Drought survival and recovery in grasses: Stress intensity and plant-plant interactions impact plant dehydration tolerance.

Plant dehydration tolerance confers drought survival in grasses, but the mortality thresholds according to soil water content (SWC), vapour pressure deficit (VPD) and plant-plant interactions are little explored. We compared the dehydration dynamics of leaf meristems, which are the key surviving organs, plant mortality, and recovery of Mediterranean and temperate populations of two perennial grass species, Dactylis glomerata and Festuca arundinacea, grown in monocultures and mixtures under a low-VPD (1.5 kPa) versus a high-VPD drought (2.2 kPa). The lethal drought index (LD50 ), that is, SWC associated with 50% plant mortality, ranged from 2.87% (ψs = -1.68 MPa) to 2.19% (ψs = -4.47 MPa) and reached the lowest values under the low-VPD drought. Populations of D. glomerata were more dehydration-tolerant (lower LD50 ), survived and recovered better than F. arundinacea populations. Plant-plant interactions modified dehydration tolerance and improved post-drought recovery in mixtures compared with monocultures. Water content as low as 20.7%-36.1% in leaf meristems allowed 50% of plants to survive. We conclude that meristem dehydration causes plant mortality and that drought acclimation can increase dehydration tolerance. Genetic diversity, acclimation and plant-plant interactions are essential sources of dehydration tolerance variability to consider when predicting drought-induced mortality.

Is a seasonally reduced growth potential a convergent strategy to survive drought and frost in plants?

BACKGROUND: Plants have adapted to survive seasonal life-threatening frost and drought. However, the timing and frequency of such events are impacted by climate change, jeopardizing plant survival. Understanding better the strategies of survival to dehydration stress is therefore timely and can be enhanced by the cross-fertilization of research between disciplines (ecology, physiology), models (woody, herbaceous species) and types of stress (drought, frost). SCOPE: We build upon the 'growth-stress survival' trade-off, which underpins the identification of global plant strategies across environments along a 'fast-slow' economics spectrum. Although phenological adaptations such as dormancy are crucial to survive stress, plant global strategies along the fast-slow economic spectrum rarely integrate growth variations across seasons. We argue that the growth-stress survival trade-off can be a useful framework to identify convergent plant ecophysiological strategies to survive both frost and drought. We review evidence that reduced physiological activity, embolism resistance and dehydration tolerance of meristematic tissues are interdependent strategies that determine thresholds of mortality among plants under severe frost and drought. We show that complete dormancy, i.e. programmed growth cessation, before stress occurrence, minimizes water flows and maximizes dehydration tolerance during seasonal life-threatening stresses. We propose that incomplete dormancy, i.e. the programmed reduction of growth potential during the harshest seasons, could be an overlooked but major adaptation across plants. Quantifying stress survival in a range of non-dormant versus winter- or summer-dormant plants, should reveal to what extent incomplete to complete dormancy could represent a proxy for dehydration tolerance and stress survival. CONCLUSIONS: Our review of the strategies involved in dehydration stress survival suggests that winter and summer dormancy are insufficiently acknowledged as plant ecological strategies. Incorporating a seasonal fast-slow economics spectrum into global plant strategies improves our understanding of plant resilience to seasonal stress and refines our prevision of plant adaptation to extreme climatic events.

Can trees buffer the impact of climate change on pasture production and digestibility of Mediterranean dehesas?

Sustainability and functioning of silvopastoral ecosystems are being threatened by the forecasted warmer and drier environments in the Mediterranean region. Scattered trees of these ecosystems could potentially mitigate the impact of climate change on herbaceous plant community but this issue has not yet tested experimentally. We carried out a field manipulative experiment of increased temperature (+2-3 °C) using Open Top Chambers and rainfall reduction (30%) through rain-exclusion shelters to evaluate how net primary productivity and digestibility respond to climate change over three consecutive years, and to test whether scattered trees could buffer the effects of higher aridity in Mediterranean dehesas. First, we observed that herbaceous communities located beneath tree canopy were less productive (351 g/m2) than in open grassland (493 g/m2) but had a higher digestibility (44% and 41%, respectively), likely promoted by tree shade and the higher soil fertility of this habitat. Second, both habitats responded similarly to climate change in terms of net primary productivity, with a 33% increase under warming and a 13% decrease under reduced rainfall. In contrast, biomass digestibility decreased under increased temperatures (-7.5%), since warming enhanced the fiber and lignin content and decreased the crude protein content of aerial biomass. This warming-induced effect on biomass digestibility only occurred in open grasslands, suggesting a buffering role of trees in mitigating the impact of climate change. Third, warming did not only affect these ecosystem processes in a direct way but also indirectly via changes in plant functional composition. Our findings suggest that climate change will alter both the quantity and quality of pasture production, with expected warmer conditions increasing net primary productivity but at the expense of reducing digestibility. This negative effect of warming on digestibility might be mitigated by scattered trees, highlighting the importance of implementing strategies and suitable management to control tree density in these ecosystems.

What do you mean "functional" in ecology? Patterns versus processes.

Use of the term "functional" trait has increased exponentially in ecology. Although accounting for numerous ecological questions, this concept raises several issues. We propose that the term "functional" could be misleading because (1) no rigorous criteria exist to identify "functional" traits and (2) it suggests that only some traits ("functional" ones) can inform our understanding of species functioning, whatever the scale or discipline. Hence, the concept of "functional" trait in ecology is starting to be challenged and it remains unclear why some traits should be considered functional, whereas other traits should not. We argue that the most used "functional" traits are meaningful because they reflect important differences between populations or species, based on synchronic comparisons, that is, irrespective of time (hereafter "pattern" traits). Hence, they are useful for identifying trade-offs and strategies across large numbers of observations, usually at rather coarse scales, and are most often used in analyses of "big data." However, given that many ecological processes occur across short time scales and narrow gradients of climate and resource availability, the efficacy of these traits to inform us about these ecological processes appears questionable. We show that trait measurements that take time explicitly into account (hereafter "process" traits) differ from pattern traits because they quantify the flows of material and energy within a given environment across a defined period of time. Although pattern traits and process traits are both functional, it is important to understand the differences between the approaches. Moreover, better accounting of ontogeny, life form, plasticity, and genetic variability is required to enhance the convergence between pattern and process approaches. This revised framework allows more explicit connections between trait ecology and other biological sciences. It should enhance the study of processes at all scales in order to investigate efficiently the adaptive responses of biological organisms to climate change.

Neighbor identity affects growth and survival of Mediterranean plants under recurrent drought.

The increasing intensity and frequency of droughts predicted for the Mediterranean basin with ongoing climate change will impact plant communities and ecosystem functioning. This study investigated the effect of severe recurrent droughts and the role of the neighbor plant identity on the growth and survival of three abundant and co-existing species of a typical Mediterranean shrubland. Two juvenile plants, either of the same species or in all possible combinations of the two woody species Quercus coccifera and Cistus albidus and the perennial grass species Brachypodium retusum were grown together in rhizotrons under controlled watering regimes for two years. Compared to a treatment with only one drought cycle, three successive droughts reduced the relative growth rates (RGR) of shoots and roots in B. retusum, but not in woody species, and increased the mortality of the woody species, but not that of the grass. The survival of C. albidus and of B. retusum, but not of Q. coccifera, increased when the neighbor individual was a different species than when it was the same species. Our data suggest that both species composition and frequency of drought events will impact the dynamics of plant communities in Mediterranean shrublands under ongoing climate change. The abundance of dehydration sensitive woody species will likely decrease under more frequent drought events at the expense of dehydration-tolerant grass species, resulting in potentially strong changes in the functioning of these ecosystems.

The resilience of perennial grasses under two climate scenarios is correlated with carbohydrate metabolism in meristems.

Extreme climatic events (ECEs) such as droughts and heat waves affect ecosystem functioning and species turnover. This study investigated the effect of elevated CO2 on species' resilience to ECEs. Monoliths of intact soil and their plant communities from an upland grassland were exposed to 2050 climate scenarios with or without an ECE under ambient (390 ppm) or elevated (520 ppm) CO2. Ecophysiological traits of two perennial grasses (Dactylis glomerata and Holcus lanatus) were measured before, during, and after ECE. At similar soil water content, leaf elongation was greater under elevated CO2 for both species. The resilience of D. glomerata increased under enhanced CO2 (+60%) whereas H. lanatus mostly died during ECE. D. glomerata accumulated 30% more fructans, which were more highly polymerized, and 4-fold less sucrose than H. lanatus. The fructan concentration in leaf meristems was significantly increased under elevated CO2. Their relative abundance changed during the ECE, resulting in a more polymerized assemblage in H. lanatus and a more depolymerized assemblage in D. glomerata. The ratio of low degree of polymerization fructans to sucrose in leaf meristems was the best predictor of resilience across species. This study underlines the role of carbohydrate metabolism and the species-dependent effect of elevated CO2 on the resilience of grasses to ECE.

Evaluation of cocksfoot (Dactylis glomerata L.) population for drought survival and behavior.

Climate change models predict frequent and intense droughts in the world. Development of drought-tolerant species and cultivars is necessary to cope with such changes. Forage grass species are affected, especially in the Mediterranean region. The aim of the present study was to investigate the diversity for drought survival, summer dormancy, and productivity within a cocksfoot population. The study was conducted in Morocco, under field conditions from 2011 to 2013. 283 genotypes of cocksfoot and parents were tested, characterized for dry matter yield, heading date, plant height, senescence, summer dormancy, and drought survival. Results exhibited a large variability between traits. 79% of the population had survived after severe drought summer while 57% yielded more than both parents. Also, 63% of the progeny had an intermediate score of summer dormancy estimated by senescence score. Large variability was also noticed for heading date and plant height. Several accessions combined a high yield and persistence under severe summer drought. Which explain the significant correlation (r = 0.18, P < 0.005) founded between total dry matter accumulated in 2013 and plant survival. Accordingly, our results showed that we can rise persistent and resilient genotypes among population with a good level of biomass.

Embolism and mechanical resistances play a key role in dehydration tolerance of a perennial grass Dactylis glomerata L.

Background and Aims: More intense droughts under climate change threaten species resilience. Hydraulic strategies determine drought survival in woody plants but have been hardly studied in herbaceous species. We explored the intraspecific variability of hydraulic and morphological traits as indicators of dehydration tolerance in a perennial grass, cocksfoot (Dactylis glomerata), which has a large biogeographical distribution in Europe. Methods: Twelve populations of cocksfoot originating from Mediterranean, Temperate and Northern European areas were grown in a controlled environment in pots. Dehydration tolerance, leaf and stem anatomical traits and xylem pressure associated with 88 or 50 % loss of xylem conductance (P88, P50) were measured. Key Results: Across the 12 populations of cocksfoot, P50 ranged from -3.06 to - 6.36 MPa, while P88 ranged from -5.06 to -11.6 MPa. This large intraspecific variability of embolism thresholds corresponded with the biogeographical distribution and some key traits of the populations. In particular, P88 was correlated with dehydration tolerance (r = -0.79). The dehydration-sensitive Temperate populations exhibited the highest P88 (-6.1 MPa). The most dehydration-tolerant Mediterranean populations had the greatest leaf dry matter content and leaf fracture toughness, and the lowest P88 (-10.4 MPa). The Northern populations displayed intermediate trait values, potentially attributable to frost resistance. The thickness of metaxylem vessel walls in stems was highly correlated with P50 (r = -0.92), but no trade-off with stem lignification was observed. The relevance of the linkage between hydraulic and stomatal traits is discussed for drought survival in perennial grasses. Conclusions: Compared with woody species, the large intraspecific variability in dehydration tolerance and embolism resistance within cocksfoot has consequences for its sensitivity to climate change. To better understand adaptive strategies of herbaceous species to increasing drought and frost requires further exploration of the role of hydraulic and mechanical traits using a larger inter- and intraspecific range of species.

A unified framework of plant adaptive strategies to drought: Crossing scales and disciplines.

Plant adaptation to drought has been extensively studied at many scales from ecology to molecular biology across a large range of model species. However, the conceptual frameworks underpinning the definition of plant strategies, and the terminology used across the different disciplines and scales are not analogous. 'Drought resistance' for instance refers to plant responses as different as the maintenance of growth and productivity in crops, to the survival and recovery in perennial woody or grassland species. Therefore, this paper aims to propose a unified conceptual framework of plant adaptive strategies to drought based on a revised terminology in order to enhance comparative studies. Ecological strategies encapsulate plant adaptation to multidimensional variation in resource variability but cannot account for the dynamic and short-term responses to fluctuations in water availability. Conversely, several plant physiological strategies have been identified along the mono-dimensional gradient of water availability in a given environment. According to a revised terminology, dehydration escape, dehydration avoidance, dehydration tolerance, dormancy, and desiccation tolerance are clearly distinguishable. Their sequential expression is expressed as water deficit increases while cavitation tolerance is proposed here to be a major hydraulic strategy underpinning adaptive responses to drought of vascular plants. This continuum of physiological strategies can be interpreted in the context of the ecological trade-off between water-acquisition vs. water-conservation, since growth maintenance is associated with fast water use under moderate drought while plant survival after growth cessation is associated with slow water use under severe drought. Consequently, the distinction between 'drought resistance' and 'drought survival', is emphasized as crucial to ensure a correct interpretation of plant strategies since 'knowing when not to grow' does not confer 'drought resistance' but may well enhance 'drought survival'. This framework proposal should improve cross-fertilization between disciplines to help tackle the increasing worldwide challenges that drought poses to plant adaptation.

Are winter and summer dormancy symmetrical seasonal adaptive strategies? The case of temperate herbaceous perennials.

BACKGROUND: Dormancy in higher plants is an adaptive response enabling plant survival during the harshest seasons and has been more explored in woody species than in herbaceous species. Nevertheless, winter and summer shoot meristem dormancy are adaptive strategies that could play a major role in enhancing seasonal stress tolerance and resilience of widespread herbaceous plant communities. SCOPE: This review outlines the symmetrical aspects of winter and summer dormancy in order to better understand plant adaptation to severe stress, and highlight research priorities in a changing climate. Seasonal dormancy is a good model to explore the growth-stress survival trade-off and unravel the relationships between growth potential and stress hardiness. Although photoperiod and temperature are known to play a crucial, though reversed, role in the induction and release of both types of dormancy, the thresholds and combined effects of these environmental factors remain to be identified. The biochemical compounds involved in induction or release in winter dormancy (abscisic acid, ethylene, sugars, cytokinins and gibberellins) could be a priority research focus for summer dormancy. To address these research priorities, herbaceous species, being more tractable than woody species, are excellent model plants for which both summer and winter dormancy have been clearly identified. CONCLUSIONS: Summer and winter dormancy, although responding to inverse conditions, share many characteristics. This analogous nature can facilitate research as well as lead to insight into plant adaptations to extreme conditions and the evolution of phenological patterns of species and communities under climate change. The development of phenotypes showing reduced winter and/or enhanced summer dormancy may be expected and could improve adaptation to less predictable environmental stresses correlated with future climates. To this end, it is suggested to explore the inter- and intraspecific genotypic variability of dormancy and its plasticity according to environmental conditions to contribute to predicting and mitigating global warming.

Summer dormancy, drought survival and functional resource acquisition strategies in California perennial grasses.

BACKGROUND AND AIMS: Evidence suggests drought severity is increasing due to climate change, but strategies promoting severe drought survival in perennial grasses have been seldom explored. This is particularly true of summer dormancy, an adaptation common in summer-dry Mediterranean-type climates. In addition, though theory predicts superior drought survival results in lower potential productivity, studies rarely measure both drought survival and growth under optimal conditions. METHODS: Physiological and functional ecological approaches were integrated to quantify interspecific variation in foliar and root traits in a suite of eight California perennial grass species. In a glasshouse experiment, summer dormancy, foliar functional trait variation, and seasonal growth and phenology under non-limiting water conditions and dehydration tolerance under progressive drought were quantified. In a second glasshouse study, root functional traits were quantified under non-limiting water conditions in rhizotrons. KEY RESULTS: Summer dormancy was associated with higher dehydration tolerance, and negatively associated with traits conferring dehydration avoidance. Species with greater summer dormancy were characterized by greater springtime productivity, earlier reproduction, and a shallow and fine root system, which are indicative of dehydration escape. Summer dormancy was associated with an acquisitive, competitive functional strategy in spring, and a conservative strategy in summer. CONCLUSIONS: Both the escape and acquisitive springtime strategies observed in summer dormant perennial taxa are typically associated with annual grasses. California grasslands were once dominated by perennial species, but have been overtaken by non-native Mediterranean annual grasses, which are expected to be further favoured by climate change. Owing to functional similarity with these exotic annuals, it is suggested that native summer dormant taxa may play an important ecological role in the future of both natural and restored California grasslands.

Elevated CO2 maintains grassland net carbon uptake under a future heat and drought extreme.

Extreme climatic events (ECEs) such as droughts and heat waves are predicted to increase in intensity and frequency and impact the terrestrial carbon balance. However, we lack direct experimental evidence of how the net carbon uptake of ecosystems is affected by ECEs under future elevated atmospheric CO2 concentrations (eCO2). Taking advantage of an advanced controlled environment facility for ecosystem research (Ecotron), we simulated eCO2 and extreme cooccurring heat and drought events as projected for the 2050s and analyzed their effects on the ecosystem-level carbon and water fluxes in a C3 grassland. Our results indicate that eCO2 not only slows down the decline of ecosystem carbon uptake during the ECE but also enhances its recovery after the ECE, as mediated by increases of root growth and plant nitrogen uptake induced by the ECE. These findings indicate that, in the predicted near future climate, eCO2 could mitigate the effects of extreme droughts and heat waves on ecosystem net carbon uptake.

Combining Drought Survival via Summer Dormancy and Annual Biomass Productivity in Dactylis glomerata L.

Under Mediterranean climates, the best strategy to produce rain-fed fodder crops is to develop perennial drought resistant varieties. Summer dormancy present in native germplasm has been shown to confer a high level of survival under severe drought. Nevertheless it has also been shown to be negatively correlated with annual biomass productivity. The aim of this study was to analyze the correlations between summer dormancy and annual biomass productivity related traits and to identify quantitative trait loci (QTL) for these traits in a progeny of a summer dormant cocksfoot parent (Kasbah) and a summer active parent (Medly). A total of 283 offspring and the parents were phenotyped for summer dormancy, plant growth rate (PGR) and heading date in Morocco and for maximum leaf elongation rate (LERm) in France. The individuals were genotyped with a total of 325 markers including 59 AFLP, 64 SSR, and 202 DArT markers. The offspring exhibited a large quantitative variation for all measured traits. Summer dormancy showed a negative correlation with both PGR (-0.34 p < 0.005) and LERm (-0.27 p < 0.005). However, genotypes with both a high level of summer dormancy and a high level of PGR were detected in the progeny. One genetic map per parent was built with a total length of 377 and 423 cM for Kasbah and Medly, respectively. Both different and co-localized QTL for summer dormancy and PGR were identified. These results demonstrate that it should be possible to create summer dormant cocksfoot varieties with a high annual biomass productivity.

What functional strategies drive drought survival and recovery of perennial species from upland grassland?

BACKGROUND AND AIMS: Extreme climatic events such as severe droughts are expected to increase with climate change and to limit grassland perennity. The present study aimed to characterize the adaptive responses by which temperate herbaceous grassland species resist, survive and recover from a severe drought and to explore the relationships between plant resource use and drought resistance strategies. METHODS: Monocultures of six native perennial species from upland grasslands and one Mediterranean drought-resistant cultivar were compared under semi-controlled and non-limiting rooting depth conditions. Above- and below-ground traits were measured under irrigation in spring and during drought in summer (50 d of withholding water) in order to characterize resource use and drought resistance strategies. Plants were then rehydrated and assessed for survival (after 15 d) and recovery (after 1 year). KEY RESULTS: Dehydration avoidance through water uptake was associated with species that had deep roots (>1·2 m) and high root mass (>4 kg m(-3)). Cell membrane stability ensuring dehydration tolerance of roots and meristems was positively correlated with fructan content and negatively correlated with sucrose content. Species that survived and recovered best combined high resource acquisition in spring (leaf elongation rate >9 mm d(-1) and rooting depth >1·2 m) with both high dehydration avoidance and tolerance strategies. CONCLUSIONS: Most of the native forage species, dominant in upland grassland, were able to survive and recover from extreme drought, but with various time lags. Overall the results suggest that the wide range of interspecific functional strategies for coping with drought may enhance the resilience of upland grassland plant communities under extreme drought events.

A fungal endophyte reinforces population adaptive differentiation in its host grass species.

Hereditary symbioses between fungal endophytes and grasses are relatively recent in the history of plant life. Given < 80 million yr of co-evolution, symbioses are likely to have impacted plant microevolutionary rather than macroevolutionary processes. Therefore, we investigated the microevolutionary role of the fungal endophyte Neotyphodium lolii in the adaptive differentiation of its host species Lolium perenne. Endophyte frequency in 22 natural L. perenne populations was established across a water availability gradient. Adaptive differentiation among five populations, and between symbiotic (S) and nonsymbiotic (NS) plants, was examined in a glasshouse experiment under nonlimiting and limiting water conditions. Genetic differentiation was subsequently assessed among populations, and between S and NS individuals, using 14 simple sequence repeats (SSR). Symbiosis frequencies were positively correlated to water availability. Adaptive population differentiation occurred following a trade-off between biomass production under nonlimiting water conditions and survivorship under water stress. Endophytic symbiosis increased plant survival in xeric populations, and reinforced competitiveness in mesic populations. No genetic difference was detected between S and NS plants within populations. Therefore, we conclude that the endophyte relationship is responsible for these effects. Local adaptation of the host plant, appears to be supported by the fungal endophyte.

Water deficit and induction of summer dormancy in perennial Mediterranean grasses.

BACKGROUND AND AIMS: Summer dormancy is a trait conferring superior drought survival in Mediterranean perennial grasses. As the respective roles of environmental factors and water deficit on induction of summer dormancy are unclear, the effect of intense drought were tested under contrasting day lengths in a range of forage and native grasses. METHODS: Plants of Poa bulbosa, Dactylis glomerata 'Kasbah' and Lolium arundinaceum 'Flecha' were grown in pots (a) from winter to summer in a glasshouse and subjected to either an early or a late-spring drought period followed by a summer water deficit and (b) in controlled conditions, with long days (LD, 16 h) or short days (SD, 9 h) and either full irrigation or water deficit followed by rehydration. Leaf elongation, senescence of aerial tissues and dehydration of basal tissues were measured to assess dormancy. Endogenous abscisic acid (ABA) in basal tissues was determined by monoclonal immunoassay analysis. KEY RESULTS: Even under irrigation, cessation of leaf elongation, senescence of lamina and relative dehydration of basal tissues were triggered only by a day length longer than 13 h 30 min (late spring and LD) in plants of Poa bulbosa and Dactylis glomerata 'Kasbah' which exhibit complete dormancy. Plants of Lolium arundinaceum 'Flecha' maintained leaf growth under irrigation irrespective of the day length since its dormancy is incomplete. ABA concentrations were not higher during late-spring drought than early, and could not be associated with spring dormancy induction. In summer, ABA concentration in bulbs of the desiccation-tolerant Poa were greater than in basal tissues of other species. CONCLUSIONS: The results of both experiments tend to invalidate the hypothesis that water deficit has a role in early summer-dormancy induction in the range of tested grasses. However, a late-spring drought tends to increase plant senescence and ABA accumulation in basal tissues of forage grasses which could enhance summer drought survival.

Summer dormancy in perennial temperate grasses.

BACKGROUND AND AIMS: Dormancy has been extensively studied in plants which experience severe winter conditions but much less so in perennial herbaceous plants that must survive summer drought. This paper reviews the current knowledge on summer dormancy in both native and cultivated perennial temperate grasses originating from the Mediterranean Basin, and presents a unified terminology to describe this trait. SCOPE: Under severe drought, it is difficult to separate the responses by which plants avoid and tolerate dehydration from those associated with the expression of summer dormancy. Consequently, this type of endogenous (endo-) dormancy can be tested only in plants that are not subjected to moisture deficit. Summer dormancy can be defined by four criteria, one of which is considered optional: (1) reduction or cessation of leaf production and expansion; (2) senescence of mature foliage; (3) dehydration of surviving organs; and (4, optional) formation of resting organs. The proposed terminology recognizes two levels of summer dormancy: (a) complete dormancy, when cessation of growth is associated with full senescence of foliage and induced dehydration of leaf bases; and (b) incomplete dormancy, when leaf growth is partially inhibited and is associated with moderate levels of foliage senescence. Summer dormancy is expressed under increasing photoperiod and temperature. It is under hormonal control and usually associated with flowering and a reduction in metabolic activity in meristematic tissues. Dehydration tolerance and dormancy are independent phenomena and differ from the adaptations of resurrection plants. CONCLUSIONS: Summer dormancy has been correlated with superior survival after severe and repeated summer drought in a large range of perennial grasses. In the face of increasing aridity, this trait could be used in the development of cultivars that are able to meet agronomic and environmental goals. It is therefore important to have a better understanding of the genetic and environmental control of summer dormancy.

Seedling survival under drought differs between an annual (Hordeum vulgare) and a perennial grass (Dactylis glomerata).

• In semiarid areas, populations of cereal crop plants can be reduced by severe drought occurring during the vegetative stage, leading to subsequent serious yield reduction. This study aimed to analyse and compare survival and dehydration tolerance in two contrasting barley (Hordeum vulgare) cultivars (cvs) and in a drought resistant perennial grass, cocksfoot (Dactylis glomerata). • In pot experiments, seedlings were subjected to intensifying droughts for analysis of adaptive responses and survival following a range of final soil water potentials (SWP). • Survival rates were 100% in cocksfoot and 32% in the drought resistant barley cv. Tadmor at a SWP of -2.5 MPa. In cocksfoot, lamina water potentials decreased earlier and water content in enclosed leaf bases stabilized at low SWP. This strategy of dehydration tolerance was associated with high survival. • At moderate water deficit, barley cv. Tadmor exhibited a 5-fold greater osmotic adjustment in lamina, lower senescence and higher accumulation of dehydrins in enclosed leaf bases than the drought sensitive cv. Plaisant. However, under severe drought, water content in enclosed leaf bases declined constantly and was associated with similar mortality in both cvs. Dehydration tolerance was low in barley and not correlated with dehydrin accumulation or resistance to moderate drought.

Drought survival, summer dormancy and dehydrin accumulation in contrasting cultivars of Dactylis glomerata.

To study survival under prolonged and severe drought in the perennial grass Dactylis glomerata we compared dormant, resistant and sensitive cultivars (cvs.) in both field and glasshouse experiments. Water status, membrane stability and expression of dehydrins were assessed in the immature leaf bases, which are the last surviving organs. Analysis of leaf elongation and senescence of aerial tissues showed that dormancy was exhibited by the potentially dormant cultivar (cv.) only in the field. This cultivar exhibited a high survival rate, similar levels of dehydration and expression of a low-molecular weight (22-24 kDa) dehydrin in both drought and irrigated plants, whether fully dormant or not. At the same level of soil water deficit, there were no differences between the non-dormant drought resistant and drought sensitive cultivars in plant water status and membrane stability. However, the accumulation of dehydrins as drought progressed was markedly different between these cultivars and was associated with their contrasting survival. The possible role of the major low-molecular dehydrins in maintenance of cell integrity under dehydration is discussed with reference to both summer dormancy and survival under severe drought.

Survival and recovery of perennial forage grasses under prolonged Mediterranean drought: I. Growth, death, water relations and solute content in herbage and stubble.

Swards of Dactylis glomerata cultivars (cvs) KM2 and Lutetia and of Lolium perenne cvs Aurora and Vigor were grown under full irrigation or prolonged summer drought (80 d) in a field experiment in the South of France. After irrigation was withheld, leaf extension rates of all cvs fell by 90% within 9-12 d, and rapid scorching of laminae followed. Tiller mortality at the end of the drought was very different in the cocksfoot cvs (4% for KM2 and 76% for Lutetia) and intermediate (41%) for both ryegrass cvs. Following re-watering, rates of herbage regrowth were closely correlated with tiller survival. Measured minerals contributed c. 0·52 MPa to osmotic potential in all treatments, whereas water-soluble carbohydrates (WSC) contributed 0·25 MPa under irrigation and 0·46 MPa during drought. There was no systematic difference between the two species for summer survival under severe drought, but large differences between the cocksfoot cvs. The traits most strongly associated with superior survival were: (a) a deep root system and greater water uptake at depth; (b) low water and osmotic potentials in surviving laminae, i.e. better tolerance to dehydration; (c) large pool-size of WSC reserves (fructans having degree of polymerization >4) in entire tiller bases (stubble); (d) low accumulation of proline in stubble; (e) rapid nitrogen uptake after rewatering.