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Rising temperatures are influencing forests on many scales, with potentially strong variation vertically across forest strata. Using published research and new analyses, we evaluate how microclimate and leaf temperatures, traits, and gas exchange vary vertically in forests, shaping tree, and ecosystem ecology. In closed-canopy forests, upper canopy leaves are exposed to the highest solar radiation and evaporative demand, which can elevate leaf temperature (T

Ecophysiologists have reported a range of relationships, including intrinsic trade-offs across and within species between plant relative growth rate in high resource conditions (RGR) vs adaptation to tolerate cold or arid climates, arising from trait-based mechanisms. Few studies have considered ecotypes within a species, in which the lack of a trade-off would contribute to a wide species range and resilience to climate change. For 15 ecotypes of Arabidopsis thaliana in a common garden we tested for associations between RGR vs adaptation to cold or dry native climates and assessed hypotheses for its mediation by 15 functional traits. Ecotypes native to warmer, drier climates had higher leaf density, leaf mass per area, root mass fraction, nitrogen per leaf area and carbon isotope ratio, and lower osmotic potential at full turgor. Relative growth rate was statistically independent of the climate of the ecotype native range and of individual functional traits. The decoupling of RGR and cold or drought adaptation in Arabidopsis is consistent with multiple stress resistance and avoidance mechanisms for ecotypic climate adaptation and would contribute to the species' wide geographic range and resilience as the climate changes.

Shifts in stomatal trait distributions across contrasting environments and their linkage with ecosystem productivity at large spatial scales have been unclear. Here, we measured the maximum stomatal conductance (g), stomatal area fraction (f), and stomatal space-use efficiency (e, the ratio of g to f) of 800 plant species ranging from tropical to cold-temperate forests, and determined their values for community-weighted mean, variance, skewness, and kurtosis. We found that the community-weighted means of g and f were higher in drier sites, and thus, that drought ‘avoidance’ by water availability-driven growth pulses was the dominant mode of adaptation for communities at sites with low water availability. Additionally, the variance of g and f was also higher at arid sites, indicating greater functional niche differentiation, whereas that for e was lower, indicating the convergence in efficiency. When all other stomatal trait distributions were held constant, increasing kurtosis or decreasing skewness of g would improve ecosystem productivity, whereas f showed the opposite patterns, suggesting that the distributions of inter-related traits can play contrasting roles in regulating ecosystem productivity. These findings demonstrate the climatic trends of stomatal trait distributions and their significance in the prediction of ecosystem productivity.

International audience; 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.

25 páginas.- 8 figuras.- 2 tablas.- 66 referencias.- Additional supporting information can be found online in the Supporting Information section at the end of this article., eaf water potential (psi(leaf)), typically measured using the pressure chamber, is the most important metric of plant water status, providing high theoretical value and information content for multiple applications in quantifying critical physiological processes including drought responses. Pressure chamber measurements of psi(leaf) (psi(leafPC)) are most typical, yet, the practical complexity of the technique and of the underlying theory has led to ambiguous understanding of the conditions to optimize measurements. Consequently, specific techniques and precautions diversified across the global research community, raising questions of reliability and repeatability. Here, we surveyed specific methods of psi(leafPC) from multiple laboratories, and synthesized experiments testing common assumptions and practices in psi(leafPC) for diverse species: (i) the need for equilibration of previously transpiring leaves; (ii) leaf storage before measurement; (iii) the equilibration of psi(leaf) for leaves on bagged branches of a range of dehydration; (iv) the equilibration of psi(leaf) across the lamina for bagged leaves, and the accuracy of measuring leaves with artificially 'elongated petioles'; (v) the need in psi(leaf) measurements for bagging leaves and high humidity within the chamber; (vi) the need to avoid liquid water on leaf surfaces; (vii) the use of 'pulse' pressurization versus gradual pressurization; and (viii) variation among experimenters in psi(leafPC) determination. Based on our findings we provide a best practice protocol to maximise accuracy, and provide recommendations for ongoing species-specific tests of important assumptions in future studies., European Union's Horizon 2020 research and innovation programme under the Marie Sklodowska-Curie grant agreement No. 751918-AgroPHYS; Marie Curie Fellowship (FP7-PEOPLE-2013-IOF-624473); Austrian research agency (FWF) project P32203; National Science Foundation (Grant IOS-#1457279)

Tree size shapes forest carbon dynamics and determines how trees interact with their environment, including a changing climate. Here, we conduct the first global analysis of among-site differences in how aboveground biomass stocks and fluxes are distributed with tree size. We analyzed repeat tree censuses from 25 large-scale (4-52 ha) forest plots spanning a broad climatic range over five continents to characterize how aboveground biomass, woody productivity, and woody mortality vary with tree diameter. We examined how the median, dispersion, and skewness of these size-related distributions vary with mean annual temperature and precipitation. In warmer forests, aboveground biomass, woody productivity, and woody mortality were more broadly distributed with respect to tree size. In warmer and wetter forests, aboveground biomass and woody productivity were more right skewed, with a long tail towards large trees. Small trees (1-10 cm diameter) contributed more to productivity and mortality than to biomass, highlighting the importance of including these trees in analyses of forest dynamics. Our findings provide an improved characterization of climate-driven forest differences in the size structure of aboveground biomass and dynamics of that biomass, as well as refined benchmarks for capturing climate influences in vegetation demographic models.

Deep-water access is arguably the most effective, but under-studied, mechanism that plants employ to survive during drought. Vulnerability to embolism and hydraulic safety margins can predict mortality risk at given levels of dehydration, but deep-water access may delay plant dehydration. Here, we tested the role of deep-water access in enabling survival within a diverse tropical forest community in Panama using a novel data-model approach. We inversely estimated the effective rooting depth (ERD, as the average depth of water extraction), for 29 canopy species by linking diameter growth dynamics (1990-2015) to vapor pressure deficit, water potentials in the whole-soil column, and leaf hydraulic vulnerability curves. We validated ERD estimates against existing isotopic data of potential water-access depths. Across species, deeper ERD was associated with higher maximum stem hydraulic conductivity, greater vulnerability to xylem embolism, narrower safety margins, and lower mortality rates during extreme droughts over 35years (1981-2015) among evergreen species. Species exposure to water stress declined with deeper ERD indicating that trees compensate for water stress-related mortality risk through deep-water access. The role of deep-water access in mitigating mortality of hydraulically-vulnerable trees has important implications for our predictive understanding of forest dynamics under current and future climates.

The mangrove Avicennia marina adjusts internal salt concentrations by foliar salt secretion. Deliquescence of accumulated salt causes leaf wetting that may provide a water source for salt-secreting plants in arid coastal wetlands where high nocturnal humidity can usually support deliquescence whereas rainfall events are rare. We tested the hypotheses that salt deliquescence on leaf surfaces can drive top-down rehydration, and that such absorption of moisture from unsaturated atmospheres makes a functional contribution to dry season shoot water balances. Sap flow and water relations were monitored to assess the uptake of atmospheric water by branches during shoot wetting events under natural and manipulated microclimatic conditions. Reverse sap flow rates increased with increasing relative humidity from 70% to 89%, consistent with function of salt deliquescence in harvesting moisture from unsaturated atmospheres. Top-down rehydration elevated branch water potentials above those possible from root water uptake, subsidising transpiration rates and reducing branch vulnerability to hydraulic failure in the subsequent photoperiod. Absorption of atmospheric moisture harvested through deliquescence of salt on leaf surfaces enhances water balances of Avicennia marina growing in hypersaline wetlands under arid climatic conditions. Top-down rehydration from these frequent, low intensity wetting events contributes to prevention of carbon starvation and hydraulic failure during drought.

The effects of climate change on tropical forests will depend on how diverse tropical tree species respond to drought. Current distributions of evergreen and deciduous tree species across local and regional moisture gradients reflect their ability to tolerate drought stress, and might be explained by functional traits. We measured leaf water potential at turgor loss (i.e. ‘wilting point’; πtlp), wood density (WD) and leaf mass per area (LMA) on 50 of the most abundant tree species in central Panama. We then tested their ability to explain distributions of evergreen and deciduous species within a 50 ha plot on Barro Colorado Island and across a 70 km rainfall gradient spanning the Isthmus of Panama. Among evergreen trees, species with lower πtlp were associated with drier habitats, with πtlp explaining 28% and 32% of habitat association on local and regional scales, respectively, greatly exceeding the predictive power of WD and LMA. In contrast, πtlp did not predict habitat associations among deciduous species. Across spatial scales, πtlp is a useful indicator of habitat preference for tropical tree species that retain their leaves during periods of water stress, and holds the potential to predict vegetation responses to climate change.

Drought decreases water transport capacity of leaves and limits gas exchange, which involves reduced leaf leaf hydraulic conductance (Kleaf) in both the xylem and outside-xylem pathways. Some literature suggests that grapevines are hyper-susceptible to drought-induced xylem embolism. We combined Kleaf and gas exchange measurements, micro-computed tomography of intact leaves, and spatially explicit modeling of the outside-xylem pathways to evaluate the role of vein embolism and Kleaf in the responses of two different grapevine cultivars to drought. Cabernet Sauvignon and Chardonnay exhibited similar vulnerabilities of Kleaf and gs to dehydration, decreasing substantially prior to leaf xylem embolism. Kleaf and gs decreased by 80% for both cultivars by Ψ leaf approximately –0.7 MPa and –1.2 MPa, respectively, while leaf xylem embolism initiated around Ψ leaf = –1.25 MPa in the midribs and little to no embolism was detected in minor veins even under severe dehydration for both cultivars. Modeling results indicated that reduced membrane permeability associated with a Casparian-like band in the leaf vein bundle sheath would explain declines in Kleaf of both cultivars. We conclude that during moderate water stress, changes in the outside-xylem pathways, rather than xylem embolism, are responsible for reduced Kleaf and gs. Understanding this mechanism could help to ensure adequate carbon capture and crop performance under drought.

As climate change drives increased drought in many forested regions, mechanistic understanding of the factors conferring drought tolerance in trees is increasingly important. The dendrochronological record provides a window through which we can understand how tree size and traits shape growth responses to droughts. We analyzed tree-ring records for 12 species in a broadleaf deciduous forest in Virginia (USA) to test hypotheses for how tree height, microenvironment characteristics, and species' traits shaped drought responses across the three strongest regional droughts over a 60-yr period. Drought tolerance (resistance, recovery, and resilience) decreased with tree height, which was strongly correlated with exposure to higher solar radiation and evaporative demand. The potentially greater rooting volume of larger trees did not confer a resistance advantage, but marginally increased recovery and resilience, in sites with low topographic wetness index. Drought tolerance was greater among species whose leaves lost turgor (wilted) at more negative water potentials and experienced less shrinkage upon desiccation. The tree-ring record reveals that tree height and leaf drought tolerance traits influenced growth responses during and after significant droughts in the meteorological record. As climate change-induced droughts intensify, tall trees with drought-sensitive leaves will be most vulnerable to immediate and longer-term growth reductions.

We review possible physiological, anatomical, and evolutionary limitations that may explain the rarity of C4 photosynthesis in trees and discuss how the C4 trees in Euphorbia are exceptions to this., Since C4 photosynthesis was first discovered >50 years ago, researchers have sought to understand how this complex trait evolved from the ancestral C3 photosynthetic machinery on >60 occasions. Despite its repeated emergence across the plant kingdom, C4 photosynthesis is notably rare in trees, with true C4 trees only existing in Euphorbia. Here we consider aspects of the C4 trait that could limit but not preclude the evolution of a C4 tree, including reduced quantum yield, increased energetic demand, reduced adaptive plasticity, evolutionary constraints, and a new theory that the passive symplastic phloem loading mechanism observed in trees, combined with difficulties in maintaining sugar and water transport over a long pathlength, could make C4 photosynthesis largely incompatible with the tree lifeform. We conclude that the transition to a tree habit within C4 lineages as well as the emergence of C4 photosynthesis within pre-existing trees would both face a series of challenges that together explain the global rarity of C4 photosynthesis in trees. The C4 trees in Euphorbia are therefore exceptional in how they have circumvented every potential barrier to the rare C4 tree lifeform.

Few studies have explored how - within the same system - clonality and positive plant-plant interactions might interact to regulate plant community composition. Canopy-dwelling epiphytes in species-rich forests provide an ideal system for studying this because many epiphytic vascular plants undertake clonal growth and because vascular epiphytes colonize canopy habitats after the formation of nonvascular epiphyte (i.e. bryophyte and lichen) mats. We investigated how clonal integration of seven dominant vascular epiphytes influenced inter-specific interactions between vascular epiphytes and nonvascular epiphytes in a subtropical montane moist forest in southwest China. Both clonal integration and environmental buffering from nonvascular epiphytes increased survival and growth of vascular epiphytes. The benefits of clonal integration for vascular epiphytes were higher when nonvascular epiphytes were removed. Similarly, facilitation from nonvascular epiphytes played a more important role when clonal integration of vascular epiphytes was eliminated. Overall, clonal integration had greater benefits than inter-specific facilitation. This study provides novel evidence for interactive effects of clonality and facilitation between vascular and nonvascular species, and has implications for our understanding of a wide range of ecosystems where both high levels of clonality and facilitation are expected to occur.

Given increasing water deficits across numerous ecosystems world-wide, it is urgent to understand the sequence of failure of leaf function during dehydration. We assessed dehydration-induced losses of rehydration capacity and maximum quantum yield of the photosystem II (Fv /Fm ) in the leaves of 10 diverse angiosperm species, and tested when these occurred relative to turgor loss, declines of stomatal conductance gs , and hydraulic conductance Kleaf , including xylem and outside xylem pathways for the same study plants. We resolved the sequences of relative water content and leaf water potential Ψleaf thresholds of functional impairment. On average, losses of leaf rehydration capacity occurred at dehydration beyond 50% declines of gs , Kleaf and turgor loss point. Losses of Fv /Fm occurred after much stronger dehydration and were not recovered with leaf rehydration. Across species, tissue dehydration thresholds were intercorrelated, suggesting trait co-selection. Thresholds for each type of functional decline were much less variable across species in terms of relative water content than Ψleaf . The stomatal and leaf hydraulic systems show early functional declines before cell integrity is lost. Substantial damage to the photochemical apparatus occurs at extreme dehydration, after complete stomatal closure, and seems to be irreversible.

Leaf density links the resistance to mechanical and hydraulic stress in Mediterranean shrubs as it is associated with the water potential at turgor loss and the moduli of elasticity and strength. Understanding the patterns of hydraulic and mechanical trait variation in vascular plants is critical to predicting species’ stress tolerance. Although previous work has shown that hydraulic and mechanical traits are decoupled in stems, there is little information available for leaves, which are organs more diversified in structure, function, and possibly drought tolerance strategies across habitats. We tested for coordination between leaf hydraulic traits related to drought tolerance and the mechanical resistance of leaves, for 17 shrub species from the arid and semiarid vegetation of the California Floristic Province. Bayesian and phylogenetic correlations showed that across species, hydraulic and mechanical traits both had strong associations with the water potential at turgor loss, and with leaf tissue density. However, leaf maximum hydraulic conductance and the water potential at 50% and 80% loss of hydraulic conductance were statistically independent of two key mechanical traits, the leaf modulus of elasticity and leaf structural strength. Our results suggest that leaf biomechanical traits, which reflect construction costs and contribute to leaf longevity, are decoupled from hydraulic capacity and safety. The independence of hydraulic and mechanical protection in leaves enables a wide range of trait combinations in leaves, which would increase their adaptive potential across ecosystems.

Understanding how plants acclimate to drought is crucial for predicting future vulnerability, yet seasonal acclimation of traits that improve drought tolerance in trees remains poorly resolved. We hypothesized that dry season acclimation of leaf and stem traits influencing shoot water storage and hydraulic capacitance would mitigate the drought-associated risks of reduced gas exchange and hydraulic failure in the mangrove Sonneratia alba. By late dry season, availability of stored water had shifted within leaves and between leaves and stems. While whole shoot capacitance remained stable, the symplastic fraction of leaf water increased 86%, leaf capacitance increased 104% and stem capacitance declined 80%. Despite declining plant water potentials, leaf and whole plant hydraulic conductance remained unchanged, and midday assimilation rates increased. Further, the available leaf water between the minimum water potential observed and that corresponding to 50% loss of stem conductance increased 111%. Shifting availability of pools of water, within and between organs, maintained leaf water available to buffer periods of increased photosynthesis and losses in stem hydraulic conductivity, mitigating risks of carbon depletion and hydraulic failure during atmospheric drought. Seasonal changes in access to tissue and organ water may have an important role in drought acclimation and avoidance.

In 1992, the Union of Concerned Scientists shared their 'World Scientists'Warning to Humanity' with governmental leaders worldwide, calling for immediate action to halt the environmental degradation that threatens the systems that support life on Earth. A follow-up 'Second Warning' was released in 2017, with over 15 000 scientists as signatories, describing the lack of progress in adopting the sustainable practices necessary to safeguard the biosphere. In their 'SecondWarning', Ripple and colleagues provided 13 'diverse and effective steps humanity can take to transition to sustainability.' Here, we discuss how the field of conservation physiology can contribute to six of these goals: (i) prioritizing connected, well-managed reserves; (ii) halting the conversion of native habitats to maintain ecosystemservices; (iii) restoring native plant communities; (iv) rewilding regions with native species; (v) developing policy instruments; and (vi) increasing outdoor education, societal engagement and reverence for nature. Throughout, we focus our recommendations on specific aspects of physiological function while acknowledging that the exact traits that will be useful in each context are oftenstill being determined and refined. However, for each goal, we include a short case study to illustrate a specific physiological trait or group of traits that is already being utilized in that context. We conclude with suggestions for how conservation physiologists can broaden the impact of their science aimed at accomplishing the goals of the 'Second Warning'. Overall, we provide an overview of how conservation physiology can contribute to addressing the grand socio-environmental challenges of our time. National Science Foundation MacroSystems Biology program [DEB-DEB-1340856]; National Science FoundationNational Science Foundation (NSF) [IOS-1755055, 1951244, 2017949]; FutureMARES (Climate Change and Future Marine Ecosystem Services and Biodiversity, EU H2020) [869300] Published version K.R.H. was supported by the National Science Foundation MacroSystems Biology program (DEB-DEB-1340856). W.A.H. was supported by the National Science Foundation (grant #IOS-1755055). M.A.P. received partial support from FutureMARES (Climate Change and Future Marine Ecosystem Services and Biodiversity, EU H2020, award no. 869300). L.S. is funded by the National Science Foundation (grants 1951244 and 2017949).

Hydraulic conductance is recognized as a major determinant of gas exchange and productivity. However, whether this also applies to seedlings, a critically important stage for vegetation regeneration, has been largely unknown. We analyzed the hydraulic and stomatal conductance of leaves and shoots for 6-wk-old Acer pseudoplatanus seedlings emerging in different lowland and treeline habitats and under glasshouse conditions, respectively, as well as on 9-, 15- and 18-wk-old plants, and related findings to leaf and xylem anatomical traits. Treeline seedlings had higher leaf area-specific shoot hydraulic conductance (Kshoot-L ), and stomatal conductance (gs ), associated with wider xylem conduits, lower leaf area and higher stomatal density than lowland and glasshouse-grown plants. Across the first 18 wk of development, seedlings increased four-fold in absolute shoot hydraulic conductance (Kshoot ) and declined by half in Kshoot-L , with correlated shifts in xylem and leaf anatomy. Distal leaves had higher leaf hydraulic conductance (Kleaf ) and gs compared to basal leaves. Seedlings show strong variation across growth environments and ontogenetic shifts in hydraulic and anatomical parameters. Across growth sites, ontogenetic stages and leaf orders, gs was tightly correlated with Kshoot-L and Kleaf , balancing hydraulic supply with demand for the earliest stages of seedling establishment.

Clarifying the mechanisms of leaf and whole plant drought responses is critical to predict the impacts of ongoing climate change. The loss of rehydration capacity has been used for decades as a metric of leaf dehydration tolerance but has not been compared with other aspects of drought tolerance. We refined methods for quantifying the percent loss of rehydration capacity (PLRC), and for 18 Southern California woody species, we determined the relative water content and leaf water potential at PLRC of 10%, 25%, and 50%, and, additionally, the PLRC at important stages of dehydration including stomatal closure and turgor loss. On average, PLRC of 10% occurred below turgor loss point and at similar water status to 80% decline of stomatal conductance. As hypothesized, the sensitivity to loss of leaf rehydration capacity varied across species, leaf habits, and ecosystems and correlated with other drought tolerance traits, including the turgor loss point and structural traits including leaf mass per area. A new database of PLRC for 89 species from the global literature indicated greater leaf rehydration capacity in ecosystems with lower growing season moisture availability, indicating an adaptive role of leaf cell dehydration tolerance within the complex of drought tolerance traits.

The lignification of the leaf vein bundle sheath (BS) has been observed in many species and would reduce conductance from xylem to mesophyll. We hypothesized that lignification of the BS in lower-order veins would provide benefits for water delivery through the vein hierarchy but that the lignification of higher-order veins would limit transport capacity from xylem to mesophyll and leaf hydraulic conductance (Kleaf ). We further hypothesized that BS lignification would mediate the relationship of Kleaf to vein length per area. We analysed the dependence of Kleaf , and its light response, on the lignification of the BS across vein orders for 11 angiosperm tree species. Eight of 11 species had lignin deposits in the BS of the midrib, and two species additionally only in their secondary veins, and for six species up to their minor veins. Species with lignification of minor veins had a lower hydraulic conductance of xylem and outside-xylem pathways and lower Kleaf . Kleaf could be strongly predicted by vein length per area and highest lignified vein order (R2 = .69). The light-response of Kleaf was statistically independent of BS lignification. The lignification of the BS is an important determinant of species variation in leaf and thus whole plant water transport.

Reanalysis of published experimental data shows that in dehydrating leaves ABA accumulation is linked with reduction of cell volume rather than turgor, providing clues toward signaling mechanisms.

Leaf structure and water relations were studied in a temperate population of Avicennia marina subsp. australasica along a natural salinity gradient [28 to 49 parts per thousand (ppt)] and compared with two subspecies grown naturally in similar soil salinities to those of subsp. australasica but under different climates: subsp. eucalyptifolia (salinity 30 ppt, wet tropics) and subsp. marina (salinity 46 ppt, arid tropics). Leaf thickness, leaf dry mass per area and water content increased with salinity and aridity. Turgor loss point declined with increase in soil salinity, driven mainly by differences in osmotic potential at full turgor. Nevertheless, a high modulus of elasticity (e) contributed to maintenance of high cell hydration at turgor loss point. Despite similarity among leaves in leaf water storage capacitance, total leaf water storage increased with increasing salinity and aridity. The time that stored water alone could sustain an evaporation rate of 1 mmol m-2 s-1 ranged from 77 to 126 min from subspecies eucalyptifolia to ssp. marina, respectively. Achieving full leaf hydration or turgor would require water from sources other than the roots, emphasizing the importance of multiple water sources to growth and survival of Avicennia marina across gradients in salinity and aridity.

Leaf hydraulic supply is crucial to maintaining open stomata for CO2 capture and plant growth. During drought-induced dehydration, the leaf hydraulic conductance (Kleaf) declines, which contributes to stomatal closure and, eventually, to leaf death. Previous studies have tended to attribute the decline of Kleaf to embolism in the leaf vein xylem. We visualized at high resolution and quantified experimentally the hydraulic vulnerability of xylem and outside-xylem pathways and modeled their respective influences on plant water transport. Evidence from all approaches indicated that the decline of Kleaf during dehydration arose first and foremost due to the vulnerability of outside-xylem tissues. In vivo x-ray microcomputed tomography of dehydrating leaves of four diverse angiosperm species showed that, at the turgor loss point, only small fractions of leaf vein xylem conduits were embolized, and substantial xylem embolism arose only under severe dehydration. Experiments on an expanded set of eight angiosperm species showed that outside-xylem hydraulic vulnerability explained 75% to 100% of Kleaf decline across the range of dehydration from mild water stress to beyond turgor loss point. Spatially explicit modeling of leaf water transport pointed to a role for reduced membrane conductivity consistent with published data for cells and tissues. Plant-scale modeling suggested that outside-xylem hydraulic vulnerability can protect the xylem from tensions that would induce embolism and disruption of water transport under mild to moderate soil and atmospheric droughts. These findings pinpoint outside-xylem tissues as a central locus for the control of leaf and plant water transport during progressive drought.

The significance of shoot surface water uptake (SSWU) has been debated, and it would depend on the range of conditions under which it occurs. We hypothesized that the decline of leaf hydraulic conductance (Kleaf ) in response to dehydration may be recovered through SSWU, and that the hydraulic conductance to SSWU (Ksurf ) declines with dehydration. We quantified effects of leaf dehydration on Ksurf and effects of SSWU on recovery of Kleaf in dehydrated leaves of Avicennia marina. SSWU led to overnight recovery of Kleaf , with recovery retracing the same path as loss of Kleaf in response to dehydration. SSWU declined with dehydration. By contrast, Ksurf declined with rehydration time but not with dehydration. Our results showed a role of SSWU in the recovery of leaf hydraulic conductance and revealed that SSWU is sensitive to leaf hydration status. The prevalence of SSWU in vegetation suggests an important role for atmospheric water sources in maintenance of leaf hydraulic function, with implications for plant responses to changing environments.

Salinity tolerance in plant species varies widely due to adaptation and acclimation processes at the cellular and whole-plant scales. In mangroves, extreme substrate salinity induces hydraulic failure and ion excess toxicity and reduces growth and survival, thus suggesting a potentially critical role for physiological acclimation to salinity. We tested the hypothesis that osmotic adjustment, a key type of plasticity that mitigates salinity shock, would take place in coordination with declines in whole-plant hydraulic conductance in a common garden experiment using saplings of three mangrove species with different salinity tolerances (Avicennia germinans L., Rhizophora mangle L. and Laguncularia racemosa (L.) C.F. Gaertn., ordered from higher to lower salinity tolerance). For each mangrove species, four salinity treatments (1, 10, 30 and 50 practical salinity units) were established and the time trajectories were determined for leaf osmotic potential (Ψs), stomatal conductance (gs), whole-plant hydraulic conductance (Kplant) and predawn disequilibrium between xylem and substrate water potentials (Ψpdd). We expected that, for all three species, salinity increments would result in coordinated declines in Ψs, gs and Kplant, and that the Ψpdd would increase with substrate salinity and time of exposure. In concordance with our predictions, reductions in substrate water potential promoted a coordinated decline in Ψs, gs and Kplant, whereas the Ψpdd increased substantially during the first 4 days but dissipated after 7 days, indicating a time lag for equilibration after a change in substratum salinity. Our results show that mangroves confront and partially ameliorate acute salinity stress via simultaneous reductions in Ψs, gs and Kplant, thus developing synergistic physiological responses at the cell and whole-plant scales.

Equations for stomatal density and maximum theoretical stomatal conductance as functions of stomatal initiation rate, epidermal cell size, and stomatal size enable scaling from development to flux.

Photosynthesis in bryophytes and lycophytes has received less attention than terrestrial plant groups. In particular, few studies have addressed the nonstomatal diffusion conductance to CO2 gnsd of these plant groups. Their lower photosynthetic rate per leaf mass area at any given nitrogen concentration compared with vascular plants suggested a stronger limitation by CO2 diffusion. We hypothesized that bryophyte and lycophyte photosynthesis is largely limited by low gnsd . Here, we studied CO2 diffusion inside the photosynthetic tissues and its relationships with photosynthesis and anatomical parameters in bryophyte and lycophyte species in Antarctica, Australia, Estonia, Hawaii and Spain. On average, lycophytes and, specially, bryophytes had the lowest photosynthetic rates and nonstomatal diffusion conductance reported for terrestrial plants. These low values are related to their very thick cell walls and their low exposure of chloroplasts to cell perimeter. We conclude that the reason why bryophytes lie at the lower end of the leaf economics spectrum is their strong nonstomatal diffusion conductance limitation to photosynthesis, which is driven by their specific anatomical characteristics.

The influence of the dynamics of leaf hydraulic conductance (K-leaf) diurnally and during dehydration on stomatal conductance and photosynthesis remains unclear. Using the model species Arabidopsis (Arabidopsis thaliana ecotype Columbia-0), we applied a multitiered approach including physiological measurements, high-resolution x-ray microcomputed tomography, and modeling at a range of scales to characterize (1) K-leaf decline during dehydration; (2) its basis in the hydraulic conductances of leaf xylem and outside-xylem pathways (K-ox); (3) the dependence of its dynamics on irradiance; (4) its impact on diurnal patterns of stomatal conductance and photosynthetic rate; and (5) its influence on gas exchange and survival under simulated drought regimes. Arabidopsis leaves showed strong vulnerability to dehydration diurnally in both gas exchange and hydraulic conductance, despite lack of xylem embolism or conduit collapse above the turgor loss point, indicating a pronounced sensitivity of K-ox to dehydration. K(leaf )increased under higher irradiance in well-hydrated leaves across the full range of water potential, but no shift in K-leaf vulnerability was observed. Modeling indicated that responses to dehydration and irradiance are likely attributable to changes in membrane permeability and that a dynamic K-ox would contribute strongly to stomatal closure, improving performance, survival, and efficient water use during drought. These findings for Columbia-0 provide a baseline for assessing variation across genotypes in hydraulic traits and their influence on gas exchange during dehydration.

Leaves are arguably the most complex and important physicobiological systems in the ecosphere. Yet, water transport outside the leaf xylem remains poorly understood, despite its impacts on stomatal function and photosynthesis. We applied anatomical measurements from 14 diverse species to a novel model of water flow in an areole (the smallest region bounded by minor veins) to predict the impact of anatomical variation across species on outside-xylem hydraulic conductance (Kox). Several predictions verified previous correlational studies: (1) vein length per unit area is the strongest anatomical determinant of Kox, due to effects on hydraulic pathlength and bundle sheath (BS) surface area; (2) palisade mesophyll remains well hydrated in hypostomatous species, which may benefit photosynthesis, (3) BS extensions enhance Kox; and (4) the upper and lower epidermis are hydraulically sequestered from one another despite their proximity. Our findings also provided novel insights: (5) the BS contributes a minority of outside-xylem resistance; (6) vapor transport contributes up to two-thirds of Kox; (7) Kox is strongly enhanced by the proximity of veins to lower epidermis; and (8) Kox is strongly influenced by spongy mesophyll anatomy, decreasing with protoplast size and increasing with airspace fraction and cell wall thickness. Correlations between anatomy and Kox across species sometimes diverged from predicted causal effects, demonstrating the need for integrative models to resolve causation. For example, (9) Kox was enhanced far more in heterobaric species than predicted by their having BS extensions. Our approach provides detailed insights into the role of anatomical variation in leaf function.

Leaf hydraulic conductance (Kleaf ) quantifies the capacity of a leaf to transport liquid water and is a major constraint on light-saturated stomatal conductance (gs ) and photosynthetic rate (Amax ). Few studies have tested the plasticity of Kleaf and anatomy across growth light environments. These provided conflicting results. The Hawaiian lobeliads are an excellent system to examine plasticity, given the striking diversity in the light regimes they occupy, and their correspondingly wide range of Amax , allowing maximal carbon gain for success in given environments. We measured Kleaf , Amax , gs and leaf anatomical and structural traits, focusing on six species of lobeliads grown in a common garden under two irradiances (300/800 μmol photons m(-2) s(-1) ). We tested hypotheses for light-induced plasticity in each trait based on expectations from optimality. Kleaf , Amax , and gs differed strongly among species. Sun/shade plasticity was observed in Kleaf , Amax, and numerous traits relating to lamina and xylem anatomy, venation, and composition, but gs was not plastic with growth irradiance. Species native to higher irradiance showed greater hydraulic plasticity. Our results demonstrate that a wide set of leaf hydraulic, stomatal, photosynthetic, anatomical, and structural traits tend to shift together during plasticity and adaptation to diverse light regimes, optimizing performance from low to high irradiance.

Resolving the drivers of hydraulic decline during drought is crucial for understanding drought tolerance in crops and natural ecosystems. In the past 15 years, studies of the decline of leaf hydraulic conductance (Kleaf) have supported a major role in controlling plant drought responses. We analyzed the variation in Kleaf decline with dehydration in a global database of 310 species, providing novel insights into its underlying mechanisms, its co-ordination with stem hydraulics, its influence on gas exchange and drought tolerance, and its linkage with species ecological distributions. Kleaf vulnerability varied strongly within and across lineages, growth forms, and biomes. A critical literature review indicates that changes in hydraulic conductance outside the xylem with dehydration drive the overall decline of Kleaf. We demonstrate a significant leaf hydraulic safety-efficiency trade-off across angiosperm species and discuss the importance of the large variation around this trend. Leaves tend to be more vulnerable than stems, with their vulnerabilities co-ordinated across species, and importantly linked with adaptation across biomes. We hypothesize a novel framework to explain diversity across species in the co-ordination of Kleaf and gas exchange during dehydration. These findings reflect considerable recent progress, yet new tools for measurement, visualization, and modeling will result in ongoing discoveries important across fields in plant biology.

© 2017 American Society of Plant Biologists. All rights reserved. The sites of evaporation within leaves are unknown, but they have drawn attention for decades due to their perceived implications for many factors, including patterns of leaf isotopic enrichment, the maintenance of mesophyll water status, stomatal regulation, and the interpretation of measured stomatal and leaf hydraulic conductances. We used a spatially explicit model of coupled water and heat transport outside the xylem, MOFLO 2.0, to map the distribution of net evaporation across leaf tissues in relation to anatomy and environmental parameters. Our results corroborate earlier predictions that most evaporation occurs from the epidermis at low light and moderate humidity but that the mesophyll contributes substantially when the leaf center is warmed by light absorption, and more so under high humidity. We also found that the bundle sheath provides a significant minority of evaporation (15% in darkness and 18% in high light), that the vertical center of amphistomatous leaves supports net condensation, and that vertical temperature gradients caused by light absorption vary over 10-fold across species, reaching 0.3°C. We show that several hypotheses that depend on the evaporating sites require revision in light of our findings, including that experimental measurements of stomatal and hydraulic conductances should be affected directly by changes in the location of the evaporating sites. We propose a new conceptual model that accounts for mixed-phase water transport outside the xylem. These conclusions have far-reaching implications for inferences in leaf hydraulics, gas exchange, water use, and isotope physiology.

A recent study found that cutting shoots under water while xylem was under tension (which has been the standard protocol for the past few decades) could produce artefactual embolisms inside the xylem, overestimating hydraulic vulnerability relative to shoots cut under water after relaxing xylem tension (Wheeler et al. 2013). That study also raised the possibility that such a 'Wheeler effect' might occur in studies of leaf hydraulic vulnerability. We tested for such an effect for four species by applying a modified vacuum pump method to leaves with minor veins severed, to construct leaf xylem hydraulic vulnerability curves. We tested for an impact on leaf xylem hydraulic conductance (Kx ) of cutting the petiole and minor veins under water for dehydrated leaves with xylem under tension compared with dehydrated leaves after previously relaxing xylem tension. Our results showed no significant 'cutting artefact' for leaf xylem. The lack of an effect for leaves could not be explained by narrower or shorter xylem conduits, and may be due to lesser mechanical stress imposed when cutting leaf petioles, and/or to rapid refilling of emboli in petioles. These findings provide the first validation of previous measurements of leaf hydraulic vulnerability against this potential artefact.

Two highly contrasting variables summarizing the efficiency of transport of materials within the leaf are recognized as playing central roles in determining gas exchange and plant performance. This paper summarizes current approaches for the measurement of mesophyll conductance to CO2 (g m) and leaf hydraulic conductance (K leaf) and addresses the physiological integration of these parameters. First, the most common methods to determine g m and K leaf are summarized. Next, novel data compilation is analysed, which indicates that, across diverse species, g m is strongly linked with gas exchange parameters such as net CO2 assimilation (A area) and stomatal conductance (g s), and with K leaf, independently of leaf vein length per leaf area. Based on their parallel responses to a number of environmental variables, this review proposes that g m is linked to the outside-xylem but not to the xylem component of K leaf. Further, a mechanistic hypothesis is proposed to explain the interactions among all these and other physiological parameters. Finally, the possibility of estimating g m based on this hypothesis was tested using a regression analysis and a neurofuzzy logic approach. These approaches enabled the estimation of g m of given species from K leaf and leaf mass per area, providing a higher predictive power than from either parameter alone. The possibility of estimating g m from measured K leaf or vice-versa would result in a rapid increase in available data. Studies in which g m, K leaf, and leaf mass per area are simultaneously determined are needed in order to confirm and strengthen predictive and explanatory models for these parameters and importantly improve resolution of the integrated hydraulic-stomatal-photosynthetic system.

Leaf vein traits are implicated in the determination of gas exchange rates and plant performance. These traits are increasingly considered as causal factors affecting the 'leaf economic spectrum' (LES), which includes the light-saturated rate of photosynthesis, dark respiration, foliar nitrogen concentration, leaf dry mass per area (LMA) and leaf longevity. This article reviews the support for two contrasting hypotheses regarding a key vein trait, vein length per unit leaf area (VLA). Recently, Blonder et al. (2011, 2013) proposed that vein traits, including VLA, can be described as the 'origin' of the LES by structurally determining LMA and leaf thickness, and thereby vein traits would predict LES traits according to specific equations. Careful re-examination of leaf anatomy, published datasets, and a newly compiled global database for diverse species did not support the 'vein origin' hypothesis, and moreover showed that the apparent power of those equations to predict LES traits arose from circularity. This review provides a 'flux trait network' hypothesis for the effects of vein traits on the LES and on plant performance, based on a synthesis of the previous literature. According to this hypothesis, VLA, while virtually independent of LMA, strongly influences hydraulic conductance, and thus stomatal conductance and photosynthetic rate. We also review (i) the specific physiological roles of VLA; (ii) the role of leaf major veins in influencing LES traits; and (iii) the role of VLA in determining photosynthetic rate per leaf dry mass and plant relative growth rate. A clear understanding of leaf vein traits provides a new perspective on plant function independently of the LES and can enhance the ability to explain and predict whole plant performance under dynamic conditions, with applications towards breeding improved crop varieties.

Author(s): Sack, Lawren; Buckley, Thomas N; Scoffoni, Christine | Abstract: As plant tissues dehydrate, water transport efficiency declines, a process typically attributed to air obstruction (embolism) in the xylem. Trifilo et al. (pages 5029-5039) dissect leaf hydraulic vulnerability and show that both xylem and living tissues may be important. If confirmed and clarified, an important role for outsidexylem hydraulic decline will change our understanding of how plants transport water and control biosphere carbon and water fluxes.

© 2016 John Wiley & Sons Ltd It was shown over 40 years ago that plants maximize carbon gain for a given rate of water loss if stomatal conductance, gs, varies in response to external and internal conditions such that the marginal carbon revenue of water, ∂A/∂E, remains constant over time. This theory has long held promise for understanding the physiological ecology of water use and for informing models of plant-atmosphere interactions. Full realization of this potential hinges on three questions: (i) Are analytical approximations adequate for applying the theory at diurnal time scales? (ii) At what time scale is it realistic and appropriate to apply the theory? (iii) How should gs vary to maximize growth over long time scales? We review the current state of understanding for each of these questions and describe future research frontiers. In particular, we show that analytical solutions represent the theory quite poorly, especially when boundary layer or mesophyll resistances are significant; that diurnal variations in hydraulic conductance may help or hinder maintenance of ∂A/∂E, and the matter requires further study; and that optimal diurnal responses are distinct from optimal long-term variations in gs, which emerge from optimal shifts in carbon partitioning at the whole-plant scale.

Water plays a central role in plant biology and the efficiency of water transport throughout the plant affects both photosynthetic rate and growth, an influence that scales up deterministically to the productivity of terrestrial ecosystems. Moreover, hydraulic traits mediate the ways in which plants interact with their abiotic and biotic environment. At landscape to global scale, plant hydraulic traits are important in describing the function of ecological communities and ecosystems. Plant hydraulics is increasingly recognized as a central hub within a network by which plant biology is connected to palaeobiology, agronomy, climatology, forestry, community and ecosystem ecology and earth-system science. Such grand challenges as anticipating and mitigating the impacts of climate change, and improving the security and sustainability of our food supply rely on our fundamental knowledge of how water behaves in the cells, tissues, organs, bodies and diverse communities of plants. A workshop, 'Emerging Frontiers in Plant Hydraulics' supported by the National Science Foundation, was held in Washington DC, 2015 to promote open discussion of new ideas, controversies regarding measurements and analyses, and especially, the potential for expansion of up-scaled and down-scaled inter-disciplinary research, and the strengthening of connections between plant hydraulic research, allied fields and global modelling efforts.

© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust Ecosystems worldwide are facing increasingly severe and prolonged droughts during which hydraulic failure from drought-induced embolism can lead to organ or whole plant death. Understanding the determinants of xylem failure across species is especially critical in leaves, the engines of plant growth. If the vulnerability segmentation hypothesis holds within leaves, higher order veins that are most terminal in the plant hydraulic system should be more susceptible to embolism to protect the rest of the water transport system. Increased vulnerability in the higher order veins would also be consistent with these experiencing the greatest tensions in the plant xylem network. To test this hypothesis, we combined X-ray micro-computed tomography imaging, hydraulic experiments, cross-sectional anatomy and 3D physiological modelling to investigate how embolisms spread throughout petioles and vein orders during leaf dehydration in relation to conduit dimensions. Decline of leaf xylem hydraulic conductance (Kx) during dehydration was driven by embolism initiating in petioles and midribs across all species, and Kx vulnerability was strongly correlated with petiole and midrib conduit dimensions. Our simulations showed no significant impact of conduit collapse on Kx decline. We found xylem conduit dimensions play a major role in determining the susceptibility of the leaf water transport system during strong leaf dehydration.

The leaf is a hydraulic bottleneck, accounting for a large part of plant resistance. Thus, the leaf hydraulic conductance (Kleaf) is of key importance in determining stomatal conductance (gs) and rates of gas exchange. Previous studies showed that Kleaf is dynamic with leaf water status and irradiance. For four species, we tested the combined impacts of these factors on Kleaf and on gs. We determined responses of Kleaf and gs to declining leaf water potential (Yleaf) under low and high irradiance ( 900 mmol photons m -2 s -1 photosynthetically active radiation, respectively). We hypothesized greater Kleaf vulnerability under high irradiance. We also hypothesized that Kleaf and gs would be similar in their responses to either light or dehydration: similar light-responses of Kleaf and gs would stabilize Yleaf across irradiances for leaves transpiring at a given vapour pressure deficit, and similar dehydration responses would arise from the control of stomata by Yleaf or a correlated signal. For all four species, the Kleaf light response declined from full hydration to turgor loss point. The Kleaf and gs differed strongly in their light- and dehydration responses, supporting optimization of hydraulic transport across irradiances, and semi-independent, flexible regulation of liquid and vapour phase water transport with leaf water status.

We determined effects of venation traits on hydraulic conductance of phyllodes (foliage), using an array of Acacia s.str. species with diverse phyllode morphologies as the source of variation. Measurements were made on phyllodes from 44 species, grown in common gardens but originating from different positions along a precipitation gradient. Kphyllode varied 18-fold and was positively correlated with primary nerve hydraulic conductance, and with primary nerve (vein) density but not with minor nerve density, in contrast with previous studies of true leaves in other dicotyledons. Phyllodes with higher primary nerve density also had greater mass per area (PMA) and larger bundle sheath extensions (BSEs) from their minor nerves. We suggest that higher primary nerve conductivity and density may decrease the distance travelled in the high-resistance extraxylem pathways of the phyllode. Further, larger BSEs may increase the area available for dispersion of water from the xylem to the extra-xylem tissue. High PMA phyllodes were more common in acacias from areas receiving lower annual precipitation. Maximizing efficient water movement through phyllodes may be more important where rainfall is meagre and infrequent, explaining relationships between nerve patterns and the climates of origin in Australian phyllodinous Acacia.

Responses of leaf and shoot hydraulic conductance to light quality were examined on shoots of silver birch (Betula pendula), cut from lower (‘shade position’) and upper thirds of the crowns (‘sun position’) of trees growing in a natural temperate forest stand. Hydraulic conductances of leaf blades (Klb), petioles (KP) and branches (i.e. leafless stem;KB) were determined using a high pressure flow meter in steady state mode. The shoots were exposed to photosynthetic photon flux density of 200–250 mmol m -2 s -1 using white, blue or red light. Klb depended significantly on both light quality and canopy position (P < 0.001), KB on canopy position (P < 0.001) and exposure time (P = 0.014), and none of the three factors had effect on KP. The highest values of Klb were recorded under the blue light (3.63 and 3.13 ¥ 10 -4 kg m -2 MPa -1 s -1 for the sun and shade leaves, respectively), intermediate values under white light (3.37 and 2.46 ¥ 10 -4 kg m -2 MPa -1 s -1 , respectively) and lowest

Summary • Plant hydraulic architecture has been studied extensively, yet we know little about how hydraulic properties relate to species’ life history strategies, such as drought and shade tolerance. The prevailing theories seem contradictory. • We measured the sapwood (Ks) and leaf (Kl) hydraulic conductivities of 40 coexisting tree species in a Bolivian dry forest, and examined associations with functional stem and leaf traits and indices of species’ drought (dry-season leaf water potential) and shade (juvenile crown exposure) tolerance. • Hydraulic properties varied across species and between life-history groups (pioneers vs shade-tolerant, and deciduous vs evergreen species). In addition to the expected negative correlation of Kl with drought tolerance, we found a strong, negative correlation between Kl and species’ shade tolerance. Across species, Ks and Kl were negatively correlated with wood density and positively with maximum vessel length. Consequently, drought and shade tolerance scaled similarly with hydraulic properties, wood density and leaf dry matter content. We found that deciduous species also had traits conferring efficient water transport relative to evergreen species. • Hydraulic properties varied across species, corresponding to the classical tradeoff between hydraulic efficiency and safety, which for these dry forest trees resulted in coordinated drought and shade tolerance across species rather than the frequently hypothesized trade-off.

Bundle sheath extensions (BSEs) are key features of leaf structure with currently little-understood functions. To test the hypothesis that BSEs reduce the hydraulic resistance from the bundle sheath to the epidermis (r be) and thereby accelerate hydropassive stomatal movements, we compared stomatal responses with reduced humidity and leaf excision among 20 species with heterobaric or homobaric leaves and herbaceous or woody life forms. We hypothesized that low r be due to the presence of BSEs would increase the rate of stomatal opening (V) during transient wrong-way responses, but more so during wrong-way responses to excision (V e) than humidity (V h), thus increasing the ratio of V e to V h. We predicted the same trends for herbaceous relative to woody species given greater hydraulic resistance in woody species. We found that V e, V h, and their ratio were 2.3 to 4.4 times greater in heterobaric than homobaric leaves and 2.0 to 3.1 times greater in herbaceous than woody species. To assess possible causes for these differences, we simulated these experiments in a dynamic compartment/resistance model, which predicted larger V e and V e/V h in leaves with smaller r be. These results support the hypothesis that BSEs reduce r be. Comparison of our data and simulations suggested that r be is approximately 4 to 16 times larger in homobaric than heterobaric leaves. Our study provides new evidence that variations in the distribution of hydraulic resistance within the leaf and plant are central to understanding dynamic stomatal responses to water status and their ecological correlates and that BSEs play several key roles in the functional ecology of heterobaric leaves.

Hawaiian endemic tree Acacia koa is a model for heteroblasty with bipinnately compound leaves and phyllodes. Previous studies suggested three hypotheses for their functional differentiation: an advantage of leaves for early growth or shade tolerance, and an advantage of phyllodes for drought tolerance.We tested the ability of these hypotheses to explain differences between leaf types for potted plants in 104 physiological and morphological traits, including gas exchange, structure and composition, hydraulic conductance, and responses to varying light, intercellular CO2, vapour pressure deficit (VPD) and drought. Leaf types were similar in numerous traits including stomatal pore area per leaf area, leaf area-based gas exchange rates and cuticular conductance. Each hypothesis was directly supported by key differences in function. Leaves had higher mass-based gas exchange rates, while the water storage tissue in phyllodes contributed to greater capacitance per area; phyllodes also showed stronger stomatal closure at high VPD, and higher maximum hydraulic conductance per area, with stronger decline during desiccation and recovery with rehydration. While no single hypothesis completely explained the differences between leaf types, together the three hypotheses explained 91% of differences.These findings indicate that the heteroblasty confers multiple benefits, realized across different developmental stages and environmental contexts.

Cavitation resistance is a critical determinant of drought tolerance in tropical tree species, but little is known of its association with life history strategies, particularly for seasonal dry forests, a system critically driven by variation in water availability. We analysed vulnerability curves for saplings of 13 tropical dry forest tree species differing in life history and leaf phenology. We examined how vulnerability to cavitation (P50) related to dry season leaf water potentials and stem and leaf traits. P50-values ranged from −0.8 to −6.2 MPa, with pioneers on average 38% more vulnerable to cavitation than shade-tolerants. Vulnerability to cavitation was related to structural traits conferring tissue stress vulnerability, being negatively correlated with wood density, and surprisingly maximum vessel length. Vulnerability to cavitation was negatively related to the Huber-value and leaf dry matter content, and positively with leaf size. It was not related to SLA. We found a strong trade-off between cavitation resistance and hydraulic efficiency. Most species in the field were operating at leaf water potentials well above their P50, but pioneers and deciduous species had smaller hydraulic safety margins than shade-tolerants and evergreens. A trade-off between hydraulic safety and efficiency underlies ecological differentiation across these tropical dry forest tree species.

Inter- and intraspecific variation in hydraulic traits was investigated in nine Cordia (Boraginaceae) species growing in three tropical rainforests differing in mean annual precipitation (MAP). Interspecific variation was examined for the different Cordia species found at each site, and intraspecific variation was studied in populations of the widespread species Cordia alliodora across the three sites. Strong intra- and interspecific variation were observed in vulnerability to drought-induced embolism. Species growing at drier sites were more resistant to embolism than those growing at moister sites; the same pattern was observed for populations of C. alliodora. By contrast, traits related to hydraulic capacity, including stem xylem vessel diameter, sapwood specific conductivity (K(s)) and leaf specific conductivity (K(L)), varied strongly but independently of MAP. For C. alliodora, xylem anatomy, K(s), K(L) and Huber value varied little across sites, with K(s) and K(L) being consistently high relative to other Cordia species. A constitutively high hydraulic capacity coupled with plastic or genotypic adjustment in vulnerability to embolism and leaf water relations would contribute to the ability of C. alliodora to establish and compete across a wide precipitation gradient.

Summary: Cycads are the most ancient lineage of living seed plants, but the design of their leaves has received little study. We tested whether cycad leaves are governed by the same fundamental design principles previously established for ferns, conifers and angiosperms, and characterized the uniqueness of this relict lineage in foliar trait relationships. Leaf structure, photosynthesis, hydraulics and nutrient composition were studied in 33 cycad species from nine genera and three families growing in two botanical gardens. Cycads varied greatly in leaf structure and physiology. Similarly to other lineages, light-saturated photosynthetic rate per mass (Am) was related negatively to leaf mass per area and positively to foliar concentrations of chlorophyll, nitrogen (N), phosphorus and iron, but unlike angiosperms, leaf photosynthetic rate was not associated with leaf hydraulic conductance. Cycads had lower photosynthetic N use efficiency and higher photosynthetic performance relative to hydraulic capacity compared with other lineages. These findings extend the relationships shown for foliar traits in angiosperms to the cycads. This functional convergence supports the modern synthetic understanding of leaf design, with common constraints operating across lineages, even as they highlight exceptional aspects of the biology of this key relict lineage. Fil: Zhang, Yong Jiang. Chinese Academy of Sciences; República de China. Harvard University; Estados Unidos Fil: Cao, Kun Fang. Guangxi University; China Fil: Sack, Lawren. University of California at Los Angeles; Estados Unidos Fil: Li, Nan. Chinese Academy of Sciences; República de China Fil: Wei, Xue Mei. Chinese Academy of Sciences; República de China Fil: Goldstein, Guillermo Hernan. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Ecología, Genética y Evolución de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Ecología, Genética y Evolución de Buenos Aires; Argentina

We compiled a global database for leaf, stem and root biomass representing c. 11 000 records for c. 1200 herbaceous and woody species grown under either controlled or field conditions. We used this data set to analyse allometric relationships and fractional biomass distribution to leaves, stems and roots. We tested whether allometric scaling exponents are generally constant across plant sizes as predicted by metabolic scaling theory, or whether instead they change dynamically with plant size. We also quantified interspecific variation in biomass distribution among plant families and functional groups. Across all species combined, leaf vs stem and leaf vs root scaling exponents decreased from c. 1.00 for small plants to c. 0.60 for the largest trees considered. Evergreens had substantially higher leaf mass fractions (LMFs) than deciduous species, whereas graminoids maintained higher root mass fractions (RMFs) than eudicotyledonous herbs. These patterns do not support the hypothesis of fixed allometric exponents. Rather, continuous shifts in allometric exponents with plant size during ontogeny and evolution are the norm. Across seed plants, variation in biomass distribution among species is related more to function than phylogeny. We propose that the higher LMF of evergreens at least partly compensates for their relatively low leaf area : leaf mass ratio. © 2015 New Phytologist Trust.