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.

In many regions of the world, such as Southern Europe and most Mediterranean areas, the frequency and magnitude of droughts and heat waves are expected to increase under global warming and will challenge the sustainability of both native and sown grasslands. To analyze the adaptive strategies of species, genotypes and cultivars, we aim both (i) to understand the composition and functioning of natural grasslands and (ii) to propose ideotypes of cultivars and optimal composition for mixtures of species/genotypes under water deficit and high temperatures. This review presents a conceptual framework to analyze adaptive responses of perennial herbaceous species, starting from resistance to moderate drought with growth maintenance (dehydration avoidance and tolerance of lamina) to growth cessation and survival of plants under severe stress (dehydration avoidance and tolerance of meristems). The most discriminating functional traits vary according to these contrasting strategies because of a trade-off between resistance to moderate moisture deficit and survival of intense drought. Consequently it is crucial to measure the traits of interest in the right organs and as a function of soil water use, in order to avoid misleading interpretations of plant responses. Furthermore, collaboration between ecologists, eco-physiologists, and agronomists is required to study the combination of plant strategies in natural grasslands as only this will provide the necessary rules for species and cultivars or ecotypes assemblage. This 'agro-ecological' approach aims to identify and enhance functional complementarity and limit competition within the multi-specific or multi-genotypic material associated in mixtures since using plant biodiversity should contribute to improving grassland resistance and resilience. (Résumé d'auteur

Due to the shortage of information on summer dormancy in tall fescue (Festuca arundinacea, syn. Lolium arundinaceum), we tested the response of 2 cultivars of differing dormancy expression and growth stage to a range of summer moisture conditions, including full irrigation, drought, and a simulated mid-summer storm and analysed whether traits associated with summer dormancy conferred better survival under severe field drought. Autumn-sown reproductive and younger, spring-sown plants of 2 cultivars, claimed to exhibit contrasting summer dormancy, were established and then tested in summer 2002 under either long drought, drought + simulated mid-summer storm, or full irrigation. The autumn-sown reproductive plants of cv. Flecha exhibited traits that can be associated with partial summer dormancy since under summer irrigation they reduced aerial growth significantly and exhibited earlier herbage senescence. Moreover, cv. Flecha used 35% less soil water over the first summer. However, the water status of leaf bases of young vegetative tillers of both cultivars was similar under irrigation and also throughout most of the drought (leaf potential and water content maintained over −4 MPa and at approx. 1 g H 2 O/g DM, respectively). The summer-active cv. Demeter did not stop leaf elongation even in drought and produced twice as much biomass as Flecha under irrigation. Cultivar Demeter responded to the simulated storm with a decline in dehydrin expression in leaf bases, whereas no decline occurred in Flecha, presumably because it remained partially dormant. The younger, spring-sown swards of both cultivars had similar biomass production under summer irrigation but whereas Demeter regrew in response to the simulated storm, cv. Flecha did not, indicating that dormancy, although only partially expressed, was reinforced by summer drought. In all trials, cv. Flecha out-yielded Demeter in autumn regrowth. In particular, the severe drought in 2003 caused a 25% loss of the basal cover in cv. Demeter, whereas Flecha fully maintained its sward allowing it to produce a higher post-drought autumn yield. This work links summer dormancy with higher persistence over long, dry summers.

The search to improve drought survival in temperate perennial grasses has led to a renewed interest in summer dormancy and how to quantify it. This endogenously controlled trait, found in some temperate perennial grasses, is associated with drought that normally occurs in summer. While cessation of leaf growth and senescence of herbage occurs in all grasses in response to drought, it is under summer irrigation that these same responses are observed only in summer-dormant germplasm and hence the trait can be identified in germplasm. Across the spectrum from completely summer-dormant to non-dormant, there is a range of expression. Our objective here is to highlight differences in characteristics of indices which measure summer dormancy and to identify aspects for incorporation into a superior index for use in measuring this trait. The experimental program comprised three field trials that compared 6 cultivars and a fourth that assessed a larger group of 12 cultivars of the same three species, cocksfoot (Dactylis glomerata L.), tall fescue (Festuca arundinacea Schreb.), and phalaris (Phalaris aquatica L.). Seasonal herbage yield and foliage senescence were measured under three summer watering regimes: complete drought, mid-summer storm, and full irrigation at Mauguio, France. Different indices were calculated to compare against the approach which evaluates senescence under drought. The key outcomes are as follows. (1) The assessment of summer dormancy needs to be viewed as the plant response to a period of non-limiting water supply over summer. It makes little difference whether this is produced by full summer irrigation or a mid-summer simulated storm after a drought. Assessment of this trait under conditions of unbroken drought is discouraged because it can result in false scores. (2) The determination of summer dormancy intensity under full summer irrigation is most appropriate for the intensive study of the dynamics of dormancy expression over the entire summer. A simulated mid-summer storm within a drought gives an instantaneous view of dormancy intensity at a specific observation date and may be well adapted to the requirements of plant breeding. These methods are complementary. (3) Summer dormancy intensity can be assessed either by measuring herbage production or by a visual assessment of the level of herbage senescence. (4) An index of summer dormancy based on comparing irrigated summer herbage yield of any cultivar with that of a high, summer-yielding, non-dormant control cultivar was able to provide a reliable score of dormancy intensity. This index functions across a range of cultivars and species of perennial grasses. Further refinement of the index is needed to identify ‘standard’ high and low summer-dormant populations.

A series of trials to increase understanding of the summer dormancy trait in Dactylis glomerata was conducted. Autumn-sown reproductive and younger, spring-sown plants of 2 drought-resistant cultivars, contrasting for summer dormancy, were established and then tested in summer 2002 under long drought, drought + midsummer storm, or full irrigation. The autumn-sown reproductive plants of cv. Kasbah were summer dormant under all moisture regimes and exhibited the characteristic traits including growth cessation, rapid herbage senescence, and dehydration of surviving organs (−6.7 MPa). Cultivar Kasbah used 8% less soil water over the summer and also began to rehydrate its leaf bases from conserved soil water before the drought broke. The non-dormant cv. Medly grew for 10 days longer under drought and whenever moisture was applied; Medly also responded to the storm with a decline in dehydrin expression in leaf bases, whereas no decline occurred in Kasbah, presumably because it remained dormant and therefore much drier. The irrigated, younger, spring-sown swards of cv. Kasbah had restrained growth and produced only about 25% of the herbage of cv. Medly. Drought reduced activity and growth of young plants of both cultivars, but whereas Medly regrew in response to the storm, cv. Kasbah did not, indicating that dormancy, although only partially expressed after spring sowing, was reinforced by summer drought. A longer drought in 2003 caused a 22% loss of the basal cover in cv. Medly, whereas Kasbah fully maintained its sward and therefore produced a higher post-drought autumn yield. This work confirms summer dormancy as a powerful trait for improving persistence over long, dry summers.

The three cool-season perennial forage grasses cocksfoot/orchardgrass, Dactylis glomerata L., tall fescue, Festuca arundinacea Schreb. syn. Lolium arundinaceum (Schreb.) Darbysh., and phalaris/harding grass, Phalaris aquatica L., are of major economic and ecological importance in regions with summer-dry environments. This review considers the constraints that these species are likely to experience under current and predicted increase of droughts due to climate change scenarios in south-eastern Australia, the southern Great Plains of USA and the Western Mediterranean Basin. The review identifies research required to maximise the development and use of C3 cool-season grasses with enhanced resilience to drought while considering the concern of some regulators that these grasses may be potential weeds. The state of knowledge of factors influencing plant drought survival and therefore recovery after stress and long-term persistence is discussed in the light of adaptive strategies. The major research needs identified to enhance traits conferring drought survival include (1) increasing the depth and density of grass root systems to strengthen dehydration avoidance; (2) exploring the biochemical, molecular and hydraulic bases of dehydration tolerance and improving techniques to measure this trait; (3) breaking the trade-off between summer dormancy and forage yield potential and improving understanding of environmental, biochemical and genetic controls over summer dormancy; (4) identifying non-toxic endophyte strains compatible with summer-dormant cultivars of tall fescue to enhance drought survival; and (5) enhancing seed production capability of new cultivars as well as the development of agronomic management packages for promoting stable mixtures combining perennial grasses and legumes. The weed potential of newly introduced summer-dormant cultivars is concluded to be minor. The research directions proposed here should improve pasture grass resilience and forage crop sustainability in Mediterranean and temperate summer-dry environments under the future drier and warmer conditions associated with climate change.

Dual-purpose cereals have been important for increasing the flexibility and profitability of mixed farming enterprises in southern Australia, providing winter feed when pasture dry matter production is low, and then recovering to produce grain. A perennial dual-purpose cereal could confer additional economic and environmental benefits. We establish that, at the end of a second growth season, selected perennial cereals were able to achieve up to 10-fold greater below-ground biomass than a resown annual wheat. We review and expand the data on available, diverse, perennial, wheat-derived germplasm, confirming that perenniality is achievable but that further improvements are essential through targeted breeding.Although not yet commercially deployable, the grain yields and dry matter production of the best performing lines approach the benchmarks predicted to achieve profitability. On reviewing the genomic composition of the most promising wheatderived perennials, we conclude that the best near-term prospect of a productive breeding program for a perennial, wheatderived cereal will utilise a diploid, perennial donor species, and the most promising one thus far is Thinopyrum elongatum. Furthermore, the breeding should be aimed at complete wheat-Th. elongatum amphiploids, a hybrid synthetic crop analogous to triticale. We advocate the generation of many primary amphiploids involving a diversity of Th. elongatum accessions and a diversity of adapted annual wheat cultivars. Primary perennial amphiploids would be inter-crossed and advanced with heavy, early-generation selection for traits such as semi-dwarf plant height, non-shattering heads, large seed size and good self-fertility, followed by later generation selection for robust perenniality, days to flowering, grain yield, forage yield, stability of grain yield across seasons, and disease resistance.

Perennial grasslands provide numerous agroenvironmental benefits due to continuous soil cover. In Mediterranean areas, chronic summer drought is expected to increase as a result of climate changes. Plant adaptations that protect meristematic tissues include dehydration delay, dehydration tolerance, and summer dormancy. Summer dormancy can only be reliably tested in plants not subjected to water deficit. Under summer irrigation, complete dormancy is manifested by cessation of growth in association with full senescence of foliage and induced dehydration of leaf bases. Incomplete dormancy occurs when leaf growth is partially constrained and associated with moderate levels of foliage senescence. Summer dormancy is under hormonal control and is induced under increasing photoperiod and temperature. Recent results show that drought cannot induce summer dormancy under early‐spring short days, although a water deficit under late‐spring long days reinforces it and could enhance drought survival. Dehydration tolerance and dormancy are independent phenomena. Summer dormancy has been correlated with superior survival after severe summer droughts in many perennial grass species. This trait has potential for improving cultivars able to meet agronomic and environmental goals.