Search

Allometric relationships among the dimensions of leaves and their cells hold across diverse eudicotyledons, but have remained untested in the leaves of grasses. We hypothesised that geometric (proportional) allometries of cell sizes across tissues and of leaf dimensions would arise due to the coordination of cell development and that of cell functions such as water, nutrient and energy transport, and that cell sizes across tissues would be associated with light-saturated photosynthetic rate. We tested predictions across 27 globally distributed C 3 and C 4 grass species grown in a common garden. We found positive relationships among average cell sizes within and across tissues, and of cell sizes with leaf dimensions. Grass leaf anatomical allometries were similar to those of eudicots, with exceptions consistent with the fewer cell layers and narrower form of grass leaves, and the specialised roles of epidermis and bundle sheath in storage and leaf movement. Across species, mean cell sizes in each tissue were associated with light-saturated photosynthetic rate per leaf mass, supporting the functional coordination of cell sizes. These findings highlight the generality of evolutionary allometries within the grass lineage and their interlinkage with coordinated development and function., (© 2023 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

A surge of papers have reported low leaf vulnerability to xylem embolism during drought. Here, we focus on the less studied, and more sensitive, outside-xylem leaf hydraulic responses to multiple internal and external conditions. Studies of 34 species have resolved substantial vulnerability to dehydration of the outside-xylem pathways, and studies of leaf hydraulic responses to light also implicate dynamic outside-xylem responses. Detailed experiments suggest these dynamic responses arise at least in part from strong control of radial water movement across the vein bundle sheath. While leaf xylem vulnerability may influence leaf and plant survival during extreme drought, outside-xylem dynamic responses are important for the control and resilience of water transport and leaf water status for gas exchange and growth., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

Intra-specific trait variation (ITV) plays a role in processes at a wide range of scales from organs to ecosystems across climate gradients. Yet, ITV remains rarely quantified for many ecophysiological traits typically assessed for species means, such as pressure volume (PV) curve parameters including osmotic potential at full turgor and modulus of elasticity, which are important in plant water relations. We defined a baseline "reference ITV" (ITV ref ) as the variation among fully exposed, mature sun leaves of replicate individuals of a given species grown in similar, well-watered conditions, representing the conservative sampling design commonly used for species-level ecophysiological traits. We hypothesized that PV parameters would show low ITV ref relative to other leaf morphological traits, and that their intraspecific relationships would be similar to those previously established across species and proposed to arise from biophysical constraints. In a database of novel and published PV curves and additional leaf structural traits for 50 diverse species, we found low ITV ref for PV parameters relative to other morphological traits, and strong intraspecific relationships among PV traits. Simulation modeling showed that conservative ITV ref enables the use of species-mean PV parameters for scaling up from spectroscopic measurements of leaf water content to enable sensing of leaf water potential., (© 2023 Scandinavian Plant Physiology Society.)

Leaf water potential (ψ 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 ψ leaf (ψ 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 ψ leafPC from multiple laboratories, and synthesized experiments testing common assumptions and practices in ψ leafPC for diverse species: (i) the need for equilibration of previously transpiring leaves; (ii) leaf storage before measurement; (iii) the equilibration of ψ leaf for leaves on bagged branches of a range of dehydration; (iv) the equilibration of ψ leaf across the lamina for bagged leaves, and the accuracy of measuring leaves with artificially 'elongated petioles'; (v) the need in ψ 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 ψ 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., (© 2022 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

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'; π tlp ), 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 π tlp were associated with drier habitats, with π tlp explaining 28% and 32% of habitat association on local and regional scales, respectively, greatly exceeding the predictive power of WD and LMA. In contrast, π tlp did not predict habitat associations among deciduous species. Across spatial scales, π tlp 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., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

One of the most notable ecological trends-described more than 2,300  years ago by Theophrastus-is the association of small leaves with dry and cold climates, which has recently been recognized for eudicotyledonous plants at a global scale 1-3 . For eudicotyledons, this pattern has been attributed to the fact that small leaves have a thinner boundary layer that helps to avoid extreme leaf temperatures 4 and their leaf development results in vein traits that improve water transport under cold or dry climates 5,6 . However, the global distribution of leaf size and its adaptive basis have not been tested in the grasses, which represent a diverse lineage that is distinct in leaf morphology and that contributes 33% of terrestrial primary productivity (including the bulk of crop production) 7 . Here we demonstrate that grasses have shorter and narrower leaves under colder and drier climates worldwide. We show that small grass leaves have thermal advantages and vein development that contrast with those of eudicotyledons, but that also explain the abundance of small leaves in cold and dry climates. The worldwide distribution of leaf size in grasses exemplifies how biophysical and developmental processes result in convergence across major lineages in adaptation to climate globally, and highlights the importance of leaf size and venation architecture for grass performance in past, present and future ecosystems.

Functional traits are frequently used to evaluate plant adaptation across environments. Yet, traits tend to have multiple functions and interactions, which cannot be accounted for in traditional correlation analyses. Plant trait networks (PTNs) clarify complex relationships among traits, enable the calculation of metrics for the topology of trait coordination and the importance of given traits in PTNs, and how they shift across communities. Recent studies of PTNs provide new insights into some important topics, including trait dimensionality, trait spectra (including the leaf economic spectrum), stoichiometric principles, and the variation of phenotypic integration along gradients of resource availability. PTNs provide improved resolution of the multiple dimensions of plant adaptation across scales and responses to shifting resources, disturbance regimes, and global change., (Copyright © 2020 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

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 (K leaf ) in response to dehydration may be recovered through SSWU, and that the hydraulic conductance to SSWU (K surf ) declines with dehydration. We quantified effects of leaf dehydration on K surf and effects of SSWU on recovery of K leaf in dehydrated leaves of Avicennia marina. SSWU led to overnight recovery of K leaf , with recovery retracing the same path as loss of K leaf in response to dehydration. SSWU declined with dehydration. By contrast, K surf 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., (© 2019 Australian National University New Phytologist © 2019 New Phytologist Trust.)

Stomata, the microvalves on leaf surfaces, exert major influences across scales, from plant growth and productivity to global carbon and water cycling. Stomatal opening enables leaf photosynthesis, and plant growth and water use, whereas plant survival of drought depends on stomatal closure. Here we report that stomatal function is constrained by a safety-efficiency trade-off, such that species with greater stomatal conductance under high water availability (g max ) show greater sensitivity to closure during leaf dehydration, i.e., a higher leaf water potential at which stomatal conductance is reduced by 50% (Ψ gs50 ). The g max - Ψ gs50 trade-off and its mechanistic basis is supported by experiments on leaves of California woody species, and in analyses of previous studies of the responses of diverse flowering plant species around the world. Linking the two fundamental key roles of stomata-the enabling of gas exchange, and the first defense against drought-this trade-off constrains the rates of water use and the drought sensitivity of leaves, with potential impacts on ecosystems.

Leaves are a nexus for the exchange of water, carbon, and energy between terrestrial plants and the atmosphere. Research in recent decades has highlighted the critical importance of the underlying biophysical and anatomical determinants of CO 2 and H 2 O transport, but a quantitative understanding of how detailed 3D leaf anatomy mediates within-leaf transport has been hindered by the lack of a consensus framework for analyzing or simulating transport and its spatial and temporal dynamics realistically, and by the difficulty of measuring within-leaf transport at the appropriate scales. We discuss how recent technological advancements now make a spatially explicit 3D leaf analysis possible, through new imaging and modeling tools that will allow us to address long-standing questions related to plant carbon-water exchange., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

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., (© 2018 American Society of Plant Biologists. All Rights Reserved.)

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., (© 2018 John Wiley & Sons Ltd.)

Premise of the Study: Studies across diverse species have established theory for the contribution of leaf traits to plant drought tolerance. For example, species in more arid climates tend to have smaller leaves of higher vein density, higher leaf mass per area, and more negative osmotic potential at turgor loss point (π TLP ). However, few studies have tested these associations for species within a given lineage that have diversified across an aridity gradient., Methods: We analyzed the anatomy and physiology of 10 Ceanothus (Rhamnaceae) species grown in a common garden for variation between and within "wet" and "dry" subgenera (Ceanothus and Cerastes, respectively) and analyzed a database for 35 species for leaf size and leaf mass per area (LMA). We used a phylogenetic generalized least squares approach to test hypothesized relationships among traits, and of traits with climatic aridity in the native range. We also tested for allometric relationships among anatomical traits., Key Results: Leaf form, anatomy, and drought tolerance varied strongly among species within and between subgenera. Cerastes species had specialized anatomy including hypodermis and encrypted stomata that may confer superior water storage and retention. The osmotic potentials at turgor loss point (π TLP ) and full turgor (π o ) showed evolutionary correlations with the aridity index (AI) and precipitation of the 10 species' native distributions, and LMA with potential evapotranspiration for the 35 species in the larger database. We found an allometric correlation between upper and lower epidermal cell wall thicknesses, but other anatomical traits diversified independently., Conclusions: Leaf traits and drought tolerance evolved within and across lineages of Ceanothus consistently with climatic distributions. The π TLP has signal to indicate the evolution of drought tolerance within small clades., (© 2018 Botanical Society of America.)

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 (K leaf ). We further hypothesized that BS lignification would mediate the relationship of K leaf to vein length per area. We analysed the dependence of K leaf , 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 K leaf . K leaf could be strongly predicted by vein length per area and highest lignified vein order (R 2  = .69). The light-response of K leaf 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., (© 2017 John Wiley & Sons Ltd.)

Leaf size varies by over a 100,000-fold among species worldwide. Although 19th-century plant geographers noted that the wet tropics harbor plants with exceptionally large leaves, the latitudinal gradient of leaf size has not been well quantified nor the key climatic drivers convincingly identified. Here, we characterize worldwide patterns in leaf size. Large-leaved species predominate in wet, hot, sunny environments; small-leaved species typify hot, sunny environments only in arid conditions; small leaves are also found in high latitudes and elevations. By modeling the balance of leaf energy inputs and outputs, we show that daytime and nighttime leaf-to-air temperature differences are key to geographic gradients in leaf size. This knowledge can enrich "next-generation" vegetation models in which leaf temperature and water use during photosynthesis play key roles., (Copyright © 2017, American Association for the Advancement of Science.)

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 (ε) 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., (© 2017 John Wiley & Sons Ltd.)

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., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Leaf dry mass per unit leaf area (LMA) is a central trait in ecology, but its anatomical and compositional basis has been unclear. An explicit mathematical and physical framework for quantifying the cell and tissue determinants of LMA will enable tests of their influence on species, communities and ecosystems. We present an approach to explaining LMA from the numbers, dimensions and mass densities of leaf cells and tissues, which provided unprecedented explanatory power for 11 broadleaved woody angiosperm species diverse in LMA (33-262 g m -2 ; R 2  = 0.94; P < 0.001). Across these diverse species, and in a larger comparison of evergreen vs. deciduous angiosperms, high LMA resulted principally from larger cell sizes, greater major vein allocation, greater numbers of mesophyll cell layers and higher cell mass densities. This explicit approach enables relating leaf anatomy and composition to a wide range of processes in physiological, evolutionary, community and macroecology., (© 2017 John Wiley & Sons Ltd/CNRS.)

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., (© 2017 American Society of Plant Biologists. All Rights Reserved.)

We tested for a tradeoff across species between plant maximum photosynthetic rate and the ability to maintain photosynthesis under adverse conditions in the unfavorable season. Such a trade-off would be consistent with the observed trade-off between maximum speed and endurance in athletes and some animals that has been explained by cost-benefit theory. This trend would have importance for the general understanding of leaf design, and would simplify models of annual leaf carbon relations. We tested for such a trade-off using a database analysis across vascular plants and using an experimental approach for 29 cycad species, representing an ancient plant lineage with diversified evergreen leaves. In both tests, a higher photosynthetic rate per mass or per area in the favorable season was associated with a stronger absolute or percent decline in the unfavorable season. We resolved a possible mechanism based on biomechanics and nitrogen allocation; cycads with high leaf toughness (leaf mass per area) and higher investment in leaf construction than in physiological function (C/N ratio) tended to have lower warm season photosynthesis but less depression in the cool season. We propose that this trade-off, consistent with cost-benefit theory, represents a significant physio-phenological constraint on the diversity and seasonal dynamics of photosynthetic rate., Competing Interests: The authors declare no competing financial interests.

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 (K x ) during dehydration was driven by embolism initiating in petioles and midribs across all species, and K x vulnerability was strongly correlated with petiole and midrib conduit dimensions. Our simulations showed no significant impact of conduit collapse on K x decline. We found xylem conduit dimensions play a major role in determining the susceptibility of the leaf water transport system during strong leaf dehydration., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

Leaf hydraulic supply is crucial to maintaining open stomata for CO 2 capture and plant growth. During drought-induced dehydration, the leaf hydraulic conductance (K leaf ) declines, which contributes to stomatal closure and, eventually, to leaf death. Previous studies have tended to attribute the decline of K leaf 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 K leaf 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 K leaf 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., (© 2017 The author(s). All Rights Reserved.)

Pan-tropically, liana density increases with decreasing rainfall and increasing seasonality. This pattern has led to the hypothesis that lianas display a growth advantage over trees under dry conditions. However, the physiological mechanisms underpinning this hypothesis remain elusive. A key trait influencing leaf and plant drought tolerance is the leaf water potential at turgor loss point (π tlp ). π tlp adjusts under drier conditions and this contributes to improved leaf drought tolerance. For co-occurring Amazonian tree (n = 247) and liana (n = 57) individuals measured during the dry and the wet seasons, lianas showed a stronger osmotic adjustment than trees. Liana leaves were less drought-tolerant than trees in the wet season, but reached similar drought tolerances during the dry season. Stronger osmotic adjustment in lianas would contribute to turgor maintenance, a critical prerequisite for carbon uptake and growth, and to the success of lianas relative to trees in growth under drier conditions., (© 2017 The Author(s).)

Leaf venation is diverse across plant species and has practical applications from paleobotany to modern agriculture. However, the impact of vein traits on plant performance has not yet been tested in a model system such as Arabidopsis thaliana. Previous studies analysed cotyledons of A. thaliana vein mutants and identified visible differences in their vein systems from the wild type (WT). We measured leaf hydraulic conductance (Kleaf ), vein traits, and xylem and mesophyll anatomy for A. thaliana WT (Col-0) and four vein mutants (dot3-111 and dot3-134, and cvp1-3 and cvp2-1). Mutant true leaves did not possess the qualitative venation anomalies previously shown in the cotyledons, but varied quantitatively in vein traits and leaf anatomy across genotypes. The WT had significantly higher mean Kleaf . Across all genotypes, there was a strong correlation of Kleaf with traits related to hydraulic conductance across the bundle sheath, as influenced by the number and radial diameter of bundle sheath cells and vein length per area. These findings support the hypothesis that vein traits influence Kleaf , indicating the usefulness of this mutant system for testing theory that was primarily established comparatively across species, and supports a strong role for the bundle sheath in influencing Kleaf ., (© 2015 John Wiley & Sons Ltd.)

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., (© 2015 American Society of Plant Biologists. All Rights Reserved.)

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., (© 2015 The Authors New Phytologist © 2015 New Phytologist Trust.)

Unlabelled: •, Premise of the Study: The diverse early Eocene flora from Laguna del Hunco (LH) in Patagonia, Argentina has many nearest living relatives (NLRs) in Australasia but few in South America, indicating the differential survival of an ancient, trans-Antarctic rainforest biome. To better understand this significant biogeographic pattern, we used detailed comparisons of leaf size and floristics to quantify the legacy of LH across a large network of Australian rainforest-plot assemblages.•, Methods: We applied vein scaling, a new method for estimating the original areas of fragmented leaves. We then compared leaf size and floristics at LH with living Australian assemblages and tabulated the climates of those where NLRs occur, along with additional data on climatic ranges of "ex-Australian" NLRs that survive outside of Australia.•, Key Results: Vein scaling estimated areas as accurately as leaf-size classes. Applying vein scaling to fossil fragments increased the grand mean area of LH by 450 mm(2), recovering more originally large leaves. The paleoflora has a majority of microphyll leaves, comparable to leaf litter in subtropical Australian forests, which also have the greatest floristic similarity to LH. Tropical Australian assemblages also share many taxa with LH, and ex-Australian NLRs mostly inhabit cool, wet montane habitats no longer present in Australia.•, Conclusions: Vein scaling is valuable for improving the resolution of fossil leaf-size distributions by including fragmented specimens. The legacy of LH is evident not only in subtropical and tropical Australia but also in tropical montane Australasia and Southeast Asia, reflecting the disparate histories of surviving Gondwanan lineages., (© 2015 Botanical Society of America, Inc.)

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., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

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., (© 2014 John Wiley & Sons Ltd.)

Leaf vein length per unit leaf area (VLA; also known as vein density) is an important determinant of water and sugar transport, photosynthetic function, and biomechanical support. A range of software methods are in use to visualize and measure vein systems in cleared leaf images; typically, users locate veins by digital tracing, but recent articles introduced software by which users can locate veins using thresholding (i.e. based on the contrasting of veins in the image). Based on the use of this method, a recent study argued against the existence of a fixed VLA value for a given leaf, proposing instead that VLA increases with the magnification of the image due to intrinsic properties of the vein system, and recommended that future measurements use a common, low image magnification for measurements. We tested these claims with new measurements using the software LEAFGUI in comparison with digital tracing using ImageJ software. We found that the apparent increase of VLA with magnification was an artifact of (1) using low-quality and low-magnification images and (2) errors in the algorithms of LEAFGUI. Given the use of images of sufficient magnification and quality, and analysis with error-free software, the VLA can be measured precisely and accurately. These findings point to important principles for improving the quantity and quality of important information gathered from leaf vein systems., (© 2014 American Society of Plant Biologists. All Rights Reserved.)

Coordination of water movement among plant organs is important for understanding plant water use strategies. The hydraulic segmentation hypothesis (HSH) proposes that hydraulic conductance in shorter lived, 'expendable' organs such as leaves and longer lived, more 'expensive' organs such as stems may be decoupled, with resistance in leaves acting as a bottleneck or 'safety valve'. We tested the HSH in woody species from a Mediterranean-type ecosystem by measuring leaf hydraulic conductance (Kleaf) and stem hydraulic conductivity (KS). We also investigated whether leaves function as safety valves by relating Kleaf and the hydraulic safety margin (stem water potential minus the water potential at which 50% of conductivity is lost (Ψstem-Ψ50)). We also examined related plant traits including the operating range of water potentials, wood density, leaf mass per area, and leaf area to sapwood area ratio to provide insight into whole-plant water use strategies. For hydrated shoots, Kleaf was negatively correlated with KS , supporting the HSH. Additionally, Kleaf was positively correlated with the hydraulic safety margin and negatively correlated with the leaf area to sapwood area ratio. Consistent with the HSH, our data indicate that leaves may act as control valves for species with high KS , or a low safety margin. This critical role of leaves appears to contribute importantly to plant ecological specialization in a drought-prone environment., (© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.)

Leaf shrinkage with dehydration has attracted attention for over 100 years, especially as it becomes visibly extreme during drought. However, little has been known of its correlation with physiology. Computer simulations of the leaf hydraulic system showed that a reduction of hydraulic conductance of the mesophyll pathways outside the xylem would cause a strong decline of leaf hydraulic conductance (K(leaf)). For 14 diverse species, we tested the hypothesis that shrinkage during dehydration (i.e. in whole leaf, cell and airspace thickness, and leaf area) is associated with reduction in K(leaf) at declining leaf water potential (Ψ(leaf)). We tested hypotheses for the linkage of leaf shrinkage with structural and physiological water relations parameters, including modulus of elasticity, osmotic pressure at full turgor, turgor loss point (TLP), and cuticular conductance. Species originating from moist habitats showed substantial shrinkage during dehydration before reaching TLP, in contrast with species originating from dry habitats. Across species, the decline of K(leaf) with mild dehydration (i.e. the initial slope of the K(leaf) versus Ψ(leaf) curve) correlated with the decline of leaf thickness (the slope of the leaf thickness versus Ψ(leaf) curve), as expected based on predictions from computer simulations. Leaf thickness shrinkage before TLP correlated across species with lower modulus of elasticity and with less negative osmotic pressure at full turgor, as did leaf area shrinkage between full turgor and oven desiccation. These findings point to a role for leaf shrinkage in hydraulic decline during mild dehydration, with potential impacts on drought adaptation for cells and leaves, influencing plant ecological distributions.

Premise of the Study: Allometric relationships among the dimensions of leaf cells, cell walls, and tissues, and whole-leaf thickness and area are likely to have key implications for leaf construction and function, but have remained virtually untested, despite the explosion of interest in allometric analysis of numerous plant properties at larger scales. •, Methods: Using leaf transverse cross sections and light microscopy, we measured leaf dimensions, tissue thicknesses, mesophyll and xylem cell sizes, and cell wall thicknesses for 14 diverse angiosperm species of wet and dry habitats and tested hypothesized allometric relationships based on geometric scaling due to development and/or function. •, Key Results: We found strong novel allometries relating the dimensions of cells, cell walls, tissues, and gross leaf form. Cell sizes and cell wall thicknesses tended to scale isometrically across mesophyll tissues within the leaf, such that species with large cells or thick cell walls in one tissue had these also in the other tissues; however, leaf vein xylem conduit sizes were independent of those of other cell types. We also found strong geometric scaling of cell wall thicknesses with cell sizes throughout the mesophyll, but not in the leaf vein xylem. Further, leaf thickness scaled with cell sizes, cell wall thicknesses and the thicknesses of component mesophyll tissues, but leaf area was independent of anatomical traits across species. •, Conclusions: These novel allometries suggest design rules operating at the smallest scales of leaf construction and the possibility of applying these relationships to better characterizing the basis for differences among species in leaf form and functional traits.

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 physiology determines the carbon acquisition of the whole plant, but there can be considerable variation in physiology and carbon acquisition within individual leaves. Alocasia macrorrhiza (L.) Schott is an herbaceous species that can develop very large leaves of up to 1 m in length. However, little is known about the hydraulic and photosynthetic design of such giant leaves. Based on previous studies of smaller leaves, and on the greater surface area for trait variation in large leaves, we hypothesized that A. macrorrhiza leaves would exhibit significant heterogeneity in structure and function. We found evidence of reduced hydraulic supply and demand in the outer leaf regions; leaf mass per area, chlorophyll concentration, and guard cell length decreased, as did stomatal conductance, net photosynthetic rate and quantum efficiency of photosystem II. This heterogeneity in physiology was opposite to that expected from a thinner boundary layer at the leaf edge, which would have led to greater rates of gas exchange. Leaf temperature was 8.8°C higher in the outer than in the central region in the afternoon, consistent with reduced stomatal conductance and transpiration caused by a hydraulic limitation to the outer lamina. The reduced stomatal conductance in the outer regions would explain the observed homogeneous distribution of leaf water potential across the leaf surface. These findings indicate substantial heterogeneity in gas exchange across the leaf surface in large leaves, greater than that reported for smaller-leafed species, though the observed structural differences across the lamina were within the range reported for smaller-leafed species. Future work will determine whether the challenge of transporting water to the outer regions can limit leaf size for plants experiencing drought, and whether the heterogeneity of function across the leaf surface represents a particular disadvantage for large simple leaves that might explain their global rarity, even in resource-rich environments.

The design and function of leaf venation are important to plant performance, with key implications for the distribution and productivity of ecosystems, and applications in paleobiology, agriculture and technology. We synthesize classical concepts and the recent literature on a wide range of aspects of leaf venation. We describe 10 major structural features that contribute to multiple key functions, and scale up to leaf and plant performance. We describe the development and plasticity of leaf venation and its adaptation across environments globally, and a new global data compilation indicating trends relating vein length per unit area to climate, growth form and habitat worldwide. We synthesize the evolution of vein traits in the major plant lineages throughout paleohistory, highlighting the multiple origins of individual traits. We summarize the strikingly diverse current applications of leaf vein research in multiple fields of science and industry. A unified core understanding will enable an increasing range of plant biologists to incorporate leaf venation into their research., (© 2013 The Authors New Phytologist © 2013 New Phytologist Trust.)

Invasive species are expected to cluster on the "high-return" end of the leaf economic spectrum, displaying leaf traits consistent with higher carbon assimilation relative to native species. Intra-leaf nitrogen (N) allocation should support these physiological differences; however, N biochemistry has not been examined in more than a few invasive species. We measured 34 leaf traits including seven leaf N pools for five native and five invasive species from Hawaii under low irradiance to mimic the forest understory environment. We found several trait differences between native and invasive species. In particular, invasive species showed preferential N allocation to metabolism (amino acids) rather than photosynthetic light reactions (membrane-bound protein) by comparison with native species. The soluble protein concentration did not vary between groups. Under these low irradiance conditions, native species had higher light-saturated photosynthetic rates, possibly as a consequence of a greater investment in membrane-bound protein. Invasive species may succeed by employing a wide range of N allocation mechanisms, including higher amino acid production for fast growth under high irradiance or storage of N in leaves as soluble protein or amino acids.

Strong latitudinal patterns in leaf form are well documented in floristic comparisons and palaeobotanical studies. However, there is little agreement about their functional significance; in fact, it is still unknown to what degree these patterns were generated by repeated evolutionary adaptation. We analysed leaf form in the woody angiosperm clade Viburnum (Adoxaceae) and document evolutionarily correlated shifts in leafing habit, leaf margin morphology, leaf shape and climate. Multiple independent shifts between tropical and temperate forest habitats have repeatedly been accompanied by a change between evergreen, elliptical leaves with entire margins and deciduous, more rounded leaves with toothed or lobed margins. These consistent shifts in Viburnum support repeated evolutionary adaptation as a major determinant of the global correlation between leaf form and mean annual temperature. Our results provide a new theoretical grounding for the inference of past climates using fossil leaf assemblages.

Leaf size and venation show remarkable diversity across dicotyledons, and are key determinants of plant adaptation in ecosystems past and present. Here we present global scaling relationships of venation traits with leaf size. Across a new database for 485 globally distributed species, larger leaves had major veins of larger diameter, but lower length per leaf area, whereas minor vein traits were independent of leaf size. These scaling relationships allow estimation of intact leaf size from fragments, to improve hindcasting of past climate and biodiversity from fossil remains. The vein scaling relationships can be explained by a uniquely synthetic model for leaf anatomy and development derived from published data for numerous species. Vein scaling relationships can explain the global biogeographical trend for smaller leaves in drier areas, the greater construction cost of larger leaves and the ability of angiosperms to develop larger and more densely vascularised lamina to outcompete earlier-evolved plant lineages.

Increasing drought is one of the most critical challenges facing species and ecosystems worldwide, and improved theory and practices are needed for quantification of species tolerances. Leaf water potential at turgor loss, or wilting (π(tlp) ), is classically recognised as a major physiological determinant of plant water stress response. However, the cellular basis of π(tlp) and its importance for predicting ecological drought tolerance have been controversial. A meta-analysis of 317 species from 72 studies showed that π(tlp) was strongly correlated with water availability within and across biomes, indicating power for anticipating drought responses. We derived new equations giving both π(tlp) and relative water content at turgor loss point (RWC(tlp) ) as explicit functions of osmotic potential at full turgor (π(o) ) and bulk modulus of elasticity (ε). Sensitivity analyses and meta-analyses showed that π(o) is the major driver of π(tlp) . In contrast, ε plays no direct role in driving drought tolerance within or across species, but sclerophylly and elastic adjustments act to maintain RWC(tlp,) preventing cell dehydration, and additionally protect against nutrient, mechanical and herbivory stresses independent of drought tolerance. These findings clarify biogeographic trends and the underlying basis of drought tolerance parameters with applications in comparative assessments of species and ecosystems worldwide., (© 2012 Blackwell Publishing Ltd/CNRS.)

Leaf hydraulic conductance (K(leaf)) is a major determinant of photosynthetic rate in well-watered and drought-stressed plants. Previous work assessed the decline of K(leaf) with decreasing leaf water potential (Ψ(leaf)), most typically using rehydration kinetics methods, and found that species varied in the shape of their vulnerability curve, and that hydraulic vulnerability correlated with other leaf functional traits and with drought sensitivity. These findings were tested and extended, using a new steady-state evaporative flux method under high irradiance, and the function for the vulnerability curve of each species was determined individually using maximum likelihood for 10 species varying strongly in drought tolerance. Additionally, the ability of excised leaves to recover in K(leaf) with rehydration was assessed, and a new theoretical framework was developed to estimate how rehydration of measured leaves may affect estimation of hydraulic parameters. As hypothesized, species differed in their vulnerability function. Drought-tolerant species showed shallow linear declines and more negative Ψ(leaf) at 80% loss of K(leaf) (P(80)), whereas drought-sensitive species showed steeper, non-linear declines, and less negative P(80). Across species, the maximum K(leaf) was independent of hydraulic vulnerability. Recovery of K(leaf) after 1 h rehydration of leaves dehydrated below their turgor loss point occurred only for four of 10 species. Across species without recovery, a more negative P(80) correlated with the ability to maintain K(leaf) through both dehydration and rehydration. These findings indicate that resistance to K(leaf) decline is important not only in maintaining open stomata during the onset of drought, but also in enabling sustained function during drought recovery.

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 (K(lb) ), petioles (K(P) ) and branches (i.e. leafless stem; K(B) ) were determined using a high pressure flow meter in steady state mode. The shoots were exposed to photosynthetic photon flux density of 200-250 µmol m⁻² s⁻¹ using white, blue or red light. K(lb) depended significantly on both light quality and canopy position (P<0.001), K(B) on canopy position (P<0.001) and exposure time (P=0.014), and none of the three factors had effect on K(P) . The highest values of K(lb) 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 values under red light (2.83 and 2.02×10⁻⁴ kg m⁻² MPa⁻¹ s⁻¹, respectively). Light quality has an important impact on leaf hydraulic properties, independently of light intensity or of total light energy, and the specific light receptors involved in this response require identification. Given that natural canopy shade depletes blue and red light, K(lb) may be decreased both by reduced fluence and shifts in light spectra, indicating the need for studies of the natural heterogeneity of K(lb) within and under canopies, and its impacts on gas exchange., (© 2011 Blackwell Publishing Ltd.)

Across plant species, leaves vary enormously in their size and their venation architecture, of which one major function is to replace water lost to transpiration. The leaf hydraulic conductance (K(leaf)) represents the capacity of the transport system to deliver water, allowing stomata to remain open for photosynthesis. Previous studies showed that K(leaf) relates to vein density (vein length per area). Additionally, venation architecture determines the sensitivity of K(leaf) to damage; severing the midrib caused K(leaf) and gas exchange to decline, with lesser impacts in leaves with higher major vein density that provided more numerous water flow pathways around the damaged vein. Because xylem embolism during dehydration also reduces K(leaf), we hypothesized that higher major vein density would also reduce hydraulic vulnerability. Smaller leaves, which generally have higher major vein density, would thus have lower hydraulic vulnerability. Tests using simulations with a spatially explicit model confirmed that smaller leaves with higher major vein density were more tolerant of major vein embolism. Additionally, for 10 species ranging strongly in drought tolerance, hydraulic vulnerability, determined as the leaf water potential at 50% and 80% loss of K(leaf), was lower with greater major vein density and smaller leaf size (|r| = 0.85-0.90; P < 0.01). These relationships were independent of other aspects of physiological and morphological drought tolerance. These findings point to a new functional role of venation architecture and small leaf size in drought tolerance, potentially contributing to well-known biogeographic trends in leaf size.

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., (© 2010 Blackwell Publishing Ltd.)

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 CO(2) , 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., (© 2010 Blackwell Publishing Ltd.)

Leaf venation architecture is tremendously diverse across plant species. Understanding the hydraulic functions of given venation traits can clarify the organization of the vascular system and its adaptation to environment. Using a spatially explicit model (the program K&#95;leaf), we subjected realistic simulated leaves to modifications and calculated the impacts on xylem and leaf hydraulic conductance (K(x) and K(leaf), respectively), important traits in determining photosynthesis and growth. We tested the sensitivity of leaves to altered vein order conductivities (1) in the absence or (2) presence of hierarchical vein architecture, (3) to major vein tapering, and (4) to modification of vein densities (length/leaf area). The K(x) and K(leaf) increased with individual vein order conductivities and densities; for hierarchical venation systems, the greatest impact was from increases in vein conductivity for lower vein orders and increases in density for higher vein orders. Individual vein order conductivities were colimiting of K(x) and K(leaf), as were their densities, but the effects of vein conductivities and densities were orthogonal. Both vein hierarchy and vein tapering increased K(x) relative to xylem construction cost. These results highlight the important consequences of venation traits for the economics, ecology, and evolution of plant transport capacity.

Mosses are an understudied group of plants that can potentially confirm or expand principles of plant function described for tracheophytes, from which they diverge strongly in structure. We quantified 35 physiological and morphological traits from cell-, leaf- and canopy-level, for 10 ground-, trunk- and branch-dwelling Hawaiian species. We hypothesized that trait values would reflect the distinctive growth form and slow growth of mosses, but also that trait correlations would be analogous to those of tracheophytes. The moss species had low leaf mass per area and low gas exchange rate. Unlike for tracheophytes, light-saturated photosynthetic rate per mass (A(mass)) did not correlate with habitat irradiance. Other photosynthetic parameters and structural traits were aligned with microhabitat irradiance, driving an inter-correlation of traits including leaf area, cell size, cell wall thickness, and canopy density. In addition, we found a coordination of traits linked with structural allocation, including costa size, canopy height and A(mass). Across species, A(mass) and nitrogen concentration correlated negatively with canopy mass per area, analogous to linkages found for the 'leaf economic spectrum', with canopy mass per area replacing leaf mass per area. Despite divergence of mosses and tracheophytes in leaf size and function, analogous trait coordination has arisen during ecological differentiation.