Water-limited Plants Research Articles

Combined effects of drought and simulated pathogen attack on root exudation rates of tomatoes

Background and aimsFood production is threatened by direct climate change effects including drought. Indirect effects, including changes in plant-pathogen dynamics and increased susceptibility to pathogens, further exacerbate the risks. Root exudation, which plays a crucial role in plant defence against drought and pathogens, is influenced by both water stress and pathogens. However, the interactive effects of these abiotic and biotic factors are rarely studied.MethodsWe conducted a controlled environment experiment to investigate the effects of moderate drought and simulated pathogen attack (using pipecolic acid, an inducer of systemic acquired resistance) on the rates of root exudation of total organic carbon (TOC) and total nitrogen (TN) of four tomato cultivars grown in potting soil.ResultsDrought increased the exudation of TOC and TN per unit of root area, while pipecolic acid did not have any significant effect. Furthermore, there was no interaction observed between the abiotic and biotic factors. However, due to the reduction in plant and root biomass caused by drought, the total exudation per plant remained similar between control and water-limited plants. Additionally, pipecolic acid reduced the carbon-to-nitrogen ratio of exudates and increased the total exudation of TN.ConclusionThe increased exudation observed in drought-stressed plants may serve as a strategy to maintain root and rhizosphere activity despite reduced root growth. Notably, the impact of drought differed among the tested cultivars, highlighting their diverse levels of drought tolerance. This emphasises the importance of preserving a wide range of crop cultivars to ensure food security under increasing drought.

Influence of Water Limitation and Provenance on Reproductive Traits in a Common Garden of Frangula alnus Mill.

Drought periods during the growing season will increase and intensify in Western Europe due to climate change. To better understand the consequences for woody perennials, we restricted watering of potted cuttings of Frangula alnus Mill. in a common garden setting in Belgium during the growing season of 2020. We focused on the responses of three provenances (Belgian, Italian and Swedish) for several reproductive traits in the year of the water limitation. F. alnus, as a shrub species, bears fruits fast. It can blossom on current season’s growth and can therefore produce ripened berries continuously during several months. The total berry count across the whole growing season was much lower in the water-limited plants, independent of the provenance. The local Belgian provenance produced more ripened berries in total than the Italian and Swedish provenances, both in the water-limited plants as in the control plants. Maximal berry production occurred in July. The control plants from the Belgian and Italian provenances displayed a second lower maximum in August. Mainly the Swedish provenance displayed a clear advancement of the maximum berry production among the water-limited plants in comparison to the control plants. Slight differences were detected in the average stone count per berry and in the average stone weight, with both traits displaying a single maximum through time. The Swedish provenance displayed the highest average stone count per berry and the Belgian provenance had the lightest average stone weight, both likely attributable to local adaptation. Remarkably, both stone traits were not affected by the water limitation. Results are discussed in the context of several drought response mechanisms including drought escape, drought avoidance, compensation growth, growth/reproduction trade-off and seed size/number trade-off.

The circadian night depression of photosynthesis analyzed in a herb, Pulmonaria vallarsae. Day/night quantitative relationships

Although many photosynthesis related processes are known to be controlled by the circadian system, consequent changes in photosynthetic activities are poorly understood. Photosynthesis was investigated during the daily cycle by chlorophyll fluorescence using a PAM fluorometer in Pulmonaria vallarsae subsp. apennina, an understory herb. A standard test consists of a light induction pretreatment followed by light response curve (LRC). Comparison of the major diagnostic parameters collected during day and night showed a nocturnal drop of photosynthetic responses, more evident in water-limited plants and consisting of: (i) strong reduction of flash-induced fluorescence peaks (FIP), maximum linear electron transport rate (Jmax, ETREM) and effective PSII quantum yield (ΦPSII); (ii) strong enhancement of nonphotochemical quenching (NPQ) and (iii) little or no change in photochemical quenching qP, maximum quantum yield of linear electron transport (Φ), and shape of LRC (θ). A remarkable feature of day/night LRCs at moderate to high irradiance was their linear-parallel course in double-reciprocal plots. Photosynthesis was also monitored in plants subjected to 2–3 days of continuous darkness (“long night”). In such conditions, plants exhibited high but declining peaks of photosynthetic activity during subjective days and a low, constant value with elevated NPQ during subjective night tests. The photosynthetic parameters recorded in subjective days in artificial darkness resembled those under natural day conditions. On the basis of the evidence, we suggest a circadian component and a biochemical feedback inhibition to explain the night depression of photosynthesis in P. vallarsae.

Competition for water and species coexistence in phenologically structured annual plant communities.

Both competition for water and phenological variation are important determinants of plant community structure, but ecologists lack a synthetic theory for how they affect coexistence outcomes. We developed an analytically tractable model of water competition for Mediterranean annual communities and demonstrated that variation in phenology alone can maintain high diversity in spatially homogenous assemblages of water-limited plants. We modelled a system where all water arrives early in the season and species vary in their ability to grow under drying conditions. As a consequence, species differ in growing season length and compete by shortening the growing season of their competitors. This model replicates and offers mechanistic explanations for patterns observed in empirical studies of how phenology influences coexistence among Mediterranean annuals. Additionally, we found that a decreasing, concave-up trade-off between growth rate and access to water can maintain high diversity under simple but realistic assumptions. High diversity is possible because: (1) later plants escape competition after their earlier season competitors have gone to seed and (2) early-season species are more than compensated for their shortened growing season by a growth rate advantage. Together, these mechanisms provide an explanation for how phenologically variable annual plant species might coexist when competing only for water.

Alternative developmental and transcriptomic responses to host plant water limitation in a butterfly metapopulation.

Predicting how climate change affects biotic interactions poses a challenge. Plant-insect herbivore interactions are particularly sensitive to climate change, as climate-induced changes in plant quality cascade into the performance of insect herbivores. Whereas the immediate survival of herbivore individuals depends on plastic responses to climate change-induced nutritional stress, long-term population persistence via evolutionary adaptation requires genetic variation for these responses. To assess the prospects for population persistence under climate change, it is therefore crucial to characterize response mechanisms to climate change-induced stressors, and quantify their variability in natural populations. Here, we test developmental and transcriptomic responses to water limitation-induced host plant quality change in a Glanville fritillary butterfly (Melitaea cinxia) metapopulation. We combine nuclear magnetic resonance spectroscopy on the plant metabolome, larval developmental assays and an RNA sequencing analysis of the larval transcriptome. We observed that responses to feeding on water-limited plants, in which amino acids and aromatic compounds are enriched, showed marked variation within the metapopulation, with individuals of some families performing better on control and others on water-limited plants. The transcriptomic responses were concordant with the developmental responses: families exhibiting opposite developmental responses also produced opposite transcriptomic responses (e.g. in growth-associated transcripts). The divergent responses in both larval development and transcriptome are associated with differences between families in amino acid catabolism and storage protein production. The results reveal intrapopulation variability in plasticity, suggesting that the Finnish M. cinxia metapopulation harbours potential for buffering against drought-induced changes in host plant quality.

Local adaptation mediates floral responses to water limitation in an annual wildflower.

Understanding how environmental stress affects the strength of mutualisms is critically important given observed and projected environmental changes. In particular, the frequency and duration of drought have been increasing worldwide. We investigated how water availability affects plant traits that mediate a pollination mutualism. For butterfly-pollinated Phlox drummondii, we determined how moisture availability affects flower size, nectar volume, and nectar sugar amount. Furthermore, we explored the role that local adaptation may play in responses to moisture availability by including plants collected from regions that differ in aridity. Finally, we determined whether responses of plant populations to selection may differ under drought by calculating heritability of traits under control and dry conditions. Flower size was generally smaller in dry plants than in control plants. Early in the treatment period, nectar volume and sugar were higher in dry plants than in control plants for plants from both arid and wetter regions. With prolonged dry treatment, nectar volume and sugar remained higher only in plants from the arid region. Heritability of floral traits was lower for water-limited plants than for control plants. Plant investment into pollination mutualisms under environmental stress may depend on the extent to which populations are already locally adapted to such conditions, suggesting that mutualism may remain strong, at least in arid regions. However, decreases in heritability under water-limitation suggest that responses to selection imposed by pollinators may be low, even if drought-adapted plants maintain production of rewards to pollinators.

Daily osmotic adjustments in stem may be good predictors of water stress intensity in poplar

Drought events impair the carbon and water balances in plants. Climate changes highlight the importance to understand the limits of woody species to reallocate carbon in different processes and the mechanisms driving the osmotic adjustments during the day under stress. In this frame, the aim of this work was to investigate the plant capability to shift energy among competing sinks and preserve the osmotic balance during the day under severe short periods of water deficit. The role of carbohydrates as osmolytes as well as energy sources was investigated in poplar plants. Results highlighted that during water deficit soluble sugars, derived both from the new synthetised carbon and starch degradation, were principally convoyed in the bark. This increase in carbohydrates allowed the maintenance of a water reserve used during the day to prevent a water decrease within the xylem. The decrease of xylem sap osmotic potential during the night, driven by an increase of K, Ca, and fructose (+0.46, 0.52, and 0.26 mg ml−1 in water limited plants after 8 days of withholding water, respectively), probably further attracted water into the xylem. This response mechanism increased at higher water deficit intensity. The little variations in carbohydrates and mineral elements within the leaves highlighted the main role of sinks rather than sources in the early response to water deficit.

Predicting Longitudinal Traits Derived from High-Throughput Phenomics in Contrasting Environments Using Genomic Legendre Polynomials and B-Splines

Recent advancements in phenomics coupled with increased output from sequencing technologies can create the platform needed to rapidly increase abiotic stress tolerance of crops, which increasingly face productivity challenges due to climate change. In particular, high-throughput phenotyping (HTP) enables researchers to generate large-scale data with temporal resolution. Recently, a random regression model (RRM) was used to model a longitudinal rice projected shoot area (PSA) dataset in an optimal growth environment. However, the utility of RRM is still unknown for phenotypic trajectories obtained from stress environments. Here, we sought to apply RRM to forecast the rice PSA in control and water-limited conditions under various longitudinal cross-validation scenarios. To this end, genomic Legendre polynomials and B-spline basis functions were used to capture PSA trajectories. Prediction accuracy declined slightly for the water-limited plants compared to control plants. Overall, RRM delivered reasonable prediction performance and yielded better prediction than the baseline multi-trait model. The difference between the results obtained using Legendre polynomials and that using B-splines was small; however, the former yielded a higher prediction accuracy. Prediction accuracy for forecasting the last five time points was highest when the entire trajectory from earlier growth stages was used to train the basis functions. Our results suggested that it was possible to decrease phenotyping frequency by only phenotyping every other day in order to reduce costs while minimizing the loss of prediction accuracy. This is the first study showing that RRM could be used to model changes in growth over time under abiotic stress conditions.

Association of Shoot and Root Responses to Water Deficit in Young Faba Bean (Vicia faba L.) Plants.

Water deficit may occur at any stage of plant growth, with any intensity and duration. Phenotypic acclimation and the mechanism of adaptation vary with the evolutionary background of germplasm accessions and their stage of growth. Faba bean is considered sensitive to various kinds of drought. Hence, we conducted a greenhouse experiment in rhizotrons under contrasting watering regimes to explore shoot and root traits and drought avoidance mechanisms in young faba bean plants. Eight accessions were investigated for shoot and root morphological and physiological responses in two watering conditions with four replications. Pre-germinated seedlings were transplanted into rhizotron boxes filled with either air-dried or moist peat. The water-limited plants received 50-ml water at transplanting and another 50-ml water 4 days later, then no water was given until the end of the experimental period, 24 days after transplanting. The well-watered plants received 100 ml of water every 12 h throughout the experimental period. Root, stem, and leaf dry mass, their mass fractions, their dry matter contents, apparent specific root length and density, stomatal conductance, SPAD value, and Fv/Fm were recorded. Water deficit resulted in 3–4-fold reductions in shoot biomass, root biomass, and stomatal conductance along with 1.2–1.4-fold increases in leaf and stem dry matter content and SPAD values. Total dry mass and apparent root length density showed accession by treatment interactions. Accessions DS70622, DS11320, and ILB938/2 shared relatively high values of total dry mass and low values of stomatal conductance under water deficit but differed in root distribution parameters. In both treatments, DS70622 was characterized by finer roots that were distributed in both depth and width, whereas DS11320 and ILB938/2 produced less densely growing, thicker roots. French accession Mélodie/2 was susceptible to drought in the vegetative phase, in contrast to previous results from the flowering phase, showing the importance of timing of drought stress on the measured response. Syrian accession DS70622 explored the maximum root volume and maintained its dry matter production, with the difference from the other accessions being particularly large in the water-limited treatment, so it is a valuable source of traits for avoiding transient drought.

Coordinated plasticity maintains hydraulic safety in sunflower leaves.

The xylem cavitation threshold water potential establishes a hydraulic limit on the ability of woody species to survive in water-limiting environments, but herbs may be more plastic in terms of their ability to adapt to drying conditions. Here, we examined the capacity of sunflower (Helianthus annuus L.) leaves to adapt to reduced water availability by modifying the sensitivity of xylem and stomata to soil water deficit. We found that sunflower plants grown under water-limited conditions significantly adjusted leaf osmotic potential, which was linked to a prolongation of stomatal opening as soil dried and a reduced sensitivity of photosynthesis to water-stress-induced damage. At the same time, the vulnerability of midrib xylem to water-stress-induced cavitation was observed to be highly responsive to growth conditions, with water-limited plants producing conduits with thicker cell walls which were more resistant to xylem cavitation. Coordinated plasticity in osmotic potential and xylem vulnerability enabled water-limited sunflowers to safely extract water from the soil, while protecting leaf xylem against embolism. High plasticity in sunflower xylem contrasts with data from woody plants and may suggest an alternative strategy in herbs.

Severe water deficit restricts biomass production of Lolium perenne L. and Trifolium repens L. and causes foliar nitrogen but not carbohydrate limitation

AimsWe investigated whether drought-induced impairment of grassland species can be explained directly by plant water deficit or by water-driven limitation of nitrogen (N) and/or carbohydrate sources.MethodsIn a field experiment, a severe drought treatment was applied on monocultures of Lolium perenne L. (cv. Alligator) (Lp) and Trifolium repens L. (cv. Hebe) (Tr) by using rainout shelters excluding all precipitation, and effects were compared to a rainfed control. Three species-fertiliser treatments were set up, crossed with the drought treatment. The two species were fertilised equally with N (200 kg N ha−1 year−1), and an additional high N fertilisation treatment was established for L. perenne (LphighN, 500 kg N ha−1 year−1).ResultsSevere soil water deficit led to significantly lower leaf water potentials in all species-fertiliser treatments (P < 0.001) down to approximately −1.2 MPa and, on average, to a 79% reduction in living plant biomass above 7 cm harvest height (P < 0.001), indicating strong plant water deficits. Under the drought treatment, living plant biomass above 7 cm did not differ among species-fertiliser treatments. Plant-available soil N was 84% lower (P ≤ 0.01) and plant N concentrations were 24% less (P < 0.001) under the drought than under the rainfed control treatment, with Lp always being more N limited than LphighN and Tr. Nitrate concentrations in water-limited plants were generally very low (< 0.85 mg g−1 dry matter), whereas non-structural carbohydrates were distinctly greater under the drought treatment in Lp (+62%), LphighN (+46%), and Tr (+18%).ConclusionsRestricted biomass production of these forage species under severe drought can primarily be explained by plant water deficits and secondarily by drought-induced limitation of N supply. However, growth seems not to be limited by carbohydrate source activity, as carbohydrates accumulated with water deficiency.

Relationship of leaf oxygen and carbon isotopic composition with transpiration efficiency in the C4 grasses Setaria viridis and Setaria italica.

Leaf carbon and oxygen isotope ratios can potentially provide a time-integrated proxy for stomatal conductance (gs) and transpiration rate (E), and can be used to estimate transpiration efficiency (TE). In this study, we found significant relationships of bulk leaf carbon isotopic signature (δ13CBL) and bulk leaf oxygen enrichment above source water (Δ18OBL) with gas exchange and TE in the model C4 grasses Setaria viridis and S. italica. Leaf δ13C had strong relationships with E, gs, water use, biomass, and TE. Additionally, the consistent difference in δ13CBL between well-watered and water-limited plants suggests that δ13CBL is effective in separating C4 plants with different availability of water. Alternatively, the use of Δ18OBL as a proxy for E and TE in S. viridis and S. italica was problematic. First, the oxygen isotopic composition of source water, used to calculate leaf water enrichment (Δ18OLW), was variable with time and differed across water treatments. Second, water limitations changed leaf size and masked the relationship of Δ18OLW and Δ18OBL with E. Therefore, the data collected here suggest that δ13CBL but not Δ18OBL may be an effective proxy for TE in C4 grasses.

Mile-a-minute weed (Persicaria perfoliata) and weevil (Rhinoncomimus latipes) response to varying moisture and temperature conditions

The combined effects of herbivory and water stress on growth and reproduction of mile-a-minute weed (Persicaria perfoliata (L.) H. Gross) were investigated in greenhouse trials over two years, with well-watered or water-limited plants either exposed or not exposed to herbivory by the mile-a-minute weevil (Rhinoncomimus latipes Korotyaev). Moisture limitation and weevil herbivory significantly reduced the number of seeds produced by P. perfoliata, with the fewest seeds produced when both factors were present. Seed weight was reduced by moisture limitation and weevil herbivory the second year, and seed viability was reduced by herbivory both years. Plant biomass was lower both years under conditions of water limitation, with an additional effect of herbivory the second year. Well-watered plants the second year also produced substantially more weevils than water-limited plants by the end of the season. Results are consistent with field observations suggesting that years of high rainfall allow resurgence of P. perfoliata plant populations that were previously suppressed by R. latipes. An additional environmental chamber trial assessed the interaction between the weed and weevil at two different temperatures. Here, plant mortality occurred only at the higher temperature with weevil herbivory, suggesting that herbivory has a greater negative effect on P. perfoliata under warm conditions. Additional studies on temperature effects are needed for a more complete understanding of interactions between P. perfoliata and R. latipes under different abiotic conditions.

Dryland Riparian Ecosystems in the American Southwest: Sensitivity and Resilience to Climatic Extremes

Drylands make up over 40% of the earth’s land surface (Noy-Meir 1973; Reynolds and others 2007). In these arid and semiarid landscapes, surface and subsurface water flows create mesic riparian environments (Kingsford 2006; Stromberg and Tellman 2009). Dryland riparian areas sustain water-limited plants and animals that cannot withstand upland conditions year-round, thereby supporting regional diversity (Sabo and others 2005; Lite and others 2005). Dryland rivers are also characterized by occasional large, high-intensity floods that rework the bed geomorphology and increase riparian heterogeneity. This combined variation in water availability and flood disturbance creates spatiotemporally complex and unique landscapes (Soykan and Sabo 2009; Soykan and others 2012). The hydrology of desert riparian ecosystems has been extensively altered by human activities including stream diversion and groundwater extraction, and is expected to undergo further change in response to changing climates (Patten 1998). Globally, increasing temperatures associated with climate change are expected to intensify the water cycle, leading to greater rates of evapotranspiration, more water in the atmosphere, and more-frequent intense storms (Huntington 2006; IPCC 2007). More intense storms are likely to increase flood intensity over the next 50 years (Dominguez and others 2012). In the American Southwest in particular, warmer sea surface temperatures could spur stronger and more-frequent El Nino-associated winter storms (Garfin and Lenart 2007). Perhaps of greater importance are projected increases in aridity. Average precipitation is expected to increase at a global scale, but some regions, including the American Southwest, are expected to experience substantial decreases in average annual rainfall as well as increases in frequency and intensity of droughts (Seager and others 2007). As aridity increases, streams and riparian zones undergo declines in surface flow and depth to groundwater; there are declines, as well, in the extent of river segments with perennial (vs. intermittent or ephemeral) stream flow. Given that rivers and their riparian corridors function as keystone elements of arid and semiarid landscapes, climate change will have far reaching effects on regional biota (Stromberg and others 2010). Nevertheless, the extent of biotic change for riparian ecosystems in coming decades will be tempered by the fact that for desert rivers, climatic and hydrologic variation is the norm (Milly and Received 30 May 2012; accepted 13 September 2012

Physiological and growth responses to water deficits in cultivated strawberry (Fragaria × ananassa) and in one of its progenitors, Fragaria chiloensis

Cultivation of strawberry (Fragaria×ananassa) requires irrigation. Improving crop water use efficiency (WUE) is important for future production. Fragaria chiloensis, a progenitor of cultivated strawberry, grows in sandy soils, and may prove useful in breeding for improved WUE. Little, however, is known about variation in drought tolerance within this species. This research explores drought tolerance in a range of F. chiloensis and F.×ananassa genotypes. Four cultivars of F.×ananassa and four accessions of F. chiloensis were compared when well watered, and when subjected to a water deficit (65% of evapotranspiration). New leaf production, stomatal conductance, and photosynthetic rate were significantly reduced under water deficit, and also significantly differed between genotypes. A significant interaction of genotype and irrigation was found for transpiration rate, leaf area and dry mass, production of runners, predawn water potential, a measure of transpiration efficiency (shoot biomass produced per litre water transpired), and carbon isotope composition, indicating that some genotypes were more severely affected by water deficit than others. The South American F. chiloensis accession ‘Manzanar Alto’ had a similar rate of transpiration to the commercial cultivars, but the remaining (North American) F. chiloensis accessions used far less water than the F.×ananassa. Well-watered F. chiloensis plants used less water than water-limited plants of the F.×ananassa cultivar ‘Florence’. Transpiration efficiency of the F. chiloensis accession ‘BSP14’ was improved by water deficit: this was the only genotype not to show a reduction in leaf area and dry mass under water deficit. Greater drought resistance in three F. chiloensis accessions compared to F.×ananassa results from a conservative vegetative growth strategy, reducing loss of water.

Thermal imaging and carbon isotope composition indicate variation amongst strawberry ( Fragaria × ananassa) cultivars in stomatal conductance and water use efficiency

Cultivars with low stomatal conductance ( g s) may show high water use efficiency (WUE) under drought conditions, but under optimal conditions low g s may result in low vigour. A combination of thermal imaging and carbon isotope composition (δ 13C) analysis offers potential for screening simultaneously for both high g s and high WUE. Ten cultivars of strawberry ( Fragaria × ananassa Duch.) were grown in well watered or water limited conditions. Thermal images were taken of the plants, with various approaches to determine the optimal protocol for detecting variation in g s, including use of reference leaves, grids to maintain leaves horizontal, and collection of meteorological data in synchrony with thermal images. δ 13C of leaves, fruit, and crowns was determined. An index of g s derived from the temperature of horizontal leaves and the temperature of wet and dry references showed significant differences between cultivars and between well watered and water limited plants, as did g s estimated from leaf temperature, the temperature of a dry reference, and humidity. Thermal imaging indicated low g s in ‘Elsanta’ and ‘Totem’ and relatively high g s in well watered ‘Elvira’, ‘Florence’ and ‘Cambridge Favourite’. δ 13C of all plant material was higher in water limited than well watered plants and showed significant differences between cultivars. In one experiment leaf δ 13C indicated lowest WUE in ‘Elvira’ and highest WUE in ‘Totem’. δ 13C was inversely correlated with an index of g s derived from thermal imaging. Although the results indicate substantial variation in g s and WUE between cultivars, generally all cultivars responded to water deficit by lowering g s and hence increasing WUE.

Sucrose feeding reverses shade-induced kernel losses in maize

Water limitations can inhibit photosynthesis and change gene expression in ways that diminish or prevent reproductive development in plants. Sucrose fed to the plants can reverse the effects. To test whether the reversal acts generally by replacing the losses from photosynthesis, sucrose was fed to the stems of shaded maize plants (Zea mays) during reproductive development. Shading was adjusted to mimic the inhibition of photosynthesis around the time of pollination in water-limited plants. Glucose and starch were imaged and quantified in the female florets. Sucrose was infused into the stems to vary the sugar flux to the ovaries. Ovaries normally grew rapidly and contained large amounts of glucose and starch, with a glucose gradient favouring glucose movement into the developing ovary. Shade inhibited photosynthesis and diminished ovary and kernel size, weight, and glucose and starch contents compared with controls. The glucose gradient became small. Sucrose fed to the stem reversed these losses, and kernels were as large as the controls. Despite similar inhibition of photosynthesis, the depletion of ovary glucose and starch was not as severe in shade as during a comparable water deficit. Ovary abortion prevalent during water deficits did not occur in the shade. It is suggested that this difference may have been caused by more translocation in shade than during the water deficit, which prevented low sugar contents necessary to trigger an up-regulation of senescence genes known to be involved in abortion. Nevertheless, sucrose feeding reversed kernel size losses and it is concluded that feeding acted generally to replace diminished photosynthetic activity.

Effect of water availability on leaf water isotopic enrichment in beech seedlings shows limitations of current fractionation models

Current models of leaf water enrichment predict that the differences between isotopic enrichment of water at the site of evaporation (Delta(e)) and mean lamina leaf water enrichment (Delta(L)) depend on transpiration rates (E), modulated by the scaled effective length (L) of water isotope movement in the leaf. However, variations in leaf parameters in response to changing environmental conditions might cause changes in the water path and thus L. We measured the diel course of Delta(L) for (18)O and (2)H in beech seedlings under well-watered and water-limited conditions. We applied evaporative enrichment models of increasing complexity to predict Delta(e) and Delta(L), and estimated L from model fits. Water-limited plants showed moderate drought stress, with lower stomatal conductance, E and stem water potential than the control. Despite having double E, the divergence between Delta(e) and Delta(L) was lower in well-watered than in water-limited plants, and thus, L should have changed to counteract differences in E. Indeed, L was about threefold higher in water-limited plants, regardless of the models used. We conclude that L changes with plant water status far beyond the variations explained by water content and other measured variables, thus limiting the use of current evaporative models under changing environmental conditions.

Elevated CO 2 alleviates the impact of drought on barley improving water status by lowering stomatal conductance and delaying its effects on photosynthesis

We analysed the impact of elevated CO 2 on water relations, water use efficiency and photosynthetic gas exchange in barley ( Hordeum vulgare L.) under wet and drying soil conditions. Soil moisture was less depleted under elevated compared to ambient [CO 2]. Elevated CO 2 had no significant effect on the water relations of irrigated plants, except on whole plant hydraulic conductance, which was markedly decreased at elevated compared to ambient CO 2 concentrations. The values of relative water content, water potential and osmotic potential were higher under elevated CO 2 during the entire drought period. The better water status of water-limited plants grown at elevated CO 2 was the result of stomatal control rather than of osmotic adjustment. Despite the low stomatal conductance produced by elevated CO 2, net photosynthesis was higher under elevated than ambient CO 2 concentrations. With water shortage, photosynthesis was maintained for longer at higher rates under elevated CO 2. The reduction of stomatal conductance and therefore transpiration, and the enhancement of carbon assimilation by elevated CO 2, increased instantaneous and whole plant water use efficiency in both irrigated and droughted plants. Thus, the metabolism of barley plants grown under elevated CO 2 and moderate or mild water deficit conditions is benefited by increased photosynthesis and lower transpiration. The reduction in plant water use results in a marked increase in soil water content which delays the onset and severity of water deficit.

QUANTITATIVE TRAIT LOCI AFFECTING δ13C AND RESPONSE TO DIFFERENTIAL WATER AVAILIBILITY IN ARABIDOPSIS THALIANA

Phenotypic plasticity is an important response mechanism of plants to environmental heterogeneity. Here, we explored the genetic basis of plastic responses of Arabidopsis thaliana to water deficit by experimentally mapping quantitative trait loci (QTL) in two recombinant inbred populations (Cvi × Ler and Ler × Col). We detected genetic variation and significant genotype-by-environment interactions for many traits related to water use. We also mapped 26 QTL, including six for carbon isotope composition (δ13C). Negative genetic correlations between fruit length and fruit production as well as between flowering time and branch production were corroborated by QTL colocalization, suggesting these correlations are due to pleiotropy or physical linkage. Water-limited plants were more apically dominant with greater root:shoot ratios and higher δ13C (higher water-use efficiency) when compared to well-watered plants. Many of the QTL effects for these traits interacted significantly with the irrigation treatment, suggesting that the observed phenotypic plasticity is genetically based. We specifically searched for epistatic (QTL-QTL) interactions using a two-dimensional genome scan, which allowed us to detect epistasis regardless of additive genetic effects. We found several significant QTL-QTL interactions including three that exhibited environmental dependence. These results provide preliminary evidence for proposed genetic mechanisms underlying phenotypic plasticity.