Differences in xylem and leaf hydraulic traits explain differences in drought tolerance among mature Amazon rainforest trees

Considerable uncertainty surrounds the impacts of anthropogenic climate change on the composition and structure of Amazon forests. Building upon results from two large‐scale ecosystem drought experiments in the eastern Brazilian Amazon that observed increases in mortality rates among some tree species but not others, in this study we investigate the physiological traits underpinning these differential demographic responses. Xylem pressure at 50% conductivity (xylem‐P₅₀), leaf turgor loss point (TLP), cellular osmotic potential (π_o), and cellular bulk modulus of elasticity (ε), all traits mechanistically linked to drought tolerance, were measured on upper canopy branches and leaves of mature trees from selected species growing at the two drought experiment sites. Each species was placed a priori into one of four plant functional type (PFT) categories: drought‐tolerant versus drought‐intolerant based on observed mortality rates, and subdivided into early‐ versus late‐successional based on wood density. We tested the hypotheses that the measured traits would be significantly different between the four PFTs and that they would be spatially conserved across the two experimental sites. Xylem‐P₅₀, TLP, and π_o, but not ε, occurred at significantly higher water potentials for the drought‐intolerant PFT compared to the drought‐tolerant PFT; however, there were no significant differences between the early‐ and late‐successional PFTs. These results suggest that these three traits are important for determining drought tolerance, and are largely independent of wood density—a trait commonly associated with successional status. Differences in these physiological traits that occurred between the drought‐tolerant and drought‐intolerant PFTs were conserved between the two research sites, even though they had different soil types and dry‐season lengths. This more detailed understanding of how xylem and leaf hydraulic traits vary between co‐occuring drought‐tolerant and drought‐intolerant tropical tree species promises to facilitate a much‐needed improvement in the representation of plant hydraulics within terrestrial ecosystem and biosphere models, which will enhance our ability to make robust predictions of how future changes in climate will affect tropical forests.     

Global analysis of plasticity in turgor loss point,a key drought tolerance trait

Many species face increasing drought under climate change. Plasticity has been predicted to strongly influence species' drought responses, but broad patterns in plasticity have not been examined for key drought tolerance traits, including turgor loss or ‘wilting’ point (π_tlp). As soil dries, plants shift π_tlp by accumulating solutes (i.e. ‘osmotic adjustment’). We conducted the first global analysis of plasticity in Δπ_tlp and related traits for 283 wild and crop species in ecosystems worldwide. Δπ_tlp was widely prevalent but moderate (?0.44 MPa), accounting for 16% of post‐drought π_tlp. Thus, pre‐drought π_tlp was a considerably stronger predictor of post‐drought π_tlp across species of wild plants. For cultivars of certain crops Δπ_tlp accounted for major differences in post‐drought π_tlp. Climate was correlated with pre‐ and post‐drought π_tlp, but not Δπ_tlp. Thus, despite the wide prevalence of plasticity, π_tlp measured in one season can reliably characterise most species' constitutive drought tolerances and distributions relative to water supply.     

鼎湖山季风常绿阔叶林两种优势树种的生境适应研究:叶片功能性状和水力结构特征

植物在长期的进化和发展过程中,通过与环境相互作用形成了适应环境的形态结构及生理特征,反映了植物适应环境的生态策略;在森林群落中,地形和土壤等的变异常导致生境的异质性,从而直接或者间接影响物种的分布格局。因此,在生境异质性较强的森林群落中,植物物种分布格局与其生态适应策略有何关系,是值得关注的问题。该文以鼎湖山南亚热带季风常绿阔叶林20 hm2监测样地为平台,针对两种常绿优势树种罗伞树(Ardisia quinquegona)和光叶山黄皮(Aidia canthioides),对比研究了两种植物的叶片功能性状和水力结构特征在山脊、山坡、山谷三种不同生境中的生态适应策略,以阐明物种分布格局与其生态适应策略的关系。结果表明:罗伞树主要是通过调整叶面积(LA)、木材密度(WD)及渗透调节来适应不同生境;光叶山黄皮主要通过调整比叶面积(SLA)、WD及渗透调节,采取养分有效保存(低SLA,高干物质含量)及慢生长高存活的策略以适应不同生境,适应环境能力更强,尤其是在山脊和山坡生境;而且影响两个树种叶片功能性状和水力结构的主导土壤因子有所不同。研究结果说明罗伞树和光叶山黄皮对山脊和山坡生境比山谷更为适应,但在叶片功能性状和水力结构特征方面的生境适应策略不同。     

Global meta-analysis shows that relationships of leaf mass per area with species shade tolerance depend on leaf habit and ontogeny

It was predicted that relationships of leaf mass per area (LMA) with juvenile shade tolerance will depend on leaf habit, and on whether species are compared at a common age as young seedlings, or at a common size as saplings. A meta-analysis of 47 comparative studies (372 species) was used to test predictions, and the effect of light environment on this relationship. The LMA of evergreens was positively correlated with shade tolerance, irrespective of ontogeny or light environment. The LMA of young seedlings (相似文献   

The impact of herbivore exclusion on forb diversity: Comparing species and functional responses during a drought

Rainfall and herbivory shape savannah herbaceous communities, but these disturbances are being altered globally. To assess potential consequences of such alterations, we evaluated herbivore effects on species and functional diversity during an episodic drought in a sodic savannah using data collected from long-term herbivore exclosures in the Kruger National Park, South Africa. Herbaceous life forms are rarely acknowledged as distinct functional entities. Moreover, the functional ecology of forbs remains elusive. Here, we present disturbances–responses by forbs separately from grasses. We hypothesised that combinations of intense utilisation and drought would be associated with low diversity and high dominance at species and functional levels for both life forms. Contrary to our hypothesis, low forb and grass diversity was associated with long-term herbivore exclusion, which exceeded expected undesirable effects of intense utilisation and drought. Grasses responded less sensitively, suggesting that forbs respond dynamically to changes in herbivore assemblage when these alterations are combined with drought. Consistent with patterns in savannah systems, forbs contributed significantly to species and functional trait diversity. High forb diversity is suggested to enhance resilience of this nutrient-rich ecosystem against declines in its functioning when subjected to drought and alterations in herbivory.     

Correlation of drought traits and the predictability of osmotic potential at full leaf turgor in vegetation from New Zealand

Scientists do not know precisely how severe will be the impact of climate change on species. Evidence suggests that for some species, their future distributions might be jeopardized by local extinctions and drought‐induced tree mortality. Thus, we require models capable of estimating drought tolerance across many species. We can approach this goal by assessing functional traits. The trait osmotic potential at full turgor, π_O, is potentially a good drought indicator; however, few studies address its importance as a drought‐tolerance predictor and it is difficult to measure in the field with accuracy. In this work, we aim to answer the questions: which drought traits correlate with π_O?; do morpho‐anatomical traits correlate with π_O?; and which trees and shrubs are more (or less) vulnerable to drought? To achieve this aim, we assessed physiological and morpho‐anatomical traits for 14 native species from New Zealand forests. We included leaf‐ and wood‐related traits, π_O, water potential and stomatal conductance. We examined how these traits correlate with π_O and sought to generate models to predict π_O as a function of other traits. We tested 33 different models and evaluated them using Akaike's information criterion. Unfortunately, none of the morpho‐anatomical traits correlated well with π_O. Instead, water potential correlated most strongly with π_O. None of the models using only morpho‐anatomical traits produced plausible results. The model with the best predictive performance incorporated the effects of both morpho‐anatomical and physiological traits: water potential and wood saturated water content. Of the species analysed, and based on their π_O response, Lophozonia menziesii was considered the most vulnerable to drought stress, whereas Plagianthus regius was the least vulnerable. Our findings imply that it is potentially valuable to keep exploring the use of π_O as a drought indicator and that the effort required to measure some physiological traits, such as water potential, may be essential to consider plant drought responses and to predict π_O.     

Leaf turgor loss point at full hydration for 41 native and introduced tree and shrub species from Central Europe

过去几年,中欧的森林遭受了由气候变化引起的直接后果—干旱。所有这些森林都有着悠久的经营历史,重新种植受损森林的责 任落到了土地所有者的身上。因此,土地所有者和政府正在寻找适合在树木灭绝地区重新种植的树种。事实上,很好的了解物种的抗旱性 对当前树木的重新种植非常关键。我们在41种原产于或引进于中欧的木本树种中确定了一个被广泛认可的叶片耐旱特性(在充分水合作用 下膨压损失点的叶片水势,即π_tlp)。采用渗透快速评价法测定了暴露于阳光下的叶片在充分水合作用下的渗透势(π_osm),并将其转化为π_tlp。 本地种的平均π_tlp为−2.33 ± 0.33 MPa。引进树种Aesculus hypocastania的π_tlp为−1.70 ± 0.11 MPa。引进树种Pseudotsuga menzesii的π_tlp值 为−3.02 ± 0.14 MPa,耐旱性最强。绝对值较低的负值πtlp表示抗旱性较低,绝对值较高的负值π_tlp表示抗旱性较高。例如,两种本地物种 Illex aquifolium和Alnus glustinosa是潮湿生境的自然伴生物种,其π_tlp值分别为−1.75 ± 0.02 MPa和−1.76 ± 0.03 MPa。     

Vulnerability to xylem cavitation of Hakea species (Proteaceae) from a range of biomes and life histories predicted by climatic niche

Background and AimsExtreme drought conditions across the globe are impacting biodiversity, with serious implications for the persistence of native species. However, quantitative data on physiological tolerance are not available for diverse flora to inform conservation management. We quantified physiological resistance to cavitation in the diverse Hakea genus (Proteaceae) to test predictions based on climatic origin, life history and functional traits.MethodsWe sampled terminal branches of replicate plants of 16 species in a common garden. Xylem cavitation was induced in branches under varying water potentials (tension) in a centrifuge, and the tension generating 50 % loss of conductivity (stem P₅₀) was characterized as a metric for cavitation resistance. The same branches were used to estimate plant functional traits, including wood density, specific leaf area and Huber value (sap flow area to leaf area ratio).Key ResultsThere was significant variation in stem P₅₀ among species, which was negatively associated with the species climate origin (rainfall and aridity). Cavitation resistance did not differ among life histories; however, a drought avoidance strategy with terete leaf form and greater Huber value may be important for species to colonize and persist in the arid biome.ConclusionsThis study highlights climate (rainfall and aridity), rather than life history and functional traits, as the key predictor of variation in cavitation resistance (stem P₅₀). Rainfall for species origin was the best predictor of cavitation resistance, explaining variation in stem P₅₀, which appears to be a major determinant of species distribution. This study also indicates that stem P₅₀ is an adaptive trait, genetically determined, and hence reliable and robust for predicting species vulnerability to climate change. Our findings will contribute to future prediction of species vulnerability to drought and adaptive management under climate change.     

Environmental and plant community determinants of species loss following nitrogen enrichment

Global energy use and food production have increased nitrogen inputs to ecosystems worldwide, impacting plant community diversity, composition, and function. Previous studies show considerable variation across terrestrial herbaceous ecosystems in the magnitude of species loss following nitrogen (N) enrichment. What controls this variation remains unknown. We present results from 23 N-addition experiments across North America, representing a range of climatic, soil and plant community properties, to determine conditions that lead to greater diversity decline. Species loss in these communities ranged from 0 to 65% of control richness. Using hierarchical structural equation modelling, we found greater species loss in communities with a lower soil cation exchange capacity, colder regional temperature, and larger production increase following N addition, independent of initial species richness, plant productivity, and the relative abundance of most plant functional groups. Our results indicate sensitivity to N addition is co-determined by environmental conditions and production responsiveness, which overwhelm the effects of initial community structure and composition.     

Intraspecific variation of drought adaptation in brittlebush: leaf pubescence and timing of leaf loss vary with rainfall

Reflective leaf pubescence of the desert shrub Encelia farinosa (brittlebrush) reduces leaf temperature and plant water loss, and is considered adaptive in xeric environments. Yet, little is known about intraspecific variation in this trait. Among three populations in the northern range of E. farinosa, which span a very broad precipitation gradient, both leaf absorptance variation and differences in the timing of drought-induced leaf loss were broadly associated with climatic variability. Where mean annual rainfall was greatest, drought-induced leaf loss was earliest, but these plants also had higher population-level mean leaf absorptance values. Higher absorptance increases the relative dependence on latent heat transfer (transpirational cooling), but it also provides greater instantaneous carbon assimilation. Plants at the driest site reached lower leaf absorptance values and maintained leaves longer into the drought period. Lower leaf absorptance reduces water consumption, and extended leaf longevity may buffer against the unpredictability of growing conditions experienced in the driest site. These observations are consistent with a trade-off scenario in which plants from wetter regions might trade off water conservation for higher instantaneous carbon gain, whereas plants from drier regions reduce water consumption and extend leaf longevity to maintain photosynthetic activity in the face of unpredictable growing conditions. Received: 2 April 1997 / Accepted: 11 August 1997     

Anatomical changes with needle length are correlated with leaf structural and physiological traits across five Pinus species

The genus Pinus has wide geographical range and includes species that are the most economically valued among forest trees worldwide. Pine needle length varies greatly among species, but the effects of needle length on anatomy, function, and coordination and trade‐offs among traits are poorly understood. We examined variation in leaf morphological, anatomical, mechanical, chemical, and physiological characteristics among five southern pine species: Pinus echinata, Pinus elliottii, Pinus palustris, Pinus taeda, and Pinus virginiana. We found that increasing needle length contributed to a trade‐off between the relative fractions of support versus photosynthetic tissue (mesophyll) across species. From the shortest (7 cm) to the longest (36 cm) needles, mechanical tissue fraction increased by 50%, whereas needle dry density decreased by 21%, revealing multiple adjustments to a greater need for mechanical support in longer needles. We also found a fourfold increase in leaf hydraulic conductance over the range of needle length across species, associated with weaker upward trends in stomatal conductance and photosynthetic capacity. Our results suggest that the leaf size strongly influences their anatomical traits, which, in turn, are reflected in leaf mechanical support and physiological capacity.     

The global distribution of ecosystems in a world without fire

This paper is the first global study of the extent to which fire determines global vegetation patterns by preventing ecosystems from achieving the potential height, biomass and dominant functional types expected under the ambient climate (climate potential). To determine climate potential, we simulated vegetation without fire using a dynamic global-vegetation model. Model results were tested against fire exclusion studies from different parts of the world. Simulated dominant growth forms and tree cover were compared with satellite-derived land- and tree-cover maps. Simulations were generally consistent with results of fire exclusion studies in southern Africa and elsewhere. Comparison of global 'fire off' simulations with landcover and treecover maps show that vast areas of humid C(4) grasslands and savannas, especially in South America and Africa, have the climate potential to form forests. These are the most frequently burnt ecosystems in the world. Without fire, closed forests would double from 27% to 56% of vegetated grid cells, mostly at the expense of C(4) plants but also of C(3) shrubs and grasses in cooler climates. C(4) grasses began spreading 6-8 Ma, long before human influence on fire regimes. Our results suggest that fire was a major factor in their spread into forested regions, splitting biotas into fire tolerant and intolerant taxa.     

A meta-analysis on morphological,physiological and biochemical responses of plants with PGPR inoculation under drought stress

Plant growth-promoting rhizobacteria (PGPR) can help plants to resist drought stress. However, the mechanisms of how PGPR inoculation affect plant status under drought remain incompletely understood. We performed a meta-analysis of plant response to PGPR inoculation by compiling data from 57 PGPR-inoculation studies, including 2, 387 paired observations on morphological, physiological and biochemical parameters under drought and well-watered conditions. We compare the PGPR effect on plants performances among different groups of controls and treatments. Our results reveal that PGPR enables plants to restore themselves from drought-stressed to near a well-watered state, and that C4 plants recover better from drought stress than C3 plants. Furthermore, PGPR is more effective underdrought than well-watered conditions in increasing plant biomass, enhancing photosynthesis and inhibiting oxidant damage, and the responses of C4 plants to the PGPR effect was stronger than that of C3 plants under drought conditions. Additionally, PGPR belonging to different taxa and PGPR with different functional traits have varying degrees of drought-resistance effects on plants. These results are important to improve our understanding of the PGPR beneficial effects on enhanced drought-resistance of plants.     

No trade-off between trichome production and tolerance to leaf and inflorescence damage in a natural population of Arabidopsis lyrata

Aims Early models of plant defense conceived resistance and tolerance to herbivore damage as mutually exclusive strategies. Support for this idea has been equivocal and studies on these two strategies are still needed to understand the evolution of defenses in natural populations. In Arabidopsis lyrata, the production of trichomes, a documented resistance trait, has been associated with a fitness cost in the absence of herbivores. We examined whether trichome production is also associated with reduced tolerance to simulated herbivore damage.Methods We conducted a field experiment in a natural Swedish population of A. lyrata where we inflicted leaf (0 vs. 50% of the area of each leaf removed) and inflorescence damage (0 vs. 50% of inflorescences removed) to trichome-producing and glabrous plants in a factorial design. We examined the response (survival, growth and reproduction) of the plants to the imposed damage over 2 years.Important findings Trichome-producing plants were not less tolerant than glabrous plants to simulated herbivore damage (no significant morph × leaf damage or morph × inflorescence damage interactions). Inflorescence and leaf damage had independent negative effects on the performance of damaged plants. Leaf damage reduced rosette size the year of damage, but effects on reproductive output in the year of damage, and on survival and reproductive performance the following year were weak and not statistically significant. Inflorescence damage significantly reduced the number of flowers, fruits and seeds the year of damage, but not in the following year. Irrespective of morph, the study population was more tolerant to leaf than to inflorescence damage. The results indicated no trade-off between trichome production and tolerance, suggesting that these two defense mechanisms have the potential to evolve independently in this A. lyrata population.     

Beyond isohydricity: The role of environmental variability in determining plant drought responses

Despite the appeal of the iso/anisohydric framework for classifying plant drought responses, recent studies have shown that such classifications can be strongly affected by a plant's environment. Here, we present measured in situ drought responses to demonstrate that apparent isohydricity can be conflated with environmental conditions that vary over space and time. In particular, we (a) use data from an oak species (Quercus douglasii) during the 2012–2015 extreme drought in California to demonstrate how temporal and spatial variability in the environment can influence plant water potential dynamics, masking the role of traits; (b) explain how these environmental variations might arise from climatic, topographic, and edaphic variability; (c) illustrate, through a “common garden” thought experiment, how existing trait‐based or response‐based isohydricity metrics can be confounded by these environmental variations, leading to Type‐1 (false positive) and Type‐2 (false negative) errors; and (d) advocate for the use of model‐based approaches for formulating alternate classification schemes. Building on recent insights from greenhouse and vineyard studies, we offer additional evidence across multiple field sites to demonstrate the importance of spatial and temporal drivers of plants' apparent isohydricity. This evidence challenges the use of isohydricity indices, per se, to characterize plant water relations at the global scale.     

Greater growth stability of trees in marginal habitats suggests a patchy pattern of population loss and retention in response to increased drought at the rear edge

Species rear range edges are predicted to retract as climate warms, yet evidence of population persistence is accumulating. Accounting for this disparity is essential to enable prediction and planning for species’ range retractions. At the Mediterranean edge of European beech‐dominated temperate forest, we tested the hypothesis that individual performance should decline at the limit of the species’ ecological tolerance in response to increased drought. We sampled 40 populations in a crossed factor design of geographical and ecological marginality and assessed tree growth resilience and decline in response to recent drought. Drought impacts occurred across the rear edge, but tree growth stability was unexpectedly high in geographically isolated marginal habitat and lower than anticipated in the species’ continuous range and better‐quality habitat. Our findings demonstrate that, at the rear edge, range shifts will be highly uneven and characterised by reduction in population density with local population retention rather than abrupt range retractions.     

Interactions of drought and shade effects on seedlings of four Quercus species: physiological and structural leaf responses

Here, we investigated the physiological and structural leaf responses of seedlings of two evergreen and two deciduous Quercus species, grown in a glasshouse and subjected to contrasted conditions of light (low, medium and high irradiance) and water (continuous watering vs 2-months drought). The impact of drought on photosynthetic rate was strongest in high irradiance, while the impact of shade on photosynthetic rate was strongest with high water supply, contradicting the hypothesis of allocation trade-off. Multivariate causal models were evaluated using d-sep method. The model that best fitted the dataset proposed that the variation in specific leaf area affects photosynthetic rate and leaf nitrogen concentration, and this trait determines stomatal conductance, which also affects photosynthetic rate. Shade conditions seemed to ameliorate, or at least not aggravate, the drought impact on oak seedlings, therefore, the drought response on leaf performance depended on the light environment.