Limited effects of xylem anatomy on embolism resistance in cycad leaves.

Drought-induced xylem embolism is a primary cause of plant mortality. Although c. 70% of cycads are threatened by extinction and extant cycads diversified during a period of increasing aridification, the vulnerability of cycads to embolism spread has been overlooked. We quantified the vulnerability to drought-induced embolism, pressure-volume curves, in situ water potentials, and a suite of xylem anatomical traits of leaf pinnae and rachises for 20 cycad species. We tested whether anatomical traits were linked to hydraulic safety in cycads. Compared with other major vascular plant clades, cycads exhibited similar embolism resistance to angiosperms and pteridophytes but were more vulnerable to embolism than noncycad gymnosperms. All 20 cycads had both tracheids and vessels, the proportions of which were unrelated to embolism resistance. Only vessel pit membrane fraction was positively correlated to embolism resistance, contrary to angiosperms. Water potential at turgor loss was significantly correlated to embolism resistance among cycads. Our results show that cycads exhibit low resistance to xylem embolism and that xylem anatomical traits - particularly vessels - may influence embolism resistance together with tracheids. This study highlights the importance of understanding the mechanisms of drought resistance in evolutionarily unique and threatened lineages like the cycads.

Water status dynamics and drought tolerance of juvenile European beech, Douglas fir and Norway spruce trees as dependent on neighborhood and nitrogen supply.

To increase the resilience of forests to drought and other hazards, foresters are increasingly planting mixed stands. This requires knowledge about the drought response of tree species in pure and mixed-culture neighborhoods. In addition, drought frequently interacts with continued atmospheric nitrogen (N) deposition. To disentangle these factors for European beech, Norway spruce and Douglas fir, we conducted a replicated 3-factorial sapling growth experiment with three moisture levels, (high, medium, and low), two N levels (high and ambient), and pure and mixed-culture neighborhoods. We measured biomass, stomatal conductance (GS), shoot water potential (at predawn: ΨPD, midday, and turgor loss point: ΨTLP), branch xylem embolism resistance (Ψ50) and minimum epidermal conductance (Gmin). The three species differed most with respect to Gmin (10-fold higher in beech than in the conifers), hydroscape area (larger in beech), and the time elapsed to reach stomatal closure (TΨGS90) and ΨTLP (TTLP; shorter in beech), while Ψ50 and ΨTLP were remarkably similar. Neighborhood (pure vs mixed-culture) influenced biomass production, water status and hydraulic traits, notably GS (higher in Douglas fir, but lower in spruce and beech, in mixtures than pure culture), hydraulic safety margin (smaller for beech in mixtures), and TΨGS90 and TTLP (shorter for spruce in mixture). High N generally increased GS, but no consistent N effects on leaf water status and hydraulic traits were detected, suggesting that neighbor identity had a larger effect on plant water relations than N availability. We conclude that both tree neighborhood and N availability modulate the drought response of beech, spruce, and Douglas fir. Species mixing can alleviate the drought stress of some species, but often by disadvantaging other species. Thus, our study suggests that stabilizing and building resilience of production forests against a drier and warmer climate may depend primarily on the right species choice; species mixing can support the agenda.

Plasticity of wood and leaf traits related to hydraulic efficiency and safety is linked to evaporative demand and not soil moisture in rubber (Hevea brasiliensis).

The predicted increase of drought intensity in South-East Asia has raised concern about the sustainability of rubber (Hevea brasiliensis Müll. Arg.) cultivation. In order to quantify the degree of phenotypic plasticity in this important tree crop species, we analysed a set of wood and leaf traits related to the hydraulic safety and efficiency in PB260 clones from eight small-holder plantations in Jambi province, Indonesia, representing a gradient in local microclimatic and edaphic conditions. Across plots, branch embolism resistance (P50) ranged from -2.14 to -2.58 MPa. The P50 and P88 values declined, and the hydraulic safety margin increased, with an increase in the mean annual vapour pressure deficit (VPD). Among leaf traits, only the changes in specific leaf area were related to the differences in evaporative demand. These variations of hydraulic trait values were not related to soil moisture levels. We did not find a trade-off between hydraulic safety and efficiency, but vessel density (VD) emerged as a major trait associated with both safety and efficiency. The VD, and not vessel diameter, was closely related to P50 and P88 as well as to specific hydraulic conductivity, the lumen-to-sapwood area ratio and the vessel grouping index. In conclusion, our results demonstrate some degree of phenotypic plasticity in wood traits related to hydraulic safety in this tropical tree species, but this is only in response to the local changes in evaporative demand and not soil moisture. Given that VPD may increasingly limit plant growth in a warmer world, our results provide evidence of hydraulic trait changes in response to a rising evaporative demand.

Drought survival in conifer species is related to the time required to cross the stomatal safety margin.

The regulation of water loss and the spread of xylem embolism have mostly been considered separately. The development of an integrated approach taking into account the temporal dynamics and relative contributions of these mechanisms to plant drought responses is urgently needed. Do conifer species native to mesic and xeric environments display different hydraulic strategies and temporal sequences under drought? A dry-down experiment was performed on seedlings of four conifer species differing in embolism resistance, from drought-sensitive to extremely drought-resistant species. A set of traits related to drought survival was measured, including turgor loss point, stomatal closure, minimum leaf conductance, and xylem embolism resistance. All species reached full stomatal closure before the onset of embolism, with all but the most drought-sensitive species presenting large stomatal safety margins, demonstrating that highly drought-resistant species do not keep their stomata open under drought conditions. Plant dry-down time to death was significantly influenced by the xylem embolism threshold, stomatal safety margin, and minimum leaf conductance, and was best explained by the newly introduced stomatal margin retention index (SMRIΨ50) which reflects the time required to cross the stomatal safety margin. The SMRIΨ50 may become a key tool for the characterization of interspecific drought survival variability in trees.

Assessing the agreement between the pneumatic and the flow-centrifuge method for estimating xylem safety in temperate diffuse-porous tree species.

The increasing frequency of global change-type droughts has created a need for fast, accurate and widely applicable techniques for estimating xylem embolism resistance to improve forecasts of future forest changes. We used data from 12 diffuse-porous temperate tree species covering a wide range of xylem safety to compare the pneumatic and flow-centrifuge method, two rapid methods used for constructing xylem vulnerability curves. We evaluated the agreement between parameters estimated with both methods and the sensitivity of pneumatic measurements to the duration of air discharge (AD) measurements. There was close agreement between xylem water potentials at 50% air discharged (PAD), estimated with the Pneumatron, and 50% loss of hydraulic conductivity (PLC), estimated with the flow-centrifuge method (mean signed deviation: 0.12 MPa, Pearson correlation: 0.96 after 15 s of gas extraction). However, the relationship between the estimated slopes was more variable, resulting in lower agreement in the xylem water potential at 12% and 88% PAD/PLC. The agreement between the two methods was not affected by species-specific vessel length distributions. All pneumatic parameters were sensitive to AD time. Overall agreement was highest at relatively short AD times, with an optimum at 16 s. Our results highlight the value of the Pneumatron as an easy and reliable tool to estimate 50% embolism thresholds for a wide range of diffuse-porous temperate angiosperms. Further, our study provides a set of useful metrics for methodological comparisons of vulnerability curves in terms of systematic and random deviations, as well as overall agreement.

Hydraulic traits and photosynthesis are coordinated with trunk sapwood capacitance in tropical tree species.

Water stored in trunk sapwood is vital for the canopy to maintain its physiological function under high transpiration demands. Little is known regarding the anatomical properties that contribute to the hydraulic capacitance of tree trunks and whether trunk capacitance is correlated with the hydraulic and gas exchange traits of canopy branches. We examined sapwood capacitance, xylem anatomical characteristics of tree trunks, embolism resistance, the minimal xylem water potential of canopy branches, leaf photosynthesis and stomatal conductance in 22 species from a tropical seasonal rainforest and savanna. The results showed that the mean trunk sapwood capacitance did not differ between the two biomes. Capacitance was closely related to the fiber lumen fraction and fiber wall reinforcement and not to the axial and ray parenchyma fractions. Additionally, it was positively correlated with the theoretical hydraulic conductivity of a trunk and the specific hydraulic conductivity of branches, and showed a trade-off with branch embolism resistance. Species with a high trunk sapwood capacitance maintained less negative canopy water potentials in the dry season, but higher leaf photosynthetic rates and stomatal conductance in the wet season. This study provides a functional link among trunk sapwood capacitance, xylem anatomy, canopy hydraulics and photosynthesis in tropical trees.

Abscisic acid acts essentially on stomata, not on the xylem, to improve drought resistance in tomato.

Drought resistance is essential for plant production under water-limiting environments. Abscisic acid (ABA) plays a critical role in stomata but its impact on hydraulic function beyond the stomata is far less studied. We selected genotypes differing in their ability to accumulate ABA to investigate its role in drought-induced dysfunction. All genotypes exhibited similar leaf and stem embolism resistance regardless of differences in ABA levels. Their leaf hydraulic resistance was also similar. Differences were only observed between the two extreme genotypes: sitiens (sit; a strong ABA-deficient mutant) and sp12 (a transgenic line that constitutively overaccumulates ABA), where the water potential inducing 50% embolism was 0.25 MPa lower in sp12 than in sit. Maximum stomatal and minimum leaf conductances were considerably lower in plants with higher ABA (wild type [WT] and sp12) than in ABA-deficient mutants. Variations in gas exchange across genotypes were associated with ABA levels and differences in stomatal density and size. The lower water loss in plants with higher ABA meant that lethal water potentials associated with embolism occurred later during drought in sp12 plants, followed by WT, and then by the ABA-deficient mutants. Therefore, the primary pathway by which ABA enhances drought resistance is via declines in water loss, which delays dehydration and hydraulic dysfunction.

Functional traits and water-transport strategies of woody species in an insular environment in a tropical forest.

PREMISE: Plants survive in habitats with limited resource availability and contrasting environments by responding to variation in environmental factors through morphophysiological traits related to species performance in different ecosystems. However, how different plant strategies influence the megadiversity of tropical species has remained a knowledge gap. METHODS: We analyzed variations in 27 morphophysiological traits of leaves and secondary xylem in Erythroxylum pulchrum and Tapirira guianensis, which have the highest absolute dominance in these physiognomies and occur together in areas of restinga and dense ombrophilous forest to infer water-transport strategies of Atlantic Forest woody plants. RESULTS: The two species presented different sets of morphophysiological traits, strategies to avoid embolism and ensure water transport, in different phytophysiognomies. Tapirira guianensis showed possible adaptations influenced by phytophysiognomy, while E. pulchrum showed less variation in the set of characteristics between different phytophysiognomies. CONCLUSIONS: Our results provide essential tools to understand how the environment can modulate morphofunctional traits and how each species adjusts differently to adapt to different phytophysiognomies. In this sense, the results for these species reveal new species-specific responses in the tropical forest. Such knowledge is a prerequisite to predict future development of the most vulnerable forests as climate changes.

Anatomical traits related to leaf and branch hydraulic functioning on Amazonian savanna plants.

Amazonian savannas are isolated patches of open habitats found within the extensive matrix of Amazonian tropical forests. There remains limited evidence on how Amazonian plants from savannas differ in the traits related to drought resistance and water loss control. Previous studies have reported several xeromorphic characteristics of Amazonian savanna plants at the leaf and branch levels that are linked to soil, solar radiation, rainfall and seasonality. How anatomical features relate to plant hydraulic functioning in this ecosystem is less known and instrumental if we want to accurately model transitions in trait states between alternative vegetation in Amazonia. In this context, we combined studies of anatomical and hydraulic traits to understand the structure-function relationships of leaf and wood xylem in plants of Amazonian savannas. We measured 22 leaf, wood and hydraulic traits, including embolism resistance (as P50), Hydraulic Safety Margin (HSM) and isotope-based water use efficiency (WUE), for the seven woody species that account for 75% of the biomass of a typical Amazonian savanna on rocky outcrops in the state of Mato Grosso, Brazil. Few anatomical traits are related to hydraulic traits. Our findings showed wide variation exists among the seven species studied here in resistance to embolism, water use efficiency and structural anatomy, suggesting no unique dominant functional plant strategy to occupy an Amazonian savanna. We found wide variation in resistance to embolism (-1.6 ± 0.1 MPa and -5.0 ± 0.5 MPa) with species that are less efficient in water use (e.g. Kielmeyera rubriflora, Macairea radula, Simarouba versicolor, Parkia cachimboensis and Maprounea guianensis) showing higher stomatal conductance potential, supporting xylem functioning with leaf succulence and/or safer wood anatomical structures and that species that are more efficient in water use (e.g. Norantea guianensis and Alchornea discolor) can exhibit riskier hydraulic strategies. Our results provide a deeper understanding of how branch and leaf structural traits combine to allow for different hydraulic strategies among coexisting plants. In Amazonian savannas, this may mean investing in buffering water loss (e.g. succulence) at leaf level or safer structures (e.g. thicker pit membranes) and architectures (e.g. vessel grouping) in their branch xylem.

Thresholds for persistent leaf photochemical damage predict plant drought resilience in a tropical rainforest.

Water stress can cause declines in plant function that persist after rehydration. Recent work has defined 'resilience' traits characterizing leaf resistance to persistent damage from drought, but whether these traits predict resilience in whole-plant function is unknown. It is also unknown whether the coordination between resilience and 'resistance' - the ability to maintain function during drought - observed globally occurs within ecosystems. For eight rainforest species, we dehydrated and subsequently rehydrated leaves, and measured water stress thresholds for declines in rehydration capacity and maximum quantum yield of photosystem II (Fv /Fm ). We tested correlations with embolism resistance and dry season water potentials (ΨMD ), and calculated safety margins for damage (ΨMD - thresholds) and tested correlations with drought resilience in sap flow and growth. Ψ thresholds for persistent declines in Fv /Fm , indicating resilience, were positively correlated with ΨMD and thresholds for leaf vein embolism. Safety margins for persistent declines in Fv /Fm , but not rehydration capacity, were positively correlated with drought resilience in sap flow. Correlations between resistance and resilience suggest that species' differences in performance during drought are perpetuated after drought, potentially accelerating shifts in forest composition. Resilience to photochemical damage emerged as a promising functional trait to characterize whole-plant drought resilience.

Tree growth is correlated with hydraulic efficiency and safety across 22 tree species in a subtropical karst forest.

Karst forests are habitats in which access to soil water can be challenging for plants. Therefore, safe and efficient xylem water transport and large internal water storage may benefit tree growth. In this study, we selected 22 tree species from a primary subtropical karst forest in southern China and measured their xylem anatomical traits, saturated water content (SWC), hydraulic conductivity (Ks) and embolism resistance (P50). Additionally, we monitored growth of diameter at breast height (DBH) in 440 individual trees of various sizes over three consecutive years. Our objective was to analyze the relationships between xylem structure, hydraulic efficiency, safety, water storage and growth of karst tree species. The results showed significant differences in structure but not in hydraulic traits between deciduous and evergreen species. Larger vessel diameter, paratracheal parenchyma and higher SWC were correlated with higher Ks. Embolism resistance was not correlated with the studied anatomical traits, and no tradeoff with Ks was observed. In small trees (5-15 cm DBH), diameter growth rate (DGR) was independent of hydraulic traits. In large trees (>15 cm DBH), higher Ks and more negative P50 accounted for higher DGR. From lower to greater embolism resistance, the size-growth relationship shifted from growth deceleration to acceleration with increasing tree size in eight of the 22 species. Our study highlights the vital contributions of xylem hydraulic efficiency and safety to growth rate and dynamics in karst tree species; therefore, we strongly recommend their integration into trait-based forest dynamic models.

Addressing controversies in the xylem embolism resistance-vessel diameter relationship.

Although xylem embolism is a key process during drought-induced tree mortality, its relationship to wood anatomy remains debated. While the functional link between bordered pits and embolism resistance is known, there is no direct, mechanistic explanation for the traditional assumption that wider vessels are more vulnerable than narrow ones. We used data from 20 temperate broad-leaved tree species to study the inter- and intraspecific relationship of water potential at 50% loss of conductivity (P50 ) with hydraulically weighted vessel diameter (Dh ) and tested its link to pit membrane thickness (TPM ) and specific conductivity (Ks ) on species level. Embolism-resistant species had thick pit membranes and narrow vessels. While Dh was weakly associated with TPM , the P50 -Dh relationship remained highly significant after accounting for TPM . The interspecific pattern between P50 and Dh was mirrored by a link between P50 and Ks , but there was no evidence for an intraspecific relationship. Our results provide robust evidence for an interspecific P50 -Dh relationship across our species. As a potential cause for the inconsistencies in published P50 -Dh relationships, our analysis suggests differences in the range of trait values covered, and the level of data aggregation (species, tree or sample level) studied.

Is a seasonally reduced growth potential a convergent strategy to survive drought and frost in plants?

BACKGROUND: Plants have adapted to survive seasonal life-threatening frost and drought. However, the timing and frequency of such events are impacted by climate change, jeopardizing plant survival. Understanding better the strategies of survival to dehydration stress is therefore timely and can be enhanced by the cross-fertilization of research between disciplines (ecology, physiology), models (woody, herbaceous species) and types of stress (drought, frost). SCOPE: We build upon the 'growth-stress survival' trade-off, which underpins the identification of global plant strategies across environments along a 'fast-slow' economics spectrum. Although phenological adaptations such as dormancy are crucial to survive stress, plant global strategies along the fast-slow economic spectrum rarely integrate growth variations across seasons. We argue that the growth-stress survival trade-off can be a useful framework to identify convergent plant ecophysiological strategies to survive both frost and drought. We review evidence that reduced physiological activity, embolism resistance and dehydration tolerance of meristematic tissues are interdependent strategies that determine thresholds of mortality among plants under severe frost and drought. We show that complete dormancy, i.e. programmed growth cessation, before stress occurrence, minimizes water flows and maximizes dehydration tolerance during seasonal life-threatening stresses. We propose that incomplete dormancy, i.e. the programmed reduction of growth potential during the harshest seasons, could be an overlooked but major adaptation across plants. Quantifying stress survival in a range of non-dormant versus winter- or summer-dormant plants, should reveal to what extent incomplete to complete dormancy could represent a proxy for dehydration tolerance and stress survival. CONCLUSIONS: Our review of the strategies involved in dehydration stress survival suggests that winter and summer dormancy are insufficiently acknowledged as plant ecological strategies. Incorporating a seasonal fast-slow economics spectrum into global plant strategies improves our understanding of plant resilience to seasonal stress and refines our prevision of plant adaptation to extreme climatic events.

Stem hydraulic conductivity and embolism resistance of Quercus species are associated with their climatic niche.

The hydraulic traits of a plant species may reflect its climate adaptations. Southwest China is considered as a biodiversity hotpot of the genus Quercus (oak). However, the hydraulic adaptations of Asian oaks to their climate niches remain unclear. Ten common garden-grown oak species with distinct natural distributions in eastern Asia were used to determine their stem xylem embolism resistance (water potential at 50% loss of hydraulic conductivity, P50), stem hydraulic efficiency (vessel anatomy and sapwood specific hydraulic conductivity (Ks)) and leaf anatomical traits. We also compiled four key functional traits: wood density, hydraulic-weighted vessel diameter, Ks and P50 data for 31 oak species from previous literature. We analyzed the relationship between hydraulic traits and climatic factors over the native ranges of 41 oak species. Our results revealed that the 10 Asian oak species, which are mainly distributed in humid subtropical habitats, possessed a stem xylem with low embolism resistance and moderate hydraulic efficiency. The deciduous and evergreen species of the 10 Asian oaks differed in the stem and leaf traits related to hydraulic efficiency. Ks differed significantly between the two phenological groups (deciduous and evergreens) in the 41-oak dataset. No significant difference in P50 between the two groups was found for the 10 Asian oaks or the 41-oak dataset. The oak species that can distribute in arid habitats possessed a stem xylem with high embolism resistance. Ks negatively related to the humidity of the native range of the 10 Asian oaks, but showed no trend when assessing the entire global oak dataset. Our study suggests that stem hydraulic conductivity and embolism resistance in Quercus species are shaped by their climate niche. Our findings assist predictions of oak drought resistance with future climate changes for oak forest management.

Embolism resistance explains mortality and recovery of five subtropical evergreen broadleaf trees to persistent drought.

Subtropical evergreen broadleaf forests (SEBF) are experiencing and expected to suffer more frequent and severe drought events. However, how the hydraulic traits directly link to the mortality and recovery of SEBF trees remains unclear. In this study, we conducted a drought-rewatering experiment on tree seedlings of five dominant species to investigate how the hydraulic traits were related to tree mortality and the resistance and recovery of photosynthesis (A) and transpiration (E) under different drought severities. Species with greater embolism resistance (P50 ) survived longer than those with a weaker P50 . However, there was no general hydraulic threshold associated with tree mortality, with the lethal hydraulic failure varying from 64% to 93% loss of conductance. The photosynthesis and transpiration of tree species with a greater P50 were more resistant to and recovered faster from drought than those with lower P50 . Other plant traits could not explain the interspecific variation in tree mortality and drought resistance and recovery. These results highlight the unique importance of embolism resistance in driving carbon and water processes under persistent drought across different trees in SEBFs. The absence of multiple efficient drought strategies in SEBF seedlings implies the difficulty of natural seedling regeneration under future droughts, which often occurs after destructive disturbances (e.g., extreme drought events and typhoon), suggesting that this biome may be highly vulnerable to co-occurring climate extremes.

Drought response in Arabidopsis displays synergistic coordination between stems and leaves.

The synergy between drought-responsive traits across different organs is crucial in the whole-plant mechanism influencing drought resilience. These organ interactions, however, are poorly understood, limiting our understanding of drought response strategies at the whole-plant level. Therefore, we need more integrative studies, especially on herbaceous species that represent many important food crops but remain underexplored in their drought response. We investigated inflorescence stems and rosette leaves of six Arabidopsis thaliana genotypes with contrasting drought tolerance, and combined anatomical observations with hydraulic measurements and gene expression studies to assess differences in drought response. The soc1ful double mutant was the most drought-tolerant genotype based on its synergistic combination of low stomatal conductance, largest stomatal safety margin, more stable leaf water potential during non-watering, reduced transcript levels of drought stress marker genes, and reduced loss of chlorophyll content in leaves, in combination with stems showing the highest embolism resistance, most pronounced lignification, and thickest intervessel pit membranes. In contrast, the most sensitive Cvi ecotype shows the opposite extreme of the same set of traits. The remaining four genotypes show variations in this drought syndrome. Our results reveal that anatomical, ecophysiological, and molecular adaptations across organs are intertwined, and multiple (differentially combined) strategies can be applied to acquire a certain level of drought tolerance.

Mutually inclusive mechanisms of drought-induced tree mortality.

Unprecedented tree dieback across Central Europe caused by recent global change-type drought events highlights the need for a better mechanistic understanding of drought-induced tree mortality. Although numerous physiological risk factors have been identified, the importance of two principal mechanisms, hydraulic failure and carbon starvation, is still debated. It further remains largely unresolved how the local neighborhood composition affects individual mortality risk. We studied 9435 young trees of 12 temperate species planted in a diversity experiment in 2013 to assess how hydraulic traits, carbon dynamics, pest infestation, tree height and neighborhood competition influence individual mortality risk. Following the most extreme global change-type drought since record in 2018, one third of these trees died. Across species, hydraulic safety margins (HSMs) were negatively and a shift towards a higher sugar fraction in the non-structural carbohydrate (NSC) pool positively associated with mortality risk. Moreover, trees infested by bark beetles had a higher mortality risk, and taller trees a lower mortality risk. Most neighborhood interactions were beneficial, although neighborhood effects were highly species-specific. Species that suffered more from drought, especially Larix spp. and Betula spp., tended to increase the survival probability of their neighbors and vice versa. While severe tissue dehydration marks the final stage of drought-induced tree mortality, we show that hydraulic failure is interrelated with a series of other, mutually inclusive processes. These include shifts in NSC pools driven by osmotic adjustment and/or starch depletion as well as pest infestation and are modulated by the size and species identity of a tree and its neighbors. A more holistic view that accounts for multiple causes of drought-induced tree mortality is required to improve predictions of trends in global forest dynamics and to identify mutually beneficial species combinations.

Soil water availability and branch age explain variability in xylem safety of European beech in Central Europe.

Xylem embolism resistance has been identified as a key trait with a causal relation to drought-induced tree mortality, but not much is known about its intra-specific trait variability (ITV) in dependence on environmental variation. We measured xylem safety and efficiency in 300 European beech (Fagus sylvatica L.) trees across 30 sites in Central Europe, covering a precipitation reduction from 886 to 522 mm year-1. A broad range of variables that might affect embolism resistance in mature trees, including climatic and soil water availability, competition, and branch age, were examined. The average P50 value varied by up to 1 MPa between sites. Neither climatic aridity nor structural variables had a significant influence on P50. However, P50 was less negative for trees with a higher soil water storage capacity, and positively related to branch age, while specific conductivity (Ks) was not significantly associated with either of these variables. The greatest part of the ITV for xylem safety and efficiency was attributed to random variability within populations. We conclude that the influence of site water availability on P50 and Ks is low in European beech, and that the high degree of within-population variability for P50, partly due to variation in branch age, hampers the identification of a clear environmental signal.

Stem and leaf xylem of angiosperm trees experiences minimal embolism in temperate forests during two consecutive summers with moderate drought.

Drought events may increase the likelihood that the plant water transport system becomes interrupted by embolism. Yet our knowledge about the temporal frequency of xylem embolism in the field is frequently lacking, as it requires detailed, long-term measurements. We measured xylem embolism resistance and midday xylem water potentials during the consecutive summers of 2019 and 2020 to estimate maximum levels of embolism in leaf and stem xylem of ten temperate angiosperm tree species. We also studied vessel and pit membrane characteristics based on light and electron microscopy to corroborate potential differences in embolism resistance between leaves and stems. Apart from A. pseudoplatanus and Q. petraea, eight species experienced minimum xylem water potentials that were close to or below those required to initiate embolism. Water potentials corresponding to ca. 12% loss of hydraulic conductivity (PLC) could occur in six species, while considerable levels of embolism around 50% PLC were limited to B. pendula and C. avellana. There was a general agreement in embolism resistance between stems and leaves, with leaves being equally or more resistant than stems. Also, xylem embolism resistance was significantly correlated to intervessel pit membrane thickness (TPM ) for stems, but not to vessel diameter and total intervessel pit membrane surface area of a vessel. Our data indicate that low amounts of embolism occur in most species during moderate summer drought, and that considerable levels of embolism are uncommon. Moreover, our experimental and TPM data show that leaf xylem is generally no more vulnerable than stem xylem.

Hydraulic variability of three temperate broadleaf tree species along a water availability gradient in central Europe.

Plant hydraulic traits are key for understanding and predicting tree drought responses. Information about the degree of the traits' intra-specific variability may guide the selection of drought-resistant genotypes and is crucial for trait-based modelling approaches. For the three temperate minor broadleaf tree species Acer platanoides, Carpinus betulus and Tilia cordata, we measured xylem embolism resistance (P50 ), leaf turgor loss point (PTLP ), specific hydraulic conductivity (KS ), Huber values (HVs), and hydraulic safety margins in adult trees across a precipitation gradient. We further quantified trait variability on different organizational levels (inter-specific to within-canopy variation), and analysed its relationship to climatic and soil water availability. Although we observed a certain intra-specific trait variability (ITV) in safety-related traits (P50 , PTLP ) with higher within-tree and between-tree than between populations variability, the magnitude was small compared to inter-specific differences, which explained 78.4% and 58.3% of the variance in P50 and PTLP , respectively. In contrast, efficiency-related traits (KS , HV) showed a high ITV both within populations and within the crowns of single trees. Surprisingly, the observed ITV of all traits was neither driven by climatic nor soil water availability. In conclusion, the high degree of conservatism in safety-related traits highlights their potential for trait-based modelling approaches.