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1.
Mechanisms of plant survival and mortality during drought: why do some plants survive while others succumb to drought? 总被引:13,自引:1,他引:13
McDowell N Pockman WT Allen CD Breshears DD Cobb N Kolb T Plaut J Sperry J West A Williams DG Yepez EA 《The New phytologist》2008,178(4):719-739
Severe droughts have been associated with regional-scale forest mortality worldwide. Climate change is expected to exacerbate regional mortality events; however, prediction remains difficult because the physiological mechanisms underlying drought survival and mortality are poorly understood. We developed a hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality. Multiple mechanisms may cause mortality during drought. A common mechanism for plants with isohydric regulation of water status results from avoidance of drought-induced hydraulic failure via stomatal closure, resulting in carbon starvation and a cascade of downstream effects such as reduced resistance to biotic agents. Mortality by hydraulic failure per se may occur for isohydric seedlings or trees near their maximum height. Although anisohydric plants are relatively drought-tolerant, they are predisposed to hydraulic failure because they operate with narrower hydraulic safety margins during drought. Elevated temperatures should exacerbate carbon starvation and hydraulic failure. Biotic agents may amplify and be amplified by drought-induced plant stress. Wet multidecadal climate oscillations may increase plant susceptibility to drought-induced mortality by stimulating shifts in hydraulic architecture, effectively predisposing plants to water stress. Climate warming and increased frequency of extreme events will probably cause increased regional mortality episodes. Isohydric and anisohydric water potential regulation may partition species between survival and mortality, and, as such, incorporating this hydraulic framework may be effective for modeling plant survival and mortality under future climate conditions. 相似文献
2.
Kimberly A. Novick Alexandra G. Konings Pierre Gentine 《Plant, cell & environment》2019,42(6):1802-1815
Over the past decade, the concept of isohydry or anisohydry, which describes the link between soil water potential (ΨS), leaf water potential (ΨL), and stomatal conductance (gs), has soared in popularity. However, its utility has recently been questioned, and a surprising lack of coordination between the dynamics of ΨL and gs across biomes has been reported. Here, we offer a more expanded view of the isohydricity concept that considers effects of vapour pressure deficit (VPD) and leaf area index (AL) on the apparent sensitivities of ΨL and gs to drought. After validating the model with tree‐ and ecosystem‐scale data, we find that within a site, isohydricity is a strong predictor of limitations to stomatal function, though variation in VPD and leaf area, among other factors, can challenge its diagnosis. Across sites, the theory predicts that the degree of isohydricity is a good predictor of the sensitivity of gs to declining soil water in the absence of confounding effects from other drivers. However, if VPD effects are significant, they alone are sufficient to decouple the dynamics of ΨL and gs entirely. We conclude with a set of practical recommendations for future applications of the isohydricity framework within and across sites. 相似文献
3.
We merge concepts from stomatal optimization theory and cohesion–tension theory to examine the dynamics of three mechanisms that are potentially limiting to leaf‐level gas exchange in trees during drought: (1) a ‘demand limitation’ driven by an assumption of optimal stomatal functioning; (2) ‘hydraulic limitation’ of water movement from the roots to the leaves; and (3) ‘non‐stomatal’ limitations imposed by declining leaf water status within the leaf. Model results suggest that species‐specific ‘economics’ of stomatal behaviour may play an important role in differentiating species along the continuum of isohydric to anisohydric behaviour; specifically, we show that non‐stomatal and demand limitations may reduce stomatal conductance and increase leaf water potential, promoting wide safety margins characteristic of isohydric species. We used model results to develop a diagnostic framework to identify the most likely limiting mechanism to stomatal functioning during drought and showed that many of those features were commonly observed in field observations of tree water use dynamics. Direct comparisons of modelled and measured stomatal conductance further indicated that non‐stomatal and demand limitations reproduced observed patterns of tree water use well for an isohydric species but that a hydraulic limitation likely applies in the case of an anisohydric species. 相似文献
4.
Isohydric and anisohydric regulations of plant water status have been observed over several decades of field, glasshouse and laboratory studies, yet the functional significance and mechanism of both remain obscure. We studied the seasonal trends in plant water status and hydraulic properties in a natural stand of Eucalyptus gomphocephala through cycles of varying environmental moisture (rainfall, groundwater depth, evaporative demand) in order to test for isohydry and to provide physiological information for the mechanistic interpretation of seasonal trends in plant water status. Over a 16 month period of monitoring, spanning two summers, midday leaf water potential (psi(leaf)) correlated with predawn psi(leaf), which was correlated with water table depth below ground level, which in turn was correlated with total monthly rainfall. Eucalyptus gomphocephala was therefore not seasonally isohydric. Despite strong stomatal down-regulation of transpiration rate in response to increasing evaporative demand, this was insufficient to prevent midday psi(leaf) from falling to levels below -2 MPa in the driest month, well into the region likely to induce xylem air embolisms, based on xylem vulnerability curves obtained in the study. However, even though midday psi(leaf) varied by over 1.2 MPa across seasons, the hydrodynamic (transpiration-induced) water potential gradient from roots to shoots (delta psi(plant)), measured as the difference between predawn and midday psi(leaf), was relatively constant across seasons, averaging 0.67 MPa. This unusual pattern of hydraulic regulation, referred to here as isohydrodynamic, is explained by a hydromechanical stomatal control model where plant hydraulic conductance is dependent on transpiration rate. 相似文献
5.
Patrick J. Mitchell Anthony P. O'Grady Elizabeth A. Pinkard Timothy J. Brodribb Stefan K. Arndt Chris J. Blackman Remko A. Duursma Rod J. Fensham David W. Hilbert Craig R. Nitschke Jaymie Norris Stephen H. Roxburgh Katinka X. Ruthrof David T. Tissue 《Global Change Biology》2016,22(5):1677-1689
The surge in global efforts to understand the causes and consequences of drought on forest ecosystems has tended to focus on specific impacts such as mortality. We propose an ecoclimatic framework that takes a broader view of the ecological relevance of water deficits, linking elements of exposure and resilience to cumulative impacts on a range of ecosystem processes. This ecoclimatic framework is underpinned by two hypotheses: (i) exposure to water deficit can be represented probabilistically and used to estimate exposure thresholds across different vegetation types or ecosystems; and (ii) the cumulative impact of a series of water deficit events is defined by attributes governing the resistance and recovery of the affected processes. We present case studies comprising Pinus edulis and Eucalyptus globulus, tree species with contrasting ecological strategies, which demonstrate how links between exposure and resilience can be examined within our proposed framework. These examples reveal how climatic thresholds can be defined along a continuum of vegetation functional responses to water deficit regimes. The strength of this framework lies in identifying climatic thresholds on vegetation function in the absence of more complete mechanistic understanding, thereby guiding the formulation, application and benchmarking of more detailed modelling. 相似文献
6.
Water scarcity is a critical limitation for agricultural systems. Two different water management strategies have evolved in plants: an isohydric strategy and an anisohydric strategy. Isohydric plants maintain a constant midday leaf water potential (Ψleaf) when water is abundant, as well as under drought conditions, by reducing stomatal conductance as necessary to limit transpiration. Anisohydric plants have more variable Ψleaf and keep their stomata open and photosynthetic rates high for longer periods, even in the presence of decreasing leaf water potential. This risk-taking behavior of anisohydric plants might be beneficial when water is abundant, as well as under moderately stressful conditions. However, under conditions of intense drought, this behavior might endanger the plant. We will discuss the advantages and disadvantages of these two water-usage strategies and their effects on the plant’s ability to tolerate abiotic and biotic stress. The involvement of plant tonoplast AQPs in this process will also be discussed. 相似文献
7.
William R. L. Anderegg Adam Wolf Adriana Arango‐Velez Brendan Choat Daniel J. Chmura Steven Jansen Thomas Kolb Shan Li Frederick C. Meinzer Pilar Pita Víctor Resco de Dios John S. Sperry Brett T. Wolfe Stephen Pacala 《Ecology letters》2018,21(7):968-977
Stomatal response to environmental conditions forms the backbone of all ecosystem and carbon cycle models, but is largely based on empirical relationships. Evolutionary theories of stomatal behaviour are critical for guarding against prediction errors of empirical models under future climates. Longstanding theory holds that stomata maximise fitness by acting to maintain constant marginal water use efficiency over a given time horizon, but a recent evolutionary theory proposes that stomata instead maximise carbon gain minus carbon costs/risk of hydraulic damage. Using data from 34 species that span global forest biomes, we find that the recent carbon‐maximisation optimisation theory is widely supported, revealing that the evolution of stomatal regulation has not been primarily driven by attainment of constant marginal water use efficiency. Optimal control of stomata to manage hydraulic risk is likely to have significant consequences for ecosystem fluxes during drought, which is critical given projected intensification of the global hydrological cycle. 相似文献
8.
Lee Turin Dickman Nate G. McDowell Charlotte Grossiord Adam D. Collins Brett T. Wolfe Matteo Detto S. Joseph Wright Jos A. Medina‐Vega Devin Goodsman Alistair Rogers Shawn P. Serbin Jin Wu Kim S. Ely Sean T. Michaletz Chonggang Xu Lara Kueppers Jeffrey Q. Chambers 《Plant, cell & environment》2019,42(5):1705-1714
Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short‐ and long‐term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015–2016 El Niño to examine how short‐ and long‐term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P = 0.32, branch P = 0.30) nor across the rainfall gradient (leaf P = 0.91, branch P = 0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests. 相似文献
9.
Bradford P. Wilcox 《Ecohydrology》2010,3(1):126-130
As a result of human activities, forests and rangelands across the globe have undergone dramatic changes that have fundamentally altered ecosystem processes. Examples of these kinds of transformational changes include increasingly hot and extensive forest fires, die‐off over vast areas of forest from insect infestations, large‐scale encroachment of rangelands by woody plants and non‐native invasive plants, and desertification. These changes have accelerated in pace, scale and magnitude in recent decades and have the potential to alter water, energy, and biogeochemical cycles in important but not fully understood ways. The related disciplines of ecohydrology and watershed management are being shaped and transformed by the need to understand the ecohydrological consequences of transformative landscape change as well as the need to mitigate and manage for these changes. Copyright © 2010 John Wiley & Sons, Ltd. 相似文献
10.
Improving plant stress tolerance and yield production: is the tonoplast aquaporin SlTIP2;2 a key to isohydric to anisohydric conversion? 总被引:2,自引:0,他引:2
Nir Sade Basia J. Vinocur Alex Diber Arava Shatil Gil Ronen Hagit Nissan Rony Wallach Hagai Karchi Menachem Moshelion 《The New phytologist》2009,181(3):651-661
11.
Root sugar accumulation was studied in two grapevine varieties contrasting in tolerance to water stress. During a 10‐day water withholding treatment, the drought‐tolerant variety, Grenache, sustained less negative predawn and midday leaf water potentials as well as root water potential compared with the sensitive variety, Semillon. Grenache vines also maintained lower stomatal conductance and transpiration than Semillon vines throughout the drying period. In both varieties there was accumulation of sucrose in the roots and concentrations were inversely correlated to leaf and root water status. In both Grenache and Semillon, elevated root osmolality was associated with decreased soil moisture indicating that sugar accumulation may play a role in osmotic protection. Petiole xylem sap abscisic acid (ABA) concentrations increased with water deficit in both varieties and were highest for vines with the most negative root and predawn leaf water potentials. Furthermore, root sucrose concentrations were positively correlated with leaf xylem sap ABA concentrations, indicative of integration between carbohydrate metabolism and the ABA signalling system. Similar root sugar accumulation patterns between the two varieties, however, demonstrate that other factors are likely influencing the ability of the drought‐tolerant variety to remain hydrated. 相似文献
12.
13.
Increasing drought is one of the most critical challenges facing species and ecosystems worldwide, and improved theory and practices are needed for quantification of species tolerances. Leaf water potential at turgor loss, or wilting (π(tlp) ), is classically recognised as a major physiological determinant of plant water stress response. However, the cellular basis of π(tlp) and its importance for predicting ecological drought tolerance have been controversial. A meta-analysis of 317 species from 72 studies showed that π(tlp) was strongly correlated with water availability within and across biomes, indicating power for anticipating drought responses. We derived new equations giving both π(tlp) and relative water content at turgor loss point (RWC(tlp) ) as explicit functions of osmotic potential at full turgor (π(o) ) and bulk modulus of elasticity (ε). Sensitivity analyses and meta-analyses showed that π(o) is the major driver of π(tlp) . In contrast, ε plays no direct role in driving drought tolerance within or across species, but sclerophylly and elastic adjustments act to maintain RWC(tlp,) preventing cell dehydration, and additionally protect against nutrient, mechanical and herbivory stresses independent of drought tolerance. These findings clarify biogeographic trends and the underlying basis of drought tolerance parameters with applications in comparative assessments of species and ecosystems worldwide. 相似文献
14.
Anna T. Trugman Leander D. L. Anderegg John S. Sperry Yujie Wang Martin Venturas William R. L. Anderegg 《Global Change Biology》2019,25(12):4008-4021
Plant functional traits provide a link in process‐based vegetation models between plant‐level physiology and ecosystem‐level responses. Recent advances in physiological understanding and computational efficiency have allowed for the incorporation of plant hydraulic processes in large‐scale vegetation models. However, a more mechanistic representation of water limitation that determines ecosystem responses to plant water stress necessitates a re‐evaluation of trait‐based constraints for plant carbon allocation, particularly allocation to leaf area. In this review, we examine model representations of plant allocation to leaves, which is often empirically set by plant functional type‐specific allometric relationships. We analyze the evolution of the representation of leaf allocation in models of different scales and complexities. We show the impacts of leaf allocation strategy on plant carbon uptake in the context of recent advancements in modeling hydraulic processes. Finally, we posit that deriving allometry from first principles using mechanistic hydraulic processes is possible and should become standard practice, rather than using prescribed allometries. The representation of allocation as an emergent property of scarce resource constraints is likely to be critical to representing how global change processes impact future ecosystem dynamics and carbon fluxes and may reduce the number of poorly constrained parameters in vegetation models. 相似文献
15.
Polar Bear Pass, located on Bathurst Island (75°40′N, 98°30′W) in the Canadian High Arctic, is a large ecologically important, low‐lying wetland that receives water inputs from streams in the surrounding hillslopes. Most of these hillslope streams are gravel‐bed streams; however, some streambeds are moss lined. One of these vegetated streams was selected to determine how vegetation influences run‐off processes (e.g. water storage and streamflow) in comparison with a similar‐sized gravel‐bed hillslope stream. During the 2009 and 2010 field seasons, snow cover, melt and streamflow measurements were collected in both streams. Frost table and soil moisture conditions were monitored weekly along four cross‐sectional transects in each stream basin, and vegetation inventories were conducted in July 2010. The active layer developed more quickly in 2010 than in 2009, yet the thaw depth remained thinner in the vegetated stream in both years. In 2009, prolonged rainfall of low intensity and meltwater inputs from long‐lasting valley snowbeds maintained streamflow. Warmer and drier conditions in 2010 caused the stream to eventually cease flow, only regenerating after intense rainfall. Moss plays an important role in modifying streamflow later in the season when low‐flow conditions persist, acting as a barrier to streamflow as the moisture deficit of the vegetation must be satisfied before streamflow can regenerate. Results indicate that there is a threshold that determines whether vegetation promotes or hinders streamflow and that this factor will have important implications in response to changing climate conditions for this site and other arctic landscapes. Copyright © 2015 John Wiley & Sons, Ltd. 相似文献
16.
Cleiton B. Eller Fernanda de V. Barros Paulo R.L. Bittencourt Lucy Rowland Maurizio Mencuccini Rafael S. Oliveira 《Plant, cell & environment》2018,41(3):548-562
Faster growth in tropical trees is usually associated with higher mortality rates, but the mechanisms underlying this relationship are poorly understood. In this study, we investigate how tree growth patterns are linked with environmental conditions and hydraulic traits, by monitoring the cambial growth of 9 tropical cloud forest tree species coupled with numerical simulations using an optimization model. We find that fast‐growing trees have lower xylem safety margins than slow‐growing trees and this pattern is not necessarily linked to differences in stomatal behaviour or environmental conditions when growth occurs. Instead, fast‐growing trees have xylem vessels that are more vulnerable to cavitation and lower density wood. We propose the growth ‐ xylem vulnerability trade‐off represents a wood hydraulic economics spectrum similar to the classic leaf economic spectrum, and show through numerical simulations that this trade‐off can emerge from the coordination between growth rates, wood density, and xylem vulnerability to cavitation. Our results suggest that vulnerability to hydraulic failure might be related with the growth‐mortality trade‐off in tropical trees, determining important life history differences. These findings are important in furthering our understanding of xylem hydraulic functioning and its implications on plant carbon economy. 相似文献
17.
18.
The feasibility of two hypothetical mechanisms for the stomatal response to humidity was evaluated by identifying theoretical constraints on these mechanisms and by analysing timecourses of stomatal aperture following a step change in humidity. The two hypothetical mechanisms, which allow guard cell turgor pressure to overcome the epidermal mechanical advantage, are: (1) active regulation of guard cell osmotic pressure, requiring no hydraulic disequilibrium between guard and epidermal cells, and (2) a substantial hydraulic resistance between guard and epidermal cells, resulting in hydraulic disequilibrium between them. Numerical simulations of the system are made possible by recently published empirical relationships between guard cell pressure and volume and between stomatal aperture, guard cell turgor pressure, and epidermal cell turgor pressure; these data allow the hypothetical control variables to be inferred from stomatal aperture and evaporative demand, given physical assumptions that characterize either hypothesis. We show that hypothesis (1) predicts that steady‐state πg is monotonically related to transpiration rate, whereas hypothesis (2) suggests that the relationship between transpiration rate and the steady‐state guard to epidermal cell hydraulic resistance may be either positive or negative, and that this resistance must change substantially during the transient phase of the stomatal response to humidity. 相似文献
19.
《Journal of Plant Interactions》2013,8(1):59-66
Abstract From the measurements of the profiles of hydraulic conductance and water potential from soil through to the leaf system in fully established melon plants, the limits to water flow set by coupling of hydraulic conductance (k) with water relation parameters was evaluated in the laboratory using high pressure flow device (HPFM) and evaporative flux method (EF). The rootstock Arava was grafted onto self, and onto two genotypes (AR57 and AR82) using side and V graft types, and there was an ungrafted control. Hydraulic transport efficiency was estimated from measurements of evaporative flux (transpiration rate) and leaf water potential (ψL) measured between pre-dawn and sunset during the growth cycle. Measured parameters to characterize the hydraulic efficiency (architecture) of the vascular system of melon were normalized to areas of leaves and stem cross section; this enabled the examination of their physiological and ecological functions. The effects of rootstock genotype were more marked on graft union and scion water relations. Differences in the magnitudes of water relation parameters of hydraulic conductance, water potential (lwp) and evaporative water loss (EF) were detected. AR/RS82 side grafted exhibited high EF and Kh despite its lower leaf water potential compared to AR/RS57 V grafted. Self grafting (Arava/Arava grafts) in melon seems to improve water relations and xylem water transport efficiency. Parameters describing the hydraulic efficiency (architecture) of vascular system of melon plants were described in relation to plant attributes. The expression of hydraulic conductance of the root and shoot system relative to plant attributes did not eliminate differences in the magnitudes of conductance elements in tomato and melon. Differences obtained among the different melon grafts in whole plant leaf and stem area specific hydraulic conductance (Kl) indicate the carbon efficiency and hence the cost of resource allocation to areas of root surface and leaves. The role of plant water relations in root-shoot communications and whole plant regulation of water flux are inferred from this study. 相似文献
20.
J. C. Ramalho Z. S. Zlatev A. E. Leitão I. P. Pais A. S. Fortunato F. C. Lidon 《Plant biology (Stuttgart, Germany)》2014,16(1):133-146
The impact of moderate water deficit on the photosynthetic apparatus of three Phaseolus vulgaris L. cultivars, Plovdiv 10 (P10), Dobrudjanski Ran (DR) and Prelom (Prel), was investigated. Water shortage had less impact on leaf hydration, RWC (predawn and midday) and predawn water potential in Prel. RWC and Ψp were more reduced in P10, while there was no osmotic adjustment in any cultivar. Although drought drastically reduced stomatal opening in P10 and DR, reduced Amax indicated non‐stomatal limitations that contributed to the negligible Pn. These limitations were on potential thylakoid electron transport rates of PSI and II, pointing to photosystem functioning as a major limiting step in photosynthesis. This agrees with decreases in actual photochemical efficiency of PSII (Fv′/Fm′), quantum yield of photosynthetic non‐cyclic electron transport (?e) and energy‐driven photochemical events (qP), although the impact on these parameters would also include down‐regulation processes. When compared to DR, Prel retained a higher functional state of the photosynthetic machinery, justifying reduced need for photoprotective mechanisms (non‐photochemical quenching, zeaxanthin, lutein, β‐carotene) and maintenance of the balance between energy capture and dissipative pigments. The highest increases in fructose, glucose, arabinose and sorbitol in Prel might be related to tolerance to a lower oxidative state. All cultivars had reduced Amax due to daytime stomatal closure in well‐watered conditions. Under moderate drought, Prel had highest tolerance, higher leaf hydration and maintenance of important photochemical use of energy. However, water shortage caused appreciable non‐stomatal limitations to photosynthesis linked to regulation/imbalance at the metabolic level (and growth) in all cultivars. This included damage, as reflected in decreased potential photosystem functioning, pointing to higher sensitivity of photosynthesis to drought than is commonly assumed. 相似文献