Stomatal responses to humidity, soil moisture and other factors that influence plant water status are critical drivers of photosynthesis, productivity, water yield, ecohydrology and climate forcing, ...yet we still lack a thorough mechanistic understanding of these responses. Here I review historical and recent advances in stomatal water relations. Clear evidence now implicates a metabolically mediated response to leaf water status (‘hydroactive feedback’) in stomatal responses to evaporative demand and soil drought, possibly involving abscisic acid production in leaves. Other hypothetical mechanisms involving vapor and heat transport within leaves may contribute to humidity, light and temperature responses, but require further theoretical clarification and experimental validation. Variation and dynamics in hydraulic conductance, particularly within leaves, may contribute to water status responses. Continuing research to fully resolve mechanisms of stomatal responses to water status should focus on several areas: validating and quantifying the mechanism of leaf-based hydroactive feedback, identifying where in leaves water status is actively sensed, clarifying the role of leaf vapor and energy transport in humidity and temperature responses, and verifying foundational but minimally replicated results of stomatal hydromechanics across species. Clarity on these matters promises to deliver modelers with a tractable and reliable mechanistic model of stomatal responses to water status.
Water movement from the xylem to stomata is poorly understood. There is still no consensus about whether apoplastic or symplastic pathways are more important, and recent work suggests vapour ...diffusion may also play a role. The objective of this study was to estimate the proportions of hydraulic conductance outside the bundle sheath contributed by apoplastic, symplastic and gas phase pathways, using a novel analytical framework based on measurable anatomical and biophysical parameters. The calculations presented here suggest that apoplastic pathways provide the majority of conductance outside the bundle sheath under most conditions, whereas symplastic pathways contribute only a small proportion. The contributions of apoplastic and gas phase pathways vary depending on several critical but poorly known or highly variable parameters namely, the effective Poiseuille radius for apoplastic bulk flow, the thickness of cell walls and vertical temperature gradients within the leaf. The gas phase conductance should increase strongly as the leaf centre becomes warmer than the epidermis – providing up to 44% of vertical water transport for a temperature gradient of 0.2 K. These results may help to explain how leaf water transport is influenced by light absorption, temperature and differences in leaf anatomy among species.
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.
Summary
Photosynthetic capacity per unit irradiance is greater, and the marginal carbon revenue of water (∂A/∂E) is smaller, in shaded leaves than sunlit leaves, apparently contradicting optimization ...theory. I tested the hypothesis that these patterns arise from optimal carbon partitioning subject to biophysical constraints on leaf water potential.
In a whole plant model with two canopy modules, I adjusted carbon partitioning, nitrogen partitioning and leaf water potential to maximize carbon profit or canopy photosynthesis, and recorded how gas exchange parameters compared between shaded and sunlit modules in the optimum.
The model predicted that photosynthetic capacity per unit irradiance should be larger, and ∂A/∂E smaller, in shaded modules compared to sunlit modules. This was attributable partly to radiation‐driven differences in evaporative demand, and partly to differences in hydraulic conductance arising from the need to balance marginal returns on stem carbon investment between modules. The model verified, however, that invariance in the marginal carbon revenue of N (∂A/∂N) is in fact optimal.
The Cowan–Farquhar optimality solution (invariance of ∂A/∂E) does not apply to spatial variation within a canopy. The resulting variation in carbon–water economy explains differences in capacity per unit irradiance, reconciling optimization theory with observations.
Recent decades have been characterized by increasing temperatures worldwide, resulting in an exponential climb in vapor pressure deficit (VPD). VPD has been identified as an increasingly important ...driver of plant functioning in terrestrial biomes and has been established as a major contributor in recent drought-induced plant mortality independent of other drivers associated with climate change. Despite this, few studies have isolated the physiological response of plant functioning to high VPD, thus limiting our understanding and ability to predict future impacts on terrestrial ecosystems. An abundance of evidence suggests that stomatal conductance declines under high VPD and transpiration increases in most species up until a given VPD threshold, leading to a cascade of subsequent impacts including reduced photosynthesis and growth, and higher risks of carbon starvation and hydraulic failure. Incorporation of photosynthetic and hydraulic traits in ‘next-generation’ land-surface models has the greatest potential for improved prediction of VPD responses at the plant- and global-scale, and will yield more mechanistic simulations of plant responses to a changing climate. By providing a fully integrated framework and evaluation of the impacts of high VPD on plant function, improvements in forecasting and long-term projections of climate impacts can be made.
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.
Abstract
Plants often experience multiple stresses in a given day or season, and it is self-evident that given functional traits can provide tolerances of multiple stresses. Yet, the multiple ...functions of individual traits are rarely explicitly considered in ecology and evolution due to a lack of a quantitative framework. We present a theory for considering the combined importance of the several functions that a single trait can contribute to alleviating multiple stresses. We derive five inter-related general predictions: (1) that trait multifunctionality is overall highly beneficial to fitness; (2) that species possessing multifunctional traits should increase in abundance and in niche breadth; (3) that traits are typically optimized for multiple functions and thus can be far from optimal for individual functions; (4) that the relative importance of each function of a multifunctional trait depends on the environment; and (5) that traits will be often “co-opted” for additional functions during evolution and community assembly. We demonstrate how the theory can be applied quantitatively by examining the multiple functions of leaf trichomes (hairs) using heuristic model simulations, substantiating the general principles. We identify avenues for further development and applications of the theory of trait multifunctionality in ecology and evolution.
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) Koxis strongly enhanced by the proximity of veins to lower epidermis; and (8) Koxis strongly influenced by spongy mesophyll anatomy, decreasing with protoplast size and increasing with airspace fraction and cell wall thickness. Correlations between anatomy and Koxacross species sometimes diverged from predicted causal effects, demonstrating the need for integrative models to resolve causation. For example, (9) Koxwas 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.
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