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.     

Non-hydraulic signals from maize roots in drying soil: inhibition of leaf elongation but not stomatal conductance

Conditions of soil drying and plant growth that lead to non-hydraulic inhibition of leaf elongation and stomatal conductance in maize (Zea mays L.) were investigated using plants grown with their root systems divided between two containers. The soil in one container was allowed to dry while the other container was kept well-watered. Soil drying resulted in a maximum 35% inhibition of leaf elongation rate which occurred during the light hours, with no measurable decline in leaf water potential (_w). Leaf area was 15% less than in control plants after 18 d of soil drying. The inhibition of elongation was observed only when the soil _w declined to below that of the leaves and, thus, the drying soil no longer contributed to transpiration. However, midday root _w in the dry container (-0.29 MPa) remained much higher than that of the surrounding soil (-1.0 MPa) after 15 d of drying, indicating that the roots in drying soil were rehydrated in the dark.To prove that the inhibition of leaf elongation was not caused by undetectable changes in leaf water status as a result of loss of half the watergathering capacity, one-half of the root system of control plants was excised. This treatment had no effect on leaf elongation or stomatal conductance. The inhibition of leaf elongation was also not explained by reductions in nutrient supply.Soil drying had no effect on stomatal conductance despite variations in the rate or extent of soild drying, light, humidity or nutrition. The results indicate that non-hydraulic inhibition of leaf elongation may act to conserve water as the soil dries before the occurrence of shoot water deficits.Symbol _w water potential Contribution from the Missouri Agricultural Experiment Station, Journal Series No. 10881     

Sequence of drought response of maize seedlings in drying soil

Leaf elongation in monocotyledonous plants is sensitive to drought. To better understand the sequence of events in plants subjected to soil drying, leaf elongation and transpiration of maize seedlings ( Zea mays L.) of 4 cultivars were monitored continuously and the diurnal courses of the root and leaf water relations were determined. Results from this study indicate the following sequence of drought response: Leaf elongation decreased before changes in the leaf water relations of non‐growing zones of leaf blades were detected and before transpiration decreased. Reductions in leaf elongation preceded changes in the root water potential (ψ_w). Root ψ_w was not a very sensitive indicator of soil dryness, whereas the root osmotic potential (ψ_s) and root turgor (ψ_p) were more sensitive indicators. The earliest events observed in drying soil were a significant increase in the largest root diameter class (1 720 to 1 960 gm) and a decrease in leaf elongation ( P = 0.08) 2 days after withholding water. Significant increases in root length were observed 2 days later. Soil drying increased the number of fine roots with diameters of <240 µm. Slight increases in soil strength did not affect leaf elongation in the drying soil.     

Quantification of water uptake by arbuscular mycorrhizal hyphae and its significance for leaf growth, water relations, and gas exchange of barley subjected to drought stress

Arbuscular mycorrhizal fungi alleviate drought stress in their host plants via the direct uptake and transfer of water and nutrients through the fungal hyphae to the host plants. To quantify the contribution of the hyphae to plant water uptake, a new split-root hyphae system was designed and employed on barley grown in loamy soil inoculated with Glomus intraradices under well-watered and drought conditions in a growth chamber with a 14-h light period and a constant temperature (15 degrees C; day/night). Drought conditions were initiated 21 days after sowing, with a total of eight 7-day drying cycles applied. Leaf water relations, net photosynthesis rates, and stomatal conductance were measured at the end of each drying cycle. Plants were harvested 90 days after sowing. Compared to the control treatment, the leaf elongation rate and the dry weight of the shoots and roots were reduced in all plants under drought conditions. However, drought resistance was comparatively increased in the mycorrhizal host plants, which suffered smaller decreases in leaf elongation, net photosynthetic rate, stomatal conductance, and turgor pressure compared to the non-mycorrhizal plants. Quantification of the contribution of the arbuscular mycorrhizal hyphae to root water uptake showed that, compared to the non-mycorrhizal treatment, 4 % of water in the hyphal compartment was transferred to the root compartment through the arbuscular mycorrhizal hyphae under drought conditions. This indicates that there is indeed transport of water by the arbuscular mycorrhizal hyphae under drought conditions. Although only a small amount of water transport from the hyphal compartment was detected, the much higher hyphal density found in the root compartment than in the hyphal compartment suggests that a larger amount of water uptake by the arbuscular mycorrhizal hyphae may occur in the root compartment.     

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     

Changes in the concentration of ABA in xylem sap as a function of changing soil water status can account for changes in leaf conductance and growth

Abstract. Maize seedlings ( Zea mays L. John Innes F1 hybrid) were grown in a greenhouse in l-m-long tubes of soil. When the plants were well established, water was withheld from half of the tubes. Control plants were watered every day during the 20-d experimental period. The soil drying treatment resulted in a substantial restriction of stomatal conductance and a limitation in shoot growth, even though there was no detectable difference in the water relations of watered and unwatered plants. From day 7 of the soil drying treatment, xylem ABA concentrations (measured using the sap exuded from detopped plants) were substantially increased in unwatered plants compared to values recorded with sap from plants watered every day. Measurements of water potential through the profile of unwatered soil suggest that xylem ABA concentrations reflects the extent of soil drying. Leaf ABA content was a much less sensitive indicator of the effect of soil drying and during the whole of experimental period there was no significant difference between ABA concentration in leaves of well watered and unwatered plants. In a second set of experiments, ABA was fed to part of the roots of potted maize plants to manipulate xylem ABA concentration. These manipulations suggested that the increases in ABA concentration in xylem sap, which resulted from soil drying, were adequate to explain the observed variation in stomatal conductance and might also explain the restriction in leaf growth rate. These results are discussed in the light of recent work which suggests that stomatal responses to soil drying are partly attributable to an as-yet unidentified inhibitor of stomatal opening.     

An Ecophysiological Comparison of Measurements of the Diurnal Rhythm of the Leaf Elongation and Changes of the Leaf Thickness of Salt-resistant Dicotyledonae and Monocotyledonae

Rozema, J., Arp, W., van Diggelen, J., Kok, E. and Letschert,J. 1987. An ecophysiological comparison of measurements of thediurnal rhythm of the leaf elongation and changes of the leafthickness of salt-resistant Dicotyledonae and Monocotyledonae.—J.exp. Bot. 38: 442–453. The continuous measurement of leaf elongation and leaf thicknesswith the use of a rotation potentiometer set up revealed a rapidand sensitive reaction of halophytic plants to conditions affectingthe plant's water relations. At increased salinity (450 molm^–3 NaCl) the rate of leaf elongation decreased both inAster tripolium and in Sparlina anghca. Increased shrinkageduring the day and a long period for recovery swelling at nightin leaves of Aster iripolium at increased salinity illustratesthat water shortage is part of the cause of salinity-inducedgrowth reduction. All dicotyledonous species analysed (Aster tripolium, A triplexhastata, A. littoralis, Suaeda maritima and Beta vulgaris) showeda day/night ratio of the leaf elongation rate lower than 1,while this ratio was higher than or equal to 1 in Monocotyledons(Spartina anglica, Juncus gerardii, J. maritimus, Festuca rubrassp. litoralis, Elymus pycnanthus). With the exception of Triglochinmaritima none of the monocotyledonous halophytes tested (Sparlinaanglica, Juncus gerardii, J. maritimus, Festuca rubra ssp. litoralis,Elymus pycnanthus) exhibited a diurnal rhythm of leaf thicknesschanges, such as was observed for all dicotyledonous speciesstudied (Aster tripolium, Atriplex hastata, A. littoralis, Salicorniabrachyslachya, Suaeda maritima, Glaux maritima, Odontites vernassp. serotina). The diurnal pattern of the leaf elongation rateand the leaf thickness changes can be explained by variationof photosynthetic rate and transpiration water losses by stomatalclosure in the dark and opening in the light such as shown forthe dicotyledon species Glaux maritima. This difference betweendicot and monocot species in diurnal variation of the leaf elongationrate and leaf thickness may partly be explained in terms ofthe different position of the growth zone and possibly by adifference in elasticity of the tissue of halophytic monocotyledonsand dicotyledons. The consequences of these differences arediscussed. Key words: Leaf elongation rate, leaf thickness, water relations, salt resistance, Dicotyledonae, Monocotyledonae     

Diurnal growth of tall fescue leaf blades : I. Spatial distribution of growth, deposition of water, and assimilate import in the elongation zone

Tall fescue leaf blades elongate at near constant rates during most of the light and dark periods of the diurnal cycle, with the dark rate being higher by 60 to 65%. Our objective was to determine relationships among diurnal rates of leaf elongation, deposition of water and deposition of dry matter (DM) into the elongation zone. Two separate experiments were conducted, both with a 15-hour photoperiod and constant 21°C at the growth zone. Increased rates of leaf elongation in darkness were due to proportionally increased rates of elongation of 4-millimeter segments of the elongation zone. Length of the total elongation zone was 30 millimeters in both light and darkness. The spatial distribution of water contents in the elongation zone varied little during the diurnal cycle. Thus, dark stimulation of leaf elongation rate (+65%) and of water deposition (+77%) into elongation zones were similar. Water content per unit leaf length increased by 50% between the basal and distal limits of the elongation zone, indicating that tissue also grew in the lateral and vertical dimensions. Longitudinal growth of tissue, however, allowed 5 to 7 times more water deposition into the elongation zone than growth in cross-sectional area. This relationship was similar in light and darkness. In both light and darkness net rates of DM deposition (microgram per millimeter leaf length per hour) increased from the zone of cell division towards the region of most active elongation, 10 to 15 millimeters from the ligule, then decreased towards the distal end of the elongation zone. Net DM deposition rates (microgram per hour) integrated over the 30-millimeter elongation zone were similar during light and darkness. Thus, DM in the elongation zone was diluted during darkness as a result of increased water deposition. Net DM deposition rates at and above the distal end of the elongation zone were clearly positive during the light, but were close to zero or negative in darkness. Thus, DM deposition into the elongation zone and the adjacent recently expanded tissue was differentially affected in the diurnal cycle, DM deposition occurred in both tissues in light, but was restricted to the elongation zone in darkness.     

Response of Fructan to Water Deficit in Growing Leaves of Tall Fescue

Changes in dry matter and water-soluble carbohydrate components, especially fructan, were examined in the basal 25 mm of expanding leaf blades of tall fescue (Festuca arundinacea Schreb.) to assess their roles in plant response to water deficit. Water was withheld from vegetative plants grown in soil in controlled-environment chambers. As stress progressed, leaf elongation rate decreased sooner in the light period than it did in the dark period. The decrease in growth rate in the dark period was associated with a decrease in local relative elongation rates and a shortening of the elongation zone from about 25 mm (control) to 15 mm. Dry matter content of the leaf base increased 23% during stress, due mainly to increased water-soluble carbohydrate near the ligule and to increased water-soluble, carbohydrate-free dry matter at distal positions. Sucrose content increased 258% in the leaf base, but especially (over 4-fold) within 10 mm of the ligule. Hexose content increased 187% in the leaf base. Content of total fructan decreased to 69% of control, mostly in regions farther from the ligule. Fructan hydrolysis could account for the hexose accumulated. Stress caused the osmotic potential to decrease throughout the leaf base, but more toward the ligule. With stress there was 70% less direct contribution of low-degree-of-polymerization fructan to osmotic potential in the leaf base, but that for sucrose and hexose increased 96 and 67%, respectively. Thus, fructan metabolism is involved but fructan itself contributes only indirectly to osmotic adjustment.     

Xylem tension affects growth-induced water potential and daily elongation of maize leaves

Diurnal rates of leaf elongation vary in maize (Zea mays L.) and are characterized by a decline each afternoon. The cause of the afternoon decline was investigated. When the atmospheric environment was held constant in a controlled environment, and water and nutrients were adequately supplied to the soil or the roots in solution, the decline persisted and indicated that the cause was internal. Inside the plants, xylem fluxes of water and solutes were essentially constant during the day. However, the forces moving these components changed. Tensions rose in the xylem, and gradients of growth-induced water potentials decreased in the surrounding growing tissues of the leaf. These potentials, measured with isopiestic thermocouple psychrometry, changed because the roots became less conductive to water as the day progressed. The increased tensions were reversed by applying pressure to the soil/root system, which rehydrated the leaf. Afternoon elongation immediately recovered to rapid morning rates. The rapid morning rates did not respond to soil/root pressurization. It was concluded that increased xylem tension in the afternoon diminished the gradients in growth-induced water potential and thus inhibited elongation. Because increased tensions cause a similar but larger inhibition of elongation if maize dehydrates, these hydraulics are crucial for shaping the growth-induced water potential and thus the rates of leaf elongation in maize over the entire spectrum of water availability.     

Diurnal cycles of leaf extension in unsalinized and salinized Beta vulgaris

Abstract. Continuous high resolution measurement of sugar beet leaf extension over 5 d in growth chambers showed average leaf extension rates (LER) in darkness to be from three to six times those in light for plants growing in non-salinized media. The changes in LER in light-dark transitions occurred within seconds, a response which was more rapid than stomatal opening or closing. When the growth medium was salinized to 100 mol m⁻³ NaCl, LER's were reduced by about 50% in darkness and 90% in light, markedly increasing the ratio of dark to light LER.
A 2-d episode of root-zone salinity imposed midway through a 5-d period of measurement decreased LER and produced higher leaf temperatures. LER and diurnal leaf temperature patterns reverted to their pre-salinized levels when root-zone salinity was removed. Thus, the effects of short episodes of high sodium chloride in the growth medium appear to be reversible, suggesting a water stress mechanism of growth reduction rather than toxicity effects of salt.     

Chilling stress response of postemergent cotton seedlings

Early season development of cotton is often impaired by sudden episodes of chilling temperature. We determined the chilling response specific to postemergent 13-day-old cotton (Gossypium hirsutum L. cv. Coker 100A-glandless) seedlings. Seedlings were gradually chilled during the dark period and rewarmed during the night-to-day transition. For some chilled plants, the soil temperature was maintained at control level. Plant growth, water relations and net photosynthesis (P(n)) were analyzed after one or three chilling cycles and after 3 days of recovery. Three chilling cycles led to lower relative growth rate (RGR) compared with controls during the recovery period, especially for plants with chilled shoots and roots. Treatment differences in RGR were associated with net assimilation rate rather than specific leaf area. Both chilling treatments led to loss of leaf turgor during the night-to-day transition; this effect was greater for plants with chilled compared with warm roots. Chilling-induced water stress was associated with accumulation of the osmolyte glycine betaine to the same extent for both chilling treatments. Inhibition of P(n) during chilling was related to both stomatal and non-stomatal effects. P(n) fully recovered after seedlings were returned to control conditions for 3 days. We conclude that leaf expansion during the night-to-day transition was a significant factor determining the magnitude of the chilling response of postemergent cotton seedlings.     

Occurrence of Inverted Water Potential Gradients between Soil and Bean Roots

Measurements with a pressure chamber were made of the xylem water potential of leaves, shoots and roots from bean plants (Pkaseolus vulgaris L. cv. Processor) grown with a 12 hour dark period and natural or artificial light conditions during the day. The water potentials were measured at the end of a dark period and during the light period. Measurements taken at the end of the dark period indicated normal potential gradients within the soil/plant system (leaf < shoot < root < soil), when the matric potential of soil water was relatively high (above ?0.02 bar), and the gradients then also remained normal during the day (natural light). When the soil water potential was ?1 bar or lower in the morning, however, the root xylem water potential was higher than the soil water potential; at very low soil water potentials (< ?4 bar) it remained higher during most of the day. In this case also leaf and shoot xylem water potentials were higher than the soil water potential in the early morning, although decreasing rapidly in daylight. Under artificial light, both leaf and root water potentials were higher than the soil water potential throughout the whole diurnal cycle when the latter potential was below ?4 bar. From measurements of stomatal diffusion resistance, transpiration, relative water content of leaves and of changes in the matric potential of soil water, it was concluded that when the matric potential of soil water was low, water could be taken up by the plant against a water potential gradient. Because leaf xylem water potential was always lower than root xylem water potential, the mechanism involved in the inversion of water potential gradient must be localized in the roots, and probably related to ion uptake. Symbols and abbreviations used in the text: Ψ: Plant water potential (thermocouple psychrometer); Ψx: Xylem water potential (pressure chamber); Ψs: Osmotic potential of xylem sap; Ψm: Matric potential of soil water; RWC: Relative water content.     

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     

LIGHT,TEMPERATURE AND SALINITY EFFECTS ON GROWTH,LEAF ANATOMY AND PHOTOSYNTHESIS OF DISTICHLIS SPICATA (L.) GREENE

The effects of light, temperature, and salinity on growth, net CO₂ exchange and leaf anatomy of Distichlis spicata were investigated in controlled environment chambers. When plants were grown at low light, growth rates were significantly reduced by high substrate salinity or low temperature. However, when plants were grown at high light, growth rates were not significantly affected by temperature or salinity. The capacity for high light to overcome depressed growth at high salinity cannot be explained completely by rates of net photosynthesis, since high salinity caused decreases in net photosynthesis at all environmental conditions. This salinity-induced decrease in net photosynthesis was caused largely by stomatal closure, although plants grown at low temperature and low light showed significant increases in internal leaf resistance to CO₂ exchange. Increased salinity resulted in generally thicker leaves with lower stomatal density but no significant differences in the ratio of mesophyll cell surface area to leaf area. Salinity and light during growth did not significantly affect rates of dark respiration. The mechanisms by which Distichlis spicata tolerates salt appear to be closely coulpled to the utilization of light energy. Salt-induced leaf succulence is of questionable importance to gas exchange at high salinity in this C₄ species.     

Root-to-shoot signalling when soil moisture is heterogeneous: increasing the proportion of root biomass in drying soil inhibits leaf growth and increases leaf abscisic acid concentration

To determine whether root-to-shoot signalling of soil moisture heterogeneity depended on root distribution, wild-type (WT) and abscisic acid (ABA)-deficient (Az34) barley (Hordeum vulgare) plants were grown in split pots into which different numbers of seminal roots were inserted. After establishment, all plants received the same irrigation volumes, with one pot watered (w) and the other allowed to dry the soil (d), imposing three treatments (1 d: 3 w, 2 d: 2 w, 3 d: 1 w) that differed in the number of seminal roots exposed to drying soil. Root distribution did not affect leaf water relations and had no sustained effect on plant evapotranspiration (ET). In both genotypes, leaf elongation was less and leaf ABA concentrations were higher in plants with more roots in drying soil, with leaf ABA concentrations and water potentials 30% and 0.2 MPa higher, respectively, in WT plants. Whole-pot soil drying increased xylem ABA concentrations, but maximum values obtained when leaf growth had virtually ceased (100 nm in Az34, 330 nm in WT) had minimal effects (<40% leaf growth inhibition) when xylem supplied to detached shoots. Although ABA may not regulate leaf growth in vivo, genetic variation in foliar ABA concentration in the field may indicate different root distributions between upper (drier) and lower (wetter) soil layers.     

Light-Dependent Short-Term Modulations of Elongation in Rice Plants

The elongation of rice shoots (Oryza saliva L.) exposed to variouslight conditions was measured using angular movement transducers.Plants growing in continuous light responded to a sudden increasein light fluence rate with a pronounced decrease in the rateof elongation. The growth rate returned to the initial steadystate within 15-60 min, depending on the difference in fluencerate between the continuous and the supplemental light. Whenthe fluence rate was reduced to its original level, the elongationrate rose transiently before returning to the initial value.The inhibition of elongation was fluence dependent, and blueand red light were more effective in reducing the elongationrate than was green or infrared irradiation. Irradiation ofa single tiller affected also other tillers of the same plantwhich were shielded from the light. Measurements of transpirationand water uptake by whole plants and of leaf temperatures indicatedthat the transient changes in growth rates were caused by perturbationsin the water balance of the plant. (Received December 5, 1988; Accepted February 7, 1989)     

Diurnal Changes in Maize Leaf Photosynthesis : III. Leaf Elongation Rate in Relation to Carbohydrates and Activities of Sucrose Metabolizing Enzymes in Elongating Leaf Tissue

Maize (Zea mays L. cv. Pioneer 3184) leaf elongation rate was measured diurnally and was related to diurnal changes in the activities of sucrose metabolizing enzymes and carbohydrate content in the elongating portion of the leaf. The rate of leaf elongation was greatest at midday (1300 hours) and was coincident with the maximum assimilate export rate from the distal portion of the leaf. Leaf elongation during the light period accounted for 70% of the total observed increase in leaf length per 24 hour period. Pronounced diurnal fluctuations were observed in the activities of acid and neutral invertase and sucrose phosphate synthase. Maximum activities of sucrose phosphate synthase and acid invertase were observed at 0900 hours, after which activity declined rapidly. The activity of sucrose phosphate synthase was substantially lower than that observed in maize leaf source tissue. Neutral invertase activity was greatest at midday (1200 hours) and was correlated positively with diurnal changes in leaf elongation rate. There was no significant change in the activity of sucrose synthase over the light/dark cycle. Sucrose accumulation rate increased during a period when leaf elongation rate was maximal and beginning to decline. Maximum sucrose concentration was observed at 1500 hours, when the activities of sucrose metabolizing enzymes were low. At no time was there a significant accumulation of hexose sugars. The rate of starch accumulation increased after the maximum sucrose concentration was observed, continuing until the end of the light period. There was no delay in the onset of starch mobilization at the beginning of the dark period, and essentially all of the starch was depleted by the end of the night. Mobilization of starch in the elongating tissue at night could account for a significant proportion of the calculated increase in the tissue dry weight due to growth. Collectively, the results suggested that leaf growth may be controlled by the activities of certain sucrose metabolizing enzymes and may be coordinated with assimilate export from the distal, source portion of the leaf. Results are discussed with reference to diurnal photoassimilation and export in the distal, source portion of the leaf.     

Effects of water stress on gas exchange and water relations of a succulent epiphyte,Kalanchoë uniflora

Summary Measurements of gas exchange, xylem tension and nocturnal malate synthesis were conducted with well-watered and droughted plants of Kalanchoë uniflora. Corresponding results were obtained with plants grown in 9 h and 12 h photoperiods. In well-watered plants, 50 to 90% of total CO₂-uptake occurred during the light period. Nocturnal CO₂-uptake and malate synthesis were higher and respiration rate was lower in old leaves (leaf pairs 6 to 10) compared to young leaves (leaf pairs 1 to 5). Within four days of drought distinct physiological changes occurred. Gas exchange during the light period decreased and CO₂-uptake during the dark period increased. Nocturnal malate synthesis significantly increased in young leaves.Respiration rate decreased during periods of drought, this decrease being more pronounced in young leaves compared to old leaves. Restriction of gas exchange during the light period resulted in a decrease of transpiration ratio from more than 100 to about 20. The difference between osmotic pressure and xylem tension decreased in young leaves, indicating a reduction in bulk leaf turgor-pressure.We conclude that both the CAM-enhancement in young leaves and the decrease of respiration rate are responsible for the increase of nocturnal CO₂-uptake during water stress. During short drought periods, which frequently occur in humid habitats, the observed physiological changes result in a marked reduction of water loss while net CO₂-uptake is maintained. This might be relevant for plant growth in the natural habitat.Abbreviations LP light period - DP dark period - CAM crassulacean acid metabolism     

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.