Are ABA,ethylene or their interaction involved in the response of leaf growth to soil water deficit? An analysis using naturally occurring variation or genetic transformation of ABA production in maize

The role of abscisic acid (ABA) and its possible interaction with ethylene in mediating leaf elongation response to soil water deficit are a matter of controversy. To address this question, we used a set of maize genotypes with various levels of ABA either due to natural variability or to genetic transformation targeted on NCED/VP14, a key enzyme of ABA synthesis. The transgenic lines yielded less strong phenotypes than available mutants, making it possible to use them under normal growing conditions. We focused on leaf elongation during night periods in order to avoid the confounding effect of ABA on leaf water status. Our results suggest that over a wide range, internal ABA level (measured in both leaf extracts or xylem sap) has no clear effect on leaf elongation response to soil water deficit, except in the case of an antisense line presenting the strongest reduction in ABA accumulation that showed a slight maintenance of leaf elongation during water deficit. Leaf ethylene production rate was variable and not related to water deficit except in the ABA-deficient transgenic lines where it was increased by water deficit on average but not systematically. Moreover, variability in ethylene production rate was not linked to variability in elongation rate. Our results thus suggest that neither ABA nor ethylene seems to play a major role in the control of leaf elongation response to soil water deficit.     

A common genetic determinism for sensitivities to soil water deficit and evaporative demand: meta-analysis of quantitative trait Loci and introgression lines of maize

Evaporative demand and soil water deficit equally contribute to water stress and to its effect on plant growth. We have compared the genetic architectures of the sensitivities of maize (Zea mays) leaf elongation rate with evaporative demand and soil water deficit. The former was measured via the response to leaf-to-air vapor pressure deficit in well-watered plants, the latter via the response to soil water potential in the absence of evaporative demand. Genetic analyses of each sensitivity were performed over 21 independent experiments with (1) three mapping populations, with temperate or tropical materials, (2) one population resulting from the introgression of a tropical drought-tolerant line in a temperate line, and (3) two introgression libraries genetically independent from mapping populations. A very large genetic variability was observed for both sensitivities. Some lines maintained leaf elongation at very high evaporative demand or water deficit, while others stopped elongation in mild conditions. A complex architecture arose from analyses of mapping populations, with 19 major meta-quantitative trait loci involving strong effects and/or more than one mapping population. A total of 68% of those quantitative trait loci affected sensitivities to both evaporative demand and soil water deficit. In introgressed lines, 73% of the tested genomic regions affected both sensitivities. To our knowledge, this study is the first genetic demonstration that hydraulic processes, which drive the response to evaporative demand, also have a large contribution to the genetic variability of plant growth under water deficit in a large range of genetic material.     

Plasticity to soil water deficit in Arabidopsis thaliana: dissection of leaf development into underlying growth dynamic and cellular variables reveals invisible phenotypes

Genetic variability in the plasticity of leaf area expansion in response to water deficit has been reported in Arabidopsis thaliana. Here, the objective was to identify the underlying dynamic and cellular processes involved in this variability. Twenty-five accessions were subjected to identical soil water deficit treatments. In all accessions, the plasticity of leaf production was low compared with that of individual leaf expansion. A subset of accessions was selected for further dissection of individual leaf expansion into its underlying variables: the rate and duration of leaf expansion and epidermal cell number and area. In all accessions, water deficit had opposite effects on the rate and duration of leaf expansion. The accumulation of these effects was reflected in changes in final leaf area. At the cellular level, moderate water deficits had opposite effects on cell number and cell size, but more severe ones reduced both variables. The importance of these opposing effects is highlighted by the behaviour of the accession An-1, for which the compensation between the decrease in leaf expansion rate and the increase in the duration of expansion is total. This dynamic plasticity in response to water deficit is not detectable when only final measurements are done.     

Dealing with the genotype x environment interaction via a modelling approach: a comparison of QTLs of maize leaf length or width with QTLs of model parameters

Quantitative genetics of adaptive traits is made difficult by the genotypexenvironment interaction. A classical assumption is that QTLs identified in both stressed and control conditions correspond to constitutive traits whereas those identified only in stressed treatments are stress-specific and correspond to adaptive traits. This hypothesis was tested by comparing, in the same set of experiments, two ways of analysing the genetic variability of the responses of maize leaf growth to water deficit. One QTL detection was based on raw phenotypic traits (length and width of leaf 6) of 100 recombinant inbred lines (RILs) in four experiments with either well-watered or stressing conditions in the field or in the greenhouse. Another detection followed a method proposed recently which consists of analysing intrinsic responses of the same RILs to environmental conditions, determined jointly over several experiments. QTLs of three responses were considered: (i) leaf elongation rate per unit thermal time in the absence of stress, (ii) its response to evaporative demand in well-watered plants, and (iii) its response to soil water status in the absence of evaporative demand. The QTL of leaf length differed between experiments, but colocalized in seven cases out of 13 with QTLs of the intrinsic leaf elongation rate, even in experiments with stressing conditions. No colocalization was found between QTLs of leaf length under water deficit and QTLs of responses to air or soil water status. By contrast, QTLs of leaf width colocalized in all experiments, regardless of environmental conditions. The classical method of identifying the QTL of constitutive versus adaptive traits therefore did not apply to the experiments presented here. It is suggested that identification of the QTL of parameters of response curves provides a promising alternative for dealing with the genetic variability of adaptive traits.     

Keep on growing under drought: genetic and developmental bases of the response of rosette area using a recombinant inbred line population

Variation in leaf development caused by water deficit was analysed in 120 recombinant inbred lines derived from two Arabidopsis thaliana accessions, Ler and An‐1. Main effect quantitative trait loci (QTLs) and QTLs in epistatic interactions were mapped for the responses of rosette area, leaf number and leaf 6 area to water deficit. An epistatic interaction between two QTLs affected the response of whole rosette area and individual leaf area but only with effects in well‐watered condition. A second epistatic interaction between two QTLs controlled the response of rosette area and leaf number with specific effects in the water deficit condition. These effects were validated by generating and phenotyping new appropriate lines. Accordingly, a low reduction of rosette area was observed for lines with a specific allelic combination at the two interacting QTLs. This low reduction was accompanied by an increase in leaf number with a lengthening of the vegetative phase and a low reduction in individual leaf area with low reductions in epidermal cell area and number. Statistical analyses suggested that responses of epidermal cell area and number to water deficit in individual leaves were partly caused by delay in flowering time and reduction in leaf emergence rate, respectively.     

Differential sensitivities of transpiration to evaporative demand and soil water deficit among wheat elite cultivars indicate different strategies for drought tolerance

In order to satisfy increasing wheat demands, scaling up wheat production will require boosting yield in suboptimal, drought-prone areas. Under rain-fed environments, one promising option is the identification of traits allowing for soil water conservation until the next rain episode. This can be achieved either by limiting transpiration rate (TR) of the crop to a maximum level when atmospheric drought (or VPD for vapor pressure deficit) is too high or by decreasing stomata conductance earlier in the soil drying cycle. Although promising, those strategies were never explored in wheat. A first objective of this study was to investigate the extent of the genetic variability of TR sensitivity to both VPD and soil water deficit among a group of eight elite wheat lines, which are cultivated under south Australian conditions. Those consisted of seven differentially drought-adapted lines and one check cultivar. TR responses to VPD were highly variable among genotypes, with six lines displaying a breakpoint in their TR response to VPD that ranged from 2.4 to 3.9 kPa, while two others had their TR increasing linearly as VPD increased. Transpiration response to a progressively decreasing fraction of transpirable soil water (FTSW) was investigated in those lines. A significant genetic variability in the responses among genotypes was observed. They revealed different FTSW thresholds at which transpiration started to decrease at levels ranging from 0.43 to 0.52, and different slopes for the decrease. A second objective was to investigate the existence of phenotypic correlations between the parameters characterizing transpiration sensitivities to both sources of water deficit (i.e., VPD and FTSW). Significant correlations were observed revealing that genotypes with conservative water use in their response to high VPD were also conservative in response to decreasing FTSW and that the drought tolerance of other lines might stem from an apposite strategy, invoking decreased sensitivity of TR to both sources of drought. Those findings provide new options for breeding drought tolerant lines based on this germplasm.     

Water relations and stomatal characteristics of Mediterranean plants with different growth forms and leaf habits: responses to water stress and recovery

The aim of this study was to extent the range of knowledge about water relations and stomatal responses to water stress to ten Mediterranean plants with different growth forms and leaf habits. Plants were subjected to different levels of water stress and a treatment of recovery. Stomatal attributes (stomatal density, StoD), stomatal conductance (g _s), stomatal responsiveness to water stress (SR), leaf water relations (pre-dawn and midday leaf water potential and relative water content), soil to leaf apparent hydraulic conductance (K _L) and bulk modulus of elasticity (ε) were determined. The observed wide range of water relations and stomatal characteristics was found to be partially depended on the growth form. Maximum g _s was related to StoD and the stomatal area index (SAI), while g _s evolution after water stress and recovery was highly correlated with K _L. Relationships between SR to water deficit and other morphological leaf traits, such as StoD, LMA or ε, provided no general correlations when including all species. It is concluded that a high variability is present among Mediterranean plants reflecting a continuum of leaf water relations and stomatal behaviour in response to water stress.     

Variability of traits associated with phosphorus efficiency in wild and cultivated genotypes of common bean

Genetic variation in plant growth under limited phosphorus (P) supply is necessary to obtain more productive cultivars on low P-available soils. Two pot experiments were conducted to evaluate the variability of some traits associated with efficiency of P absorption and utilization in wild and cultivated genotypes of common bean (Phaseolus vulgaris L.) under biological N2 fixation. At two P levels (20 and 80 mg P kg-1 soil, P1 and P2, respectively), 20 wild and 6 cultivated genotypes were grown in Experiment 1, and 4 wild and 27 cultivated genotypes were grown in Experiment 2. Plants were harvested at flowering, but in Experiment 1 wild accessions that did not flower were harvested at the beginning of leaf senescence. In Experiment 1, part of the genotypic variability of wild accessions was attributed to a less homogeneous ontogenetic stage at harvest, whereas in Experiment 2 some variation in biomass production was due to distinct phenologies of cultivated genotypes. Wild lines did not seem more tolerant to low P conditions, but the genotypic variation observed suggests these materials as a source of genetic diversity. Part of the variation in the root area and root efficiency ratio (total P content:root area) was compensatory, resulting in narrow genotypic differences in the total P content. The total P content and root efficiency ratio presented a wider amplitude of variation at P2 than at P1, and P uptake was more influenced by P supply than root production. Since the genotype × P level interaction was not significant for shoot biomass and shoot P concentration in Experiment 2, P utilization efficiency may be a useful selection criterion for cultivars between limited and adequate P supply. Within the sample of genetic diversity evaluated herein, there was large genotypic variability for traits related to P efficiency among wild and cultivated genotypes of common bean.     

Effect of Verticillium dahliae on Photosynthesis, Leaf Expansion and Senescence of Field-grown Sunflower

On the basis of known sunflower (Helianthus annuus L.) responsesto soil water deficit, it is proposed that the effect of thefungus Verticillium dahliae Klebahn on plant leaf area precedesand is greater than its effect on leaf photosynthesis and stomatalconductance. To test this hypothesis, we measured shoot andleaf area growth, leaf photosynthetic rate, stomatal conductanceand disease symptoms in a field experiment including hybridsof high (Sankol) and low (Dekasol 3900) susceptibility to V.dahliae. Plants inoculated with V. dahliae and controls werecompared. We also investigated the effect of V. dahliae on keycomponents of plant leaf area, leaf expansion and senescence,in inoculated and control plants of Sankol and Toba, a hybridof intermediate susceptibility to V. dahliae. Reduction in plantleaf area caused by V. dahliae was first detected 31 d afterinoculation (DAI), when visual symptoms of disease in inoculatedplants were slight (Sankol) or absent (Dekasol 3900). Reductionin leaf photosynthesis was first observed 66 DAI; stomatal conductanceand leaf dark respiration were both unaffected by V. dahliaeduring the whole experiment. In comparison with controls, V.dahliae reduced seasonal duration of plant leaf area by 25%in Dekalb 3900 and by 55% in Sankol, whereas the average reductionin leaf photosynthetic rate was 9%. In correspondence with thereduction in leaf area duration, inoculation reduced shoot drymatter of mature Sankol by 50%. In both experiments, less leafexpansion accounted for most of the early reduction in plantleaf area; as the disease progressed, increasing senescencealso contributed to reduced plant leaf area. It is concludedthat the response of sunflower to V. dahliae resembled the responseof the plant to soil water deficit: (1) plant leaf area, ratherthan leaf photosynthetic rate, accounted for the reduction ingrowth in mass; and (2) reduced leaf expansion early in theseason and faster leaf senescence in older plants accountedfor the decrease in plant leaf area. Copyright 2000 Annals ofBotany Company Helianthus annuus, Verticillium dahliae, allometry, apical dominance, drought, leaf expansion, leaf senescence, photosynthesis, stomatal conductance, growth     

PHENOPSIS, an automated platform for reproducible phenotyping of plant responses to soil water deficit in Arabidopsis thaliana permitted the identification of an accession with low sensitivity to soil water deficit

The high-throughput phenotypic analysis of Arabidopsis thaliana collections requires methodological progress and automation. Methods to impose stable and reproducible soil water deficits are presented and were used to analyse plant responses to water stress. Several potential complications and methodological difficulties were identified, including the spatial and temporal variability of micrometeorological conditions within a growth chamber, the difference in soil water depletion rates between accessions and the differences in developmental stage of accessions the same time after sowing. Solutions were found. Nine accessions were grown in four experiments in a rigorously controlled growth-chamber equipped with an automated system to control soil water content and take pictures of individual plants. One accession, An1, was unaffected by water deficit in terms of leaf number, leaf area, root growth and transpiration rate per unit leaf area. Methods developed here will help identify quantitative trait loci and genes involved in plant tolerance to water deficit.     

Cell and leaf size plasticity in Arabidopsis: what is the role of endoreduplication?

Leaf area expansion is affected by environmental conditions because of differences in cell number and/or cell size. Increases in the DNA content (ploidy) of a cell by endoreduplication are related to its size. The aim of this work was to determine how cell ploidy interacts with the regulation of cell size and with leaf area expansion. The approach used was to grow Arabidopsis thaliana plants performing increased or decreased rounds of endoreduplication under shading and water deficit. The shading and water deficit treatments reduced final leaf area and cell number; however, cell area was increased and decreased, respectively. These differences in cell size were unrelated to alterations of the endocycle, which was reduced by these treatments. The genetic modification of the extent of endoreduplication altered leaf growth responses to shading and water deficit. An increase in the extent of endoreduplication in a leaf rendered it more sensitive to the shade treatment but less sensitive to water deficit conditions. The link between the control of whole organ and individual cell expansion under different environmental conditions was demonstrated by the correlation between the plasticity of cell size and the changes in the duration of leaf expansion.     

Are source and sink strengths genetically linked in maize plants subjected to water deficit? A QTL study of the responses of leaf growth and of Anthesis-Silking Interval to water deficit

Leaf growth and Anthesis-Silking Interval (ASI) are the main determinants of source and sink strengths of maize via their relations with light interception and yield, respectively. They depend on the abilities of leaves and silks to expand under fluctuating environmental conditions, so the possibility is raised that they may have a partly common genetic determinism. This possibility was tested in a mapping population which segregates for ASI. Maximum leaf elongation rate per unit thermal time (parameter a) and the slopes of its responses to evaporative demand and soil water status (parameters b and c) were measured in greenhouse and growth chamber experiments, in two series of 120 recombinant inbred lines (RILs) studied in 2004 and 2005 with 33 RILs in common both years. ASI was measured in three and five fields under well-watered conditions and water deficit, respectively. For each RIL, the maximum elongation rate per unit thermal time was reproducible over several experiments in well-watered plants. It was accounted for by five QTLs, among which three co-localized with QTLs of ASI of well-watered plants. The alleles conferring high leaf elongation rate conferred a low ASI (high silk elongation rate). The responses of leaf elongation rate to evaporative demand and to predawn leaf water potential were linear, allowing each RIL to be characterized by the slopes of these response curves. These slopes had three QTLs in common with ASI of plants under water deficit. The allele for leaf growth maintenance was, in all cases, that for shorter ASI (maintained silk elongation rate). By contrast, other regions influencing ASI had no influence on leaf growth. These results may have profound consequences for modelling the genotype x environment interaction and for designing drought-tolerant ideotypes.     

ADAPTATION TO FINE-GRAINED ENVIRONMENTAL VARIATION: AN ANALYSIS OF WITHIN-INDIVIDUAL LEAF VARIATION IN AN ANNUAL PLANT

Recent studies of evolution in heterogeneous environments have concentrated on the role of coarse-grained environmental variation. Here I explore the potential for a modular organism to adapt to fine-grained environmental variation through within-individual variation among modules. I describe the pattern of variation among leaves of single individuals and report results of initial analyses of genetic variation for within-individual variability in leaf traits and of genetic correlations that could influence the rate of further evolution of within-individual variation of these traits. Plants from 24 paternal half-sib families were raised in growth chambers, and five traits were measured for two leaves produced by each plant. Four of the five traits differed significantly between sampling times. Genetic analyses revealed significant additive genetic variation for within-individual variation in several traits. Estimates of family mean correlations between traits expressed at different times suggest few relationships that would be expected to impede response to selection for changes in the pattern of within-individual variation in leaf traits. These results support the possibility that within-individual variation could evolve as an adaptive response to fine-grained environmental variation and suggest a need for further investigation to improve understanding of evolution in heterogeneous environments.     

Co-ordination between leaf initiation and leaf appearance in field-grown maize (Zea mays): genotypic differences in response of rates to temperature

BACKGROUND AND AIMS: In maize (Zea mays), early flowering date, which is a valuable trait for several cropping systems, is associated with the number of leaves per plant and the leaf appearance rate. Final leaf number depends upon the rate and duration of leaf initiation. The aims of this study were to analyse the genotypic variation in the response to temperature of leaf appearance rate and leaf initiation rate, and to investigate the co-ordination between these processes under field conditions. METHODS: Sixteen hybrids of different origins were grown under six contrasting environmental conditions. The number of appeared leaves was measured twice a week to estimate leaf appearance rate (leaves d(-1)). Plants were dissected at four sampling dates to determine the number of initiated leaves and estimate leaf initiation rate (leaves d(-1)). A co-ordination model was fitted between the number of initiated leaves and the number of appeared leaves. This model was validated using two independent data sets. KEY RESULTS: Significant (P < 0.05) differences were found among hybrids in the response to temperature of leaf initiation rate (plastochron) and leaf appearance rate (phyllochron). Plastochron ranged between 24.3 and 36.4 degree days (degrees Cd), with a base temperature (Tb) between 4.0 and 8.2 degrees C. Phyllochron ranged between 48.6 and 65.5 degrees Cd, with a Tb between 2.9 and 5.0 degrees C. A single co-ordination model was fitted between the two processes for all hybrids and environments (r2= 0.96, P < 0.0001), and was successfully validated (coefficient of variation < 9 %). CONCLUSIONS: This work has established the existence of genotypic variability in leaf initiation rate and leaf appearance rate in response to temperature, which is a promising result for maize breeding; and the interdependence between these processes from seedling emergence up to floral initiation.     

Nitrogen, phosphorus and light effects on growth and allocation of biomass and nutrients in wild rice

Separating plastic from ontogenetic and growth-limiting responses of plants to changes in resource availability can be challenging because there are a total of eight combinations of these three types of responses. These can, however, be uniquely distinguished on plots of root:shoot ratios against total biomass through time. We used this approach to separate ontogenetic, plastic, and growth-limiting responses of wild rice (Zizania palustris L.) to changes in nitrogen, phosphorus, and light availabilities. Relative growth rate was limited primarily by nitrogen but responded to increased light and phosphorus after nitrogen limitations were alleviated. Nitrogen addition increased relative growth rate because it simultaneously increased unit leaf rate, specific leaf area, and leaf weight ratio. Increased light did not change relative growth rate because decreased specific leaf area and leaf weight ratio compensated the increased unit leaf rate. Phosphorus did not change either relative growth rate or its underlying components. Plants responded ontogenetically to increased nitrogen and light availabilities by accelerating their developmental rate, and plastically by decreasing or increasing their root:shoot ratios, respectively. Plants did not respond either ontogenetically or plastically to increased phosphorus availability. Ontogenetic changes in growth can be separated from plastic and growth-limiting responses by plotting root:shoot ratio against total biomass in the context of the eight possible responses identified above, and also by examining how the underlying components of relative growth rate respond.     

Sources of Variation in Leaf Shape among Two Populations of Achillea Lanulosa

Achillea lanulosa has complex, highly dissected leaves that vary in shape and size along an altitudinal gradient. Plants from a high and an intermediate altitude population were clonally replicated and grown in a controlled environment at warm and cool conditions under bright light. There were genetic differences among populations and among individuals within populations in leaf size and shape. Heritabilities for leaf size and shape characters were moderate. Leaves of the lower altitude population were larger and differed from the higher altitude plants in both coarse and fine shape. Plastic response to temperature of the growth environment paralleled the genetic differentiation between low and high altitude populations. There was no apparent trade-off between genetic control over morphology and the capacity for directional plastic response to the environment. Differences in leaf dissection and size at contrasting altitudes in this species are the result of both genetic divergence among populations and of acclimative responses to local environments.     

Genotypic Differences in Leaf Water Potential, Abscisic Acid and Proline Concentrations in Spring Wheat during Drought Stress

Recent work with spring wheat has revealed significant genotypicvariation in changes of water potential and abscisic acid (ABA)concentration in response to drought Two experiments with eightspring wheat genotypes have been carried out to check the earlierwork on relationships between water potential and ABA concentrationand to examine causes of genotypic variation in the rate ofdecline of water potential during drought Changes in prolineconcentration were also studied Plants were grown in controlled environment cabinets with nutrientsolution culture and were stressed by withholding water as thefifth or sixth leaf on the main stem emerged. Plants were harvested4, 5 and 6 days after the treatment commenced and measurementsof leaf water potential, stomatal conductance, ABA and prolineconcentrations, and tissue d wts were taken. Significant genotypic variation was found in the decrease ofwater potential with time and in the slopes of linear regressionsof ABA concentration on water potential, confirming earlierresults When differences between leaf areas at the start of the treatmentwere minimised by varying the genotype sowing date significantgenotypic variation in water potentials at harvest was stillobtained. The change in water potential was significantly positivelycorrelated with shoot root d wt ratios at harvest and pre-treatmentstomatal conductances. Proline concentrations were significantly correlated with waterpotential for every genotype, although there was no clear evidenceof genotypic variation in proline concentrations at a givenwater potential The possible role of ABA concentration in drought resistanceof cereals is discussed Triticum aestivum L, spring wheat, water potential, abscisic acid, proline, drought stress     

Effects of soil water conditions on the morphology, phenology, and photosynthesis of Betula ermanii in the boreal forest

Four 2-year watering treatments were applied to Betula ermanii seedlings to investigate their responses to soil water conditions: ID [irrigated well (I) in the previous year and water deficit (D) in the current year], DD, DI, and II. RGR of the seedlings in current-year water deficit without experiencing previous-year water deficit (ID) was smaller than that of seedlings irrigated well in both years (II). Surprisingly, RGR did not differ between 2-year-water-deficited (DD) and -well-irrigated (II) treatments. There was no difference in the area-based photosynthetic rate of the late leaves, which are formed in the current-year environment, between the four water treatments, but their leaf longevity was shortened by water deficit. Area-based photosynthetic rate of the early leaves, whose buds are formed in the previous year environment, was smaller in ID than in II, but it did not differ between DD and II. The de-epoxidation state of the xanthophyll cycle was greater in ID than in DD. Leaf mass per area (leaf dry weight/leaf area) did not differ among the four water treatments. Specific root length (root total length/root dry weight) did not differ between ID and II. It was greater in DD than in II, probably resulting in increased water uptake efficiency. Photoprotective systems such as xanthophyll contents and antioxidant enzyme activities did not differ among the four treatments. Morphological responses of B. ermanii roots seem to be important as whole-plant-level responses to water deficit for maintaining RGR, in addition to leaf-level photosynthetic and phenological responses of two types of leaves. Our results partly explain how B. ermanii seedlings survive and maintain growth even under varying soil water conditions in the boreal forest.     

Impact of drought on productivity and water use efficiency in 29 genotypes of Populus deltoides x Populus nigra

We examined the relationships among productivity, water use efficiency (WUE) and drought tolerance in 29 genotypes of Populus x euramericana (Populus deltoides x Populus nigra), and investigated whether some leaf traits could be used as predictors for productivity, WUE and drought tolerance. At Orléans, France, drought was induced on one field plot by withholding water, while a second plot remained irrigated and was used as a control. Recorded variables included stem traits (e.g. biomass) and leaf structural (e.g. leaf area) and functional traits [e.g. intrinsic water use efficiency (Wi) and carbon isotope discrimination (Delta)]. Productivity and Delta displayed large genotypic variability and were not correlated. Delta scaled negatively with Wi and positively with stomatal conductance under moderate drought, suggesting that the diversity for Delta was mainly driven by stomatal conductance. Most of the productive genotypes displayed a low level of drought tolerance (i.e. a large reduction of biomass), while the less productive genotypes presented a large range of drought tolerance. The ability to increase WUE in response to water deficit was necessary but not sufficient to explain the genotypic diversity of drought tolerance.     

The Effects of Position and Temperature on the Expansion of Leaves of Vicia faba L.

The growth in area of the first eight leaves of broad bean plantswas investigated in growth room experiments. Plants were grownat either 20 or 14 °C or transferred from 20 to 14 °C.Rates of leaf appearance and unfolding increased with temperature.The duration of growth of a leaf increased with leaf numberfor the first five leaves and then remained constant The meangrowth rate declined or remained constant with increasing leafnumber Durations of growth were shorter and growth rates largerat 20 °C than at 14 °C Plants responded immediatelyto the change in temperature Final areas of leaves which expandedafter transfer from 20 to 14 °C were larger than those grownat 20 °C Vicla faba L., broad bean, leaf expansion, temperature responses