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Summary Mature leaf area (LA) is a showcase of diversity – varying enormously within and across species, and associated with the productivity and distribution of plants and ecosystems. Yet, it remains unclear how developmental processes determine variation in LA.We introduce a mathematical framework pinpointing the origin of variation in LA by quantifying six epidermal ‘developmental traits’: initial mean cell size and number (approximating values within the leaf primordium), and the maximum relative rates and durations of cell proliferation and expansion until leaf maturity. We analyzed a novel database of developmental trajectories of LA and epidermal anatomy, representing 12 eudicotyledonous species and 52 Arabidopsis experiments.Within and across species, mean primordium cell number and maximum relative cell proliferation rate were the strongest developmental determinants of LA. Trade‐offs between developmental traits, consistent with evolutionary and metabolic scaling theory, strongly constrain LA variation. These include trade‐offs between primordium cell number vs cell proliferation, primordium mean cell size vs cell expansion, and the durations vs maximum relative rates of cell proliferation and expansion. Mutant and wild‐type comparisons showed these trade‐offs have a genetic basis in Arabidopsis.Analyses of developmental traits underlying LA and its diversification highlight mechanisms for leaf evolution, and opportunities for breeding trait shifts. 

Summary Plant cuticles protect the interior tissues from ambient hazards, including desiccation, UV light, physical wear, herbivores and pathogens. Consequently, cuticle properties are shaped by evolutionary selection.We compiled a global dataset of leaf cuticle thickness (CT) and accompanying leaf traits for 1212 species, mostly angiosperms, from 293 sites representing all vegetated continents. We developed and tested 11 hypotheses concerning ecological drivers of interspecific variation in CT.CT showed clear patterning according to latitude, biome, taxonomic family, site climate and other leaf traits. Species with thick leaves and/or high leaf mass per area tended to have thicker cuticles, as did evergreen relative to deciduous woody species, and species from sites that during the growing season were warmer, had fewer frost days and lower wind speeds, and occurred at lower latitudes. CT–environment relationships were notably stronger among nonwoody than woody species.Heavy investment in cuticle may be disadvantaged at sites with high winds and frequent frosts for ‘economic’ or biomechanical reasons, or because of reduced herbivore pressure. Alternatively, cuticles may become more heavily abraded under such conditions. Robust quantification of CT–trait–environment relationships provides new insights into the multiple roles of cuticles, with additional potential use in paleo‐ecological reconstruction. 

Abstract Plant ecological strategies are shaped by numerous functional traits and their trade‐offs. Trait network analysis enables testing hypotheses for the shifting of trait correlation architecture across communities differing in climate and productivity.We built plant trait networks (PTNs) for 118 species within six communities across an aridity gradient, from forest to semi‐desert across the California Floristic Province, based on 34 leaf and wood functional traits, representing hydraulic and photosynthetic function, structure, economics and size. We developed hypotheses for the association of PTN parameters with climate and ecosystem properties, based on theory for the adaptation of species to low resource/stressful environments versus higher resource availability environments with greater potential niche differentiation. Thus, we hypothesized that across community PTNs, trait network connectivity (i.e., the degree that traits are intercorrelated) and network complexity (i.e., the number of trait modules, and the degree of trait integration among modules) would be lower for communities adapted to arid climates and higher for communities adapted to greater water availability, similarly to trends expected for phylogenetic diversity, functional richness and productivity. Further, within given PTNs, we hypothesized that traits would vary strongly in their network connectivity and that the traits most centrally connected within PTNs would be those with the least across‐species variation.Across communities from more arid to wetter climates, PTN architecture varied from less to more interconnected and complex, in association with functional richness, but PTN architecture was independent of phylogenetic diversity and ecosystem productivity. Within the community PTNs, traits with lower species variation were more interconnected.Synthesis. The responsiveness of PTN architecture to climate highlights how a wide range of traits contributes to physiological and ecological strategies with an architecture that varies among plant communities. Communities in more arid environments show a lower degree of phenotypic integration, consistent with lesser niche differentiation. Our study extends the usefulness of PTNs as an approach to quantify tradeoffs among multiple traits, providing connectivity and complexity parameters as tools that clarify plant environmental adaptation and patterns of trait associations that would influence species distributions, community assembly, and ecosystem resilience in response to climate change. 

ABSTRACT Identifying the physiological mechanisms by which plants are adapted to drought is critical to predict species responses to climate change. We measured the responses of leaf hydraulic and stomatal conductances (K_leafandg_s, respectively) to dehydration, and their association with anatomy, in seven species of CaliforniaCeanothusgrown in a common garden, including some of the most drought‐tolerant species in the semi‐arid flora. We tested for matching of maximum hydraulic supply and demand and quantified the role of decline ofK_leafin driving stomatal closure. AcrossCeanothusspecies, maximumK_leafandg_swere negatively correlated, and bothK_leafandg_sshowed steep declines with decreasing leaf water potential (i.e., a high sensitivity to dehydration). The leaf water potential at 50% decline ing_swas linked with a low ratio of maximum hydraulic supply to demand (i.e., maximumK_leaf:g_s). This sensitivity ofg_s, combined with low minimum epidermal conductance and water storage, could contribute to prolonged leaf survival under drought. The specialized anatomy of subg.Cerastesincludes trichomous stomatal crypts and pronounced hypodermis, and was associated with higher water use efficiency and water storage. Combining our data with comparative literature of other California species, species of subg. Cerastesshow traits associated with greater drought tolerance and reliance on leaf water storage relative to other California species. In addition to drought resistance mechanisms such as mechanical protection and resistance to embolism, drought avoidance mechanisms such as sensitive stomatal closure could contribute importantly to drought tolerance in dry‐climate adapted species. 

Many questions in leaf physiology, from mechanisms of water transport and stomatal responses to interpretation of gas exchange measurements, hinge on distributions of water potential among compartments in the leaf. A new tool, AquaDust, offers the possibility of quantifying those distributions at unprecedentedly small scales of organization 

Summary Grasses are exceptionally productive, yet their hydraulic adaptation is paradoxical. Among C₃grasses, a high photosynthetic rate (A_area) may depend on higher vein density (D_v) and hydraulic conductance (K_leaf). However, the higherD_vof C₄grasses suggests a hydraulic surplus, given their reduced need for highK_leafresulting from lower stomatal conductance (g_s).Combining hydraulic and photosynthetic physiological data for diverse common garden C₃and C₄species with data for 332 species from the published literature, and mechanistic modeling, we validated a framework for linkages of photosynthesis with hydraulic transport, anatomy, and adaptation to aridity.C₃and C₄grasses had similarK_leafin our common garden, but C₄grasses had higherK_leafthan C₃species in our meta‐analysis. Variation inK_leafdepended on outside‐xylem pathways. C₄grasses have highK_leaf : g_s, which modeling shows is essential to achieve their photosynthetic advantage.Across C₃grasses, higherA_areawas associated with higherK_leaf, and adaptation to aridity, whereas for C₄species, adaptation to aridity was associated with higherK_leaf : g_s. These associations are consistent with adaptation for stress avoidance.Hydraulic traits are a critical element of evolutionary and ecological success in C₃and C₄grasses and are crucial avenues for crop design and ecological forecasting. 

ABSTRACT ‘Water potential’ is the biophysically relevant measure of water status in vegetation relating to stomatal, canopy and hydraulic conductance, as well as mortality thresholds; yet, this cannot be directly related to measured and modelled fluxes of water at plot‐ to landscape‐scale without understanding its relationship with ‘water content’. The capacity for detecting vegetation water content via microwave remote sensing further increases the need to understand the link between water content and ecosystem function. In this review, we explore how the fundamental measures of water status, water potential and water content are linked at ecosystem‐scale drawing on the existing theory of pressure‐volume (PV) relationships. We define and evaluate the concept and limitations of applying PV relationships to ecosystems where the quantity of water can vary on short timescales with respect to plant water status, and over longer timescales and over larger areas due to structural changes in vegetation. As a proof of concept, plot‐scale aboveground vegetation PV curves were generated from equilibrium (e.g., predawn) water potentials and water content of the above ground biomass of nine plots, including tropical rainforest, savanna, temperate forest, and a long‐term Amazonian rainforest drought experiment. Initial findings suggest that the stored water and ecosystem capacitance scale linearly with biomass across diverse systems, while the relative values of ecosystem hydraulic capacitance and physiologically accessible water storage do not vary systematically with biomass. The bottom‐up scaling approach to ecosystem water relations identified the need to characterise the distribution of water potentials within a community and also revealed the relevance of community‐level plant tissue fractions to ecosystem water relations. We believe that this theory will be instrumental in linking our detailed understanding of biophysical processes at tissue‐scale to the scale at which land surface models operate and at which tower‐based, airborne and satellite remote sensing can provide information. 

Abstract Leaf surface conductance to water vapor and CO2 across the epidermis (gleaf) strongly determines the rates of gas exchange. Thus, clarifying the drivers of gleaf has important implications for resolving the mechanisms of photosynthetic productivity and leaf and plant responses and tolerance to drought. It is well recognized that gleaf is a function of the conductances of the stomata (gs) and of the epidermis + cuticle (gec). Yet, controversies have arisen around the relative roles of stomatal density (d) and size (s), fractional stomatal opening (α; aperture relative to maximum), and gec in determining gleaf. Resolving the importance of these drivers is critical across the range of leaf surface conductances, from strong stomatal closure under drought (gleaf,min), to typical opening for photosynthesis (gleaf,op), to maximum achievable opening (gleaf,max). We derived equations and analyzed a compiled database of published and measured data for approximately 200 species and genotypes. On average, within and across species, higher gleaf,min was determined 10 times more strongly by α and gec than by d and negligibly by s; higher gleaf,op was determined approximately equally by α (47%) and by stomatal anatomy (45% by d and 8% by s), and negligibly by gec; and higher gleaf,max was determined entirely by d. These findings clarify how diversity in stomatal functioning arises from multiple structural and physiological causes with importance shifting with context. The rising importance of d relative to α, from gleaf,min to gleaf,op, enables even species with low gleaf,min, which can retain leaves through drought, to possess high d and thereby achieve rapid gas exchange in periods of high water availability. 

Summary 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. 

Abstract The relationship between stomatal traits and environmental drivers across plant communities has important implications for ecosystem carbon and water fluxes, but it has remained unclear. Here, we measure the stomatal morphology of 4492 species-site combinations in 340 vegetation plots across China and calculate their community-weighted values for mean, variance, skewness, and kurtosis. We demonstrate a trade-off between stomatal density and size at the community level. The community-weighted mean and variance of stomatal density are mainly associated with precipitation, while that of stomatal size is mainly associated with temperature, and the skewness and kurtosis of stomatal traits are less related to climatic and soil variables. Beyond mean climate variables, stomatal trait moments also vary with climatic seasonality and extreme conditions. Our findings extend the knowledge of stomatal trait–environment relationships to the ecosystem scale, with applications in predicting future water and carbon cycles.