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• Leaf hydraulic and mesophyll CO₂ conductance are both influenced by leaf anatomical traits, however it is poorly understood how the temperature response of these conductances differs between C₄ and C₃ species with distinct leaf anatomy. • This study investigated the temperature response of leaf hydraulic conductance (K leaf), stomatal (g s) and mesophyll (g m) conductance to CO₂, and leaf anatomical traits in phylogenetically related Panicum antidotale (C₄) and P. bisulcatum (C₃) grasses. • The C₄ species had lower hydraulic conductance outside xylem (K ox) and K leaf compared with the C₃ species. However, the C₄ species had higher g m compared with the C₃ species. Traits associated with leaf water movement, K leaf and K ox, increased with temperature more in the C₃ than in the C₄ species, whereas traits related to carbon uptake, A net and g m, increased more with temperature in the C₄ than the C₃ species. • Our findings demonstrate that, in addition to a CO₂ concentrating mechanism, outside-xylem leaf anatomy in the C₄ species P. antidotale favours lower water movement through the leaf and stomata that provides an additional advantage for greater leaf carbon uptake relative to water loss with increasing leaf temperature than in the C₃ species P. bisulcatum.

Leaf hydraulic conductance (K leaf) and mesophyll conductance (g m) both represent major constraints to photosynthetic rate (A), and previous studies have suggested that K leaf and g m is correlated in leaves. However, there is scarce empirical information about their correlation. In this study, K leaf, leaf hydraulic conductance inside xylem (K x), leaf hydraulic conductance outside xylem (K ox), A, stomatal conductance (g s), g m, and anatomical and structural leaf traits in 11 Oryza genotypes were investigated to elucidate the correlation of H2O and CO2 diffusion inside leaves. All of the leaf functional and anatomical traits varied significantly among genotypes. K leaf was not correlated with the maximum theoretical stomatal conductance calculated from stomatal dimensions (g smax), and neither g s nor g smax were correlated with K x. Moreover, K ox was linearly correlated with g m and both were closely related to mesophyll structural traits. These results suggest that K leaf and g m are related to leaf anatomical and structural features, which may explain the mechanism for correlation between g m and K leaf.

The classical view that the drought-related hormone ABA simply acts locally at the guard cell level to induce stomatal closure is questioned by differences between isolated epidermis and intact leaves in stomatal response to several stimuli. We tested the hypothesis that ABA mediates, in addition to a local effect, a remote effect in planta by changing hydraulic regulation in the leaf upstream of the stomata. By gravimetry, porometry to water vapour and argon, and psychrometry, we investigated the effect of exogenous ABA on transpiration, stomatal conductance and leaf hydraulic conductance of mutants described as ABA-insensitive at the guard cell level. We show that foliar transpiration of several ABA-insensitive mutants decreases in response to ABA. We demonstrate that ABA decreases stomatal conductance and down-regulates leaf hydraulic conductance in both the wildtype Col-0 and the ABA-insensitive mutant ost2-2. We propose that ABA promotes stomatal closure in a dual way via its already known biochemical effect on guard cells and a novel, indirect hydraulic effect through a decrease in water permeability within leaf vascular tissues. Variability in sensitivity of leaf hydraulic conductance to ABA among species could provide a physiological basis to the isohydric or anisohydric behaviour.

• The hydraulic plumbing of vascular plant leaves varies considerably between major plant groups both in the spatial organization of veins, as well as their anatomical structure. • Five conifers, three ferns and 12 angiosperm trees were selected from tropical and temperate forests to investigate whether the profound differences in foliar morphology of these groups lead to correspondingly profound differences in leaf hydraulic efficiency. • We found that angiosperm leaves spanned a range of leaf hydraulic conductance from 3.9 to 36 mmol m2 s-1 MPa-1, whereas ferns (5.9-11.4 mmol m-2 s-1 MPa-1) and conifers (1.6-9.0 mmol m-2 s-1 MPa-1) were uniformly less conductive to liquid water. Leaf hydraulic conductance ( Kleaf) correlated strongly with stomatal conductance indicating an internal leaf-level regulation of liquid and vapour conductances. Photosynthetic capacity also increased with Kleaf, however, it became saturated at values of Kleaf over 20 mmol m-2 s-1 MPa-1. • The data suggest that vessels in the leaves of the angiosperms studied provide them with the flexibility to produce highly conductive leaves with correspondingly high photosynthetic capacities relative to tracheid-bearing species.

Key message Different groundwater conditions affect leaf hydraulic conductance and leaf pressure–volume parameters in Populus euphratica at the extremely arid zone in the northwest of China. Efficient water transport inside leaves constitutes a major determinant of plant function, especially in drought-stressed plants. The previous researches have reported the correlation between leaf hydraulic properties and water availability. In this study, we tested the hypothesis that water relation parameters of Populus euphratica in an extremely arid zone of China are sensitive and acclimated to groundwater depth. We measured leaf hydraulic conductance ( K leaf ) using rehydration kinetics methods (RKM), pressure–volume (P–V) curves, and leaf vulnerability curves of P. euphratica growing at four groundwater depth gradients. We also assessed the hydraulic safety margins across groundwater depth gradients. We found that K leaf–max shows an increasing trend as the groundwater depth increases, while osmotic potential at full turgor (π ft ) and turgor loss point (Ψ tlp ) exhibits a decreasing trend, suggesting that increased tolerance to drought is formed as the groundwater depth increases. Furthermore, safety margins showed positive and negative variations under different groundwater depths, indicating that P. euphratica has formed special drought survival strategies, which can be summarized as a “conservative” strategy in favorable water conditions or a “risk” strategy in severe drought stress.

• Hydraulic characteristics of pteridophyte (fern and Selaginella) foliage were investigated to determine whether the processes of water conduction and water loss are coordinated in these early vascular plants similarly to angiosperms. • Eight species of pteridophytes and associated woody angiosperms were examined from the sun and shade in a seasonally dry tropical forest. • Maximum leaf hydraulic conductivity (Kleaf) in the four pteridophytes was within the range of the sampled shade angiosperms but much lower than that of the sun-dwelling angiosperms. Hydraulic conductivity of both angiosperm and pteridophyte leaves showed a similar response to desiccation, with Kleafbecoming rapidly depressed once leaf water potential fell below a threshold. Stomatal closure in angiosperms corresponded closely with the water potential responsible for 50% loss of Kleafwhile pteridophytes were found to close stomata before Kleafdepression. • The contrasting behaviour of stomata in this sample of pteridophytes suggest that this may be an intrinsic difference between pteridophytes and angiosperms, with lower safety margins in angiosperms possibly enhancing both optimization of gas exchange and xylem investment.

• Clarifying the coordination of leaf hydraulic traits with gas exchange across closely-related species adapted to varying rainfall can provide insights into plant habitat distribution and drought adaptation. • The leaf hydraulic conductance (K leaf), stomatal conductance (g s), net assimilation (A), vein embolism and abscisic acid (ABA) concentration during dehydration were quantified, as well as pressure–volume curve traits and vein anatomy in 10 Caragana species adapted to a range of mean annual precipitation (MAP) conditions and growing in a common garden. • We found a positive correlation between Ψleaf at 50% loss of K leaf (K leaf P 50) and maximum K leaf (K leaf-max) across species. Species from low-MAP environments exhibited more negative K leaf P 50 and turgor loss point, and higher K leaf-max and leaf-specific capacity at full turgor, along with higher vein density and midrib xylem per leaf area, and a higher ratio of K leaf-max : maximum g s. Tighter stomatal control mediated by higher ABA accumulation during dehydration in these species resulted in an increase in hydraulic safety and intrinsic water use efficiency (WUEi) during drought. • Our results suggest that high hydraulic safety and efficiency combined with greater stomatal sensitivity triggered by ABA production and leading to greater WUEi provides drought tolerance in Caragana species adapted to low-MAP environments.

Given increasing water deficits across numerous ecosystems world-wide, it is urgent to understand the sequence of failure of leaf function during dehydration. We assessed dehydration-induced losses of rehydration capacity and maximum quantum yield of the photosystem II (F v/F m) in the leaves of 10 diverse angiosperm species, and tested when these occurred relative to turgor loss, declines of stomatal conductance g s, and hydraulic conductance K leaf, including xylem and outside xylem pathways for the same study plants. We resolved the sequences of relative water content and leaf water potential Ψleaf thresholds of functional impairment. On average, losses of leaf rehydration capacity occurred at dehydration beyond 50% declines of g s, K leaf and turgor loss point. Losses of F v/F m occurred after much stronger dehydration and were not recovered with leaf rehydration. Across species, tissue dehydration thresholds were intercorrelated, suggesting trait co-selection. Thresholds for each type of functional decline were much less variable across species in terms of relative water content than Ψleaf. The stomatal and leaf hydraulic systems show early functional declines before cell integrity is lost. Substantial damage to the photochemical apparatus occurs at extreme dehydration, after complete stomatal closure, and seems to be irreversible.

Climate change is expected to exacerbate drought for many plants, making drought tolerance a key driver of species and ecosystem responses. Plant drought tolerance is determined by multiple traits, but the relationships among traits, either within individual plants or across species, have not been evaluated for general patterns across plant diversity. We synthesized the published data for stomatal closure, wilting, declines in hydraulic conductivity in the leaves, stems, and roots, and plant mortality for 262 woody angiosperm and 48 gymnosperm species. We evaluated the correlations among the drought tolerance traits across species, and the general sequence of water potential thresholds for these traits within individual plants. The trait correlations across species provide a framework for predicting plant responses to a wide range of water stress from one or two sampled traits, increasing the ability to rapidly characterize drought tolerance across diverse species. Analyzing these correlations also identified correlations among the leaf and stem hydraulic traits and the wilting point, or turgor loss point, beyond those expected from shared ancestry or independent associations with water stress alone. Further, on average, the angiosperm species generally exhibited a sequence of drought tolerance traits that is expected to limit severe tissue damage during drought, such as wilting and substantial stem embolism. This synthesis of the relationships among the drought tolerance traits provides crucial, empirically supported insight into representing variation in multiple traits in models of plant and ecosystem responses to drought.

In order to study the role of PIP1 aquaporins in leaf water and CO2 transport, several lines of PIP1-deficient transgenic Populus tremula x alba were generated using a reverse genetic approach. These transgenic lines displayed no visible developmental or morphological phenotypes when grown under conditions of no water stress. Major photosynthetic parameters were also not affected by PIP1 down regulation. However, low levels of PIP1 expression resulted in greater leaf hydraulic resistance (an increase of 27%), which effectively implicated PIP1 role in water transport. Additionally, the expression level of PIP1 genes in the various transgenic lines was correlated with reductions in mesophyll conductance to CO2 (gm), suggesting that in poplar, these aquaporins influenced membrane permeability to CO2. Overall, although analysis showed that PIP1 genes contributed to the mass transfer of water and CO2 in poplar leaves, their down-regulation did not dramatically impair the physiological needs of this fast growing tree when cultivated under conditions of no stress.