Water Relations of White Clover (Trifolium repens): Water Potential Gradients and Plant Morphology

Water potential, osmotic potential, pressure potential and relativewater content were measured in stolons and leaves of white cloverplants grown under a range of conditions of water supply andevaporative demand. The importance of adventitious roots fromthe nodes was examined. Gradients along stolons were alwaysextremely small, of the order of only 01 MPa. Stolon up waterpotential was representative of plant water status regardlessof stolon length, presence/absence of nodal roots, degree ofwater stress and evaporative demand. It is concluded that waterconduction along stolons was very good. Gradients were foundto exist along petioles; they may have a greater resistanceto water flow than stolons. The relationship between water fluxand stem anatomy, and the importance of differential flow ratesthrough stolons and petioles to plant behaviour during waterstress, are discussed. Trifolium repens L., white clover, water relations     

The Growth and Carbon Allocation Patterns of White Clover (Trifolium repens L.) Plants of Contrasting Branching Structure

The growth, morphology and carbon allocation patterns of F1progeny white clover (Trifolium repens L.) plants selected foreither low (‘LBF’) or high (‘HBF’) frequencyof stolon branching were compared in two controlled-environmentexperiments. Selections from within both a small-leaved (‘GrasslandsTahora’) and a large-leaved (‘Grasslands Kopu’)clover cultivar were compared, and plants were grown under arelatively lenient defoliation treatment (expt 1) or under threelevels of defoliation seventy (expt 2). Carbon allocation patternswere measured by ¹⁴CO₂ pulse-chase labelling using fully unfoldedleaves on the main (parent) stolon. LBF and HBF displayed consistent differences in the selectedcharacter though, within cultivars, the difference between selectionswas most pronounced for Kopu. The selections developed fundamentallydifferent branching structures resulting from differences inbranching frequency, with total branch weight per plant averaging122 mg for LBF and 399 mg for HBF (mean of both experiments).More C moved from parent stolon leaves to branches in HBF thanin LBF (mean 22.6% vs. 15.1% respectively of the ¹⁴C exportedfrom source leaves). More C also moved to stolon tissue in HBF,but, counterbalancing this and the difference in allocationto branches, less moved to developing leaves and roots on theparent stolon itself compared to LBF. However, the total weightof developing leaves and roots per parent stolon was generallygreater in HBF than in LBF, probably reflecting greater C importby these sinks from the higher number of branches present perplant in the former selection. HBF plants were consistentlylarger at harvest than LBF plants. There were no defoliationtreatment x selection interactions in C allocation patternsin expt 2. The implications of the results for plant performancein grazed pastures are discussed. Branching, carbon translocation, defoliation, growth, morphology, Trifolium repens, white clover     

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     

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     

Implication of ABA and proline on cell membrane injury of water deficit stressed barley seedlings

The aim of this work was to examine the ability of ABA and proline to counteract the deleterious effect of water deficit stress on cell membrane injuries. Six-day-old seedlings of two barley genotypes (cv. Aramir, line R567) were treated with ABA (2·10⁻⁴ M) or proline (0.1 M) for 24 h, and then subjected to osmotic stress for 24h, by immersing their roots in polyethylene glycol (PEG 6000) solution of osmotic potential of −1.0 MPa and −1.5 MPa or by submerging the leaf pieces in PEG solution of osmotic potential of −1.6 MPa. Pretreatment of plants with ABA and proline caused an increase of free proline level in the leaves. Plants treated with ABA exhibited a lower membrane injury index under water stress conditions than those untreated even when no effect of this hormone on RWC in the leaves of stressed plants was observed. Pretreatment of plants with proline prevented to some extent membrane damage in leaves of the stressed seedlings, but only in the case when stress was imposed to roots. Improvement in water status of leaves was also observed in seedlings pretreatment with proline. The protective effect of both ABA and proline was more pronounced in line R567 that exhibited higher membrane injury under water deficit stress conditions.     

An inland and a coastal population of the Mediterranean xero-halophyte species Atriplex halimus L. differ in their ability to accumulate proline and glycinebetaine in response to salinity and water stress

Soil salinity and drought compromise water uptake and lead toosmotic adjustment in xero-halophyte plant species. These importantenvironmental constraints may also have specific effects onplant physiology. Stress-induced accumulation of osmocompatiblesolutes was analysed in two Tunisian populations of the Mediteraneanshrub Atriplex halimus L.—plants originating from a salt-affectedcoastal site (Monastir) or from a non-saline semi-arid area(Sbikha)—were exposed to nutrient solution containingeither low (40 mM) or high (160 mM) doses of NaCl or 15% polyethyleneglycol. The low NaCl dose stimulated plant growth in both populations.Plants from Monastir were more resistant to high salinity andexhibited a greater ability to produce glycinebetaine in responseto salt stress. Conversely, plants from Sbikha were more resistantto water stress and displayed a higher rate of proline accumulation.Proline accumulated as early as 24 h after stress impositionand such accumulation was reversible. By contrast, glycinebetaineconcentration culminated after 10 d of stress and did not decreaseafter the stress relief. The highest salt resistance of Monastirplants was not due to a lower rate of Na⁺ absorption; plantsfrom this population exhibited a higher stomatal conductanceand a prodigal water-use strategy leading to lower water-useefficiency than plants from Sbikha. Exogenous application ofproline (1 mM) improved the level of drought resistance in Monastirplants through a decrease in oxidative stress quantified bythe malondialdehyde concentration, while the exogenous applicationof glycinebetaine improved the salinity resistance of Sbikhaplants through a positive effect on photosystem II efficiency. Key words: Atriplex halimus, glycinebetaine, halophyte, NaCl, osmotic adjustment, proline, salinity, water stress     

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     

Role of proline and leaf expansion rate in the recovery of stressed white clover leaves with increased phosphorus concentration

Osmotic adjustment (OA) and increased cell-wall extensibility required for expansive leaf growth are well defined components of adaptation to water stress in dry soil, which might interact with soil phosphorus (P) concentration and defoliation frequency for intensively grazed white clover in legume-based pastures. Experiments were conducted with frequently and infrequently defoliated mini-swards of white clover growing in dry soil with low and high P concentrations. The higher yielding high-P plants were able to dry the soil to greater soil water suctions; their leaves had lower water potential values, yet they showed fewer water stress symptoms and underwent a more complete recovery from the water stress symptoms on rewatering, than the low-P plants. High- P plants had greater OA, proline concentration and leaf expansion rate. On the other hand, low-P plants showed an increased osmotic concentration when there was no change in the total solute content per unit of leaf d. wt, indicating more loss of water from the leaf tissue. The key measures that appeared to be directly associated with plant recovery over a short period following water stress were increased proline concentration and leaf expansion rate, probably resulting from increased cell-wall extensibility rather than increased production of cells for the high-P plants.     

Water Content-Potential Relationship in Soya Bean: Changes in Component Potentials for Mature and Immature Leaves under Field Conditions

Changes in turgor and osmotic potentials of soya bean leaves(Glycine max.) with changes in water content were measured throughouta season using the pressure-volume technique. Two distinct reponsesto water loss were found. When water was expressed from leavesin the pressure chamber their osmotic behavior was describedby a concentration effect based on the osmotic volume. The osmoticfraction of the total water content averaged 0·72 and0·84 for mature and immature leaves, respectively. Thechanges in turgor pressure in the chamber were described bya volumetric modulus of elasticity which increased linearlywith turgor pressure. The changes in total potential at highturgor pressures were almost exclusively due to changes in turgordue to the high modulus (high tissue rigidity) in that range.Responses were different, however, for leaves drying in thefield. For these, the osmotic changes were always large anddominated by solute adjustment. Diurnal changes in osmotic potentialwere as much as 5 bars (500 kPa), or around 50 per cent, andwere about the same magnitude as the changes in turgor pressurefor both mature and immature leaves. The elastic modulus atthe time of sampling showed the normal turgor dependence forimmature leaves but for mature leaves the initial modulus wasapparently constant at about 180 bars. The different behaviourin the pressure bomb and the field is interpreted in terms ofa rate dependence for turgor and osmotic response to water loss.     

Comparison of Osmotic Adjustment Responses to Water and Temperature Stresses in Spring Wheat and Sudangrass

This study explores the mechanisms of osmotic adjustment bycomparing the growth of spring wheat and sudangrass, which exhibitdifferent degrees of osmotic adjustment, under soil water andtemperature stresses. Leaf water potential ( ₁), osmotic potential(), and rate of leaf area growth of spring wheat and sudangrassseedlings were measured at combinations of five soil water potentials,from -0·03 to -0·25 MPa, and six root temperatures,from 14 to 36°C. Spring wheat exhibit little osmotic adjustment.The leaf osmotic potential was not affected by either soil wateror root temperature stress. Osmotic potential of sudangrassdecreased in parallel with the decreasing leaf water potentialas a result of osmotic adjustment. As soil water potential decreasedfrom -0·03 to -0·25 MPa, the rates of growth andphotosynthesis of spring wheat both decreased by about 30%.For sudangrass with the same range of soil water potential,the photosynthesis rate decreased by only 10% while the leafarea growth rate decreased by 49%. We introduce a dimensionlessindex (R) to quantify the degree to which environmental stressesalter the balance between production of photosynthates and theiruse for growth. The index, R, is equal to 1 when stress reducesgrowth and photosynthesis by the same degree, i.e. the balancebetween production and consumption of photosynthate is not disturbed.R is smaller than 1 when growth is reduced more than photosynthesis.R was equal to 1 for spring wheat where there was no osmoticadjustment. For sudangrass, R decreased from 1 to 0·25as osmotic potential decreased from -1·10 to -1·63MPa. These findings lead to the hypothesis that osmotic adjustmentcould result from an imbalance between production, consumptionand translocation of photosynthates under stressed conditions.Copyright1993, 1999 Academic Press Osmotic adjustment, water stress, root temperature     

Expression of dehydrins under heat stress and their relationship with water relations of sugarcane leaves

The heat stress-induced dehydrin proteins (DHNs) expression and their relationship with the water relations of sugarcane (Saccharum officinarum L.) leaves were studied. Sugarcane seedlings were subjected to heat stress (day/night temperature of 40/35 °C) under relative humidity 60/65 % to avoid aerial desiccation and determinations made at 4, 12, 24, 36, 48, 60 and 72 h. The leaves showed a sharp decline in the water and osmotic potentials, and relative water content during first 12 h of heat stress, but a regain in their values in 24 h. The pressure potential (ψ_p) decreased initially but increased later and approached control leaves. The increase in ψ_p was tightly correlated to the accumulation of free proline, glycinebetaine and soluble sugars, indicating their possible involvement in the osmotic adjustment under heat stress. Immunological detection revealed the expression of three DHNs with an apparent molecular mass of 21, 23 and 27 kDa under heat stress (48 to 72 h) and their expression was independent of the changes in the water relations of leaves.     

Rapid Osmotic Adjustment in Detached Wheat Leaves

Osmotic adjustment is induced in detached wheat leaves by rapiddrying to a relative water content below 0·65, followedby re-saturation. Quantitatively, the response to this treatmentis comparable to the maximum of adjustment obtained with pottedplants at the same developmental stage. Low temperatures duringdrying and re-saturation of the leaves reduce the adjustmentresponse. We conclude that drought stress serves as a triggeronly, while the metabolic events lowering the osmotic potentialare favoured by high or intermediate water contents. Triticum durum L., durum wheat, rapid dehydration, osmotic adjustment, pressure-volume curves     

Ammonium-induced proline and sucrose accumulation, and their significance in antioxidative activity and osmotic adjustment

Excess of ammonia generates oxidative and osmotic stress, and results in an accumulation of compatible solutes. The aim of this study was to investigate the physiological significance of excess ammonium-induced proline and sucrose accumulation on antioxidative activity and osmotic adjustment. The detached leaves of white clover (Trifolium repense L.) were fed with 0, 10, 50, 100, and 200 mM NH₄Cl, and the contribution of proline and sucrose to osmotic adjustment and their relationship with antioxidative enzymes activity were assessed. A gradual decline of relative water content and osmotic potential (Ψ_π) with increasing NH₄Cl feeding level was accompanied by an increase in ammonia concentration. Significant accumulation of proline and sucrose was observed when NH₄Cl was fed over 100 mM compared with control (0 mM NH₄Cl). The increase in enzyme activity was significant only at 200 mM for ascorbate peroxidase (APOD) and over 100 mM NH₄Cl for guaiacol peroxidase (GPOD) and catalase (CAT). The contribution of proline and sucrose to osmotic adjustment over 100 mM, where proline and sucrose accumulation was more important, maintained at control levels or significantly decreased. The content of proline and sucrose as affected by NH₄Cl feeding level was positively related with the activity of APOD, GPOD, and CAT. These results suggest that proline and sucrose accumulation induced by the excess of ammonium has a more influential role in antioxidative activity rather than osmotic adjustment.     

The Effect of Grain Number per Ear (Sink Size) on Source Activity and its Water-Relations in Wheat

Blum, A., Mayer, J. and Golan, G. 1988. The effect of grainnumber per ear (sink size) on source activity and its water-relationsin wheat.–J. exp. Bot. 39: 106–114. Work was done to evaluate the nature of sink-source relationshipsin wheat (Triticum aestivum L.), when the strength of the sinkwas modified by the removal of half of the grain from the earat about anthesis. The main hypothesis was that sink-sourcerelationship would be modified by water stress and that a weakersink would improve the drought resistance of the source. Two experiments were performed. The first experiment evaluatedthe effect of de-graining in two wheat varieties grown in thefield. The second experiment (in the greenhouse) evaluated theeffect of de-graining in plants subjected to water stress afteranthesis by immersing the root system in a solution of polyethyleneglycol (6000), as compared with non-stressed controls. In bothexperiments measurements were performed after de-graining toprovide data on leaf gas exchange, leaf water potential, osmoticadjustment of leaves and ears (greenhouse), the percent of stemweight loss as an index of stem reserve mobilization, finalroot weight (greenhouse) and ear weight components. De-graining caused a decrease in flag leaf stomatal conductance,carbon exchange rate (CER) and transpiration and an increasein flag leaf water potential. These effects were stronger withwater stress. De-graining did not affect osmotic adjustmentin the flag leaf but induced better adjustment in glumes andawns. De-graining decreased the percent of stem weight lossand increased final root weight, especially under drought stress. A weaker sink was, therefore, considered to improve plant droughtresistance in terms of the maintenance of higher leaf waterpotential, a larger root, a better osmotic adjustment in theear and, possibly, increased flag leaf longevity. The ‘cost’of this improved drought resistance was in reduced flag leafCER and reduced stem (and root?) reserve mobilization. Key words: Drought resistance, carbon exchange rate, stomata, transpiration, osmotic adjustment, leaf water potential, root, awns, yield     

The Effect of Defoliation on the Nitrogen Economy of White Clover: Regrowth and the Remobilization of Plant Organic Nitrogen

Single plants of white clover (Trifolium repens) were establishedfrom stolon cuttings rooted in acid-washed silver sand. Allplants were inoculated with Rhizobium trifolii, and receivednutrient solution containing 0·5 mg ¹⁵N as either ammoniumor nitrate weekly for 12 weeks (i.e. 6 mg ¹⁵N in total). Plantswere then leniently defoliated or left intact, and the labelledN supply was replaced with unlabelled N. Lenient defoliationremoved fully expanded leaves only, leaving immature leaveswhich accounted for 50–55% of the total; growing pointnumbers were not reduced. Nodules, leaves and growing pointswere counted over the following 21 d period, and d. wts, N contents,and ¹⁵N enrichments of individual plant organs were determined. Defoliated plants had fewer nodules, but numbers of growingpoints were unaffected by defoliation. The rates of both leafemergence and expansion were accelerated in defoliated plants;in consequence the number of young leaves remained less thanin intact plants until day 21. Total dry matter (DM) and N accumulationwere less in defoliated plants, and a greater proportion oftotal plant DM was invested in roots. About 97 % of plant totalN was derived from fixed atmospheric N, but there was incompletemixing of fixed and mineral N within the plant. Relatively moremineral N was incorporated into roots, whereas there was relativelymore fixed N in nodules. There was isotopic evidence that Nwas remobilized from root and stolon tissue for leaf regrowthafter defoliation; approximately 2 % of plant N turned overdaily in the 7-d period after defoliation, and this contributedabout 50% of the N increment in leaf tissue. White clover, Trifolium repens L. cv. SI84, lenient defoliation, N economy, regrowth, N remobilization     

Understanding osmotic stress tolerance in leaves and nodules of two <Emphasis Type="Italic">Phaseolus vulgaris</Emphasis> cultivars with contrasting drought tolerance

Drought and salinity are environmental constraints that affect crop yields worldwide. In nature, both stresses are multifaceted problems that are usually associated with other adverse circumstances which limit plant performance such as water shortage and nutrient deficits. In order to assess common features of both stresses, the effects of mannitol-induced osmotic stress were monitored using two Phaseolus vulgaris cultivars, Cv. ‘Flamingo’ (tolerant) and Cv. ‘Coco Blanc’ (sensitive) which differed in their drought and salinity tolerance. Growth, water relations, organic and inorganic compound accumulation and soluble protein contents were measured in leaves and nodules of these N₂-fixing plants. The aim of the present study was to check whether osmotic stress tolerance is associated with accumulation of some of these compounds either in leaves, nodules or both organs. At the whole-plant level, Cv. ‘Flamingo’ showed a better maintenance of plant biomass and shoot water status. At the cell level, this was related to a better osmotic adjustment ability both in leaves and nodules and also to a better adjustment of the cell wall elasticity. At the metabolic level, the contrasting accumulation of the different amino acids in nodules of each cultivar suggested that amino acids pathways can be regulated to different degrees under stress conditions. At the metabolic level, it seems that symbiosis in the sink organ (the nodule) plays a crucial role in conferring drought and salinity tolerance in the common bean.     

The Effect of Water Stress on the Vegetative Growth of White Clover (Trifolium repens L): Comparison of Long-term Water Deficit and a Short-term Developing Water Stress

White clover plants were subjected to either a short-term developingwater stress or long-term stable levels of water deficit on‘water stress columns’. The short-term stress reducedplant water status to –2?0 MPa over 15 d. The water stresscolumns imposed only mild levels of water stress (a reductionof 0?35 MPa in leaf water potential for the more severe treatment)but these were maintained for several weeks. The absolute growthof plants on the control columns was maintained throughout theexperimental period. Vegetative growth was measured. Stolon, petiole, and laminagrowth were all reduced to some extent when plants were grownsymbiotically. The two regimes gave comparable results. Whennitrate was supplied there was no effect of water stress. Aconsiderably reduced absolute growth rate did not result ina similar decrease in final organ size. Stolon growth was mostreduced by water stress. Leaf death during water stress wasas important as changes in growth in determining final dry matteryield. Consequently, the yield of petiole and lamina from plantsgrown without supplied nitrate on the water stress columns waslower than that of stolon at the end of the treatment period. The merits of the water stress column system for imposing long-termwater deficit are discussed. Key words: Trifolium repens, white clover, water stress, vegetative growth     

Freezing Tolerance and Solute Changes in Contrasting Genotypes of Lolium perenne L. Acclimated to Cold and Drought

Clonal ramets of 12 contrasting genotypes of Lolium perenneL. were grown in sand or soil-based compost and maintained underwell-watered conditions at 20/15°C or acclimated to lowtemperature (2°C) or to a restricted water supply. Freezingtolerance was measured as LT₅₀ following exposure to sub-zerotemperatures in a freezing tank. Measurements were also madeof osmotic potential, water-soluble carbohydrates, free proline,free amino acids, and minerals in entire tillers. Acclimationto both drought and cold lowered LT₅₀, induced osmotic adjustment,and increased concentrations of proline and amino acids, Rootingmedium had little effect on LT₅₀, but caused large differencesin osmotic potential and in proline and amino-acid concentrations.There was considerable genetic diversity for all charactersmeasured, except for mineral contents. There was, however, norelationship between LT₅₀ and osmotic potential or solute contentthat was consistent across the three sources of variation (growingmedium, acclimation, genotype). Furthermore, the diverse genotypicvalues of cold-induced freezing tolerance were not correlatedwith those of drought-induced tolerance. It is concluded thatmore precise measurements are needed of the partitioning ofsolutes during acclimation and of the sensitivity of differentorgans and tissues to freezing.Copyright 1993, 1999 AcademicPress Perennial ryegrass, hardening, acclimation, osmotic potential, solute potential, carbohydrates, proline     

Factors Affecting Polyamine Accumulation in Barley (Hordeum vulgare L.) Leaf Sections during Osmotic Stress

Turner, L. B. and Stewart, G. R. 1988. Factors affecting polyamineaccumulation in barley (Hordeum vulgare L.) leaf sections duringosmotic stress.-J. exp. Bot. 39: 311–316. Polyamine concentrations in peeled leaf sections of Hordeumvulgare were unaffected by decreases in leaf water potentialif osmotic adjustment took place and leaf turgor was maintained.Putrescine accumulation occurred concomitantly with a decreasein leaf turgor. Increases in the order of 3-to 4-fold were observed.An apparently greater putrescine accumulation (7-fold) occurredwhen leaf sections were osmotically stressed in the presenceof exogenous phosphate ions. This was the result of water lossfrom the tissue and the large decline which occurred in theputrescine levels of control tissue sections incubated withphosphate ions. Putrescine accumulation was at its maximum after4 h osmotic stress. In contrast, proline accumulation took placebetween 4 h and 24 h after the imposition of osmotic stress. Key words: Hordeum vulgare, osmotic stress, polyamine     

The Production of Leaves and Stolon Branches on Established White Clover Cuttings in Relation to Temperature and Soil Moisture in the Field

Successive sets of cuttings of three white clover genotypeswere raised in a 15 °C growth room and transferred to thefield at 14 d intervals over the course of a year. Rates ofleaf appearance (leaves per stolon growing point per unit time)were found to be closely correlated with 10 cm soil temperatures.Petiole lengths, and weights of the lamina+petiole increasedin May and decreased towards the end of August, but also exhibiteda marked response to a mid-season water deficit. In the conditionsof the experiment (i.e. in the absence of competition from neighbouringplants) the vast majority of axillary buds developed into visiblebranches at all times of the year. There was, however, an increasein the nodal age at which bud development was first observedin winter. Deferred bud development was also observed, particularlyin one genotype, during periods characterized by dry soil surfaceconditions. The results are discussed in relation to observedpatterns of stolon branching in sward conditions. White clover, Trifolium repens, axillary bud development, branching, growing points, leaf appearance rate, petiole length, soil moisture, soil temperature