference in balancing pressure was greater at high (2.0 kPa) than low (1.2 kPa) atmospheric vapour pressure deficit (VPD). Pressurizing the roots did not prevent the decline in leaf elongation rate induced by withholding N at either high or low VPD. Thus low shoot water status did not limit leaf growth of N-deprived plants.     

Cell Shape in Leaves of Drought-stressed Barley Examined by Low Temperature Scanning Electron Microscopy

Cells in leaves of well-watered and slowly drought-stressedbarley seedlings were examined by low temperature scanning electronmicroscopy, when the leaves were turgid, when just wilting,and when sufficiently stressed to prevent either regain of turgor(leaf blades) or regrowth (leaf sheath bases) after rewatering.Deformation of the cell surface was a major response to cellvolume reduction during stress. Folds occurred in the wallsof cells in leaf blades which were just wilting. In severelystressed and damaged plants a range of cell shapes and deformationsoccurred characteristic of a particular cell type and oftenunlike the control cell shape. Cell shape, drought, frost, barley, Hordeum