A tale of two pumps: Blue light and abscisic acid alter Arabidopsis leaf hydraulics via bundle sheath cell H + -ATPases.

The bundle sheath cell (BSC) layer tightly enveloping the xylem throughout the leaf is recognized as a major signal-perceiving "valve" in series with stomata, regulating leaf hydraulic conductance (Kleaf) and thereby radial water flow via the transpiring leaf. The BSC blue light (BL) signaling pathway increases Kleaf and the underlying BSC water permeability. Here, we explored the hypothesis that BSCs also harbor a Kleaf-downregulating signaling pathway related to the stress phytohormone abscisic acid (ABA). We employed fluorescence imaging of xylem sap in detached leaves and BSC protoplasts from different genotypes of Arabidopsis (Arabidopsis thaliana) plants, using pH and membrane potential probes to monitor physiological responses to ABA and BL in combination with pharmacological agents. We found that BL-enhanced Kleaf required elevated BSC cytosolic Ca2+. ABA inhibited BL-activated xylem-sap-acidifying BSC H + -ATPase AHA2 (Arabidopsis H + -ATPase 2), resulting in depolarized BSCs and alkalinized xylem sap. ABA also stimulated BSC vacuolar H + -ATPase (VHA), which alkalinized the BSC cytosol. Each pump stimulation, AHA2 by BL and VHA by ABA (under BL), also required Ca2+. ABA stimulated VHA in the dark depending on Ca2+, but only in an alkaline external medium. Taken together with earlier findings on the pH sensitivity of BSC osmotic water permeability (i.e., aquaporin activity), our results suggest a Ca2+-dependent and pH-mediated causative link between the BL- and ABA-regulated activities of two BSC H + -ATPases and Kleaf.

Leaf diffusional capacity largely contributes to the reduced photosynthesis in rice plants under magnesium deficiency.

Numerous studies have clarified the impacts of magnesium (Mg) on leaf photosynthesis from the perspectives of protein synthesis, enzymes activation and carbohydrate partitioning. However, it still remains largely unknown how stomatal and mesophyll conductances (gs and gm, respectively) are regulated by Mg. In the present study, leaf gas exchanges, leaf hydraulic parameters, leaf structural traits and cell wall composition were examined in rice plants grown under high and low Mg treatments to elucidate the impacts of Mg on gs and gm. Our results showed that reduction of leaf photosynthesis under Mg deficiency was mainly caused by the decreased gm, followed by reduced leaf biochemical capacity and gs, and leaf outside-xylem hydraulic conductance (Kox) was the major factor restricting gs under Mg deficiency. Moreover, increased leaf hemicellulose, lignin and pectin contents and decreased cell wall effective porosity were observed in low Mg plants relative to high Mg plants. These results suggest that Kox and cell wall composition play important roles in regulating gs and gm, respectively, in rice plants under Mg shortages.

Stomatal response to VPD is not triggered by changes in soil-leaf hydraulic conductance in Arabidopsis or Callitris.

Stomatal closure under high VPDL (leaf to air vapour pressure deficit) is a primary means by which plants prevent large excursions in transpiration rate and leaf water potential (Ψleaf) that could lead to tissue damage. Yet, the drivers of this response remain controversial. Changes in Ψleaf appear to drive stomatal VPDL response, but many argue that dynamic changes in soil-to-leaf hydraulic conductance (Ks-l) make an important contribution to this response pathway, even in well-hydrated soils. Here, we examined whether the regulation of whole plant stomatal conductance (gc) in response to typical changes in daytime VPDL is influenced by dynamic changes in Ks-l. We use well-watered plants of two species with contrasting ecological and physiological features: the herbaceous Arabidopsis thaliana (ecotype Columbia-0) and the dry forest conifer Callitris rhomboidea. The dynamics of Ks-l and gc were continuously monitored by combining concurrent in situ measurements of Ψleaf using an open optical dendrometer and whole plant transpiration using a balance. Large changes in VPDL were imposed to induce stomatal closure and observe the impact on Ks-l. In both species, gc was observed to decline substantially as VPDL increased, while Ks-l remained stable. Our finding suggests that stomatal regulation of transpiration is not contingent on a decrease in Ks-l. Static Ks-l provides a much simpler explanation for transpiration control in hydrated plants and enables simplified modelling and new methods for monitoring plant water use in the field.

Illuminating plant water dynamics: the role of light in leaf hydraulic regulation.

Light intensity and quality influence photosynthesis directly but also have an indirect effect by increasing stomatal apertures and enhancing gas exchange. Consequently, in areas such as the upper canopy, a high water demand for transpiration and temperature regulation is created. This paper explores how light intensity and the natural high Blue-Light (BL) : Red-Light (RL) ratio in these areas, is important for controlling leaf hydraulic conductance (Kleaf ) by BL signal transduction, increasing water permeability in cells surrounding the vascular tissue, in supporting the enormous water demands. Conversely, shaded inner-canopy areas receive less radiation, have lower water and cooling demands, and exhibit reduced Kleaf due to diminished intensity and BL induction. Intriguingly, shaded leaves display higher water-use efficiency (compared with upper-canopy) due to decreased transpiration and cooling requirements while the presence of RL supports photosynthesis.

Increased stomatal conductance and leaf biochemical capacity, not mesophyll conductance, contributing to the enhanced photosynthesis in Oryza plants during domestication.

MAIN CONCLUSION: Leaf biochemical capacity and the ratio of leaf biochemical capacity to stomatal conductance are promising to enhance leaf photosynthetic rate and water use efficiency in rice plants, respectively. Domestication may have great impact on crop photosynthetic rate, which has not been fully understood, especially from the perspective of stomatal conductance, mesophyll conductance, and leaf biochemical capacity simultaneously. In this study, we constructed a database consisting of 141 and 92 sets of data from wild and cultivated rice, respectively, including leaf gas exchange parameters, hydraulic conductance, structural traits, and nitrogen content. We found that, compared to wild rice, enhanced leaf photosynthetic rate in cultivated rice was mainly resulted by the increased stomatal conductance and leaf biochemical capacity, rather than mesophyll conductance. The unchanged mesophyll conductance observed during domestication suggested that it might have been optimized during plant evolution in wild rice. Additionally, the positive relationship between stomatal density and stomatal conductance disappeared in Oryza plants during domestication, suggesting that stomatal conductance in cultivated rice is less restricted by stomatal density, compared to wild rice. Moreover, in both wild and cultivated rice, leaf photosynthetic rate was mainly determined by leaf biochemical capacity, rather than stomatal conductance and mesophyll conductance. This study highlighted the important role of stomatal conductance and leaf biochemical capacity in improvement of leaf photosynthetic rate in rice during domestication. Leaf biochemical capacity and the ratio of leaf biochemical capacity to stomatal conductance should be further investigated to enhance leaf photosynthetic rate and water use efficiency in rice plants, respectively.

Prospects and perspectives: inferring physiological and regulatory targets for CAM from molecular and modelling approaches.

BACKGROUND AND SCOPE: This review summarizes recent advances in our understanding of Crassulacean Acid Metabolism (CAM) by integrating evolutionary, ecological, physiological, metabolic and molecular perspectives. A number of key control loops which moderate the expression of CAM phases, and their metabolic and molecular control, are explored. These include nocturnal stomatal opening, activation of phosphoenolpyruvate carboxylase by a specific protein kinase, interactions with circadian clock control, as well as daytime decarboxylation and activation of Rubisco. The vacuolar storage and release of malic acid and the interplay between the supply and demand for carbohydrate reserves are also key metabolic control points. FUTURE OPPORTUNITIES: We identify open questions and opportunities, with experimentation informed by top-down molecular modelling approaches allied with bottom-up mechanistic modelling systems. For example, mining transcriptomic datasets using high-speed systems approaches will help to identify targets for future genetic manipulation experiments to define the regulation of CAM (whether circadian or metabolic control). We emphasize that inferences arising from computational approaches or advanced nuclear sequencing techniques can identify potential genes and transcription factors as regulatory targets. However, these outputs then require systematic evaluation, using genetic manipulation in key model organisms over a developmental progression, combining gene silencing and metabolic flux analysis and modelling to define functionality across the CAM day-night cycle. From an evolutionary perspective, the origins and function of CAM succulents and responses to water deficits are set against the mesophyll and hydraulic limitations imposed by cell and tissue succulence in contrasting morphological lineages. We highlight the interplay between traits across shoots (3D vein density, mesophyll conductance and cell shrinkage) and roots (xylem embolism and segmentation). Thus, molecular, biophysical and biochemical processes help to curtail water losses and exploit rapid rehydration during restorative rain events. In the face of a changing climate, we hope such approaches will stimulate opportunities for future research.

Hydraulic segmentation explains differences in loss of branch conductance caused by fire.

The hydraulic death hypothesis suggests that fires kill trees by damaging the plant's hydraulic continuum in addition to stem cambium. A corollary to this hypothesis is that plants that survive fires possess 'pyrohydraulic' traits that prevent heat-induced embolism formation in the xylem and aid post-fire survival. We examine whether hydraulic segmentation within stem xylem may act as such a trait. To do so, we measured the percentage loss of conductance (PLC) and vulnerability to embolism axially along segments of branches exposed to heat plumes in two differing species, fire-tolerant Eucalyptus cladocalyx F. Muell and fire-sensitive Kiggelaria africana L., testing model predictions that fire-tolerant species would exhibit higher degrees of hydraulic segmentation (greater PLC in the distal parts of the branch than the basal) than fire-intolerant species (similar PLC between segments). Following exposure to a heat plume, K. africana suffered between 73 and 84% loss of conductance in all branch segments, whereas E. cladocalyx had 73% loss of conductance in whole branches, including the distal tips, falling to 29% in the most basal part of the branch. There was no evidence for differences in resistance segmentation between the species, and there was limited evidence for differences in distal vulnerability to embolism across the branches. Hydraulic segmentation in E. cladocalyx may enable it to resprout effectively post-fire with a functional hydraulic system. The lack of hydraulic segmentation in K. africana reveals the need to understand possible trade-offs associated with hydraulic segmentation in long-lived woody species with respect to drought and fire.

Exogenous Calcium Induces Different Hydraulic Strategies in Response to Osmotic Stress in Maize Seedlings.

Recent discoveries regarding the signal molecules involved in abiotic stresses require integration into the field of plant hydraulic property research. Although calcium (Ca) is an important second messenger involved in numerous complex, abiotic stress-induced signaling pathways, it remains unclear how exogenous calcium mediates cellular signaling to promote plant drought resistance. We investigated the effects of calcium on the water balance and hydraulic properties in maize seedlings (Zea mays L.) under osmotic stress simulated by 10% (m/v) PEG-6000 in a hydroponic culture. The osmotic stress dramatically decreased the photosynthetic rate, transpiration rate, stomatal conductance, leaf water content, and root water absorption. However, the short-term (2 h) and long-term (10 d) exogenous Ca2+ (CaCl2: 10 mM) treatments had different effects on the maize gas exchange parameters and leaf water status. The short-term treatment improved the leaf transpiration by inhibiting the abscisic acid (ABA) synthesis and accumulation in the leaves, generating a stronger transpiration pull and enhancing the root water absorption and axial flow path water transport by increasing the root hydraulic conductance to relieve the osmotic stress-induced inhibition. The long-term treatment induced the ABA and H2O2 accumulation in the roots and leaves. Under osmotic stress, the accumulation of ABA, H2O2, and Ca2+ rapidly repressed the transpiration and enhanced the radial flow path water transport, decreasing the water loss and improving the stress tolerance. These insights suggest a role for a judicious use of Ca fertilizer in reducing the adverse effects of drought on agricultural production.

Axial conduit widening, tree height, and height growth rate set the hydraulic transition of sapwood into heartwood.

The size-related xylem adjustments required to maintain a constant leaf-specific sapwood conductance (KLEAF) with increasing height (H) are still under discussion. Alternative hypotheses are that: (i) the conduit hydraulic diameter (Dh) at any position in the stem and/or (ii) the number of sapwood rings at stem base (NSWr) increase with H. In addition, (iii) reduced stem elongation (ΔH) increases the tip-to-base conductance through inner xylem rings, thus possibly the NSWr contributing to KLEAF. A detailed stem analysis showed that Dh increased with the distance from the ring apex (DCA) in all rings of a Picea abies and a Fagus sylvatica tree. Net of DCA effect, Dh did not increase with H. Using sapwood traits from a global dataset, NSWr increased with H, decreased with ΔH, and the mean sapwood ring width (SWrw) increased with ΔH. A numerical model based on anatomical patterns predicted the effects of H and ΔH on the conductance of inner xylem rings. Our results suggest that the sapwood/heartwood transition depends on both H and ΔH, and is set when the carbon allocation to maintenance respiration of living cells in inner sapwood rings produces a lower gain in total conductance than investing the same carbon in new vascular conduits.

Theoretical Analyses of Turgor Pressure during Stress Relaxation and Water Uptake, and after Changes in Expansive Growth Rate When Water Uptake Is Normal and Reduced.

Turgor pressure provides the force needed to stress and deform the cell walls of plants, algae, and fungi during expansive growth. However, turgor pressure plays another subtle but equally important role in expansive growth of walled cells: it connects the two biophysical processes of water uptake and wall deformation to ensure that the volumetric rates of water uptake and enlargement of the cell wall chamber are equal. In this study, the role of turgor pressure as a 'connector' is investigated analytically by employing validated and established biophysical equations. The objective is to determine the effect of 'wall loosening' on the magnitude of turgor pressure. It is known that an increase or decrease in turgor pressure and/or wall loosening rate increases or decreases the expansive growth rate, respectively. Interestingly, it is shown that an increase in the wall loosening rate decreases the turgor pressure slightly, thus reducing the effect of wall loosening on increasing the expansive growth rate. Other analyses reveal that reducing the rate of water uptake results in a larger decrease in turgor pressure with the same increase in wall loosening rate, which further reduces the effect of wall loosening on increasing the expansive growth rate.

Role of hydraulic traits in stomatal regulation of transpiration under different vapour pressure deficits across five Mediterranean tree crops.

The differential stomatal regulation of transpiration among plant species in response to water deficit is not fully understood, although several hydraulic traits have been reported to influence it. This knowledge gap is partly due to a lack of direct and concomitant experimental data on transpiration, stomatal conductance, and hydraulic traits. We measured sap flux density (Js), stomatal conductance (gs), and different hydraulic traits in five crop species. Our aim was to contribute to establishing the causal relationship between water consumption and its regulation using a hydraulic trait-based approach. The results showed that the species-specific regulation of Js by gs was overall coordinated with the functional hydraulic traits analysed. Particularly relevant was the negative and significant relationship found between the Huber value (Hv) and its functional analogue ratio between maximum Js and gs (Jsmax/gsmax) which can be understood as a compensation to maintain the hydraulic supply to the leaves. The Hv was also significantly related to the slope of the relationship between gs and Js response to vapour pressure deficit and explained most of its variability, adding up to evidence recognizing Hv as a major trait in plant water relations. Thus, a hydraulic basis for regulation of tree water use should be considered.

Irrigation with primary wastewater alters wood anatomy and composition in willow Salix miyabeana SX67.

Traditional treatment of wastewaters is a burden for local governments. Using short rotation coppice willow (SRCW) as vegetal filter has several environmental and economic benefits. Here, we investigated the effect of primary wastewater irrigation on wood structure and composition of the willow cultivar Salix miyabeana 'SX67' following two years of growth. Compared to unirrigated plants (UI), stem sections of plants irrigated with primary wastewater (WWD) showed an unexpected decrease of hydraulic conductance (KS) associated with a decrease in vessel density but not vessel diameter. The majority (86%) of vessels had diameters range groups [20-30[, [30-40[and [40-50[µm and contributed to > 75% of theoretical KS, while the group class [50-60[µm (less than 10% of vessels) still accounted for > 20% of total KS regardless irrigation treatments. WWD significantly alters the chemical composition of wood with an increase of glucan content by 9 to 16.4% and a decrease of extractives by 35.3 to 36.4% when compared to UI or to plants irrigated with potable water (PW). The fertigation did also increase the proportion of the tension wood which highly correlated with glucan content. In the context of energetic transition and mitigation of climate change, such results are of high interest since WWD effectively permit the phytofiltration of large amounts of organic contaminated effluents without impairing SRCW physiology.

Effects of a Pseudomonas Strain on the Lipid Transfer Proteins, Appoplast Barriers and Activity of Aquaporins Associated with Hydraulic Conductance of Pea Plants.

Lipid transfer proteins (LTPs) are known to be involved in suberin deposition in the Casparian bands of pea roots, thereby reinforcing apoplast barriers. Moreover, the Pseudomonas mandelii IB-Ki14 strain accelerated formation of the Casparian bands in wheat plants, although involvement of LTPs in the process was not studied. Here, we investigated the effects of P. mandelii IB-Ki14 on LTPs, formation of the Casparian bands, hydraulic conductance and activity of aquaporins (AQPs) in pea plants. RT PCR showed a 1.6-1.9-fold up-regulation of the PsLTP-coding genes and an increase in the abundance of LTP proteins in the phloem of pea roots induced by the treatment with P. mandelii IB-Ki14. The treatment was accompanied with increased deposition of suberin in the Casparian bands. Hydraulic conductance did not decrease in association with the bacterial treatment despite strengthening of the apoplast barriers. At the same time, the Fenton reagent, serving as an AQPs inhibitor, decreased hydraulic conductance to a greater extent in treated plants relative to the control group, indicating an increase in the AQP activity by the bacteria. We hypothesize that P. mandelii IB-Ki14 stimulates deposition of suberin, in the biosynthesis of which LTPs are involved, and increases aquaporin activity, which in turn prevents a decrease in hydraulic conductance due to formation of the apoplast barriers in pea roots.

Physiological response and photosynthetic recovery to an extreme drought: Evidence from plants in a dry-hot valley savanna of Southwest China.

The frequency of extreme drought events has been rising worldwide, but due to its unpredictability, how plants will respond remains poorly understood. Here, we aimed to characterize how the hydraulics and photosynthesis of savanna plants respond to extreme drought, and tested whether they can subsequently recover photosynthesis after drought. There was an extreme drought in 2019 in Southwest (SW) China. We investigated photosynthetic gas exchange, leaf-, stem-, and whole-shoot hydraulic conductance of 18 plant species with diverse leaf habits (deciduous, semi-deciduous and evergreen) and growth forms (tree and shrub) from a dry-hot valley savanna in SW China for three rainy seasons from 2019 to 2021. We also compared photosynthetic gas exchange to those of a regular year (2014). We found that leaf stomatal and hydraulic conductance and maximum photosynthetic rate were significantly lower during the drought in 2019 than in the wetter years. In 2019, all studied plants maintained stomatal conductance at their minimum level observed, which could be related to high vapor pressure deficits (VPD, >2 kPa). However, no significant difference in stem and shoot hydraulic conductance was detected across years. The reductions in leaf hydraulic conductance and stomatal regulation under extreme drought might help keep the stem hydraulic function. Stomatal conductance and photosynthesis after drought (2020 and 2021) showed comparable or even higher values compared to that of 2014, suggesting high recovery of photosynthetic gas exchange. In addition, the response of hydraulic and photosynthetic traits to extreme drought was convergent across leaf habits and growth forms. Our results will help better understand the physiological mechanism underlying the response of savanna ecosystems to climate change.

Soil-plant hydraulics explain stomatal efficiency-safety tradeoff.

The efficiency-safety tradeoff has been thoroughly investigated in plants, especially concerning their capacity to transport water and avoid embolism. Stomatal regulation is a vital plant behaviour to respond to soil and atmospheric water limitation. Recently, a stomatal efficiency-safety tradeoff was reported where plants with higher maximum stomatal conductance (gmax ) exhibited greater sensitivity to stomatal closure during soil drying, that is, less negative leaf water potential at 50% gmax (ψgs50 ). However, the underlying mechanism of this gmax -ψgs50 tradeoff remains unknown. Here, we utilized a soil-plant hydraulic model, in which stomatal closure is triggered by nonlinearity in soil-plant hydraulics, to investigate such tradeoff. Our simulations show that increasing gmax is aligned with less negative ψgs50 . Plants with higher gmax (also higher transpiration) require larger quantities of water to be moved across the rhizosphere, which results in a precipitous decrease in water potential at the soil-root interface, and therefore in the leaves. We demonstrated that the gmax -ψgs50 tradeoff can be predicted based on soil-plant hydraulics, and is impacted by plant hydraulic properties, such as plant hydraulic conductance, active root length and embolism resistance. We conclude that plants may therefore adjust their growth and/or their hydraulic properties to adapt to contrasting habitats and climate conditions.

Identifying physiological and genetic determinants of faba bean transpiration response to evaporative demand.

BACKGROUND AND AIMS: Limiting maximum transpiration rate (TR) under high vapour pressure deficit (VPD) works as a water conservation strategy. While some breeding programmes have incorporated this trait into some crops to boost yields in water-limited environments, its underlying physiological mechanisms and genetic regulation remain unknown for faba bean (Vicia faba). Thus, we aimed to identify genetic variation in the TR response to VPD in a population of faba bean recombinant inbred lines (RILs) derived from two parental lines with contrasting water use (Mélodie/2 and ILB 938/2). METHODS: Plants were grown in well-watered soil in a climate-controlled glasshouse with diurnally fluctuating VPD and light conditions. Whole plant transpiration was measured in a gas exchange chamber that tightly regulated VPD around the shoot under constant light, while whole-plant hydraulic conductance and its components (root and stem hydraulic conductance) were calculated from dividing TR by water potential gradients measured with a pressure chamber. KEY RESULTS: Although TR of Mélodie/2 increased linearly with VPD, ILB 938/2 limited its TR above 2.0 kPa. Nevertheless, Mélodie/2 had a higher leaf water potential than ILB 938/2 at both low (1.0 kPa) and high (3.2 kPa) VPD. Almost 90 % of the RILs limited their TR at high VPD with a break-point (BP) range of 1.5-3.0 kPa and about 10 % had a linear TR response to VPD. Thirteen genomic regions contributing to minimum and maximum transpiration, and whole-plant and root hydraulic conductance, were identified on chromosomes 1 and 3, while one locus associated with BP transpiration was identified on chromosome 5. CONCLUSIONS: This study provides insight into the physiological and genetic control of transpiration in faba bean and opportunities for marker-assisted selection to improve its performance in water-limited environments.

Above and belowground traits impacting transpiration decline during soil drying in 48 maize (Zea mays) genotypes.

BACKGROUND AND AIMS: Stomatal regulation allows plants to promptly respond to water stress. However, our understanding of the impact of above and belowground hydraulic traits on stomatal regulation remains incomplete. The objective of this study was to investigate how key plant hydraulic traits impact transpiration of maize during soil drying. We hypothesize that the stomatal response to soil drying is related to a loss in soil hydraulic conductivity at the root-soil interface, which in turn depends on plant hydraulic traits. METHODS: We investigate the response of 48 contrasting maize (Zea mays) genotypes to soil drying, utilizing a novel phenotyping facility. In this context, we measure the relationship between leaf water potential, soil water potential, soil water content and transpiration, as well as root, rhizosphere and aboveground plant traits. KEY RESULTS: Genotypes differed in their responsiveness to soil drying. The critical soil water potential at which plants started decreasing transpiration was related to a combination of above and belowground traits: genotypes with a higher maximum transpiration and plant hydraulic conductance as well as a smaller root and rhizosphere system closed stomata at less negative soil water potentials. CONCLUSIONS: Our results demonstrate the importance of belowground hydraulics for stomatal regulation and hence drought responsiveness during soil drying. Furthermore, this finding supports the hypothesis that stomata start to close when soil hydraulic conductivity drops at the root-soil interface.

Acid waters in tank bromeliads: Causes and potential consequences.

PREMISE: The consequences of acidity for plant performance are profound, yet the prevalence and causes of low pH in bromeliad tank water are unknown despite its functional relevance to key members of many neotropical plant communities. METHODS: We investigated tank water pH for eight bromeliad species in the field and for the widely occurring Guzmania monostachia in varying light. We compared pH changes over time between plant and artificial tanks containing a solution combined from several plants. Aquaporin transcripts were measured for field plants at two levels of pH. We investigated relationships between pH, leaf hydraulic conductance, and CO2 concentration in greenhouse plants and tested proton pump activity using a stimulator and inhibitor. RESULTS: Mean tank water pH for the eight species was 4.7 ± 0.06 and was lower for G. monostachia in higher light. The pH of the solution in artificial tanks, unlike in plants, did not decrease over time. Aquaporin transcription was higher for plants with lower pH, but leaf hydraulic conductance did not differ, suggesting that the pH did not influence water uptake. Tank pH and CO2 concentration were inversely related. Fusicoccin enhanced a decrease in tank pH, whereas orthovanadate did not. CONCLUSIONS: Guzmania monostachia acidified its tank water via leaf proton pumps, which appeared responsive to light. Low pH increased aquaporin transcripts but did not influence leaf hydraulic conductance, hence may be more relevant to nutrient uptake.

Stomatal conductance in rice leaves and panicles responds differently to abscisic acid and soil drought.

Improvement of photosynthesis in non-foliar green tissues is beneficial for enhancing crop yield. Recently, we have demonstrated that panicle stomatal conductance is a major limiting factor for photosynthesis. However, mechanisms underlying the responses of panicle stomatal conductance (gs,panicle) and photosynthesis (Apanicle) to environmental stimuli remain unknown. In the present study, the responses of gs,panicle and leaf stomatal conductance (gs,leaf) to exogenous application of abscisic acid and step-changes in vapor pressure deficit were investigated at the anthesis stage in pot-grown rice plants. Furthermore, the effects of drought on Apanicle and leaf photosynthesis (Aleaf) were examined. Smearing and xylem feeding of abscisic acid significantly decreased gs,leaf. In contrast, while smearing of abscisic acid substantially increased gs,panicle, its xylem feeding dramatically decreased gs,panicle. In addition, both gs,leaf and gs,panicle effectively responded to step changes in vapor pressure deficit. Furthermore, both Aleaf and Apanicle were sensitive to plant dehydration; however, given the lower sensitivity of panicle water potential than leaf water potential to drought, Apanicle was less sensitive to soil drought than Aleaf. These findings indicate that gs,panicle is hydropassively regulated, while panicle photosynthesis is less sensitive to drought.

Leaves as bottlenecks: The contribution of tree leaves to hydraulic resistance within the soil-plant-atmosphere continuum.

Within vascular plants, the partitioning of hydraulic resistance along the soil-to-leaf continuum affects transpiration and its response to environmental conditions. In trees, the fractional contribution of leaf hydraulic resistance (Rleaf ) to total soil-to-leaf hydraulic resistance (Rtotal ), or fRleaf (=Rleaf /Rtotal ), is thought to be large, but this has not been tested comprehensively. We compiled a multibiome data set of fRleaf using new and previously published measurements of pressure differences within trees in situ. Across 80 samples, fRleaf averaged 0.51 (95% confidence interval [CI] = 0.46-0.57) and it declined with tree height. We also used the allometric relationship between field-based measurements of soil-to-leaf hydraulic conductance and laboratory-based measurements of leaf hydraulic conductance to compute the average fRleaf for 19 tree samples, which was 0.40 (95% CI = 0.29-0.56). The in situ technique produces a more accurate descriptor of fRleaf because it accounts for dynamic leaf hydraulic conductance. Both approaches demonstrate the outsized role of leaves in controlling tree hydrodynamics. A larger fRleaf may help stems from loss of hydraulic conductance. Thus, the decline in fRleaf with tree height would contribute to greater drought vulnerability in taller trees and potentially to their observed disproportionate drought mortality.