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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.

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.

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.

Crop photosynthesis and yield are limited by slow photosynthetic induction in sunflecks. We quantified variation in induction kinetics across diverse genotypes of wheat for the first time. Following a preliminary study that hinted at wide variation in induction kinetics across 58 genotypes, we grew 10 genotypes with contrasting responses in a controlled environment and quantified induction kinetics of carboxylation capacity (V cmax) from dynamic A versus ci curves after a shift from low to high light (from 50 μmol m–2 s–1 to 1500 μmol m–2 s–1), in five flag leaves per genotype. Within-genotype median time for 95% induction (t 95) of Vcmax varied 1.8-fold, from 5.2 min to 9.5 min. Our simulations suggest that non-instantaneous induction reduces daily net carbon gain by up to 15%, and that breeding to speed up V cmax induction in the slowest of our 10 genotypes to match that in the fastest genotype could increase daily net carbon gain by up to 3.4%, particularly for leaves in mid-canopy positions (cumulative leaf area index ≤1.5 m² m–2), those that experience predominantly short-duration sunflecks, and those with high photosynthetic capacities.

Reduced stomatal conductance is a common plant response to rising atmospheric CO 2 and increases water use efficiency ( W ). At the leaf-scale, W depends on water and nitrogen availability in addition to atmospheric CO 2 . In hydroclimate models W is a key driver of rainfall, droughts, and streamflow extremes. We used global climate data to derive Aridity Indices (AI) for forests over the period 1965–2015 and synthesised those with data for nitrogen deposition and W derived from stable isotopes in tree rings. AI and atmospheric CO 2 account for most of the variance in W of trees across the globe, while cumulative nitrogen deposition has a significant effect only in regions without strong legacies of atmospheric pollution. The relation of aridity and W displays a clear discontinuity. W and AI are strongly related below a threshold value of AI ≈ 1 but are not related where AI > 1. Tree ring data emphasise that effective demarcation of water-limited from non-water-limited behaviour of stomata is critical to improving hydrological models that operate at regional to global scales. Water use efficiency is a key measure of plant responses to climate change. Here, the authors investigate its control by CO2, nitrogen deposition, and water availability using a global tree-ring dataset. They find an aridity threshold and quantify changes in control over the past 50 years.

It is clear that stomata play a critical role in regulating water loss from terrestrial vegetation. What is not clear is how this regulation is achieved. Stomata appear to respond to perturbations of many aspects of the soil-plant-atmosphere hydraulic continuum, but there is little agreement regarding the mechanism (or mechanisms) by which stomata sense such perturbations. This review discusses feedback and feedforward mechanisms by which hydraulic perturbations are putatively transduced into stomatal movements, in relation to generic empirical features of those responses. It is argued that a metabolically mediated feedback response of stomatal guard cells to the water status in their immediate vicinity ('hydro-active local feed-back') remains the best explanation for many well-known features of hydraulically related stomatal behaviour, such as transient 'wrong-way' responses and the equivalence of hydraulic supply and demand as stomatal effectors. Furthermore, many curious phenomena that appear inconsistent with feedback, such as 'apparent feedforward' humidity responses and 'isohydric' behaviour (water potential homeostasis), are in fact expected to emerge from the juxtaposition of hydro-active local feedback and the well-known hysteretic and threshold-like effect of water potential on xylem hydraulic resistance.

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.

Recent advances have improved our ability to model stomatal conductance using process- or optimality-based models, and continuing research should focus on how stomata sense leaf turgor and on how to quantify the direct carbon costs of low leaf water potential.

It has been 75 yr since leaf respiratory metabolism in the light (day respiration) was identified as a low-flux metabolic pathway that accompanies photosynthesis. In principle, it provides carbon backbones for nitrogen assimilation and evolves CO2 and thus impacts on plant carbon and nitrogen balances. However, for a long time, uncertainties have remained as to whether techniques used to measure day respiratory efflux were valid and whether day respiration responded to environmental gaseous conditions. In the past few years, significant advances have beenmade using carbon isotopes, ‘omics’ analyses and surveys of respiration rates in mesocosms or ecosystems. There is substantial evidence that day respiration should be viewed as a highly dynamic metabolic pathway that interacts with photosynthesis and photorespiration and responds to atmospheric CO2 mole fraction. The view of leaf day respiration as a constant and/or negligible parameter of net carbon exchange isnow outdated and it should now be regarded as a central actor of plant carbon-use efficiency.

Rates of change in intrinsic water use efficiency ( W ) of trees relative to those in atmospheric [CO 2 ] ( c a ) have been mostly assessed via short-term studies (e.g., leaf analysis, flux analysis) and/or step increases in c a (e.g., FACE studies). Here we use compiled data for abundances of carbon isotopes in tree stems to show that on decadal scales, rates of change ( dW/dc a ) vary with location and rainfall within the global tropics. For the period 1915–1995, and including corrections for mesophyll conductance and photorespiration, dW/dc a for drier tropical forests (receiving ~ 1000 mm rainfall) were at least twice that of the wettest (receiving ~ 4000 mm). The data also empirically confirm theorized roles of tropical forests in changes in atmospheric 13 C/ 12 C ratios (the 13 C Suess Effect). Further formal analysis of geographic variation in decade-to-century scale dW/dc a will be needed to refine current models that predict increases in carbon uptake by forests without hydrological cost. How the water use efficiency of trees changes with atmospheric CO 2 variations has mostly been studied on short time scales. Here, a newly compiled data set covering 1915 to 1995 shows how rates of change in water use efficiency vary with location and rainfall over the global tropics on a decadal scale.