Ecophysiology of two solar tracking desert winter annuals

Summary The seasonal course of water relations in field populations of two leaf solar tracking desert winter annuals was examined. Measurements were made of leaf movements in relation to leaf conductance and water potential. Malvastrum rotundifolium maintained solar tracking movements up to the wilting point of the plant (-4 MPa). Lupinus arizonicus altered its morphology through paraheliotropic leaf movements as leaf water potentials declined to-1.8 MPa. Diurnal patterns of leaf conductance showed marked seasonal trends, with gas exchange activity being restricted to early morning hours as water availability declined. Studies of potted plants showed that L. arizonicus was not able to alter its osmotic potential in response to drought, while M. rotundifolium underwent a 1.86 MPa reduction in osmotic potential. The significance of the two contrasting patterns is discussed in terms of observed plant distribution and origin.     

Stomatal response to drying soil in relation to changes in the xylem sap composition of Helianthus annuus. II. Stomatal sensitivity to abscisic acid imported from the xylem sap

Sunflower plants [Helianthus annuus L.) were subjected to soil drought. Leaf conductance declined with soil water content even when the shoot was kept turgid throughout the drying period. The concentration of abscisic acid in the xylem sap increased with decreasing soil water content. No general relation could be established between abscisic acid concentration in the xylem sap and leaf conductance due to marked differences in the sensitivity of leaf conductance of individual plants to abscisic acid from the xylem sap. The combination of these results with data from Gollan, Schurr & Schulze (1992, see pp. 551–559, this issue) reveals close connection of the effectiveness of abscisic acid as a root to shoot signal to the nutritional status of the plant.     

Non-structural carbohydrate dynamics associated with antecedent stem water potential and air temperature in a dominant desert shrub

Non-structural carbohydrates (NSCs) are necessary for plant growth and affected by plant water status, but the temporal dynamics of water stress impacts on NSC are not well understood. We evaluated how seasonal NSC concentrations varied with plant water status (predawn xylem water potential, Ψ) and air temperature (T) in the evergreen desert shrub Larrea tridentata. Aboveground sugar and starch concentrations were measured weekly or monthly for ~1.5 years on 6–12 shrubs simultaneously instrumented with automated stem psychrometers; leaf photosynthesis (A_net) was measured monthly for 1 year. Leaf sugar increased during the dry, premonsoon period, associated with lower Ψ (greater water stress) and high T. Leaf sugar accumulation coincided with declines in leaf starch and stem sugar, suggesting the prioritization of leaf sugar during low photosynthetic uptake. Leaf starch was strongly correlated with A_net and peaked during the spring and monsoon seasons, while stem starch remained relatively constant except for depletion during the monsoon. Recent photosynthate appeared sufficient to support spring growth, while monsoon growth required the remobilization of stem starch reserves. The coordinated responses of different NSC fractions to water status, photosynthesis, and growth demands suggest that NSCs serve multiple functions under extreme environmental conditions, including severe drought.     

Drought and changes in leaf orientation for two California chaparral shrubs: Ceanothus megacarpus and Ceanothus crassifolius

Summary Predawn xylem pressure potentials were measured on two California chaparral shrubs, Ceanothus megacarpus and Ceanothus crassifolius, throughout the winter and spring growing season and into the summer drought. On the days xylem pressure potentials were measured, leaf orientation measurements were made on a population of marked leaves from the same shrubs. Predawn xylem pressure potentials decreased from -0.1 MPa in both species to -7.8 and -6.6 MPa in C. megacarpus and C. crassifolius, respectively, between May and August, 1981. Leaf inclinations became more vertical during this period with the greatest change observed in C. crassifolius. This change in leaf inclination was reversible, and, in the late winter and early spring, one year old leaves became more horizontal. Leaf azimuths were random and did not change seasonally. Simulations of solar radiation interception indicated that the increase in leaf inclination associated with summer drought reduced the absorption of solar radiation in August by 6% for C. megacarpus and 20% for C. crassifolius. Standard leaf energy budget calculations suggest that steep leaf inclinations would result in slightly lower leaf temperatures and transpiration rates under summer conditions.     

Drought-induced changes in xylem pH, ionic composition, and ABA concentration act as early signals in field-grown maize (Zea mays L.).

Early signals potentially regulating leaf growth and stomatal aperture in field-grown maize (Zea mays L.) subjected to drought were investigated. Plants grown in a field lysimeter on two soil types were subjected to progressive drought during vegetative growth. Leaf ABA content, water status, extension rate, conductance, photosynthesis, nitrogen content, and xylem sap composition were measured daily. Maize responded similarly to progressive drought on both soil types. Effects on loam were less pronounced than on sand. Relative to fully-watered controls, xylem pH increased by about 0.2 units one day after withholding irrigation (DAWI) and conductivity decreased by about 0.25 mS cm(-1) 1-3 DAWI. Xylem nitrate, ammonium, and phosphate concentrations decreased by about 50% at 1-5 DAWI and potassium concentration decreased by about 50% at 7-8 DAWI. Xylem ABA concentration consistently increased by 45-70 pmol ml(-1) at 7 DAWI. Leaf extension rate decreased 5 DAWI, after the changes in xylem chemical composition had occurred. Leaf nitrogen significantly decreased 8-16 DAWI in droughted plants. Midday leaf water potential and photosynthesis were significantly decreased in droughted plants late in the drying period. Xylem nitrate concentration was the only ionic xylem sap component significantly correlated to increasing soil moisture deficit and decreasing leaf nitrogen concentration. Predawn leaf ABA content in droughted plants increased by 100-200 ng g(-1) dry weight at 7 DAWI coinciding with a decrease in stomatal conductance before any significant decrease in midday leaf water potential was observed. Based on the observed sequence, a chain of signal events is suggested eventually leading to stomatal closure and leaf surface reduction through interactive effects of reduced nitrogen supply and plant growth regulators under drought.     

Leaf and stem physiological responses to summer and winter extremes of woody species across temperate ecosystems

Winter cold limits temperate plant performance, as does summer water stress in drought‐prone ecosystems. The relative impact of seasonal extremes on plant performance has received considerable attention for individual systems. An integrated study compiling the existing literature was needed to identify overall trends. First, we conducted a meta‐analysis of the impacts of summer and winter on ecophysiology for three woody plant functional types (winter deciduous angiosperms, evergreen angiosperms and conifers), including data for 210 records from 75 studies of ecosystems with and without summer drought across the temperate zone. Second, we tested predictions by conducting a case study in a drought‐prone Mediterranean ecosystem subject to winter freezing. As indicators of physiological response of leaves and xylem to seasonal stress, we focused on stomatal conductance (g_s), percent loss of stem xylem hydraulic conductivity (PLC) and photochemical efficiency of photosystem II (F_v/F_m). Our meta‐analysis showed that in ecosystems without summer drought, g_s was higher during summer than winter. By contrast, in drought‐prone ecosystems many species maintained open stomata during winter, with potential strong consequences for plant carbon gain over the year. Further, PLC tended to increase and F_v/F_m to decrease from summer to winter for most functional types and ecosystems due to low temperatures. Overall, deciduous angiosperms were most sensitive to climatic stress. Leaf gas exchange and stem xylem hydraulics showed a coordinated seasonal response at ecosystems without summer drought. In our Mediterranean site subjected to winter freezing the species showed similar responses to those typically found for ecosystems without summer drought. We conclude that winter stress is most extreme for systems without summer drought and systems with summer drought and winter freezing, and less extreme for drought‐prone systems without freezing. In all cases the evergreen species show less pronounced seasonal responses in both leaves and stems than deciduous species.     

三种锦鸡儿属植物水力结构特征及其干旱适应策略

水分胁迫是干旱半干旱区限制植物生长的主要因素。以干旱半干旱区的3种锦鸡儿属植物为研究对象,从生态适应策略角度来分析3种锦鸡儿植物产生生态分离的原因。对三种锦鸡儿属植物茎干叶片的显微结构、生理功能(导水率、光合速率以及水分利用效率)进行测定,并统计了3种锦鸡儿植株的形态特征,如一、二级枝的直径、长度、末端叶面积。结果表明:三种锦鸡儿属植物都能形成较小的导管直径来适应旱生环境,但是在导水结构上又表现出一定的差异性。中间锦鸡儿的导管直径最小,次脉密度和最大净光合速率最大;柠条锦鸡儿的导管直径、叶片厚度和比叶重(LMA)最大。小叶锦鸡儿在导水率下降50%时的水势(P_(50))最大,水分胁迫时极易发生栓塞,但正是由于导管的栓塞降低了水分运输效率,使其在旱生环境中能够通过减少水分的供应来降低水分的丧失,从而保证自身生长的水分需求;而中间锦鸡儿则主要通过减小导管直径来适应旱生环境;柠条锦鸡儿的水分利用效率最高,抗栓塞能力最强,抗旱性最好,同时柠条锦鸡儿可以通过减少蒸腾面积来减少水分的丧失。植物的导管直径大小、叶片厚度、LMA、叶脉密度对植物导水速率、光合速率等生理功能都有一定的影响。     

Poplar vulnerability to xylem cavitation acclimates to drier soil conditions

Xylem vulnerability to cavitation differs between tree species according to their drought resistance, more xerophilous species being more resistant to xylem cavitation. Variability in xylem vulnerability to cavitation is also found within species, especially between in situ populations. The origin of this variability has not been clearly identified. Here we analyzed the response of xylem hydraulic traits of Populus tremula×Populus alba trees to three different soil water regimes. Stem xylem vulnerability was scored as the xylem water potential causing 12, 50 and 88% loss of conductivity (P₁₂, P₅₀ and P₈₈). Vulnerability to cavitation was found to acclimate to growing conditions under different levels of soil water content, with P₅₀ values of ?1.82, ?2.03 and ?2.45 MPa in well‐watered, moderately water‐stressed and severely water‐stressed poplars, respectively. The value of P₁₂, the xylem tension at which cavitation begins, was correlated with the lowest value of midday leaf water potential (ψm) experienced by each plant, the difference between the two parameters being approximately 0.5 MPa, consistent with the absence of any difference in embolism level between the different water treatments. These results support the hypothesis that vulnerability to cavitation is a critical trait for resistance to drought. The decrease in vulnerability to cavitation under growing conditions of soil drought was correlated with decreased vessel diameter, increased vessel wall thickness and a stronger bordered pit field (t/b)². The links between these parameters are discussed.     

Impact of long-term drought on xylem embolism and growth in Pinus halepensis Mill.

 The present study was carried out to elucidate the response mechanisms of 50-year-old Pinus halepensis Mill. trees to a long-term and severe drought. The amount of water available to trees was artificially restricted for 12 months by covering the soil with a plastic roof. Over the short term a direct and rapid impact of drought was evident on the water relations and gas exchanges of trees: as the soil dried out in the Spring, there was a concurrent decrease of predawn water potential; transpiration was strongly reduced by stomatal closure. Seasonal changes in the water volume fractions of twig and stem xylem were observed and interpreted as the result of cavitation and refilling in the xylem. When droughted trees recovered to a more favourable water status, refilling of embolized xylem was observed; twig predawn water potentials were still negative in the period when the embolism was reversed in the twig xylem. A few months after the removal of the covering, no differences in whole plant hydraulic resistance were observed between droughted and control trees. Needle and shoot elongation and stem radial growth were considerably reduced in droughted trees; no strategy of trees to allocate carbon preferentially to the stem conducting tissues was apparent throughout the experiment. An after-effect of the drought on growth was observed. Received: 4 August 1997 / Accepted: 1 October 1997     

Change in water loss regulation after canopy clearcut of a dominant shrub in Sahelian agrosystems, Guiera senegalensis J. F. Gmel

This paper analyzes the effect of the canopy age of Guiera senegalensis J.F. Gmel on water regulation processes and adaptative strategy to drought over a period of 2 years. The species is widespread in the agricultural Sahel. Before sowing, farmers cut back the shrubs to limit competition with crops. The stumps resprout after the millet harvest. Leaf water potential and stomatal conductance were measured in two fallows and in the two adjacent cultivated fields. Leaf transpiration rate and soil-to-leaf hydraulic conductance were deduced. The decrease in both stomatal and plant hydraulic conductance caused by seasonal drought was greater in mature shrubs than in current year resprouts. The decrease in predawn and midday leaf water potentials in response to seasonal drought was isohydrodynamic, and it was greater in mature shrubs, suggesting that current year resprouts are under less stress. In resprouts, the leaf transpiration rate stopped increasing beyond a hydraulic conductance threshold of 0.05 mol. m^?2 s^?1 MPa^?1. Vulnerability to cavitation was determined on segments of stems in the laboratory. The leaf water potential value at which stomatal closure occurred was ?2.99 ± 0.68 MPa, which corresponded to a 30 % loss in xylem conductivity. Thanks to its positive safety margin of 0.6 MPa, G. senegalensis can survive above this value. The observed strategy places G. senegalensis among the non-extreme xeric plants, leading us to suppose that this species will be vulnerable to the expected increase in regional drought.     

Effects of drought on nutrient and ABA transport in Ricinus communis

We studied the effects of variations of water flux through the plant, of diurnal variation of water flux, and of variation of vapour pressure deficit at the leaf on compensation pressure in the Passioura-type pressure chamber, the composition of the xylem sap and leaf conductance in Ricinus communis. The diurnal pattern of compensation pressure showed stress relaxation during the night hours, while stress increased during the day, when water limitation increased. Thus compensation pressure was a good measure of the momentary water status of the root throughout the day and during drought. The bulk soil water content at which predawn compensation pressure and abscisic acid concentration in the xylem sap increased and leaf conductance decreased, was high when the water usage of the plant was high. For all xylem sap constituents analysed, variations in concentrations during the day were larger than changes in mean concentrations with drought. Mean concentrations of phosphate and the pH of the xylem sap declined with drought, while nitrate concentration remained constant. When the measurement leaf was exposed to a different VPD from the rest of the plant, leaf conductance declined by 400mmol m^?2 s^?1 when compensation pressure increased by 1 MPa in all treatments. The compensation pressure needed to keep the shoot turgid, leaf conductance and the abscisic acid concentration in the xylem were linearly related. This was also the case when the highly dynamic development of stress was taken into account.     

Plant water uptake from soil through a vapor pathway

Water uptake from the soil via a vapor pathway was tested. Viburnum suspensum L. plants were divided into: (1) irrigated, (2) drought with vapor and (3) drought without vapor treatments. Each plant was placed into a larger bucket containing deuterium-labeled water as a vapor source (vapor treatment) or no water (drought and irrigation treatments). We also tested whether uptake via a vapor pathway could mitigate drought effects. Net CO₂ assimilation (A), transpiration (E) and stomatal conductance (gs) were measured daily until the first visible signs of stress. Soil water content, stem water potential (Ψ) and the stable hydrogen isotope ratio (δ²H) of soil and plant xylem water were then measured in all treatments. We show that water is taken up by plants through the vapor phase in dry soils. The δ²H values of the soil water in the vapor treatment were highly enriched compared to the background isotope ratios of the non-vapor exposed irrigated and drought treatments. Stem water δ²H values for the vapor treatment were significantly greater than those for irrigation and drought treatments not exposed to isotopically enriched vapor. In this experiment, movement of water to the plant via the vapor phase did not mitigate drought effects. A, E, plant Ψ and gs significantly decreased in the drought and vapor treatments relative to the controls, with no significant differences between vapor and drought treatments.     

Reproductive water supply is prioritized during drought in tomato

Reproductive success largely defines the fitness of plant species. Understanding how heat and drought affect plant reproduction is thus key to predicting future plant fitness under rising global temperatures. Recent work suggests reproductive tissues are highly vulnerable to water stress in perennial plants where reproductive sacrifice could preserve plant survival. However, most crop species are annuals where such a strategy would theoretically reduce fitness. We examined the reproductive strategy of tomato (Solanum lycopersicum var. Rheinlands Ruhm) to determine whether water supply to fruits is prioritized above vegetative tissues during drought. Using optical methods, we mapped xylem cavitation and tissue shrinkage in vegetative and reproductive organs during dehydration to determine the priority of water flow under acute water stress. Stems and peduncles of tomato showed significantly greater xylem cavitation resistance than leaves. This maintenance of intact water supply enabled tomato fruit to continue to expand during acute water stress, utilizing xylem water made available by tissue collapse and early cavitation of leaves. Here, tomato plants prioritize water supply to reproductive tissues, maintaining fruit development under drought conditions. These results emphasize the critical role of water transport in shaping life history and suggest a broad relevance of hydraulic prioritization in plant ecology.     

Ecophysiological responses of five emergent-wetland plants to diminished water supply: an experimental microcosm study

Freshwater wetlands are fundamentally tied to hydrology as they are often found along the boundaries between terrestrial uplands and open waters. Although wetland systems are frequently prone to extended periods of flooding and exposure, the degree of water deprivation may intensify during periods of low precipitation or drought. Therefore, the purpose of this study was to evaluate plant–water relations in five emergent macrophytes (Carex alata, Juncus effusus, Peltandra virginica, Saururus cernuus, and Justicia americana) to simulated drought conditions. Weekly evaluations of tissue water content and xylem water potential (free energy of water in xylem tissues) were conducted on plants grown in experimental microcosms over a 9-week period. Plant performance was also evaluated in each species by monitoring the changes in plant biomass, leaf area, and survival. The results indicate that J. effusus and P. virginica performed better in both flooded and moderately dry conditions, and plants that maintained higher water content in water logged soils (i.e., J. americana) were less tolerant to drying conditions. This study also illustrates the importance of periodic water withdrawal on plant performance. In general, plants that were subjected to both flooded and dry conditions responded better physiologically than plants that were either continuously flooded or received extended droughts (≥4 weeks). Therefore, provided the duration of water deficit is not extensive, short periods of water withdrawal can enhance the performance and water relations in some emergent-wetland plant species.     

Ecological relevance of minimum seasonal water potentials

The minimum seasonal water potential a plant experiences, ψ_min, provides an important measure of plant water status, as it reflects the maximum water deficit that leaves and xylem must tolerate to maintain physiological activity. ψ_min also acts as a selective force on xylem structure which, in turn, generates correlations between ψ_min and numerous hydraulic traits. This review focuses on the ecological relevance of ψ_min as a reflection of overall plant hydraulic strategy. The focus is on plant functional strategies with respect to soil drought, but we conclude with preliminary findings on the role of atmospheric drought.     

Abscisic acid in apoplastic sap can account for the restriction in leaf conductance of white lupins during moderate soil drying and after rewatering

We studied the effects of drought on leaf conductance (g) and on the concentration of abscisic acid (ABA) in the apoplastic sap of Lupinus albus L. leaves. Withholding watering for 5d resulted in complete stomatal closure and in severe leaf water deficit. Leaf water potential fully recovered immediately after rewatering, but the aftereffect of drought on stomata persisted for 2d. ABA and sucrose were quantified in pressurized leaf xylem extrudates. We assumed that the xylem sucrose concentration is negligible and hence that the presence of sucrose in leaf extrudates indicated that they were contaminated by phloem. To eliminate this interference, the concentration of ABA in leaf apoplast was estimated by extrapolation to zero sucrose concentration, using the regression between ABA and sucrose concentrations. The estimated apoplastic ABA concentration increased by 100-fold with soil drying and did not return to pre-stress values immediately following rewatering. g was closely related to the concentration of ABA in leaf apoplast. Furthermore, the feeding of exogenous ABA to leaves detached from well-watered plants brought about the same degree of depression in g as resulted from the drought-induced increase in ABA concentration. We therefore conclude that the observed changes in the concentration of ABA in leaf apoplast were quantitatively adequate to explain drought-induced stomatal closure and the delay in stomatal reopening following rewatering.     

Ambiol,spermine, and aminoethoxyvinylglycine prevent water stress and protect membranes in Pinus strobus L under drought

The ability of antistress compounds to enhance the drought tolerance of conifer seedlings was tested by feeding plant growth regulators (PGRs) to 1-year-old white pine (Pinus strobus L.), which were then subjected to either a moderate (11 day) or a more severe (16 day) drought. The following PGRs were either fed directly into the xylem or applied as a root drench: the antioxidant Ambiol (2-methyl-4-[dimethylaminomethyl]- 5-hydroxybenzimidazole dihydrochloride), the polyamine, spermine, an anti-ethylene agent, aminoethoxyvinylglycine (AVG), and the inhibitor, abscisic acid (ABA). Leaf water potentials (ƒ_l) declined in untreated seedlings when they were exposed to drought. Preconditioning with PGRs postponed water deficits and prevented membrane leakage under drought. The specific physiological adjustments observed were found to vary, depending on the type of compound. Ambiol, AVG and spermine caused transpirant rates to decline under drought. Although the antitranspirant effects of Ambiol and spermine would explain the increase in water use efficiency under drought, spermine also enhanced photosynthesis. The same compounds promoted osmotic adjustment, which would help to maintain turgor under drought. This was shown by the decline in osmotic potential at full turgor, and at zero turgor, in Ambiol and spermine-treated seedlings. Seedlings treated with Ambiol and ABA could sustain a greater water loss before turgor declined to zero. The possibility that preconditioning may help to maintain leaf physiological functioning under drought by reducing water stress and stress-ethylene production is discussed.     

Drought increases freezing tolerance of both leaves and xylem of Larrea tridentata

Drought and freezing are both known to limit desert plant distributions, but the interaction of these stressors is poorly understood. Drought may increase freezing tolerance in leaves while decreasing it in the xylem, potentially creating a mismatch between water supply and demand. To test this hypothesis, we subjected Larrea tridentata juveniles grown in a greenhouse under well‐watered or drought conditions to minimum temperatures ranging from ?8 to ?24 °C. We measured survival, leaf retention, gas exchange, cell death, freezing point depression and leaf‐specific xylem hydraulic conductance (k_l). Drought‐exposed plants exhibited smaller decreases in gas exchange after exposure to ?8 °C compared to well‐watered plants. Drought also conferred a significant positive effect on leaf, xylem and whole‐plant function following exposure to ?15 °C; drought‐exposed plants exhibited less cell death, greater leaf retention, higher k_l and higher rates of gas exchange than well‐watered plants. Both drought‐exposed and well‐watered plants experienced 100% mortality following exposure to ?24 °C. By documenting the combined effects of drought and freezing stress, our data provide insight into the mechanisms determining plant survival and performance following freezing and the potential for shifts in L. tridentata abundance and range in the face of changing temperature and precipitation regimes.     

Effects of drought stress on photosynthetic gas exchange, chlorophyll fluorescence and stem diameter of soybean plants

Changes in plant growth, photosynthetic gas exchange, chlorophyll fluorescence and stem diameter of soybean [Glycine max (L.) Merr.] plants under drought stress were studied. Total plant dry mass was reduced by 30 % compared to well-watered control plants. Leaf water potential was slightly decreased by water stress. Water stress induced daytime shrinkage and reduced night-time expansion of stem. Photosynthetic rate, stomatal conductance and transpiration rate were significantly declined by water stress, while the intercellular CO₂ concentration was changed only slightly at the initiation of stress treatment. The maximum photochemical efficiency of photosystem 2 and apparent photosynthetic electron transport rate were not changed by water stress.     

Growth and physiological responses to successional water deficit and recovery in four warm‐temperate woody species

Plant responses to drought and their subsequent rehydration can provide evidence for forest dynamics within the context of climate change. In this study, the seedlings of two native species (Vitex negundo var. heterophylla, Quercus acutissima) and two exotic species (Robinia pseudoacacia, Amorpha fruticosa) to China were selected in a greenhouse experiment. The gas exchange, stem hydraulic parameters, plant osmoprotectant contents and antioxidant activities of the seedlings that were subjected to sustained drought and rehydration (test group) as well as those of well‐irrigated seedlings (control group) were measured. The two native species exhibited a greater degree of isohydry with drought because they limited the stomatal opening timely from the onset of the drought. However, the two exotic species showed a more ‘water spender’‐like strategy with R. pseudoacacia showing anisohydric responses and A. fruticosa showing isohydrodynamic responses to drought. Severe drought significantly decreased the leaf gas exchange rates and hydraulic properties, whereas the instantaneous water use efficiency and osmoprotectant contents increased markedly. Most of the physiological parameters recovered rapidly after mild drought rehydration, but the water potential and/or supply of nonstructural carbohydrates did not recover after severe drought rehydration. The results demonstrate that the xylem hydraulic conductivity and shoot water potential jointly play a crucial role in the drought recovery of woody plants. In brief, the native species may play a dominant role in the future in warm‐temperate forests because they employ a better balance between carbon gain and water loss than the alien species under extreme drought conditions.