Osmotic Adjustment and Stomatal Response to Water Deficits in Maize

A pot experiment was carried out using five maize {Zea maysL.) cultivars under three soil moisture levels (MPa 0 to –0.05,–0.3 to –0.9 and –1.2 to –1.5) to investigatethe effects of water deficits on osmotic adjustment and stomatalconductance. The degree of leaf rolling and the sugar and nutrientconcentrations in leaf cell sap were measured. Leaf water potential and osmotic potential decreased and stomatalconductance decreased with increasing water deficits. Stomatalconductance correlated positively with leaf water potentialand osmotic potential. Degree of leaf rolling was lower in cultivarswhich maintained higher turgor. Osmotic adjustment of 0.08 to0.43 MPa was found under the lowest soil moisture level in fivecultivars used. Sugar and K were the major osmotic substancesin the maize plant. Sugar, K and Mg concentrations increasedunder water deficit, and correlated negatively with a decreasein osmotic potential. Key words: Zea mays L., leaf water relations, leaf rolling, osmotic adjustment, stomatal conductance, water deficit     

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

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

Somatic Embryogenesis and Plant Regeneration from Cultured Leaf Explants of Zea mays

Young leaf segments of Zea mays L. seedlings were cultured onMurashige and Skoog's basal nutrient medium supplemented with2 mg l^–1 2, 4-D and sub-cultured on medium containing8 mg l^–1 2,4-D. Two types of callus tissues appeared—embryogenicand non-embryogenic. The embryogenic callus tissue producednumerous somatic embryos which on transfer to media containinglow amounts of 2,4-D or ABA produced plantlets. Callus tissuesexhibited embryogenic potential for more than 1 year. Zea mays L. cv. Ageti-76, Zea mays L. cv. N-L-D-Comp., maize, leaf, callus, somatic embryogenesis, regeneration     

Kinetics of Maize Leaf Elongation: II. RESPONSES OF A NA-EXCLUDING CULTIVAR AND A NA-INCLUDING CULTIVAR TO VARYING NA/CA SALINITIES

Salinity causes physiological and morphological changes in plantsand calcium can mitigate many of these effects. In this study,the effects of salinity (75 mol m^–3 NaCl) with or withoutsupplemental Ca (10 mol m^–3) on the kinetics of maize(Zea mays L.) leaf elongation were examined using Linear VariableDifferential Transformers (LVDTs). Short-term growth responsesof two cultivars (Dekalb hybrid XL75 and Pioneer hybrid 3906)differing in salt tolerance were compared. Salinity caused animmediate reduction in the leaf elongation rate (LER). Within2 h, elongation rates had increased and reached new steady rates.Significant differences between salinity treatments with highand low Ca could be detected within the first 2 h after impositionof salinity for Dekalb hybrid XL75, but not for Pioneer hybrid3906. After 24 h, distinct differences for both cultivars weredetected. Dekalb hybrid XL75, a Na-includer, was more salt-sensitiveand responsive to supplemental Ca (10 mol m^–3) than Pioneerhybrid 3906, a Na-excluder. Turgor was not reduced 24 h aftersalinization because there was complete osmotic adjustment inthe elongation zone of the leaves. Analysis of the growth parameterslimiting LER indicated that the yield threshold (Y) was increasedfor salt-stressed plants. In addition, both the cell wall extensibilityand hydraulic conductance were reduced 24 h after salinization.Supplemental Ca increased LER of salt-stressed plants by increasinghydraulic conductance. The differences in LER of the two cultivarsunder saline conditions was attributed to differences in theincrease of Y caused by salinity. Key words: Calcium, growth, salinity, sodium, Zea mays L.     

Effect of the Osmotic Environment on Assimilate Transport in Isolated Developing Embryos of Maize (Zea mays)

Embryos obtained from developing kernels of maize (Zea maysL.) were incubated in a bathing medium, to measure the effectof the osmotic environment on the balance between uptake andrelease of assimilates by the embryo. Net efflux of sucroseand amino acids from the embryo decreased with increasing mannitolconcentration in the bathing medium and net uptake of [¹⁴C]valine increased with increasing mannitol concentration. Therole of a high osmolality of the seed apoplast in seed developmentis discussed Zea mays, maize, embryo, seed development, assimilate transport, turgor-sensitive transport     

Root and Shoot Growth of Plants Treated with Abscisic Acid

Young seedlings of Capsicum annum L., Commelina communis L.and maize (Zea mays L.) were subjected to a mild water-stressingtreatment and/or treated with abscisic acid (ABA). Plants rootedin soil received a soil-drying treatment and their leaves weresprayed with a 10–⁴ M solution of ABA. Plants grown insolution culture were stressed by the addition of polyethyleneglycol (PEG) to the rooting medium and ABA was also added tothe rooting medium, either with or without PEG. The effectsof both treatments on the growth of roots and shoots and theultimate root: shoot dry weight ratio were very similar. Shootgrowth was limited both by water stress and by ABA application;while there was some evidence that mild water stress and/orABA application may have resulted in a stimulation of root growth.More severe water stress reduced the growth of roots but theoverall effect of stress was to increase the ratio of rootsto shoots. Capsicum annum L., Commelina communis L., Zea mays L., water stress, abscisic acid     

The spontaneous growth response of maize coleoptile segments and the change in tissue 8

Decapitated segments from maize (Zea mays L.) coleoptiles orientedvertically in an upright position show a strong spontaneousgrowth response (SGR) 3 h after decapitation. The latent periodof the SGR is markedly reduced when these segments are orientedin an inverted position. Coleoptile segments with intact tipsexhibit a weak and transient SGR in the vertical upright orientation.However, in the inverted orientation, these segments show atypical SGR. The data are inconsistent with the current hypothesisthat the SGR is caused by a time-dependent increase in tissuesensitivity to auxin. The parallel increase in membrane potentialdifference and growth rate during the time-course of the SGRindicates a possible role for PM H⁺-ATPase in the establishmentof the SGR in maize coleoptile segments. Key words: Auxin, spontaneous growth response, membrane potential, plasma membrane H⁺-ATPase, Zea mays L.     

Water stress effects on leaf elongation,leaf water potential,transpiration, and nutrient uptake of rice,maize, and soybean

A pot experiment was conducted in the greenhouse to determine and compare the responses of rice (Oryza sativa L. var, IR 36), maize (Zea mays L. var. DMR-2), and soybean (Glycine max [L.] Merr. var. Clark 63) to soil water stress. Leaf elongation, dawn leaf water potential, transpiration rate, and nutrient uptake in stressed rice declined earlier than in maize and soybean. Maize and soybean, compared with rice, maintained high dawn leaf water potential for a longer period of water stress before leaf water potential. Nutrient uptake under water stress conditions was influenced more by the capacity of the roots to absorb nutrients than by transpiration. Transport of nutrients to the shoots may occur even at reduced transpiration rate It is concluded that the ability of maize and soybean to grow better than rice under water stress conditions may be due to their ability to maintain turgor as a result of the slow decline in leaf water potential brought about by low, transpiration rate and continued uptake of nutrient, especially K, which must have allowed osmotic adjustment to occur.     

Kinetics of Maize Leaf Elongation: I. INCREASED YIELD THRESHOLD LIMITS SHORT-TERM, STEADY-STATE ELONGATION RATES AFTER EXPOSURE TO SALINITY

The short-term responses of leaf elongation to salinity areinvestigated in this study. The kinetics of maize (Zea maysL.) leaf elongation were measured with Linear Variable DifferentialTransformers (LVDTs). After exposure to salinity (0 to 120 molm^–3 NaCl), leaf elongation rates (LER) declined rapidly.Within 4 h, LER had recovered and reached a new steady-statefor all salinity treatments. These rates were reduced by 10,20, and 60% of control rates by 40, 80 and 120 mol m^–3NaCl, respectively. Osmotic adjustment in the growing zone ofleaves was correlated with the recovery of LER after plant exposureto salinity. However, after 4 h of exposure, the osmolalityof the cell sap continued to increase without effect on steady-stateLER. Estimates of the apparent turgor in the growing zone indicatedthat turgor was no longer limiting LER of salt-stressed plantsafter 4 h. An in vivo technique was developed to apply a unidirectionalforce to intact growing leaves of maize to mimic increases inelongation force. Relative elongation rate (RER) were increasedby adding weights to the LVDT core to increase elongation force.Plots of RER as a function of elongation force gave estimatesof two growth coefficients: the yield threshold and the yieldingcoefficient, mL/(m + L), where m is the cell wall extensibilityand L is the hydraulic conductivity. RER as a function of elongationforce was determined immediately, 05, 4, and 21 h after plantswere salinized. Estimates of the growth coefficients indicatedthat the apparent yield threshold decreased immediately aftersalinization. However, when LER reached steady-state, the yieldthreshold of salt-stressed plants had increased above controlvalues and was the only limiting growth coefficient. There wereno significant effects of salinity on the yielding coefficients,cell wall extensibility or hydraulic conductivity. One of theadvantages of this in vivo technique over other methods is thatyield threshold, yielding coefficient, and cell wall extensibilitycan be determined without the confounding effects of woundingor osmotic stress. This technique may prove widely applicableto the study of other growth regulating factors. Key words: Salinity, leaf growth, Zea mays L     

Effects of Increasing Water Stress on Simulated Swards of Festuca arundinacea Schreb under Wind Tunnel Conditions

The effects of increasing water stress on water relations, leafconductance, leaf extension and leaf rolling of Festuca arundinaceain sward (I m²) were investigated under wind tunnel conditions.The plants were grown in a container 60 cm deep and the experimentwas conducted over a 36 d period. Upon cessation of watering(day 11), leaf extension and conductance were affected. Within8 d, the onset of leaf rolling helped to reduce transpirationand to maintain leaf water potential. Nocturnal recovery of turgor potential helped in maintainingleaf extension at a moderate level and in the final 5 d waterand osmotic potentials dropped sharply as leaf rolling becamemore acute and leaf extension stopped. The grass combines various morphological and physiological mechanismsto prevent water losses and maintain growth. Festuca arundinacea, tall fescue, wind tunnel, water stress, water potential, osmotic potential, conductance, leaf rolling, leaf extension     

Osmotic Stress as a Pregrowth Procedure for Cryopreservation 2. Water Relations and Metabolic State of Sycamore and Soybean Cell Suspensions

Addition of 6 per cent mannitol or sorbitol to liquid culturemedium decreased the water potential (_w) by –0.93 MPa( 382 ± 7 mOsm kg^–1 water). Sycamore cells grownto exponential phase in such media exhibited increased levelsof total and soluble protein and respiratory activity, but decreasedamounts of free proline. Soybean cells showed increased respiratoryactivity and free proline levels, but total protein levels remainedunaffected. Soluble protein levels were reduced under sorbitol-inducedstress. In both species osmotic stress had little effect oncell dry weight. Water relation studies indicate that sycamore cells are capableof much greater osmotic adjustment than soybean cells, and thatmannitol uptake does not contribute significantly to that adjustment. Acer pseudoplatanus L., sycamore, Glycine max L. var. Biloxi, soybean, suspension culture cells, osmotic stress, water relations, metabolism     

Timing of Kernel Development in Water-stressed Maize: Water Potentials and Abscisic Acid Concentrations

Maize (Zea mays L. cv. Pioneer 3925) subjected to post-anthesiswater stress during the first 2 weeks of kernel developmenthad lower leaf-water potentials and higher leaf-ABA concentrationsthan well-watered controls. There was a concomitant rise inABA concentration in kernel tissues 3 and 7 d after pollination(DAP), after which the concentration decreased to control levelsby 13 DAP. Kernel water potential, however, remained unchangedby the water stress. Radiolabelled ABA, fed to a leaf, was translocatedto kernels, where free ABA as well as several ABA metaboliteswere the major labelled fractions. This suggested that the stress-inducedkernel ABA was of maternal origin. Since ABA plays a putativerole in seed maturation of several crop species, and appliedABA or water stress often hastens seed development, we expectedthat a water-stress-induced rise in kernel ABA concentrationearly in grain development may serve to prematurely induce storage-productaccumulation. Zein, starch and several enzymes key to the starchsynthesis pathway followed the same course of induction throughoutthe experiment, with no difference between treatments Henceit was concluded that although water stress increased kernelABA independent of kernel water status, there was no apparenteffect of water stress or ABA on timing of early kernel developmentalprocesses. Zea mays L. cv. Pioncer 3925, maize, water stress, abscisic acid, endosperm development     

Cell Membrane Stability and Leaf Surface Wax Content as Affected by Increasing Water Deficits in Maize

A field experiment was conducted with a water-stressed treatmentand well-watered control using eight maize (Zea mays L.) cultivars.Effects of water deficits on cell membrane stability (CMS) measuredby the polyethylene glycol (PEG) test, leaf surface wax content,and relative growth rate were investigated. Cytoplasmic lipidcontent was also analysed. Cell membrane stability and leaf surface wax content increasedwith the degrees of stress. Tolerance to drought evaluated asincrease in CMS under water deficit conditions was well differentiatedbetween cultivars and was well correlated with a reduction inrelative growth rate under stress. A negative correlation wasfound between percentage injury in the PEG test and leaf surfacewax content. High phospholipid contents were observed in tissuesof drought tolerant cultivars under water deficit conditions. Key words: Cell membrane stability, cytoplasmic lipid, drought tolerance, leaf surface wax, relative growth rate     

The Comparative Drought Resistance of Landraces of Sorghum and Millet From Dry and Humid Regions

It may be that land-races of sorghum (Sorghum sp.) and millet[Pennisetum americanum (L.) Leeke] which evolved along geographicalgradients of rainfall in Africa and India, differ in their droughtresistance. Any physiological attributes found to be correlatedwith low rainfall might be important and effective characteristicsfor crop production in dry regions. Twenty land-races were chosen which evolved along geographicalgradients of rainfall, seven millets from India, six sorghumsfrom Mali, and seven sorghums from the Sudan. Races were evaluatedfor their growth potential and plant water relations under hydroponicsconditions in a growth chamber. A water stress treatment wasimposed by adding polyethylene glycol-8000 to the nutrient solution,giving a solute water potential of -0.5 MPa, compared with acontrol solution at 003 MPa. Drought resistance, in terms of relatively less growth inhibitionunder stress, was higher in races from dry regions than in racesfrom humid regions. Of all the physiological variables measured[carbon exchange rate, (CER), transpiration, transpiration ratio(CER/transpiration), leaf diffusive resistance, leaf water potentialand osmotic adjustment], only osmotic adjustment under stresswas generally correlated with average rainfall at each race'sorigin, indicating greater osmotic adjustment in land-racesfrom drier regions. Races with a greater capacity for osmoticadjustment were characterized by smaller plants with high ratesof transpiration and low rates of leaf senescence under stress. The carbon exchange rate per unit leaf area increased as liveleaf area decreased under stress due to leaf senescence. Thus,drought resistant races under stress tended to have lower CERper unit live leaf area (but not per plant) than susceptibleraces. Transpiration ratios under stress were lower in resistantthan in susceptible races, mainly because resistant races hadhigher transpiration. The results for the measured variables showed a general trendfor greater drought resistance in sorghum than in millet, indicatingthat the commonly observed adapation of the millets to dry environmentsmay be due to other factors, such as drought escape or heattolerance. Sorghum sp. Pennisetum americanum L. (Leeke), water stress, osmotic adjustment, photosynthesis, transpiration, evolution, drought resistance     

Ionic and Water Relations Responses of Two Populations of a Non-Halophyte to Salinity

Bowman, W. D. 1988. Ionic and water relations responses of twopopulations of a non-halophyte to salinity.–J. exp. Bot39: 97–105 Salinity-induced changes in the ionic and water relations inplants from two naturally-occurring populations of the C₄ non-halophyteAndropogon glomeratus were measured to detect differences inthe capacity to adjust osmotic potentials and in ion contentpotentially responsible for the osmotic adjustment Pressure-volumecurves and leaf ion content were measured in plants from twopopulations, salt marsh and inland, after long-term exposureto three salinity levels. Osmotic adjustment and decreases inthe bulk tissue elasticity occurred to a similar extent in bothpopulations with increasing salinity. Cl^– concentrationsincreased with increasing salinity in both populations, whereasleaf Na⁺ concentrations increased only in the inland population,but were higher at all salinities in the marsh population. K⁺concentrations changed little with increasing salinity. Prolineconcentrations increased only at the highest salinity level,and did not difler significantly between populations. Theseresults suggest a role for Na⁺ uptake and regulation in osmoticadjustment in the marsh population, contrasting with studiesof salt tolerance in other nonhalophytic grasses     

Salt Tolerance in the Triticeae: Growth and Solute Accumulation in Leaves of Thinopyrum bessarabicum

Gorham, J., McDonnell, E., Budrewicz, E. and Wyn Jones, R. G.1985. Salt tolerance in the Triticeae: growth and solute accumulationin leaves of Thinopyrum bessarabicum.—J. exp. Bot. 36:1021–1031. The diploid wheatgrass Thinopyrum bessarabicum was found towithstand prolonged exposure to 350 mol m^–3 NaCl in hydroponicculture. During the gradual addition of salt to the externalmedium, osmotic adjustment was rapidly achieved by the accumulationof Na and Cl. Following osmotic adjustment constant leaf Naand Cl concentrations were maintained, and K was retained ata high level. Thinopyrum bessarabicum may be described as anosmoconformer, adjusting its internal osmotic pressure to 400–500mOsmol kg^–1 above that of the external medium in hydroponicculture. Both slower shoot initiation and reduced leaf lengthcontributed to the reduced growth rates at higher salinities.Leaf width was not affected. Increasing salinity resulted inincreases in leaf concentrations of phosphate, glycinebetaine,sucrose and proline, and in decreases in the concentrationsof nitrate, sulphate, magnesium, calcium, total amino acidsand organic acids. Thinopyrum bessarabicum exhibits salt tolerancecharacters which may be useful in wheat breeding. Key words: Salt stress, solute accumulation, osmotic adjustment, Thinopyrum     

Plant Growth Analysis: Growth and Yield Component Responses to Population Density in Forage Maize

Forage maize (Zea mays L.) was grown in monocultures at populationdensities ranging from 4·9 to 11·1 plants m^–2.Data for plant growth analysis were obtained from six harvestscarried out from 21 to 115 d after planting. Conventional plantgrowth analysis indicated that improvements in forage productivityper unit land area by high population density resulted directlyfrom increased plant presence. Reduction in dry weight per shootat high population density was associated with reduced unitleaf rate. Leaf area ratio was little affected, which may implythat competition for soil nutrients or oxygen was the chiefcause of plant interference. Yield component analysis demonstratedthe increasing importance of population density treatments asa source of variation as growth progressed. Direct relationshipsbetween variation in yield per plant and variation in two yieldcomponents, stem diameter and the inverse of leaf area ratio,were demonstrated. Both conventional plant growth analysis andyield component analysis indicated complex physiological andmorphological adjustments to species population density. Plant growth analysis, yield component analysis, Zea mays L     

Influence of arbuscular mycorrhiza on lipid peroxidation and antioxidant enzyme activity of maize plants under temperature stress

The influence of the arbuscular mycorrhizal (AM) fungus, Glomus etunicatum, on characteristics of growth, membrane lipid peroxidation, osmotic adjustment, and activity of antioxidant enzymes in leaves and roots of maize (Zea mays L.) plants was studied in pot culture under temperature stress. The maize plants were placed in a sand and soil mixture under normal temperature for 6 weeks and then exposed to five different temperature treatments (5oC, 15oC, 25oC, 35oC, and 40oC) for 1 week. AM symbiosis decreased membrane relative permeability and malondialdehyde content in leaves and roots. The contents of soluble sugar content and proline in roots were higher, but leaf proline content was lower in mycorrhizal than nonmycorrhizal plants. AM colonization increased the activities of superoxide dismutase, catalase, and peroxidase in leaves and roots. The results indicate that the AM fungus is capable of alleviating the damage caused by temperature stress on maize plants by reducing membrane lipid peroxidation and membrane permeability and increasing the accumulation of osmotic adjustment compounds and antioxidant enzyme activity. Consequently, arbuscular mycorrhiza formation highly enhanced the extreme temperature tolerance of maize plant, which increased host biomass and promoted plant growth.     

Turgor Differences and Water Stress in Maize and Sorghum Leaves During Drought and Recovery

The pressure potentials (turgor pressure) in leaves of maize(Zea mays L.) and grain sorghum (Sorghum vulgare Pers.) plantssubjected to water stress in a controlled environment were estimatedfrom measurements of water and osmotic potentials. Changes inturgor pressure were larger in sorghum than in maize duringthe development of water stress and after re-watering. It issuggested that this indicates a lower cell wall elasticity insorghum than in maize. This fact may affect some of the physiologicalactivities of sorghum     

Phosphate Absorption, Fluxes, and Symplasmic Transport in Osmotically-Shocked Zea mays Roots

Osmotic shock with sequential 30 min treatments in ice-coldsaline solutions and distilled water inhibited both the subsequentuptake of orthophosphate (Pi) and its transport into the xylemof excised corn (Zea mays L.) roots. Measurements of Pi fluxeswith ³²P indicated that the decrease in net Pi uptake over a24 h period caused by osmotic shock was due primarily to delayedrecovery of Pi influx rather than to increasing efflux. Despitecomplete recovery of Pi absorption within 2–6 h aftershocking with 150–200 mM NaCl, transport to the xylemduring the subsequent 24 h only partially recovered. Leucineuptake and incorporation into protein was also markedly inhibitedby osmotic shock but both almost completely resumed controlrates within 24 h after shocking with up to 150 mM NaCl. Tetracyclineinhibited recovery of Pi uptake after NaCl treatment whereaspuromycin did not. These results with corn roots are consistentwith the hypothesis that recovery of Pi uptake activity aftermoderate osmotic shock requires de novo synthesis of membraneproteins. Incomplete recovery of Pi transport to the xylem suggeststhat osmotic shock may damage plasmodesmata. Key words: Corn, Ion uptake, Leucine uptake, NaCl, Puromycin, Tetracycline