Publications

Ectomycorrhizal fungi (EMF) are common belowground tree symbionts, supplying trees with water and nutrients. In return, large amounts of C assimilated by trees can be allocated into EMF. However, the chemical forms in which the C is transferred from trees to fungi under field conditions are mostly unknown. In this study, we aimed to unravel the fate of tree-derived C in EMF. We conducted ¹³CO₂ pulse labelling of Pinus halepensis trees in two forest sites with adjacent EMF sporocarps, combined with a non-targeted metabolomics profiling of root and sporocarp tissues. ¹³C was measured in sporocarps of Tricholoma terreum and Suillus collinitus up to 3 m from pine stems. C was assimilated in the labelled trees' needles and transferred to their roots. Starting from Day 2 after labelling, the C was transferred to adjacent sporocarps, peaking on Day 5. We identified more than 100 different labelled metabolites of different chemical groups present in roots and sporocarps. Of them, 17 were common to pine roots and both EMF species, and additional eight common to roots and one of the two EMF. The major labelled metabolites in the root tips were amino acids and tricarboxylic acid intermediates. The major labelled metabolites in sporocarps were amino acids, nucleotides, and fatty acids. We also identified labelled carbohydrates in all tissues. Labelling patterns diverged across different tissues, which can hint at how the C was transferred. Considering the young tree as a sole C source for these sporocarps, and with a diurnal assimilation of 5.4 g C, the total monthly C source is ~165 g C. On average, there were 10 sporocarps around each tree, each requiring ~1 g C. Therefore, a 10 g C investment would make 6% of total tree C allocation, and about 12% of net primary productivity. Overall, we found that this significant and ubiquitous transfer of metabolites from tree roots to EMF sporocarps is more rapid and chemically diverse than once thought. Read the free Plain Language Summary for this article on the Journal blog.

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 (Ψ
_gs50) 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.

Recent findings suggest that trees can survive high levels of drought-induced xylem embolism. In many cases, the embolism is irreversible and, therefore, can potentially affect post-drought recovery and tree function under recurring droughts. We examined the development of embolism in potted Aleppo pines, a common species in hot, dry Mediterranean habitats. We asked (1) how post-drought recovery is affected by different levels of embolism and (2) what consequences this drought-induced damage has under a recurring drought scenario. Young trees were dehydrated to target water potential (Ψ_x) values of −3.5, −5.2 and −9.5 MPa (which corresponded to ~6%, ~41% and ~76% embolism), and recovery of the surviving trees was measured over an 8-months period (i.e., embolism, leaf gas-exchange, Ψ_x). An additional group of trees was exposed to Ψ_x of −6.0 MPa, either with or without preceding drought (Ψ_x of −5.2 MPa) to test the effect of hydraulic damage during repeated drought. Trees that reached −9.5 MPa died, but none from the other groups. Embolism levels in dying trees were on average 76% of conductive xylem and no tree was dying below 62% embolism. Stomatal recovery was negatively proportional to the level of hydraulic damage sustained during drought, for at least a month after drought relief. Trees that experienced drought for the second time took longer to reach fatal Ψ_x levels than first-time dehydrating trees. Decreased stomatal conductance following drought can be seen as “drought legacy,” impeding recovery of tree functioning, but also as a safety mechanism during a consecutive drought.

Severe droughts limit tree growth and forest productivity worldwide, a phenomenon which is expected to aggravate over the next decades. However, how drought intensity and climatic conditions before and after drought events modulate tree growth resilience remains unclear, especially when considering the range-wide phenotypic variability of a tree species. We gathered 4632 Aleppo pine (Pinus halepensis Mill.) tree-ring width series from 281 sites located in 11 countries across the Mediterranean basin, representing the entire geographic and bioclimatic range of the species. For each site and year of the period 1950–2020, we quantified tree-growth resilience and its two components, resistance and recovery, to account for the impact of drought and the capacity to recover from it. Relative drought intensity of each year was assessed using SPEI (Standardized Precipitation Evapotranspiration Index), a climatic water deficit index. Generalized additive mixed models were used to explore the non-linear relationships between resilience and its two components and drought intensity, preceding and following years climatic conditions. We found that P. halepensis radial growth was highly dependent on the SPEI from September of the previous year to June of the current year. Trees growing under more arid bioclimates showed higher inter-annual growth variability and were more sensitive to drought, resulting in an increased response magnitude to pre-, during and post-drought conditions. In contrast to our expectations, drought intensity only slightly affected resilience, which was rather negatively affected by favorable preceding conditions and improved by favorable following conditions. Resilience and its components are highly dependent on preceding and following years climatic conditions, which should always be taken into account when studying growth response to drought. With the observed and predicted increase in drought frequency, duration and intensity, favorable conditions following drought episodes may become rare, thus threatening the future acclimation capacity of P. halepensis in its current distribution.

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 (Ψ_l) of all species declined significantly along the summer, reaching significantly lower Ψ_l 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.

Key message: Desert Acacia trees photosynthesize during the hot dry summer, and use stored carbon for summer growth. Trees that grow in hyper-arid environments can provide important insight into the role of carbon use and carbon storage for tree survival and growth in extreme conditions. Acacia trees, in particular, experience some of the most arid conditions in which trees can grow on the planet, enduring high temperatures, high radiation and drought. Here we measured for two years photosynthesis along the day, stem circumference growth, and seasonal changes in non-structural carbohydrates in adult Acacia trees in their natural hyper-arid habitat (Arava desert, southern Israel). The peak of net CO₂ assimilation was at midday in all seasons, even during summer when vapor pressure deficit was at maximum of 6 kPa and light intensities were at high levels (1800 µmol photons m⁻² s⁻¹). Tree growth started in the spring and increased in the hot summer season and during the dry fall season (autumn). Starch concentrations in the branches were highest in the winter and spring (17% in dry matter) decreasing in the summer and fall (7% in dry matter). Our observations indicated that carbon assimilated during the winter was stored in the branches as starch reserves, which were later used for tree growth in summer. Still, most of the growth was subsidized by concurrent assimilation during the dry season. These findings show that Acacia trees are able to photosynthesize in conditions that other trees cannot, indicating a strong potential to contribute to ecosystem carbon sequestration in warming and drying climates.

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.

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.

Safeguarding Earth’s tree diversity is a conservation priority due to the importance of trees for biodiversity and ecosystem functions and services such as carbon sequestration. Here, we improve the foundation for effective conservation of global tree diversity by analyzing a recently developed database of tree species covering 46,752 species. We quantify range protection and anthropogenic pressures for each species and develop conservation priorities across taxonomic, phylogenetic, and functional diversity dimensions. We also assess the effectiveness of several influential proposed conservation prioritization frameworks to protect the top 17% and top 50% of tree priority areas. We find that an average of 50.2% of a tree species’ range occurs in 110-km grid cells without any protected areas (PAs), with 6,377 small-range tree species fully unprotected, and that 83% of tree species experience nonnegligible human pressure across their range on average. Protecting high-priority areas for the top 17% and 50% priority thresholds would increase the average protected proportion of each tree species’ range to 65.5% and 82.6%, respectively, leaving many fewer species (2,151 and 2,010) completely unprotected. The priority areas identified for trees match well to the Global 200 Ecoregions framework, revealing that priority areas for trees would in large part also optimize protection for terrestrial biodiversity overall. Based on range estimates for >46,000 tree species, our findings show that a large proportion of tree species receive limited protection by current PAs and are under substantial human pressure. Improved protection of biodiversity overall would also strongly benefit global tree diversity.

Inter-kingdom belowground carbon (C) transfer is a significant, yet hidden, biological phenomenon, due to the complexity and highly dynamic nature of soil ecology. Among key biotic agents influencing C allocation belowground are ectomycorrhizal fungi (EMF). EMF symbiosis can extend beyond the single tree-fungus partnership to form common mycorrhizal networks (CMNs). Despite the high prevalence of CMNs in forests, little is known about the identity of the EMF transferring the C and how these in turn affect the dynamics of C transfer. Here, Pinus halepensis and Quercus calliprinos saplings growing in forest soil were labeled using a ¹³CO₂ labeling system. Repeated samplings were applied during 36 days to trace how ¹³C was distributed along the tree-fungus-tree pathway. To identify the fungal species active in the transfer, mycorrhizal fine root tips were used for DNA-stable isotope probing (SIP) with ¹³CO₂ followed by sequencing of labeled DNA. Assimilated ¹³CO₂ reached tree roots within four days and was then transferred to various EMF species. C was transferred across all four tree species combinations. While Tomentella ellisii was the primary fungal mediator between pines and oaks, Terfezia pini, Pustularia spp., and Tuber oligospermum controlled C transfer among pines. We demonstrate at a high temporal, quantitative, and taxonomic resolution, that C from EMF host trees moved into EMF and that C was transferred further to neighboring trees of similar and distinct phylogenies.

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 unprecedented heatwave which hit the Pacific northwest of North America in late June-early July 2021 impacted ecosystems and communities, yet evidence for and analysis of this impact are still missing. Here we bring a unique dataset quantifying the impact on conifer trees, which are keystone species of many northwest ecosystems. Moreover, we take advantage of this exceptional event as a broad, extreme, 'field experiment' to test a fundamental theory in plant physiology and prepare our forests for a harsher future. Overall, the data collected confirm the role of hydraulic vulnerability in drought-induced injury to trees.

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.

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.

Green walls can improve indoor air-quality by reducing concentrations of carbon dioxide (CO2) and other air pollutants. Our study focused on the spider plant, Chlorophytum comosum, and devil’s ivy, Epipremnum aureum, both common green-wall plants that have been found to be efficient CO2 absorbers. Both species have multiple variants with varying degrees of leaf green-white segmentation. Since photosynthesis depends on the concentration of leaf chlorophylls, we hypothesized that green variants are more efficient carbon absorbers than green-white variants. In addition, we tested the hypothesis that the photosynthetic rate of plants is affected by pot volume, as suggested by previous studies. We used a portable gas exchange system to determine the rate of photosynthesis of the study plants. No evidence was found for better photosynthetic performance in the green vs. green-white variants of each species. In fact, our results suggest the opposite. It was observed that a spider plants assimilated carbon more efficiently when grown in a larger pot volume. In conclusion, our study shows that in terms of carbon assimilation, green-white variants of spider plants are the better choice for indoor green walls. Their efficiency can be improved dramatically by increasing pot volume.

Ancient burial caves represent some of the most important sources of information on human history. In a world heritage site in Israel, such caves are under threat due to tree root growth penetrating from the ceilings and causing a risk of cave collapse. To facilitate historical conservation, while avoiding cutting of the mixed forest above the caves, we identified the tree species responsible for the damage, facilitating their specific cutting. Accurate identification of tree roots balances the needs to protect human structures on one side, while conserving the majority of vegetation on the other. Our approach is applicable to the management of ancient sites, as well as to urban management more broadly. Summary: Tree roots have penetrated the ceiling of burial caves in a ~1,800-years-old necropolis in the Galilee, Israel, damaging the antiquities and risking the catacombs with collapse. Root identification was needed to enable selective cutting (at the species level), facilitating the conservation of the world heritage site, while maintaining the majority of trees growing on top of the burial caves. Samples of roots penetrating the cave ceilings were collected and identified both by using DNA barcoding, which has become a standard method for the reliable identification of organisms in their natural environment, and also by traditional morpho-anatomical methods. However, woody plant species of the Mediterranean region are under-represented in DNA databases. Therefore, we added relevant species to the ITS2 database by sequencing the internal transcribed spacer (ITS)2 of the 18S-26S rDNA from sampled leaves of 19 woody species of the Mediterranean maquis. The identification of these tree species facilitated their selective removal, balancing antiquities, and nature conservation needs.

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.

Many ecologically important forest trees from dry areas have been insufficiently investigated for their ability to adapt to the challenges posed by climate change, which hampers the implementation of mitigation policies. We analyzed 14 common-garden experiments across the Mediterranean which studied the widespread thermophilic conifer Pinus halepensis and involved 157 populations categorized into five ecotypes.Ecotype-specific tree height responses to climate were applied to projected climate change (2071-2100 ad), to project potential growth patterns both locally and across the species' range.We found contrasting ecotypic sensitivities to annual precipitation but comparatively uniform responses to mean temperature, while evidence of local adaptation for tree height was limited to mesic ecotypes. We projected intriguing patterns of response range-wide, implying either height inhibition or stimulation of up to 75%, and deduced that the ecotype currently experiencing more favorable (wetter) conditions will show the largest inhibition. Extensive height reductions can be expected for coastal areas of France, Greece, Spain and northern Africa.Our findings underline the fact that intraspecific variations in sensitivity to precipitation must be considered when projecting tree height responses of dry forests to future climate. The ecotype-specific projected performances call for management activities to ensure forest resilience in the Mediterranean through, for example, tailored deployment strategies.

Land surface models (LSMs) typically use empirical functions to represent vegetation responses to soil drought. These functions largely neglect recent advances in plant ecophysiology that link xylem hydraulic functioning with stomatal responses to climate.We developed an analytical stomatal optimization model based on xylem hydraulics (SOX) to predict plant responses to drought. Coupling SOX to the Joint UK Land Environment Simulator (JULES) LSM, we conducted a global evaluation of SOX against leaf- and ecosystem-level observations.SOX simulates leaf stomatal conductance responses to climate for woody plants more accurately and parsimoniously than the existing JULES stomatal conductance model. An ecosystem-level evaluation at 70 eddy flux sites shows that SOX decreases the sensitivity of gross primary productivity (GPP) to soil moisture, which improves the model agreement with observations and increases the predicted annual GPP by 30% in relation to JULES. SOX decreases JULES root-mean-square error in GPP by up to 45% in evergreen tropical forests, and can simulate realistic patterns of canopy water potential and soil water dynamics at the studied sites.SOX provides a parsimonious way to incorporate recent advances in plant hydraulics and optimality theory into LSMs, and an alternative to empirical stress factors.

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.

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-time-period 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.

Are mature forests carbon limited? To explore this question, we exposed ca. 110-year-old, 40-m tall Picea abies trees to a 550-ppm CO₂ concentration in a mixed lowland forest in NW Switzerland. The site receives substantial soluble nitrogen (N) via atmospheric deposition, and thus, trees are unlikely N-limited. We used a construction crane to operate the free-air CO₂ release system and for canopy access. Here, we summarize the major results for growth and carbon (C) fluxes. Tissue ¹³C signals confirmed the effectiveness of the CO₂ enrichment system and permitted tracing the continuous flow of new C in trees. Tree responses were individually standardized by pre-treatment signals. Over the five experimental years, needles retained their photosynthetic capacity and absorbed up to 37% more CO₂ under elevated (E) compared to ambient (A) conditions. However, we did not detect an effect on stem radial growth, branch apical growth and needle litter production. Neither stem nor soil CO₂ efflux was stimulated under elevated CO₂. The rate at which fine roots filled soil ingrowth cores did not significantly differ between A- and E-trees. Since trees showed no stomatal responses to elevated CO₂, sap flow remained unresponsive, both in the long run as well as during short-term CO₂ on–off experiments. As a consequence, soil moisture remained unaffected. We trapped significantly more nitrate in the root sphere of E-trees suggesting a CO₂-stimulated breakdown of soil organic matter, presumably induced by extra carbohydrate exudation (‘priming’). Synthesis. The lack of a single enhanced C sink to match the increased C uptake meant a missing C sink. Increased C transport to below-ground sinks was indicated by C transfer to ectomycorrhiza and on to neighbouring trees and by increased C export to soil. We conclude that these tall Picea abies trees are not C limited at current CO₂ concentrations and further atmospheric CO₂ enrichment will have at most subtle effects on growth, despite enhanced N availability.

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 suffcient 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, confrming 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 ∼25 days) but delayed cambial growth (by ∼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 thaliana PDI 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.