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Abstract Rising atmospheric CO₂levels place terrestrial ecosystems under novel environmental conditions, and research in field settings is key to understanding how real plant communities will respond. Despite decades of progress in elevated CO₂(eCO₂) experiments, major gaps persist in our knowledge of plant responses to interacting influences of climate change, especially in areas outside North America and Western Europe.With a goal to expand access to field‐based eCO₂research, we designed, built, and tested TinyCO₂, a low‐cost field experiment for climate change research on plants. TinyCO₂features sixteen 0.62‐m²plot areas, half with ambient and half with elevated (+200 ppm) CO₂concentrations, and is suitable for short‐stature plants (≤0.5 m in height).Using a proportional‐integral control algorithm and constant sampling of air within the plots, TinyCO₂achieves consistent elevation of [CO₂] averaging +196.9 ppm. During testing, 95.1% of measured CO₂concentrations fell within 20% of the setpoint (ambient CO₂ + 200 ppm). A streamlined design and efficient use of instrumentation reduced the cost of the system to roughly one‐fifth of the cost of similar experiments from the past 30 years ($13.68 vs. $64.65 ppm⁻¹ m⁻², adjusted to 2024 USD).Our results demonstrate a system capable of precise and accurate field‐based CO₂elevation for significantly reduced cost. We envision the TinyCO₂design being implemented in a multitude of field‐based eCO₂studies, perhaps as part of a globally distributed collaborative network experiment. 

Abstract PremisePrevious studies have suggested a trade‐off between trichome density (D_t) and stomatal density (D_s) due to shared cell precursors. We clarified how, when, and why this developmental trade‐off may be overcome across species. MethodsWe derived equations to determine the developmental basis forD_tandD_sin trichome and stomatal indices (i_tandi_s) and the sizes of epidermal pavement cells (e), trichome bases (t), and stomata (s) and quantified the importance of these determinants ofD_tandD_sfor 78 California species. We compiled 17 previous studies ofD_t–D_srelationships to determine the commonness ofD_t–D_sassociations. We modeled the consequences of differentD_t–D_sassociations for plant carbon balance. ResultsOur analyses showed that higherD_twas determined by higheri_tand lowere, and higherD_sby higheri_sand lowere. Across California species, positiveD_t–D_scoordination arose due toi_t–i_scoordination and impacts of the variation ine. AD_t–D_strade‐off was found in only 30% of studies. Heuristic modeling showed that species sets would have the highest carbon balance with a positive or negative relationship or decoupling ofD_tandD_s, depending on environmental conditions. ConclusionsShared precursor cells of trichomes and stomata do not limit higher numbers of both cell types or drive a generalD_t–D_strade‐off across species. This developmental flexibility across diverse species enables differentD_t–D_sassociations according to environmental pressures. Developmental trait analysis can clarify how contrasting trait associations would arise within and across species. 

Abstract PremiseThe adaptive significance of amphistomy (stomata on both upper and lower leaf surfaces) is unresolved. A widespread association between amphistomy and open, sunny habitats suggests the adaptive benefit of amphistomy may be greatest in these contexts, but this hypothesis has not been tested experimentally. Understanding amphistomy informs its potential as a target for crop improvement and paleoenvironment reconstruction. MethodsWe developed a method to quantify “amphistomy advantage” () as the log‐ratio of photosynthesis in an amphistomatous leaf to that of the same leaf but with gas exchange blocked through the upper surface (pseudohypostomy). Humidity modulated stomatal conductance and thus enabled comparing photosynthesis at the same total stomatal conductance. We estimated and leaf traits in six coastal (open, sunny) and six montane (closed, shaded) populations of the indigenous Hawaiian species ʻilima (Sida fallax). ResultsCoastal ʻilima leaves benefit 4.04 times more from amphistomy than montane leaves. Evidence was equivocal with respect to two hypotheses: (1) that coastal leaves benefit more because they are thicker and have lower CO₂conductance through the internal airspace and (2) that they benefit more because they have similar conductance on each surface, as opposed to most conductance being through the lower surface. ConclusionsThis is the first direct experimental evidence that amphistomy increases photosynthesis, consistent with the hypothesis that parallel pathways through upper and lower mesophyll increase CO₂supply to chloroplasts. The prevalence of amphistomatous leaves in open, sunny habitats can partially be explained by the increased benefit of amphistomy in “sun” leaves, but the mechanistic basis remains uncertain. 

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. 

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. 

Abstract Plants differ widely in how soil drying affects stomatal conductance (g_s) and leaf water potential (ψ_leaf), and in the underlying physiological controls. Efforts to breed crops for drought resilience would benefit from a better understanding of these mechanisms and their diversity. We grew 12 diverse genotypes of common bean (Phaseolus vulgarisL.) and four of tepary bean (P. acutifolius;a highly drought resilient species) in the field under irrigation and post‐flowering drought, and quantified responses ofg_sandψ_leaf, and their controls (soil water potential [ψ_soil], evaporative demand [Δw] and plant hydraulic conductance [K]). We hypothesised that (i) common beans would be more “isohydric” (i.e., exhibit strong stomatal closure in drought, minimisingψ_leafdecline) than tepary beans, and that genotypes with largerψ_leafdecline (more “anisohydric”) would exhibit (ii) smaller increases in Δw, due to less suppression of evaporative cooling by stomatal closure and hence less canopy warming, but (iii) largerKdeclines due toψ_leafdecline. Contrary to our hypotheses, we found that half of the common bean genotypes were similarly anisohydric to most tepary beans; canopy temperature was cooler in isohydric genotypes leading to smaller increases in Δwin drought; and that stomatal closure andKdecline were similar in isohydric and anisohydric genotypes.g_sandψ_leafwere virtually insensitive to drought in one tepary genotype (G40068). Our results highlight the potential importance of non‐stomatal mechanisms for leaf cooling, and the variability in drought resilience traits among closely related crop legumes. 

Abstract Changes in leaf temperature are known to drive stomatal responses, because the leaf‐to‐air water vapour gradient (Δw) increases with temperature if ambient vapour pressure is held constant, and stomata respond to changes in Δw. However, the direct response of stomata to temperature (DRST; the response when Δwis held constant by adjusting ambient humidity) has been examined far less extensively. Though the meagre available data suggest the response is usually positive, results differ widely and defy broad generalisation. As a result, little is known about the DRST. This review discusses the current state of knowledge about the DRST, including numerous hypothesised biophysical mechanisms, potential implications of the response for plant adaptation, and possible impacts of the DRST on plant‐atmosphere carbon and water exchange in a changing climate. 

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. 

Synopsis Classic debates in community ecology focused on the complexities of considering an ecosystem as a super-organ or organism. New consideration of such perspectives could clarify mechanisms underlying the dynamics of forest carbon dioxide (CO2) uptake and water vapor loss, important for predicting and managing the future of Earth’s ecosystems and climate system. Here, we provide a rubric for considering ecosystem traits as aggregated, systemic, or emergent, i.e., representing the ecosystem as an aggregate of its individuals or as a metaphorical or literal super-organ or organism. We review recent approaches to scaling-up plant water relations (hydraulics) concepts developed for organs and organisms to enable and interpret measurements at ecosystem-level. We focus on three community-scale versions of water relations traits that have potential to provide mechanistic insight into climate change responses of forest CO2 and H2O gas exchange and productivity: leaf water potential (Ψcanopy), pressure volume curves (eco-PV), and hydraulic conductance (Keco). These analyses can reveal additional ecosystem-scale parameters analogous to those typically quantified for leaves or plants (e.g., wilting point and hydraulic vulnerability) that may act as thresholds in forest responses to drought, including growth cessation, mortality, and flammability. We unite these concepts in a novel framework to predict Ψcanopy and its approaching of critical thresholds during drought, using measurements of Keco and eco-PV curves. We thus delineate how the extension of water relations concepts from organ- and organism-scales can reveal the hydraulic constraints on the interaction of vegetation and climate and provide new mechanistic understanding and prediction of forest water use and productivity. 

Abstract Stomata are the gatekeepers of plant water use and must quickly respond to changes in plant water status to ensure plant survival under fluctuating environmental conditions. The mechanism for their closure is highly sensitive to disturbances in leaf water status, which makes isolating their response to declining water content difficult to characterise and to compare responses among species. Using a small‐scale non‐destructive nuclear magnetic resonance spectrometer as a leaf water content sensor, we measure the stomatal response to rapid induction of water deficit in the leaves of nine species of eucalypt from contrasting climates. We found a strong linear correlation between relative water content at 50% stomatal conductance (RWC_gs50) and mean annual temperature at the climate of origin of each species. We also show evidence for stomata to maintain control over water loss well below turgor loss point in species adapted to warmer climates and secondary increases in stomatal conductance despite declining water content. We propose that RWC_gs50is a promising trait to guide future investigations comparing stomatal responses to water deficit. It may provide a useful phenotyping trait to delineate tolerance and adaption to hot temperatures and high leaf‐to‐air vapour pressure deficits.