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.     

Integrated drought responses of black poplar: how important is phenotypic plasticity?

Climate change is expected to increase drought frequency and intensity which will threaten plant growth and survival. In such fluctuating environments, perennial plants respond with hydraulic and biomass adjustments, resulting in either tolerant or avoidant strategies. Plants' response to stress relies on their phenotypic plasticity. The goal of this study was to explore physiology of young Populus nigra in the context of a time‐limited and progressive water deficit in regard to their growth and stress response strategies. Fourteen French 1‐year‐old black poplar genotypes, geographically contrasted, were subjected to withholding water during 8 days until severe water stress. Water fluxes (i.e. leaf water potentials and stomatal conductance) were analyzed together with growth (i.e. radial and longitudinal branch growth, leaf senescence and leaf production). Phenotypic plasticity was calculated for each trait and response strategies to drought were deciphered for each genotype. Black poplar genotypes permanently were dealing with a continuum of adjusted water fluxes and growth between two extreme strategies, tolerance and avoidance. Branch growth, leaf number and leaf hydraulic potential traits had contrasted plasticities, allowing genotype characterization. The most tolerant genotype to water deficit, which maintained growth, had the lowest global phenotypic plasticity. Conversely, the most sensitive and avoidant genotype ceased growth until the season's end, had the highest plasticity level. All the remaining black poplar genotypes were close to avoidance with average levels of traits plasticity. These results underpinned the role of plasticity in black poplar response to drought and calls for its wider use into research on plants' responses to stress.     

Seasonal acclimation of leaf respiration in Eucalyptus saligna trees: impacts of elevated atmospheric CO2 and summer drought

Understanding the impacts of atmospheric [CO₂] and drought on leaf respiration (R) and its response to changes in temperature is critical to improve predictions of plant carbon‐exchange with the atmosphere, especially at higher temperatures. We quantified the effects of [CO₂]‐enrichment (+240 ppm) on seasonal shifts in the diel temperature response of R during a moderate summer drought in Eucalyptus saligna growing in whole‐tree chambers in SE Australia. Seasonal temperature acclimation of R was marked, as illustrated by: (1) a downward shift in daily temperature response curves of R in summer (relative to spring); (2)≈60% lower R measured at 20^oC (R₂₀) in summer compared with spring; and (3) homeostasis over 12 months of R measured at prevailing nighttime temperatures. R₂₀, measured during the day, was on average 30–40% higher under elevated [CO₂] compared with ambient [CO₂] across both watered and droughted trees. Drought reduced R₂₀ by≈30% in both [CO₂] treatments resulting in additive treatment effects. Although [CO₂] had no effect on seasonal acclimation, summer drought exacerbated the seasonal downward shift in temperature response curves of R. Overall, these results highlight the importance of seasonal acclimation of leaf R in trees grown under ambient‐ and elevated [CO₂] as well as under moderate drought. Hence, respiration rates may be overestimated if seasonal changes in temperature and drought are not considered when predicting future rates of forest net CO₂ exchange.     

Traits explain sorting of C4 grasses along a global precipitation gradient

1. Species distributions are closely associated with moisture availability, but the underlying mechanisms remain unresolved. Drought relations are especially important for plants such as C₄ grasses that dominate seasonally dry ecosystems. Here, we test the hypothesis that C₄ grass species sampled across global precipitation gradients show variation in survival under drought that can be explained by their traits.
2. Our experiment subjected 18 C₄ grass species to a lethal drought under controlled environmental conditions. The number of days until death was measured, along with root traits, senescence, and aspects of hydraulic function.
3. We identified two strategies: Drought‐avoiding species that stayed green as the water potential declined and drought‐tolerating species that senesced more quickly but could extend survival via drought‐tolerant meristems.
4. Plants that stay‐green for longer occupied drier habitats and had the longest survival under drought, facilitated by narrow root diameter and isohydric stomatal behavior. Plants that senesced quickly had thicker roots, an anisohydric strategy, and occupied wetter habitats.
5. Global distributions of C₄ grasses can be predicted by variation in rates of senescence, meristem survival, root traits, and stomatal strategy, showing the value of these traits for understanding plant distributions in relation to climate.

     

FACE facts hold for multiple generations; Evidence from natural CO2 springs

Rising atmospheric CO₂ concentration is a key driver of enhanced global greening, thought to account for up to 70% of increased global vegetation in recent decades. CO₂ fertilization effects have further profound implications for ecosystems, food security and biosphere‐atmosphere feedbacks. However, it is also possible that current trends will not continue, due to ecosystem level constraints and as plants acclimate to future CO₂ concentrations. Future predictions of plant response to rising [CO₂] are often validated using single‐generation short‐term FACE (Free Air CO₂ Enrichment) experiments but whether this accurately represents vegetation response over decades is unclear. The role of transgenerational plasticity and adaptation in the multigenerational response has yet to be elucidated. Here, we propose that naturally occurring high CO₂ springs provide a proxy to quantify the multigenerational and long‐term impacts of rising [CO₂] in herbaceous and woody species respectively, such that plasticity, transgenerational effects and genetic adaptation can be quantified together in these systems. In this first meta‐analysis of responses to elevated [CO₂] at natural CO₂ springs, we show that the magnitude and direction of change in eight of nine functional plant traits are consistent between spring and FACE experiments. We found increased photosynthesis (49.8% in spring experiments, comparable to 32.1% in FACE experiments) and leaf starch (58.6% spring, 84.3% FACE), decreased stomatal conductance (g_s, 27.2% spring, 21.1% FACE), leaf nitrogen content (6.3% spring, 13.3% FACE) and Specific Leaf Area (SLA, 9.7% spring, 6.0% FACE). These findings not only validate the use of these sites for studying multigenerational plant response to elevated [CO₂], but additionally suggest that long‐term positive photosynthetic response to rising [CO₂] are likely to continue as predicted by single‐generation exposure FACE experiments.     

干旱对土壤剖面不同深度土壤CO2通量的影响

由于全球气候变化,预计未来我国亚热带地区干旱频率和持续时间将会增加。森林土壤CO₂的释放是陆地生态系统碳循环的重要组成部分,然而,有关不同深度土壤CO₂通量对干旱响应的理解仍相当有限。选择武夷山针叶林（Coniferous Forest,CF）和常绿阔叶林（Evergreen Broadleaved Forest,EBF）为研究对象,于2014年6月至2015年12月,采用梯度法计算10、30 cm和50 cm深度各层土壤CO₂通量,探讨模拟干旱对其影响。结果表明：CF和EBF样地土壤CO₂浓度均随土壤深度的增加而升高。CF和EBF样地对照（CK）处理10 cm深度土壤CO₂生产量分别占总CO₂生产量的53.5%和55.7%,表明土壤CO₂生产量主要来源于浅层土壤,这可能与浅层土壤有高的有机碳含量及细根生物量主要分布区有关。干旱处理使CF和EBF样地不同深度土壤CO₂通量均显著减少。在两个样地土壤CO₂通量的温度敏感性（Q₁₀）值均随着土壤深度的增加而减少。干旱处理显著减少了CF样地浅层土壤的Q₁₀值（P=0.02）,对深层土壤影响不显著（30 cm：P=0.30;50 cm：P=0.23）;而在EBF样地干旱处理显著减少了深层土壤的Q₁₀值（30 cm：P=0.02;50 cm：P=0.01）,对浅层土壤影响不显著（P=0.32）。     

Physiological and biochemical traits of drought tolerance in Argania spinosa

The objective of this study was to understand and characterize the physiological and biochemical tolerance mechanisms of Argania spinosa under drought stress for selection tolerant ecotypes. Significant differences were observed among ecotypes in indices of leaf water status studied: stomatal conductance (g_s), predawn leaf water potential (Ψ_pd) and leaf relative water content. There was a significant decrease in these physiological traits with increasing degree of drought stress in all ecotypes. Drought stress significantly increased endogenous H₂O₂ and lipid peroxidation. Moderate and severe drought stress increased significantly the catalase, superoxide dismutase, peroxidase, polyphenoloxidase and lipoxygenase activities, depending on time. Their constitutive activities were higher in inland ecotypes than in coastal ecotypes. According to canonical discriminant analysis, the inland ecotypes were essentially distinguished from the coastal ecotypes by the following physiological and biochemical traits: Ψ_pd, g_s, polyphenol oxidase, superoxide dismutase and malonyldialdehyde. Inland ecotypes seem to be more tolerant to drought stress than coastal ecotypes.     

Tree seedling responses to in situ CO2-enrichment differ among species and depend on understorey light availability

Seedlings of six major European temperate forest tree species (Fagus sylvatica, Acer pseudoplatanus, Quercus robur, Taxus baccata, Abies alba, Pinus sylvestris) were exposed to 360, 500, and 660 μL CO₂ L^?1 in the understorey of a 120‐y‐old forest over two growing seasons. Seedlings rooted in the natural forest soil within 36 open‐top chambers (12 OTCs per CO₂ treatment), each with a different known quantum flux density (QFD) ranging from 0.36 to 2.16 mol m^?2 d^?1 (= 0.8% to 4.8% of full sun). In contrast to a frequent assumption the natural CO₂ concentration in the understorey is close to the ambient concentration in the free atmosphere during daytime. The CO₂‐effect on seedling growth differed greatly among species and was strongly codetermined by microsite‐specific QFD. Biomass production in the deep‐shade tolerant species Fagus and Taxus increased by 73% and 37% under elevated CO₂ in low QFD microsites but was not significantly different among CO₂‐treatments in high QFD microsites. The less shade‐tolerant species Acer, Quercus, and Abies showed no significant response to elevated CO₂ in low QFD microsites, but increased their biomass by 39%, 25%, and 55% in high QFD microsites. In the shade‐intolerant Pinus, seedling survival was too low for a safe conclusion. Our data showed that the largest relative responses to increasing CO₂ occurred at a comparatively small increase from 360 to 500 μL L^?1 with only small and non‐significant changes with a further increase to 660 μL L^?1. Subtle shifts in the availability of light can totally reverse interspecific differences in the CO₂ response. Given these different responses, we conclude that increasing atmospheric CO₂ is likely to induce changes in species composition of temperate forests due to altered chances of recruitment. However, these shifts will depend on light patterns in the understorey, and thus on canopy structure, disturbance patterns and forest management.     

Winners always win: growth of a wide range of plant species from low to future high CO2

Evolutionary adaptation to variation in resource supply has resulted in plant strategies that are based on trade‐offs in functional traits. Here, we investigate, for the first time across multiple species, whether such trade‐offs are also apparent in growth and morphology responses to past low, current ambient, and future high CO₂ concentrations. We grew freshly germinated seedlings of up to 28 C₃ species (16 forbs, 6 woody, and 6 grasses) in climate chambers at 160 ppm, 450 ppm, and 750 ppm CO₂. We determined biomass, allocation, SLA (specific leaf area), LAR (leaf area ratio), and RGR (relative growth rate), thereby doubling the available data on these plant responses to low CO₂. High CO₂ increased RGR by 8%; low CO₂ decreased RGR by 23%. Fast growers at ambient CO₂ had the greatest reduction in RGR at low CO₂ as they lost the benefits of a fast‐growth morphology (decoupling of RGR and LAR [leaf area ratio]). Despite these shifts species ranking on biomass and RGR was unaffected by CO₂, winners continued to win, regardless of CO_2. Unlike for other plant resources we found no trade‐offs in morphological and growth responses to CO₂ variation, changes in morphological traits were unrelated to changes in growth at low or high CO₂. Thus, changes in physiology may be more important than morphological changes in response to CO₂ variation.     

Climate and CO2 controls on global vegetation distribution at the last glacial maximum: analysis based on palaeovegetation data, biome modelling and palaeoclimate simulations

The global vegetation response to climate and atmospheric CO₂ changes between the last glacial maximum and recent times is examined using an equilibrium vegetation model (BIOME4), driven by output from 17 climate simulations from the Palaeoclimate Modelling Intercomparison Project. Features common to all of the simulations include expansion of treeless vegetation in high northern latitudes; southward displacement and fragmentation of boreal and temperate forests; and expansion of drought‐tolerant biomes in the tropics. These features are broadly consistent with pollen‐based reconstructions of vegetation distribution at the last glacial maximum. Glacial vegetation in high latitudes reflects cold and dry conditions due to the low CO₂ concentration and the presence of large continental ice sheets. The extent of drought‐tolerant vegetation in tropical and subtropical latitudes reflects a generally drier low‐latitude climate. Comparisons of the observations with BIOME4 simulations, with and without consideration of the direct physiological effect of CO₂ concentration on C₃ photosynthesis, suggest an important additional role of low CO₂ concentration in restricting the extent of forests, especially in the tropics. Global forest cover was overestimated by all models when climate change alone was used to drive BIOME4, and estimated more accurately when physiological effects of CO₂ concentration were included. This result suggests that both CO₂ effects and climate effects were important in determining glacial‐interglacial changes in vegetation. More realistic simulations of glacial vegetation and climate will need to take into account the feedback effects of these structural and physiological changes on the climate.     

Are functional traits a good predictor of global change impacts on tree species abundance dynamics in a subtropical forest?

Significant changes in the composition of tree species have been observed in various forests worldwide. We hypothesised that these changes might result from variable sensitivities of species to global change, and species sensitivities might be quantified, using functional traits. Employing long‐term (1978–2010) species abundance data of 48 tree species from a permanent subtropical forest plot, where multiple global change factors have been observed, including soil drying, we examined the relationships between temporal trends in abundance and suits of functional traits. We found that species with high photosynthesis rates, leaf phosphorus and nitrogen concentrations, specific leaf area, hydraulic conductivity, turgor loss point and predawn leaf water potential had increased in abundance, while species with opposite trait patterns had decreased. Our results demonstrate that functional traits underlie tree species abundance dynamics in response to drought stress, thus linking traits to compositional shifts in this subtropical forest under global changes.     

Elevated [CO2] does not ameliorate the negative effects of elevated temperature on drought‐induced mortality in Eucalyptus radiata seedlings

It has been reported that elevated temperature accelerates the time‐to‐mortality in plants exposed to prolonged drought, while elevated [CO₂] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO₂] and temperature on the inter‐dependent carbon and hydraulic characteristics associated with drought‐induced mortality in Eucalyptus radiata seedlings grown in two [CO₂] (400 and 640 μL L^?1) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO₂] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO₂], may be the primary contributors to drought‐induced seedling mortality under future climates.     

The combined and separate impacts of climate extremes on the current and future US rainfed maize and soybean production under elevated CO2

Heat and drought are two emerging climatic threats to the US maize and soybean production, yet their impacts on yields are collectively determined by the magnitude of climate change and rising atmospheric CO₂ concentrations. This study quantifies the combined and separate impacts of high temperature, heat and drought stresses on the current and future US rainfed maize and soybean production and for the first time characterizes spatial shifts in the relative importance of individual stress. Crop yields are simulated using the Agricultural Production Systems Simulator (APSIM), driven by high‐resolution (12 km) dynamically downscaled climate projections for 1995–2004 and 2085–2094. Results show that maize and soybean yield losses are prominent in the US Midwest by the late 21st century under both Representative Concentration Pathway (RCP) 4.5 and RCP8.5 scenarios, and the magnitude of loss highly depends on the current vulnerability and changes in climate extremes. Elevated atmospheric CO₂ partially but not completely offsets the yield gaps caused by climate extremes, and the effect is greater in soybean than in maize. Our simulations suggest that drought will continue to be the largest threat to US rainfed maize production under RCP4.5 and soybean production under both RCP scenarios, whereas high temperature and heat stress take over the dominant stress of drought on maize under RCP8.5. We also reveal that shifts in the geographic distributions of dominant stresses are characterized by the increase in concurrent stresses, especially for the US Midwest. These findings imply the importance of considering heat and drought stresses simultaneously for future agronomic adaptation and mitigation strategies, particularly for breeding programs and crop management. The modeling framework of partitioning the total effects of climate change into individual stress impacts can be applied to the study of other crops and agriculture systems.     

Suitability of Drought-Preconditioning Techniques in Mediterranean Climate

Abstract Arid and semiarid ecosystems in the Mediterranean are under high risk of desertification. Revegetation with native well‐adapted evergreen shrubs is desirable, but techniques for successful establishment of these species are not fully developed. Transplant shock is a key hurdle to plantation success. The application of a drought‐preconditioning treatment during the last months of nursery culture is a potential technique for reducing transplant shock. This technique has been widely applied in boreal habitats and humid temperate areas. Three representative Mediterranean species (Pistacia lentiscus, Quercus coccifera, and Juniperus oxycedrus seedlings) were exposed to classic drought‐preconditioning treatment consisting of reductions in the watering regime. The effects of preconditioning on seedling quality were assessed by cell water relationships (pressure–volume curves), minimal transpiration, leaf capacitance, chlorophyll fluorescence, and gas exchange. Moreover, seedlings were exposed to transplant shock (intense drought period) during which water potential (predawn and midday) and maximal photochemical efficiency were evaluated to establish seedling performance. Results showed that preconditioning did not affect cell water relationships and minimal transpiration in any of the three species. Preconditioned seedlings of P. lentiscus maintained higher water content during desiccating conditions as a consequence of an increase in leaf water content at full turgor. These changes allowed plants to maintain higher net CO₂ assimilation rates and an elevated photosystem II status, facilitating an increase in drought survival. Preconditioning improved the performance of Q. coccifera and J. oxycedrus seedlings, but these two species were much less responsive than P. lentiscus seedlings. Finally, results suggest that sensitivity to drought preconditioning may be related to drought tolerance or avoidance strategy of each species. Drought‐related strategies should be considered to optimize management scale preconditioning.     

Water use strategy affects avoidance of ozone stress by stomatal closure in Mediterranean trees—A modelling analysis

Both ozone (O₃) and drought can limit carbon fixation by forest trees. To cope with drought stress, plants have isohydric or anisohydric water use strategies. Ozone enters plant tissues through stomata. Therefore, stomatal closure can be interpreted as avoidance to O₃ stress. Here, we applied an optimization model of stomata involving water, CO₂, and O₃ flux to test whether isohydric and anisohydric strategies may affect avoidance of O₃ stress by stomatal closure in four Mediterranean tree species during drought. The data suggest that stomatal closure represents a response to avoid damage to the photosynthetic mechanisms under elevated O₃ depending on plant water use strategy. Under high-O₃ and well-watered conditions, isohydric species limited O₃ fluxes by stomatal closure, whereas anisohydric species activated a tolerance response and did not actively close stomata. Under both O₃ and drought stress, however, anisohydric species enhanced the capacity of avoidance by closing stomata to cope with the severe oxidative stress. In the late growing season, regardless of the water use strategy, the efficiency of O₃ stress avoidance decreased with leaf ageing. As a result, carbon assimilation rate was decreased by O₃ while stomata did not close enough to limit transpirational water losses.     

How do herbivorous insects respond to drought stress in trees?

Increased frequency and severity of drought, as a result of climate change, is expected to drive critical changes in plant–insect interactions that may elevate rates of tree mortality. The mechanisms that link water stress in plants to insect performance are not well understood. Here, we build on previous reviews and develop a framework that incorporates the severity and longevity of drought and captures the plant physiological adjustments that follow moderate and severe drought. Using this framework, we investigate in greater depth how insect performance responds to increasing drought severity for: (i) different feeding guilds; (ii) flush feeders and senescence feeders; (iii) specialist and generalist insect herbivores; and (iv) temperate versus tropical forest communities. We outline how intermittent and moderate drought can result in increases of carbon‐based and nitrogen‐based chemical defences, whereas long and severe drought events can result in decreases in plant secondary defence compounds. We predict that different herbivore feeding guilds will show different but predictable responses to drought events, with most feeding guilds being negatively affected by water stress, with the exception of wood borers and bark beetles during severe drought and sap‐sucking insects and leaf miners during moderate and intermittent drought. Time of feeding and host specificity are important considerations. Some insects, regardless of feeding guild, prefer to feed on younger tissues from leaf flush, whereas others are adapted to feed on senescing tissues of severely stressed trees. We argue that moderate water stress could benefit specialist insect herbivores, while generalists might prefer severe drought conditions. Current evidence suggests that insect outbreaks are shorter and more spatially restricted in tropical than in temperate forests. We suggest that future research on the impact of drought on insect communities should include (i) assessing how drought‐induced changes in various plant traits, such as secondary compound concentrations and leaf water potential, affect herbivores; (ii) food web implications for other insects and those that feed on them; and (iii) interactions between the effects on insects of increasing drought and other forms of environmental change including rising temperatures and CO₂ levels. There is a need for larger, temperate and tropical forest‐scale drought experiments to look at herbivorous insect responses and their role in tree death.     

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.