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.Key words: Poa bulbosa, Dactylis glomerata, Lolium arundinaceum, abscisic acid, water deficit, photoperiod, summer dormancy, perennial grass     

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.     

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.     

Regulation of summer dormancy by water deficit and ABA in Poa bulbosa ecotypes

BACKGROUND AND AIMS: Survival of many herbaceous species in Mediterranean habitats during the dry, hot summer depends on the induction of summer dormancy by changes in environmental conditions during the transition between the winter (growth) season to the summer (resting) season, i.e. longer days, increasing temperature and drought. In Poa bulbosa, a perennial geophytic grass, summer dormancy is induced by long days, and the induction is enhanced by high temperature. Here the induction of summer dormancy in a Mediterranean perennial grass by water deficit under non-inductive photoperiodic conditions is reported for the first time. METHODS: Plants grown under 22/16 degrees C and non-inductive short-day (9 h, SD) were subjected to water deficit (WD), applied as cycles of reduced irrigation, or sprayed with ABA solutions. They were compared with plants in which dormancy was induced by transfer from SD to inductive long-day (16 h, LD). Responses of two contrasting ecotypes, from arid and mesic habitats were compared. Dormancy relaxation in bulbs from these ecotypes and treatments was studied by comparing sprouting capacity in a wet substrate at 10 degrees C of freshly harvested bulbs to that of dry-stored bulbs at 40 degrees C. Endogenous ABA in the bulbs was determined by monoclonal immunoassay analysis. KEY RESULTS: Dormancy was induced by WD and by ABA application in plants growing under non-inductive SD. Dormancy induction by WD was associated with increased levels of ABA. Bulbs were initially deeply dormant and their sprouting capacity was very low, as in plants in which dormancy was induced by LD. Dormancy was released after 2 months dry storage at 40 degrees C in all treatments. ABA levels were not affected by dormancy relaxation. CONCLUSIONS: Summer dormancy in P. bulbosa can be induced by two alternative and probably additive pathways: (1) photoperiodic induction by long-days, and (2) water deficit. Increased levels of endogenous ABA are involved in both pathways.     

Savannah woodland vegetation in the South‐East District of South Australia: the influence of evaporative aerodynamics on the foliage structure of the understorey invaded by introduced annuals

Abstract Evaporative aerodynamics determine the foliage projective cover of the understorey of perennial tussock grasses and associated perennial herbs in the savannah woodland dominated by Eucalyptus camalduknsis on gleyed podsolic soils in the Mediterranean climate of the South‐East District of South Australia. By the mid 1940s, winter‐spring evapotranspiration from the ‘thin’ leaves (with low leaf specific weight) of introduced annual plants was depleting surface soil water and thus reducing the annual growth of the summer‐growing savannah understorey; perennial herbs between the tussock grasses were the first to succumb to this competition. During spring, the percentage of the ground covered by the savannah understorey was increased by 10% in the subhumid zone to 30% in the humid zone as the pre‐European perennial herbs between the tussock grasses were replaced by introduced annuals. Application of phosphatic fertilizer to the understorey increased the growth of introduced annuals, which formed a dense stratum during their winter‐spring growing season, increasing evapotranspiration and leading eventually to the extinction of the native perennial grasses. When the savannah understorey, invaded by introduced annuals in the mid‐1940s, was converted to improved pasture, the percentage of ground covered by the seasonal foliage was increased by 20–30%; 100% coverage of overlapping foliage resulted in the humid zone.     

Abscisic acid involvement in the induction of summer‐dormancy in Poa bulbosa, a grass geophyte

Summer‐dormancy occurs in geophytes that inhabit regions with a Mediterranean climate (mild, rainy winters and hot, dry summers). The environmental control of summer‐dormancy and the involvement of phytohormones in its induction have been little studied. Poa bulbosa L. is a perennial grass geophyte in which summer‐dormancy is induced by long days and by high temperature. Prolonged treatment with ABA (0.1‐1.0 m M ) under non‐inductive 8‐h short days (SD) resulted in cessation of leaf and tiller production and in the development of typical features of dormancy: bulbing at the base of the tillers and leaf senescence. Short‐term applications of ABA had similar effects but dormancy was transient, i.e. after a short while, leaf growth from the formed bulbs was resumed. ABA treatment of plants growing under an inductive 16‐h photoperiod (LD) enhanced the onset of dormancy. Endogenous levels of ABA in leaf blades and at the tiller base (where the bulb develops) increased markedly after the plants were transferred from SD to LD. This increase was greater in the tiller base, and concomitant with bulb maturation. High temperature (27/22 vs 22/17°C) accelerated both bulb development and ABA accumulation in leaf blades.
These results suggest that ABA plays a key role in the photoperiodic induction and development of summer‐dormancy in P. bulbosa .     

Tradeoffs between functional strategies for resource-use and drought-survival in Mediterranean rangeland species

In environments where light is not a limiting resource such as rangelands and grasslands, there is much disagreement regarding the benefits provided by rapid light capture during the growing season and the species’ ability to withstand drought during the dry period. In this study, we selected four perennial herbaceous species with contrasting resource-use strategies (acquisitive versus conservative), which were transplanted as monocultures into PVC pots to evaluate their species-specific responses to drought. The two main strategies of drought-survival (avoidance versus tolerance) were driven by distinct underlying mechanisms that allow the plant to delay or tolerate water deficit in leaves. On the one hand, plants that produced reduced leaves with lower surface area:mass ratio (lower SLA) exhibited higher values of leaf water potential (LWP) and leaf relative water content (LRWC), which could be associated to a higher ability to delay tissue dehydration in enlarged leaves. Regarding the below-ground compartment, dehydration avoidance was promoted by prolonged elongation rates of thinner roots that allow the plant to increase water uptake and accessibility during the dry period. On the other hand, dehydration tolerance was positively related with progressive foliage senescence under water deficit, which probably favored a longer survival of meristematic basal tissues. The results presented in this study suggest the existence of a trade-off between the traits favoring rapid light-acquisition and those enhancing the ability to delay leaf dehydration. Thus, the species related most closely with a resource-acquisition strategy (Bromus erectus and Potentilla neumanniana) could be considered less efficient to delay leaf dehydration than the others (Carex humilis and Festuca christiani-bernardii), as indicated by their lower values of leaf water potential (LWP) and leaf relative water content (LRWC) under identical conditions of water deficit. Our findings support evidence that there is not a single strategy to effectively cope with drought and reveal the diversity of adaptive mechanisms among coexisting species.     

Seasonal patterns of growth, dehydrins and water-soluble carbohydrates in genotypes of Dactylis glomerata varying in summer dormancy

• Background and Aims Summer dormancy in perennial grasseshas been studied inadequately, despite its potential to enhanceplant survival and persistence in Mediterranean areas. The aimof the present work was to characterize summer dormancy anddehydration tolerance in two cultivars of Dactylis glomerata(dormant ‘Kasbah’, non-dormant ‘Oasis’)and their hybrid using physiological indicators associated withthese traits. • Methods Dehydration tolerance was assessed in a glasshouseexperiment, while seasonal metabolic changes which produce putativeprotectants for drought, such as carbohydrates and dehydrinsthat might be associated with summer dormancy, were analysedin the field. • Key Results The genotypes differed in their ability tosurvive increasing soil water deficit: lethal soil water potential(_s) was –3·4 MPa for ‘Kasbah’ (althoughnon-dormant), –1·3 MPa for ‘Oasis’,and –1·6 MPa for their hybrid. In contrast, lethalwater content of apices was similar for all genotypes (approx.0·45 g H₂O g d. wt^–1), and hence the greater survivalof ‘Kasbah’ can be ascribed to better drought avoidancerather than dehydration tolerance. In autumn-sown plants, ‘Kasbah’had greatest dormancy, the hybrid was intermediate and ‘Oasis’had none. The more dormant the genotype, the lower the metabolicactivity during summer, and the earlier the activity declinedin spring. Decreased monosaccharide content was an early indicatorof dormancy induction. Accumulation of dehydrins did not correlatewith stress tolerance, but dehydrin content was a function ofthe water status of the tissues, irrespective of the soil moisture.A protein of approx. 55 kDa occurred in leaf bases of the mostdormant cultivar even in winter. • Conclusions Drought avoidance and summer dormancy arecorrelated but can be independently expressed. These traitsare heritable, allowing selection in breeding programmes.     

Involvement of the ethylene response pathway in dormancy induction in chrysanthemum

Temperature plays a significant role in the annual cycling between growth and dormancy of the herbaceous perennial chrysanthemum (Chrysanthemum morifolium Ramat.). After exposure to high summer temperatures, cool temperature triggers dormancy. The cessation of flowering and rosette formation by the cessation of elongation are characteristic of dormant plants, and can be stimulated by exogenous ethylene. Thus, the ethylene response pathway may be involved in temperature-induced dormancy of chrysanthemum. Transgenic chrysanthemums expressing a mutated ethylene receptor gene were used to assess this involvement. The transgenic lines showed reduced ethylene sensitivity: ethylene causes leaf yellowing in wild-type chrysanthemums, but leaves remained green in the transgenic lines. Extension growth and flowering of wild-type and transgenic lines varied between temperatures: at 20 degrees C, the transgenic lines showed the same stem elongation and flowering as the wild type; at cooler temperatures, the wild type formed rosettes with an inability to flower and entered dormancy, but some transgenic lines continued to elongate and flower. This supports the involvement of the ethylene response pathway in the temperature-induced dormancy of chrysanthemum. At the highest dosage of ethephon, an ethylene-releasing agent, wild-type plants formed rosettes with an inability to flower and became dormant, but one transgenic line did not. This confirms that dormancy is induced via the ethylene response pathway.     

It is risky out there: the costs of emergence and the benefits of prolonged dormancy

Prolonged dormancy is a stage in herbaceous perennial plants in which some individuals remain alive below ground for one or more growing seasons instead of emerging. Prolonged dormancy is puzzling, because foregoing opportunities for growth and reproduction seems costly. However, studies have shown that it buffers plants from the negative consequences associated with environmental stochasticity, suggesting that dormancy is a beneficial strategy to avoid the risks of stress above ground. If so, emergence during unfavorable conditions should have significant costs. Here, we test the hypothesis that emergence during times of stress has negative demographic consequences in a native perennial forb, Astragalus scaphoides, and investigate the potential underlying physiological mechanisms. We measured plant responses to a severe seasonal drought and an experimental defoliation to ask: (1) How do emergent plants respond to above-ground stress? (2) Do these responses have negative demographic consequences? and (3) Based on these responses, does stress increase the risk of emergence? Plants showed remarkable physiological tolerance to stress in the short term: high temperatures and low moisture did not have a strong effect on photosynthesis rates, and neither drought nor defoliation significantly impacted stored resources. However, stress did result in demographic costs for emergent plants relative to plants experiencing more favorable conditions. Drought resulted in decreased flowering probabilities relative to the long-term average and defoliation significantly increased mortality rates. These results demonstrate that the risk of emerging and experiencing stress entails considerable costs, supporting the hypothesis that prolonged dormancy is a beneficial strategy to avoid such risk.     

California perennial grasses are physiologically distinct from both Mediterranean annual and perennial grasses

In the Central Valley of California, native perennial grass species have been largely replaced by Eurasian annual species, while in many parts of the Mediterranean Basin native perennial grasses continue to dominate, even on disturbed or degraded sites. We assessed whether differences in summer rainfall patterns have lead to the development of different plant-water strategies between grasses from these two regions. We compared six measures of plant-water physiology for three guilds of grasses: California perennial grasses, Mediterranean perennial grasses, and Mediterranean annual grasses. Discriminant analysis distinguished between the three guilds; Mediterranean perennial grasses were characterized by a more conservative water-relations physiology than Mediterranean annual grasses, whereas California perennial grasses were in some ways intermediate between the two Mediterranean grass guilds. For individual traits, California perennial grasses were either intermediate or more like Mediterranean annuals than Mediterranean perennials. Our results suggest California perennials are more drought tolerant than Mediterranean annuals but less drought tolerant than Mediterranean perennials, despite the fact that California??s Central Valley has a more intense summer drought than the Mediterranean Basin. These patterns may help explain why Mediterranean annuals, but not Mediterranean perennials, have been more successful invaders of interior California grasslands.     

Drought tolerance of juvenile and mature leaves of a deciduous dwarf shrub Vaccinium myrtillus L. in a boreal environment

The difference between drought tolerance of juvenile and mature leaves of the winter-deciduous dwarf shrub bilberry (Vaccinium myrtillus L.) from a northern boreal environment was investigated. It was hypothesised that mature leaves are more drought sensitive than juvenile leaves. Bilberry plants were allowed to dry out by excluding irrigation when leaves were at juvenile and mature stages. Tissue water content decreased at both phenological stages, but the response was more pronounced in the mature leaves. Anthocyanin concentrations increased as the tissue water content decreased, and again this occurred to a greater extent in the mature leaves. Chlorophyll concentrations decreased only marginally at the juvenile stage, while the decrease was significant in the mature leaves. Chlorophyll degradation was enhanced by drought stress. Soluble proteins decreased and protein oxidation increased in the mature leaves, and degradation of oxidised proteins increased in the drought-stressed plants. The results suggest that leaves of bilberry are more sensitive to drought stress at the mature stage, and that drought stress accelerates senescence at the mature stage. The significance of the results is that dry periods during the juvenility of leaves are not as detrimental as they may be later in summer. In addition, the strategy of a winter-deciduous plant is obviously to protect its perennial parts from severe drought by accelerated leaf senescence at the mature stage. Therefore, the deciduous life form may provide an excellent adaptation against drought also in northern ecosystems. The role of anthocyanins in photoprotection under drought stress is also discussed.     

Contribution of arbuscular mycorrhizal fungi (AMF) to the adaptations exhibited by the deciduous shrub Anthyllis cytisoides L. under water deficit

Anthyllis cytisoides L. is highly colonized by arbuscular mycorrhizal fungi (AMF) and behaves as a drought-avoider species in the field. Our objectives were: (1) to study the response of A. cytisoides when exposed to moderate (acclimation) or severe (peak) drought and subsequent rewatering under nursery conditions; and (2) to verify if AMF improved the adaptation of A. cytisoides to stress. The soil compactness in drought-acclimated treatments increased four times compared with that of well-watered controls, which could reinforce the effects of water deficit on plant physiology. Photosynthetic rates decreased by around 50% and 70% and leaf conductance decreased by 40% and 50% in drought-acclimated non-mycorrhizal and mycorrhizal plants, respectively. Peak drought limited plant growth, accelerated leaf senescence and induced the conversion of starch into soluble sugars in the leaves of stressed plants. The accumulation of sugars could contribute to a decrease in water potential in order to achieve the required tension to let water move from soil to shoot. Mycorrhizal plants showed a two-fold higher chlorotic leaf biomass than non-mycorrhizal plants under severe drought. Moreover, mycorrhizal A. cytisoides showed enhanced epicuticular waxes on the surfaces of the remaining green leaves. Increased leaf senescence, together with wax deposition, could reduce whole plant transpiration, thus allowing mycorrhizal plants to maintain a higher leaf relative water content (50%) than non-mycorrhizal plants (35%). After drought recovery, leaf abscission in stressed mycorrhizal plants was 10 times greater than that in non-mycorrhizal plants. The results suggest that AMF conferred greater responsiveness of A. cytisoides to drought. Enhanced wax deposition and leaf senescence could be an ecological adaptation to cope with severe water deficit.     

Alteration of the phenology of leaf senescence and fall in winter deciduous species by climate change: effects on nutrient proficiency

Leaf senescence in winter deciduous species signals the transition from the active to the dormant stage. The purpose of leaf senescence is the recovery of nutrients before the leaves fall. Photoperiod and temperature are the main cues controlling leaf senescence in winter deciduous species, with water stress imposing an additional influence. Photoperiod exerts a strict control on leaf senescence at latitudes where winters are severe and temperature gains importance in the regulation as winters become less severe. On average, climatic warming will delay and drought will advance leaf senescence, but at varying degrees depending on the species. Warming and drought thus have opposite effects on the phenology of leaf senescence, and the impact of climate change will therefore depend on the relative importance of each factor in specific regions. Warming is not expected to have a strong impact on nutrient proficiency although a slower speed of leaf senescence induced by warming could facilitate a more efficient nutrient resorption. Nutrient resorption is less efficient when the leaves senesce prematurely as a consequence of water stress. The overall effects of climate change on nutrient resorption will depend on the contrasting effects of warming and drought. Changes in nutrient resorption and proficiency will impact production in the following year, at least in early spring, because the construction of new foliage relies almost exclusively on nutrients resorbed from foliage during the preceding leaf fall. Changes in the phenology of leaf senescence will thus impact carbon uptake, but also ecosystem nutrient cycling, especially if the changes are consequence of water stress.     

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.     

Within-generation and transgenerational plasticity in growth and regeneration of a subordinate annual grass in a rainfall experiment

Precipitation changes may induce shifts in plant species or life form dominance in ecosystems, making some previously subordinate species abundant. The plasticity of certain plant functional traits of these expanding subordinate species may be one possible mechanism behind their success. In this study, we tested if the subordinate winter annual grass Secale sylvestre shows plasticity in growth and reproduction in response to altered environment associated with field-scale rainfall manipulations (severe drought, moderate drought, and watering) in a semiarid grassland, and whether the maternal environment influences offspring germination or growth in a subsequent pot experiment. Compared to control plots, S. sylvestre plants grew 38% taller, and produced 32% more seeds in severe drought plots, while plants in watered plots were 17% shorter, and had 22% less seeds. Seed mass was greatest in severe drought plots. Plants growing in drought plots had offspring with enhanced juvenile shoot growth compared to the progeny whose mother plants grew in watered plots. These responses are most likely explained by the decreased cover of previously dominant perennial grasses in severe drought plots, which resulted in wetter soil compared to control and watered plots during the peak growth of S. sylvestre. We conclude that the plasticity of this subordinate annual species in response to changing environment may help to gain dominance with recurring droughts that suppress perennial grasses. Our results highlight that exploring both within-generation and transgenerational plasticity of subordinate species may lead to a better prediction of changes in plant species dominance under climate change.

     

The effects of temperature and drought on autumnal senescence and leaf shed in apple under warm,east mediterranean climate

Key message

Autumnal senescence of apple in a warm climate corresponds to accumulated degree-days beneath 22 °C. Summer drought delays senescence and enables replenishment of carbohydrate reserves. Recovery of the root system plays a key role.

Abstract

Autumnal senescence of apple (Malus domestica Borkh.), a deciduous, temperate climate species, is triggered by a rather abrupt temperature drop, down to the lower teens. Under the warmer, east Mediterranean climate of northern Israel, the temperature drop is gradual and much more moderate. Another characteristic of this climate is the complete lack of precipitation during summer. The aim of the present study was to elucidate the effects of summer drought on seasonal leaf senescence in a warm autumn. We hypothesized that summer drought delays senescence due to an increased demand for carbohydrates during autumn. The advent of autumnal senescence was followed for 3 years (2009–2011) on trees exposed to various levels of drought. Total canopy green area (effective leaf area, ELA) and hue angle were estimated periodically by means of image analysis, as a measure of leaf drop and autumnal color change. Photosynthesis, midday stem water potential, and roots’ non-structural carbohydrate contents were measured on several occasions. The time course of leaf drop followed the decline in air and soil temperatures. The rate of decline in ELA closely corresponded to accumulated degree-days beneath 22 °C in the soil, a much higher temperature threshold than previously reported for apple. Drought stress during the summer delayed leaf senescence even further, when compared with well-irrigated trees. Leaves maintained their photosynthetic functionality throughout autumn, until late December. The delayed senescence enabled replenishment of root carbohydrate reserves, which is critical for next year’s growth and fruiting. The eco-physiological significance of the findings is discussed.     

Potential growth and drought tolerance of eight desert grasses: lack of a trade-off?

Eight perennial C-4 grasses from the Jornada del Muerto Basin in southern New Mexico show five-fold differences in relative growth rates under well- watered conditions (RGR_max). In a controlled environment, we tested the hypothesis that there is an inverse relationship (trade-off) between RGR_max and the capacity of these species to tolerate drought. We examined both physiological (gas exchange) and morphological (biomass allocation, leaf properties) determinants of growth for these eight species under three steady-state drought treatments (none=control, moderate, and severe). When well watered, the grasses exhibited a large interspecific variation in growth, which was reflected in order-of-magnitude biomass differences after 5 weeks. The species had similar gas-exchange characteristics, but differed in all the measured allocation and morphological characteristics, namely tiller mass and number, root:shoot ratio, dry-matter content, and specific leaf area (SLA). Drought affected tillering, morphology, and allocation, and reduced growth by 50 and 68% (moderate and severe drought, respectively) compared to the well-watered controls. With the exception of SLA, none of these variables showed a significant species-by-treatment interaction. We calculated three indices of drought tolerance, defined as the ratio in final biomass between all the possible ”dry”/”wet” treatment pairs: severe/moderate, moderate/control, and severe/control. We found no significant correlation between these drought tolerance indices, on the one hand, and three indices of growth potential (greenhouse RGR_max, final biomass in the control treatment, and final:initial biomass ratio in controls), on the other. Based on these controlled-environment results, we hypothesize that the commonly reported correlation between plant growth potential and drought tolerance in the field may in some cases be explained by differential effects of plants on soil-water content rather than by differences in species responses to drought. Received: 10 March 1999 / Accepted: 8 November 1999     

Water conservation in<Emphasis Type="Italic"> Artemisia tridentata</Emphasis> through redistribution of precipitation

Water conservation is important for plants that maintain physiologically active foliage during prolonged periods of drought. A variety of mechanisms for water conservation exist including stomatal regulation, foliage loss, above- and below-ground allocation patterns, size of xylem vessels and leaf pubescence. Using the results of a field and simulation study with Artemisia tridentata in the Great Basin, USA, we propose an additional mechanism of water conservation that can be used by plants in arid and semi-arid environments following pulses of water availability. Precipitation redistributed more uniformly in the soil column by roots (hydraulic redistribution of water downward) slows the rate at which this water can subsequently be taken up by plants, thus prolonging water availability during periods of drought. By spreading out water more uniformly in the soil column at lower water potentials following precipitation events, water use is reduced due to lower soil conductivity. The greater remaining soil water and more uniform distribution result in higher plant predawn water potentials and transpiration rates later in the drought period. Simulation results indicate that plants can benefit during drought periods from water storage following both summer rain events (small summer pulses) and overwinter recharge (large spring pulse). This mechanism of water conservation may aid in sustaining active foliage, maintaining root-soil hydraulic connectivity, and increasing survival probability of plants which remain physiologically active during periods of drought.