Publications

Survival and growth of woody species in the Mediterranean are mainly restricted by water availability. We tested the hypothesis that Mediterranean species acclimate their xylem vulnerability and osmotic potential along a precipitation gradient. We studied five predominant co-occurring Mediterranean species; Quercus calliprinos, Pistacia palaestina, Pistacia lentiscus, Rhamnus lycioides and Phillyrea latifolia, over two summers at three sites. The driest of the sites is the distribution edge for all the five species. We measured key hydraulic and osmotic traits related to drought resistance, including resistance to embolism (Ψ50) and the seasonal dynamics of water and osmotic potentials. The leaf water potentials (Ψl) of all species declined significantly along the summer, reaching significantly lower Ψl at the end of summer in the drier sites. Surprisingly, we did not find plasticity along the drought gradient in Ψ50 or osmotic potentials. This resulted in much narrower hydraulic safety margins (HSMs) in the drier sites, where some species experienced significant embolism. Our analysis indicates that reduction in HSM to null values put Mediterranean species in embolism risk as they approach their hydraulic limit near the geographical dry edge of their distribution. Read the free Plain Language Summary for this article on the Journal blog.

Understanding the causes of the arrest of species distributions has been a fundamental question in ecology and evolution. These questions are of particular interest for trees owing to their long lifespan and sessile nature. A surge in data availability evokes a macro-ecological analysis to determine the underlying forces limiting distributions. Here we analyse the spatial distribution of >3,600 major tree species to determine geographical areas of range-edge hotspots and find drivers for their arrest. We confirmed biome edges to be strong delineators of distributions. Importantly, we identified a stronger contribution of temperate than tropical biomes to range edges, adding strength to the notion that tropical areas are centres of radiation. We subsequently identified a strong association of range-edge hotspots with steep spatial climatic gradients. We linked spatial and temporal homogeneity and high potential evapotranspiration in the tropics as the strongest predictors of this phenomenon. We propose that the poleward migration of species in light of climate change might be hindered because of steep climatic gradients.

Cavitation resistance has often been viewed as a relatively static trait, especially for stems of forest trees. Meanwhile, other hydraulic traits, such as turgor loss point (psi(tlp)) and xylem anatomy, change during the season. In this study, we hypothesized that cavitation resistance is also dynamic, changing in coordination with psi(tlp). We began with a comparison of optical vulnerability (OV), microcomputed tomography (mu CT) and cavitron methods. All three methods significantly differed in the slope of the curve,psi(12) and psi(88), but not in psi(50) (xylem pressures that cause 12%, 88%, 50% cavitation, respectively). Thus, we followed the seasonal dynamics (across 2 years) of psi(50) in Pinus halepensis under Mediterranean climate using the OV method. We found that psi(50) is a plastic trait with a reduction of approximately 1 MPa from the end of the wet season to the end of the dry season, in coordination with the dynamics of the midday xylem water potential (psi(midday)) and the psi(tlp). The observed plasticity enabled the trees to maintain a stable positive hydraulic safety margin and avoid cavitation during the long dry season. Seasonal plasticity is vital for understanding the actual risk of cavitation to plants and for modeling species' ability to tolerate harsh environments.

Abstract Deforestation of tropical forests has been a critical issue affecting climate change mitigation and biodiversity conservation. Reforestation strives to remedy this situation, yet it is futile as long as deforestation of primary forests continues. Since deforestation is partly motivated by the demand for valuable tropical wood, reforestation should focus not only on planting native tree species, but specifically on a high diversity of native tree species with high-quality wood. However, the eco-physiological information required for growing such species is limited, and their resilience to drought events is unknown. Here, we focused on four native tropical wood tree species identified as suitable for Brazil’s Atlantic Forest reforestation. Growth, carbon assimilation, water-use and xylem hydraulics were studied in seedlings of the two legume species Dalbergia nigra and Plathymenia foliolosa and the two non-legume species Cariniana legalis and Zeyheria tuberculosa. Seedlings were monitored weekly for 9 consecutive weeks, three to five weeks of which under induced drought. Growth and carbon assimilation were 25–65 per cent higher in the legume vs. non-legume species. In turn, non-legume species mostly avoided the drought by stomatal closure, producing a 50 per cent higher water-use efficiency (WUE) compared with the legume species. The average water potential at 50 per cent stomatal conductivity (Ψgs50) for legume species was −2.6 MPa, whereas for non-legume species it was −0.85 MPa. Still, each species showed a unique set of responses, indicating different growth strategies under mesic and xeric conditions. Our results indicate a divergence among legume and non-legume species, driven by a trade-off between plant productivity (carbon assimilation and growth) and plant safety (stomatal regulation and WUE). All in all, the four species of juvenile potted plants demonstrated a high capacity for recovery from drought, which supports their potential role in future reforestation under climate change.

Xylem embolism impairs hydraulic conductivity in trees and drives drought-induced mortality. While embolism has been monitored in vivo in potted plants, and research has revealed evidence of embolism in field-grown trees, continuous in situ monitoring of cavitation in forests is lacking. Seasonal patterns of embolism were monitored in branchlets of Aleppo pine (Pinus halepensis) trees growing in a dry Mediterranean forest. Optical visualization (OV) sensors were installed on terminal branches, in addition to monthly sampling for micro computed tomography scans. We detected 208 cavitation events among four trees, which represented an embolism increase from zero to c. 12% along the dry season. Virtually all the cavitation events occurred during daytime hours, with 77% occurring between 10:00 and 17:00 h. The probability for cavitation in a given hour increased as vapor pressure deficit (VPD) increased, up to a probability of 42% for cavitation when VPD > 5 kPa. The findings uniquely reveal the instantaneous environmental conditions that lead to cavitation. The increased likelihood of cavitation events under high VPD in water-stressed pines is the first empirical support for this long hypothesized relationship. Our observations suggest that low levels of embolism are common in Aleppo pine trees at the dry edge of their distribution.

Montane treelines are defined by a threshold low temperature. However, what are the dynamics when the snow-free summer growth season coincides with a 6-month seasonal drought?

We tested this fundamental question by measuring tree growth and leaf activity across elevations in Mt. Hermon (2,814 m; in Israel and Syria), where oak trees (Quercus look and Q. boissieri) form an observed treeline at 1900 m.

While in theory, individuals can be established at higher elevations (minimum daily temperature > 6.5°C for > 4 months even at the summit), soil drying and vapor pressure deficit (VPD) in summer enforces growth cessation in August, leaving only 2–3 months for tree growth. At lower elevations, Q. look is replaced by Q. cerris (1,300 m) and Q. calliprinos (1,000 m) in accompanying Q. boissieri, and growth season length (GSL) is higher due to an earlier start in April. Leaf gas exchange continues during autumn, but assimilates are no longer utilized in growth. Interestingly, the growth and activity of Q. boissieri were equivalent to that of each of the other three species across the ~ 1 km elevation gradient. A planting experiment at 2100 m showed that seedlings of the four oak species survived the cold winter and showed budding of leaves in summer, but wilted in August.

Our unique mountain site in the Eastern Mediterranean introduces a new factor to the formation of treelines, involving a drought limitation on GSL. This site presents the elevation edge for each species and the southern distribution edge for both the endemic Q. look and the broad-range Q. cerris. With ongoing warming, Q. look and Q. boissieri are slowly expanding to higher elevations, while Q. cerris is at risk of future extirpation.

Earth's forests face grave challenges in the Anthropocene, including hotter droughts increasingly associated with widespread forest die-off events. But despite the vital importance of forests to global ecosystem services, their fates in a warming world remain highly uncertain. Lacking is quantitative determination of commonality in climate anomalies associated with pulses of tree mortality-from published, field-documented mortality events-required for understanding the role of extreme climate events in overall global tree die-off patterns. Here we established a geo-referenced global database documenting climate-induced mortality events spanning all tree-supporting biomes and continents, from 154 peer-reviewed studies since 1970. Our analysis quantifies a global "hotter-drought fingerprint" from these tree-mortality sites-effectively a hotter and drier climate signal for tree mortality-across 675 locations encompassing 1,303 plots. Frequency of these observed mortality-year climate conditions strongly increases nonlinearly under projected warming. Our database also provides initial footing for further community-developed, quantitative, ground-based monitoring of global tree mortality.

The current projections of climate change might exceed the ability of European forest trees to adapt to upcoming environmental conditions. However, stomatal and leaf morphological traits could greatly influence the acclimation potential of forest tree species subjected to global warming, including the single most important forestry species in Europe, European beech. We analysed stomatal (guard cell length, stomatal density and potential conductance index) and leaf (leaf area, leaf dry weight and leaf mass per area) morphological traits of ten provenances from two provenance trials with contrasting climates between 2016 and 2020. The impact of meteorological conditions of the current and preceding year on stomatal and leaf traits was tested by linear and quadratic regressions. Ecodistance was used to capture the impact of adaptation after the transfer of provenances to new environments. Interactions of trial-provenance and trial-year factors were significant for all measured traits. Guard cell length was lowest and stomatal density was highest across beech provenances in the driest year, 2018. Adaptation was also reflected in a significant relationship between aridity ecodistance and measured traits. Moreover, the meteorological conditions of the preceding year affected the interannual variability of stomatal and leaf traits more than the meteorological conditions of the spring of the current year, suggesting the existence of plant stress memory. High intraspecific variability of stomatal and leaf traits controlled by the interaction of adaptation, acclimation and plant memory suggests a high acclimation potential of European beech provenances under future conditions of global climate change.

The ongoing global warming and associated drying are shaping the fate of forests worldwide. While processes of tree mortality are visible and studied, a decrease in forest regeneration is mostly overlooked, although equally deleterious. Populations at the edge of tree species distribution areas are at higher risk and are hence hotspots for species extinctions. Here we use a semi-arid pine forest growing at the timberline edge of forest existence as a model for forest survival under warming and drying conditions. Seedling recruitment, including seed germination, seedling survivorship, and multiyear seedling growth, were measured along six consecutive years. To pinpoint the role of drought, we designed a field experiment, manipulating stand density at three levels and grazing regimes. Seed germination was high across all studied plots, but seedling survivorship and multiyear seedling growth were near-zero. Stand density and grazing exclusion positively affected germination. Seedling survivorship was higher in wetter years. Multiyear seedling growth was stunted by grazing, and seedling height was distributed differently across different stand densities. Our data indicate that seedling survivorship during the first dry season acts as a bottleneck for forest existence at the dry and hot edge of current forest distribution. We also quantified the roles of other stressors such as shading, and highlighted the eliminating role of grazing on multiyear seedling growth. Forest regeneration should be more closely monitored in sensitive populations, as climate change-driven forest loss can happen even without mature tree mortality.

Although atmospheric CO2 concentration ([CO2]) continues to rise, the question of how tree carbon (C) allocation is affected by this change remains. Studies show that C assimilation increases under elevated CO2 (eCO(2)). Yet, no detailed study has determined the fate of the surplus C, i.e., its compartment and physiological process allocation, nor in multiple species together. In this project, we grew 2-year-old saplings of four key Mediterranean tree species (the conifers Cupressus sempervirens L. and Pinus halepensis Mill., and the broadleaf Quercus calliprinos Webb. and Ceratonia siliqua L.) to [CO2] levels of 400 or 700 p.p.m. for 6 months. We measured the allocation of C to below and aboveground growth, respiration, root exudation, storage and leaf litter. In addition, we monitored intrinsic water-use efficiency (WUE), soil moisture, soil chemistry and nutrient uptake. Net assimilation, WUE and soil nitrogen uptake significantly increased at eCO(2) across the four species. Broadleaf species showed soil water savings, which were absent in conifers. All other effects were species-specific: Cupressus had higher leaf respiration, Pinus had lower starch in branches and transiently higher exudation rate and Quercus had higher root respiration. Elevated CO2 did not affect growth or litter production. Our results are pivotal to understanding the sensitivity of tree C allocation to the change in [CO2] when water is abundant. Species-specific responses should be regarded cautiously when predicting future changes in forest function in a higher CO2 world.

Carbon (C) allocation in trees involves three different aspects, namely, the fate of C in terms of tree compartment, C compound, and physiological process (often termed C flux). A review and synthesis of the recent literature on C allocation dynamics in Mediterranean pines, with an emphasis on stress conditions, reveals some important patterns. First, C allocation to stem growth is highly plastic in phenology and magnitude, whereas C allocation to needle growth is sensitive to growing conditions (e.g. drought, competition), but genetically programmed with respect to timing and magnitude. Second, starch is a major C reserve in Mediterranean pines, providing C under limited photosynthesis for weeks and months, replenished during recovery before growth is resumed, and stored at high concentrations in the root system. Third, C respiration is dominated by the roots, while growth and litter production are dominated by the foliage. Relocation fluxes occur under summer drought, i.e. from stem to foliage, and from roots to stem. Although minor in magnitude, these C fluxes might represent an important drought acclimation mechanism for pines in the Mediterranean.

It has been assumed that mixing of species with high physiological diversity reduces competition over water and light resources, compared to single-species forests. Although several mechanisms to explain this observation have been proposed, quantification of these effects is lacking. Here we studied water-use dynamics for five tree species in a mature, mixed, evergreen, and Mediterranean forest. We use empirical measurements of key tree structural attributes including root distribution, through DNA barcoding and soil cores, tree height and biomass along with measurements of species-specific water use for two years. These measurements at the tree-scale were used to parameterize an ecosystem model of coupled water, carbon and energy fluxes (Regional Hydro Ecologic Simulation System, RHESSys). Site-scale empirical measurements showed contrasting diurnal and seasonal transpiration and sap flow curves across tree species, with year-round activity in angiosperms, and mostly wet season-activity in gymnosperms. Water-use patterns matched the rooting depth patterns, with the deep- and shallow-rooted Ceratonia and Cupressus, showing year-round and seasonal behaviors, respectively. RHESSys estimates of species-specific and stand-scale transpiration, biomass and productivity across 20 years of climate variation showed substantial differences between mixed and monoculture scenarios. Stand-scale annual net primary productivity and transpiration increased by 20–70 g C m−1 yr−1 and 40–80 mm yr−1, respectively, for mixed stands relative to average fluxes aggregated across monocultures. Model results, collaborated by field data provide evidence for niche partitioning of the soil water resource among co-habiting tree species, and demonstrate that this mechanism can facilitate higher productivity and an enhanced forest carbon sink especially in semi-arid regions.

•Tree C allocation is estimated by either isotopic or mass balance approaches.•We combine these approaches to study C allocation in five evergreen treespecies.•Within 8 days, conifers allocated 30% of the C belowground, and broadleaves 

The high-mountain ecosystems of Central Asia are a biodiversity hotspot with unique plant communities and many endemic species. Intense human pressure and global warming have caused habitat destruction in these areas and a parallel increase in the number of endangered species. Lagochilus species are key medicinal herbaceous plants native to Central Asia, many of which have been recently added to the endangered of species in Uzbekistan. To assess the climate sensitivity of Lagochilus species, we (1) located populations of six species in their native sites across Uzbekistan, and assessed their health by partitioning to ontogenetic stages along five consecutive years; (2) collected plant materials from these species, as well as from old herbarium samples (1918–1964); and (3) analyzed the carbon-13 composition in those samples, as an indicator for drouht stress. Over the course of five years (2014–2018) of continuous monitoring, fluctuations in annual precipitation in the region indicated a decrease by ∼20 %, and the fraction of young plants in each population decreased from 20–50% to 0–5 %, depending on the species. Comparing the carbon-13 composition in current and historical leaf samples showed an increase of 1.5–3.5‰ associated with a decrease in precipitation of 2–30 %, depending on the site and species. Our results show the high sensitivity of Lagochilus populations’ regeneration to drying, among six species and in sites across Uzbekistan. On a multi-decadal temporal scale, the dramatic changes in carbon-13 indicate that the response to precipitation reduction is related with drought stress. Considering the expectation for drier and hotter climate in Uzbekistan in the coming decades, conservation of Lagochilus populations should become a priority in Central Asia.

The intimate connection between roots and soil particles is a prerequisite for the continuous flow of water and nutrients from soil to trees, and carbon flow from roots to the rhizosphere. The soil-root interface has been studied in multiple laboratory and greenhouse experiments. Yet its multiple roles and their differential contributions to tree health have rarely been experimentally studied on mature trees in the field. We took advantage of mature olive tree transplanting to test the physiological effects of breakage of the soil-root interface in situ. Eight olive trees were monitored following transplantation into a site located 4 km from their native site, along two years. Additional eight trees were monitored simultaneously at the native site, as control. To decrease mortality risk, transplanted trees were heavily pruned before transplanting, and were irrigated and fertilized in their new site. Transplanted trees had ~50% lower rates of leaf photosynthesis and transpiration; ~80% lower root starch content; and ~30% higher loss of xylem conductivity, than native trees. Leaf water potential (LWP) was similar across trees, becoming more negative in the transplanted trees only in the second year following transplanting. While starch content and xylem conductivity recovered in the second year, leaf gas exchange and LWP remained significantly lower than in native trees. Soil P and K were higher under transplanted trees that remained stressed than under trees that recovered. Breakage of the soil-root interface caused a multi-system stress to trees. The lack of persistent LWP response might indicate that loss of hydraulic conductivity was driven by root, rather than aboveground, embolisms. Degradation of starch in the roots indicates an increase in belowground sinks. In the long run, recovery of starch content means no carbon limitation; yet the prevailing effects on leaf activities suggest a long-term stress unrelated with water or carbon supply.

Root exudates are part of the rhizodeposition process, which is the major source of soil organic carbon (C) released by plant roots. This flux of C is believed to have profound effects on C and nutrient cycling in ecosystems. The quantity of root exudates depends on the plant species, the period throughout the year, and external biotic and abiotic factors. Since root exudates of mature trees are difficult to collect in field conditions, very little is known about their flux, especially in water-limited ecosystems, such as the seasonally hot and dry Mediterranean maquis. Here, we collected exudates from DNA-identified roots in the forest from the gymnosperm Cupressus sempervirens L. and the evergreen angiosperm Pistacia lentiscus L. by 48-h incubations on a monthly temporal resolution throughout the year. We examined relationships of the root exudate C flux to abiotic parameters of the soil (water content, water potential, temperature) and atmosphere (vapor pressure deficit, temperature). We also studied relationships to C fluxes through the leaves as indicators of tree C balance. Root exudation rates varied significantly along the year, increasing from 6 μg C cm -2 root day-1 in both species in the wet season to 4- and 11-fold rates in Pistacia and Cupressus, respectively, in the dry season. A stepwise linear mixed-effects model showed that the three soil parameters were the most influential on exudation rates. Among biotic factors, there was a significant negative correlation of exudation rate with leaf assimilation in Cupressus and a significant negative correlation with leaf respiration in Pistacia. Our observation of enhanced exudation flux during the dry season indicates that exudation dynamics in the field are less sensitive to the low tree C availability in the dry season. The two key Mediterranean forest species seem to respond to seasonal changes in the rhizosphere such as drying and warming, and therefore invest C in the rhizosphere under seasonal drought.

Carbon reserve use is a major drought response in trees, enabling tree survival in conditions prohibiting photosynthesis. However, regulation of starch metabolism under drought at the whole-tree scale is still poorly understood. To this end, we combined measurements of nonstructural carbohydrates (NSCs), tree physiology and gene expression. The experiment was conducted outside on olive trees in pots under 90 d of seasonal spring to summer warming. Half of the trees were also subjected to limited water conditions for 28 d. Photosynthesis decreased in dehydrating trees from 19 to 0.5 µmol m−2 s−1 during the drought period. Starch degradation and mannitol production were a major drought response, with mannitol increasing to 71% and 41% out of total NSCs in shoots and roots, respectively. We identified the gene family members potentially relevant either to long-term or stress-induced carbon storage. Partitioning of expression patterns among β amylase and starch synthase family members was observed, with three β amylases possibly facilitating the rapid starch degradation under heat and drought. Our results suggest a group of stress-related, starch metabolism genes, correlated with NSC fluctuations during drought and recovery. The daily starch metabolism gene expression was different from the stress-mode starch metabolism pattern, where some genes are uniquely expressed during the stress-mode response.

The ongoing increase in human population and the subsequent freshwater demands raise conflicts regarding conservation of riparian ecosystems. Identifying anthropogenic effects on these vulnerable nature resources is crucial for preventing future damages. Here we measured tree-ring width and isotopic carbon composition (delta C-13) in stem wood of protected Platanus orientalis trees at the Kziv Nature Reserve, a Mediterranean riparian ecosystem in northern Israel. In this reserve, water was pumped for human needs during 1976-2006 from the major spring feeding the stream. We show a negative exponential correlation between stem growth and pumped water amounts. Spring drawdown had a significant negative effect on growth even in years when amounts were reduced to similar to 2.10(6)m(3) year(-1), leveling off at around 5.10(6) m(3) year(-1). Precipitation and spring volume effects on growth were exposed only after pumping stopped, further indicating its role in inhibiting tree growth. During the pumping years, stem wood delta C-13 was up to 1.8%0 higher than after pumping cessation, indicating the drought stress imposed on trees. Our results provide an unequivocal evidence for the adverse effect of water pumping on riparian Platanus orientalis tree growth. Such effects, and their related tree mortality risk, must be considered in sustainable water management planning.

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives.

The mutualistic symbiosis between forest trees and ectomycorrhizal fungi (EMF) is among the most ubiquitous and successful interactions in terrestrial ecosystems. Specific species of EMF are known to colonize specific tree species, benefitting from their carbon source, and in turn, improving their access to soil water and nutrients. EMF also form extensive mycelial networks that can link multiple root-tips of different trees. Yet the number of tree species connected by such mycelial networks, and the traffic of material across them, are just now under study. Recently we reported substantial belowground carbon transfer between Picea, Pinus, Larix and Fagus trees in a mature forest. Here, we analyze the EMF community of these same individual trees and identify the most likely taxa responsible for the observed carbon transfer. Among the nearly 1,200 EMF root-tips examined, 50%-70% belong to operational taxonomic units (OTUs) that were associated with three or four tree host species, and 90% of all OTUs were associated with at least two tree species. Sporocarp C-13 signals indicated that carbon originating from labelled Picea trees was transferred among trees through EMF networks. Interestingly, phylogenetically more closely related tree species exhibited more similar EMF communities and exchanged more carbon. Our results show that belowground carbon transfer is well orchestrated by the evolution of EMFs and tree symbiosis.

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.

Water availability is becoming a limiting factor with increasing world population that challenges global food security. Thus, we need to enhance cultivation in increasingly drier and hotter climate and prepare fruit trees for the ongoing climate change. Wild tree species might offer vital information and plant material in face of these challenges.A year-long comparative field study was conducted to investigate the mechanisms underlying drought tolerance in pear species (cultivated Pyrus communis and Pyrus pyrifolia vs. the wild Pyrus syriaca).We confirmed the hypothesis of higher drought tolerance in wild pear compared to its cultivated relative. P. syriaca xylem had fewer, narrower vessels, and lower vulnerability to embolism. It showed higher intrinsic water-sue efficiency and more robust seasonal patterns of photosynthesis, hydraulic conductivity, and PIP (plasma intrinsic protein) aquaporin expression. Across species, we identified a ubiquitous gene (PIP1:5/1:6), nine drought-inhibited genes, and two drought-induced genes (PIP1:4 and 2:6/2:7, confirming previous studies).Our study highlights the potential of using wild relatives of fruit tree species to prepare key crops to a drier and hotter future. The study of PIPS leads the way to a more focused research of the role of these cellular water channels in minimizing tree water loss under drought, while ensuring hydration of specific tissues.

The response of tree leaf gas exchange to elevated CO2 concentrations has been investigated in numerous experiments along the past 30 years. Stomatal regulation is a major plant control over leaf gas exchange, and the response to the increasing CO2 will shape the biological activity of forests in the future. Here, we collected 144 records from 57 species on stomatal conductance in CO2 manipulation experiments on trees (340-980 ppm CO2). CO2-induced stomatal downregulation was calculated as the slope of the linear regression between stomatal conductance and [CO2]. Among tree species, the slope (a) of change in stomatal conductance per 100 ppm CO(2 )increase ranged between 0 and -151, indicating stomatal downregulation, and only four species showed upregulation. There was a significant divergence between evergreen gymnosperms (a = -3.6 +/- 1.0), deciduous angiosperms (a = -16.3 +/- 3.1) and evergreen angiosperms (a = -32.8 +/- 7.1). Gymnosperms were less sensitive to CO2 changes than deciduous angiosperms even when considering only field experiments. The significant role of tree functional group in predicting CO2-induced stomatal downregulation was detected in multiple mixed-effect models, with p values ranging between 0.0002 and 0.0295. The significantly higher stomatal sensitivity to CO2 of angiosperms versus gymnosperms might be related to the overall higher stomatal conductance of angiosperms; their thinner leaves, in turn losing water faster; and the decreasing atmospheric [CO2] at the time of their taxa diversification. We conclude that species differences must be taken into account in forecasting future forest fluxes. A plain language summary is available for this article.

Drought-related tree mortality had become a widespread phenomenon in forests around the globe. This process leading to these events and its complexity is not fully understood. Trees in the dry timberline are exposed to ongoing drought, and the available water for transpiration in the soil can determine their survival chances. Recent drought years led to 5%–10% mortality in the semi-arid pine forest of Yatir (Israel). The distribution of dead trees was, however, highly heterogeneous with parts of the forest showing >80% dead trees (D plots) and others with mostly live trees (L plots). At the tree level, visible stress was associated with low pre-dawn leaf water potential at the dry season (−2.8 MPa vs. −2.3 MPa in non-stressed trees), shorter needles (5.5 vs. 7.7 mm) and lower chlorophyll content (0.6 vs. 1 mg/g dw). Trends in tree-ring widths reflected differences in stress intensity (30% narrower rings in stressed compared with unstressed trees), which could be identified 15–20 years prior to mortality. At the plot scale, no differences in topography, soil type, tree age or stand density could explain the mortality difference between the D and L plots. It could only be explained by the higher surface rock cover and in stoniness across the soil profile in the L plots. Simple bucket model simulations using the site’s long-term hydrological data supported the idea that these differences could result in higher soil water concentration (m 3 /m 3 ) in the L plots and extend the time above wilting point by several months across the long dry season. Accounting for subsurface heterogeneity may therefore critical to assessing stand-level response to drought and projecting tree survival, and can be used in management strategies in regions undergoing drying climate trends. 

Understanding which species are able to recover from drought, under what conditions, and the mechanistic processes involved, will facilitate predictions of plant mortality in response to global change. In response to drought, some species die because of embolism-induced hydraulic failure, whilst others are able to avoid mortality and recover, following rehydration. Several tree species have evolved strategies to avoid embolism, whereas others tolerate high embolism rates but can recover their hydraulic functioning upon drought relief. Here, we focus on structures and processes that might allow some plants to recover from drought stress via embolism reversal. We provide insights into how embolism repair may have evolved, anatomical and physiological features that facilitate this process, and describe possible trade-offs and related costs. Recent controversies on methods used for estimating embolism formation/repair are also discussed, providing some methodological suggestions. Although controversial, embolism repair processes are apparently based on the activity of phloem and ray/axial parenchyma. The mechanism is energetically demanding, and the costs to plants include metabolism and transport of soluble sugars, water and inorganic ions. We propose that embolism repair should be considered as a possible component of a hydraulic efficiency-safety' spectrum. We also advance a framework for vegetation models, describing how vulnerability curves may change in hydrodynamic model formulations for plants that recover from embolism.

Green roofs provide many ecosystem services such as regulation of building temperatures, reducing urban heat-island effects and draining rainwater. In addition, they are expected to reduce the high levels of CO2 concentrations in big cities. Previous CO2 fixation studies on green roofs were done by taking long-timeperiod samples using expensive equipment and with limited replication. To plan green roofs for optimal CO2 reduction, a simple method to quantify CO2 fixation rate in relation to plant species-composition is required. The method we tested is direct measurement of CO2 concentrations with a portable air-quality meter, which allows a large number of samples. Here we focus on differences in the photosynthetic effect between plots containing the local Mediterranean succulent, Sedum sediforme and plots containing various annuals. In a factorial design (presence or absence of Sedum crossed with presence or absence of annuals), we tested the effect of sedum and annual treatments on CO2 concentrations. To compare our results with a commonly used method, and to evaluate the role of the different species, we examined the photosynthetic activity at the single plant level under these treatments by using a portable gas-exchange measuring system. We found that our method can detect the effect of different green roof plots and can be used as a simple and reliable tool for green-roof planers. We found that annuals reduced CO2 concentrations, but only in the absence of Sedum. Sedum alone had no effect on CO2 concentrations. This emphasizes the importance of integrating plots with annual plants in Sedum-based green roofs. (C) 2017 Elsevier GmbH. All rights reserved.

Climate change is expected to exacerbate drought for many plants, making drought tolerance a key driver of species and ecosystem responses. Plant drought tolerance is determined by multiple traits, but the relationships among traits, either within individual plants or across species, have not been evaluated for general patterns across plant diversity. We synthesized the published data for stomatal closure, wilting, declines in hydraulic conductivity in the leaves, stems, and roots, and plant mortality for 262 woody angiosperm and 48 gymnosperm species. We evaluated the correlations among the drought tolerance traits across species, and the general sequence of water potential thresholds for these traits within individual plants. The trait correlations across species provide a framework for predicting plant responses to a wide range of water stress from one or two sampled traits, increasing the ability to rapidly characterize drought tolerance across diverse species. Analyzing these correlations also identified correlations among the leaf and stem hydraulic traits and the wilting point, or turgor loss point, beyond those expected from shared ancestry or independent associations with water stress alone. Further, on average, the angiosperm species generally exhibited a sequence of drought tolerance traits that is expected to limit severe tissue damage during drought, such as wilting and substantial stem embolism. This synthesis of the relationships among the drought tolerance traits provides crucial, empirically supported insight into representing variation in multiple traits in models of plant and ecosystem responses to drought.

In deciduous trees growing in temperate forests, bud break and growth in spring must rely on intrinsic carbon (C) reserves. Yet it is unclear whether growth and C storage occur simultaneously, and whether starch C in branches is sufficient for refoliation. To test in situ the relationships between growth, phenology and C utilization, we monitored stem growth, leaf phenology and stem and branch nonstructural carbohydrate (NSC) dynamics in three deciduous species: Carpinus betulus L., Fagus sylvatica L. and Quercus petraea (Matt.) Liebl. To quantify the role of NSC in C investment into growth, a C balance approach was applied. Across the three species, >95% of branchlet starch was consumed during bud break, confirming the importance of C reserves for refoliation in spring. The C balance calculation showed that 90% of the C investment in foliage (7.0-10.5 kg tree(-1) and 5-17 times the C needed for annual stem growth) was explained by simultaneous branchlet starch degradation. Carbon reserves were recovered sooner than expected, after leaf expansion, in parallel with stem growth. Carpinus had earlier leaf phenology (by similar to 25 days) but delayed cambial growth (by similar to 15 days) than Fagus and Quercus, the result of a competitive strategy to flush early, while having lower NSC levels.

Short-term, intense heat waves (hamsins) are common in the eastern Mediterranean region and provide an opportunity to study the resilience of forests to such events that are predicted to increase in frequency and intensity. The response of a 50-yr-old Aleppo pine (Pinus halepensis) forest to hamsin events lasting 1-7d was studied using 10yr of eddy covariance and sap flow measurements. The highest frequency of heat waves was c. four per month, coinciding with the peak productivity period (March-April). During these events, net ecosystem carbon exchange (NEE) and canopy conductance (gc ) decreased by c. 60%, but evapotranspiration (ET) showed little change. Fast recovery was also observed with fluxes reaching pre-stress values within a day following the event. NEE and gc showed a strong response to vapor pressure deficit that weakened as soil moisture decreased, while sap flow was primarily responding to changes in soil moisture. On an annual scale, heat waves reduced NEE and gross primary productivity by c. 15% and 4%, respectively. Forest resilience to short-term extreme events such as heat waves is probably a key to its survival and must be accounted for to better predict the increasing impact on productivity and survival of such events in future climates.

A chloroplast protein disulfide isomerase (PDI) was previously proposed to regulate translation of the unicellular green alga Chlamydomonas reinhardtii chloroplast psbA mRNA, encoding the D1 protein, in response to light. Here we show that AtPDI6, one of 13 Arabidopsis thalianaPDI genes, also plays a role in the chloroplast. We found that AtPDI6 is targeted and localized to the chloroplast. Interestingly, AtPDI6 knockdown plants displayed higher resistance to photoinhibition than wild-type plants when exposed to a tenfold increase in light intensity. The AtPDI6 knockdown plants also displayed a higher rate of D1 synthesis under a similar light intensity. The increased resistance to photoinhibition may not be rationalized by changes in antenna or non-photochemical quenching. Thus, the increased D1 synthesis rate, which may result in a larger proportion of active D1 under light stress, may led to the decrease in photoinhibition. These results suggest that, although the D1 synthesis rates observed in wild-type plants under high light intensities are elevated, repair can potentially occur faster. The findings implicate AtPDI6 as an attenuator of D1 synthesis, modulating photoinhibition in a light-regulated manner.

We review observational, experimental, and model results on how plants respond to extreme climatic conditions induced by changing climatic variability. Distinguishing between impacts of changing mean climatic conditions and changing climatic variability on terrestrial ecosystems is generally underrated in current studies. The goals of our review are thus (1) to identify plant processes that are vulnerable to changes in the variability of climatic variables rather than to changes in their mean, and (2) to depict/evaluate available study designs to quantify responses of plants to changing climatic variability. We find that phenology is largely affected by changing mean climate but also that impacts of climatic variability are much less studied, although potentially damaging. We note that plant water relations seem to be very vulnerable to extremes driven by changes in temperature and precipitation and that heatwaves and flooding have stronger impacts on physiological processes than changing mean climate. Moreover, interacting phenological and physiological processes are likely to further complicate plant responses to changing climatic variability. Phenological and physiological processes and their interactions culminate in even more sophisticated responses to changing mean climate and climatic variability at the species and community level. Generally, observational studies are well suited to study plant responses to changing mean climate, but less suitable to gain a mechanistic understanding of plant responses to climatic variability. Experiments seem best suited to simulate extreme events. In models, temporal resolution and model structure are crucial to capture plant responses to changing climatic variability. We highlight that a combination of experimental, observational, and/or modeling studies have the potential to overcome important caveats of the respective individual approaches.