Isotope-Based Techniques to Investigate Factors Influencing Water Use Efficiency in Pinus koraiensis Leaves during Plant Growth.

Plant water use efficiency (WUE) is a comprehensive physiological indicator of plant growth and ability to adapt to drought. However, research on the mechanisms controlling WUE during plant growth and development remains weak. Here, we studied Pinus koraiensis as a typical evergreen conifer species in Northeast China. After collecting 80 tree samples with varying diameters at breast height (DBH), we measured δ13C and δ18O as an indicator of WUE, leaf morphology (volume, dry weight, and total epidermal area), ecological stoichiometry (carbon, nitrogen, and phosphorus content), and abiotic factors (light environment, soil pH, soil water content, and soil nutrient content). Correlational analysis of these variables revealed distinct differences between smaller/younger and larger/older plants: (1) In plants with DBH less than 52 cm, δ13C was positively related to DBH, and δ18O was negatively related to DBH. Plants with DBH greater than 52 cm showed no relationship between δ13C and DBH, and δ18O was positively related to DBH. (2) In plants with DBH less than 52 cm, there was a negative correlation between δ13C and δ18O and between δ13C and leaf phosphorus content (LP), but a positive correlation between δ13C and DBH, leaf mass per area (LMA), and leaf density (LD). The slopes of DBH-δ13C, δ18O-δ13C, leaf nitrogen content (LN)-δ13C, and LMA-δ13C correlations were greater in smaller plants than large plants. (3) Structural equation modelling showed that in smaller plants, DBH had a direct positive effect on δ13C content and a direct negative effect on δ18O, and there was a direct positive effect of light environment on δ18O. In larger plants, there was a direct negative effect of light environment on δ13C and a direct positive effect of DBH on light environment, as well as a negative effect of soil nitrogen content on leaf nitrogen. In smaller plants, DBH was the most important factor influencing δ13C, followed by δ18O and soil moisture, with light and soil pH showing minimal influence. In larger plants, light environment influenced δ13C the most, followed by soil nitrogen content and soil moisture content, with leaf nitrogen and DBH contributing little. The results suggest that water use efficiency strategies of P. koraiensis vary according to growth stage, and the effects of abiotic factors and functional traits vary at different growth stages.

[Water use characteristics and the mechanism of water uptake in two typical sand-fixing species in Mu Us sandy land]. / æ¯›ä¹Œç´ å ¸åž‹å›ºæ²™æ¤ç‰©çš„æ°´åˆ†åˆ©ç”¨ç‰¹æ€§ä¸Žå¸æ°´æœºåˆ¶.

Understanding water absorption mechanisms of sand-fixing plants is important for the rational establishment of plant community structures, thereby providing a scientific basis for desertification control and the efficient utilization of water resources in sandy areas. Based on the hydrogen and oxygen isotopic compositions of precipi-tation, soil water, xylem water, and groundwater, coupled with soil water-heat dynamics, annual water consumption characteristics of vegetation, using the multi-source linear mixing model (IsoSource), we analyzed the differences in water sources between Salix psammophila and Artemisia ordosica, during winter and the growing season. We further examined the effects of groundwater depth (2 m and 10 m), soil freezing-thawing, and drought on their water utilization to elucidate water absorption mechanisms of those species. The results showed that: 1) During soil freezing-thawing period (January to March), S. psammophila mainly utilized soil water in 60-120 cm depths below the frozen layer (69.1%). In the green-up season (April and May), soil water from the 0-60 cm layers could satisfy the water demand of S. psammophila (30.9%-87.6%). During the dry period of the growing season (June), it predominantly utilized soil water at the depth of 120-160 cm (27.4%-40.8%). Over the rainy season (July and September), soil water in 0-60 cm depths provided 59.8%-67.9% of the total water required. A. ordosica, with shallow roots, could not utilize soil water after complete freezing of root zone but could overwinter by storing water in rhizomes during autumn. During the growing season, it primarily relied on 0-40 cm soil layer (23.4%-86.8%). During the dry period, it mainly utilized soil water from 40-80 cm and 80-160 cm soil layers, with utilization rates of 14.6%-74.4% and 21.8%-78.2%, respectively. 2) With decreasing groundwater depth, vegetation shifted its water absorption depth upward, with water source of S. psammophila transitioning from 120-160 cm to 60-160 cm layers, while A. ordosica shifted water absorption depth from 80-160 cm to 0-40 cm. S. psammophila's utilization of soil water is influenced by transpiration, adopting an "on-demand" approach to achieve a balance between water supply and energy conservation, whereas A. ordosica tends to utilize shallow soil water, exhibiting a higher depen-dence on water sources from a single soil layer.

Impacts of elevated CO2 levels and temperature on photosynthesis and stomatal closure along an altitudinal gradient are counteracted by the rising atmospheric vapor pressure deficit.

The efficiency of water use in plants, a critical ecophysiological parameter closely related to water and carbon cycles, is essential for understanding the interactions between plants and their environment. This study investigates the effects of ongoing climate change and increasing atmospheric CO2 concentration on intrinsic (stomata-based; iWUE) and evaporative (transpiration-based; eWUE) water use efficiency in oak trees along a naturally small altitudinal gradient (130-630 m a.s.l.) of Vihorlat Mountains (eastern Slovakia, Central Europe). To assess changes in iWUE and eWUE values over the past 60 years (1961-2020), stable carbon isotope ratios in latewood cellulose (δ13Ccell) of annually resolved tree rings were analyzed. Such an approach was sensitive enough to distinguish tree responses to growth environments at different altitudes. Our findings revealed a rising trend in iWUE, particularly in oak trees at low and middle altitudes. However, this increase was negligible at high altitudes. Warmer and drier conditions at lower altitudes likely led to significant stomatal closure and enhanced efficiency in photosynthetic CO2 uptake due to rising CO2 concentration. Conversely, the increasing intracellular-to-ambient CO2 ratio (Ci/Ca) at higher altitudes indicated lower efficiency in photosynthetic CO2 uptake. In contrast to iWUE, eWUE showed no increasing trends over the last 60 years. This suggests that the positive impacts of elevated CO2 concentrations and temperature on photosynthesis and stomatal closure are counteracted by the rising atmospheric vapor pressure deficit (VPD). These differences underscore the importance of the correct interpretation of stomata-based and transpiration-based WUEs and highlight the necessity of atmospheric VPD correction when applying tree-ring δ13C-derived WUE at ecosystem and global levels.

The effect of shrub cover on conifer water-use patterns, growth and response to precipitation variability in the southern Sierra Nevada.

As wildfires increase in size and severity, large areas of forest are undergoing substantial increases in shrub cover. In forests where water is the limiting resource, the partitioning of soil water between shrubs and young trees may determine how shrubs affect tree growth and water-stress. Here we evaluated juvenile trees (average age = 32 years) of two dominant conifer species in the southern Sierra Nevada of California (Abies concolor (white fir) and Pinus jeffreyi (Jeffrey pine)) growing in the presence or absence of shrubs. The two shrub species included Arctostaphylos patula and Ceanothus cordulatus, a nitrogen-fixing species. We analyzed the δ2H and δ18O values of xylem water for both tree and shrub species to assess how shrub cover affects the water-uptake patterns of conifers and whether there is niche partitioning between trees and shrubs. We found that growing near shrubs did not have a significant effect on the water source dynamics of either tree species, with similar source water contributions calculated for conifers growing in both the presence and absence of shrubs. Using a tree-ring analysis of growth and δ13C from 2016 to 2021, a period of high precipitation variability, we found that shrub cover had a positive effect on tree growth while decreasing carbon discrimination, which may be due to increased nitrogen availability from Ceanothus cordulatus. Overall, our results suggest that growing in the presence of shrubs does not alter the water uptake patterns of white fir and Jeffrey pine and instead may have a positive effect on the growth rates of these species during both wet and dry years.

Grass veins are leaky pipes: vessel widening in grass leaves explain variation in stomatal conductance and vessel diameter among species.

The widening of xylem vessels from tip to base of trees is an adaptation to minimize the hydraulic resistance of a long pathway. Given that parallel veins of monocot leaves do not branch hierarchically, vessels should also widen basipetally but, in addition to minimizing resistance, should also account for water volume lost to transpiration since they supply water to the lamina along their lengths, that is 'leakiness'. We measured photosynthesis, stomatal conductance, and vessel diameter at five locations along each leaf of five perennial grass species. We found that the rate of conduit widening in grass leaves was larger than the widening exponent required to minimize pathlength resistance (0.35 vs c. 0.22). Furthermore, variation in the widening exponent among species was positively correlated with maximal stomatal conductance (r2 = 0.20) and net CO2 assimilation (r2 = 0.45). These results suggest that faster rates of conduit widening (> 0.22) were associated with higher rates of water loss. Taken together, our results show that the widening exponent is linked to plant function in grass leaves and that natural selection has favored parallel vein networks that are constructed to meet transpiration requirements while minimizing hydraulic resistance within grass blades.

Water use strategies of Nitraria tangutorum in the lake-basin region of the Badain Jaran Desert.

Information regarding plant water-use strategies is essential for understanding the hydrological processes and plant survival adaptation mechanisms in desert lake basin regions. To examine the water use strategies of plants in desert lake basin areas, water uptake patterns, water use efficiency, and water potential of Nitraria tangutorum were investigated at different distances from the lake duringhe growing seasons in the lake basin regions of the Badain Jaran Desert. The results indicate that N. tangutorum primarily absorbed groundwater in May (63.8%) and August (53.5%), relied on deep soil water in June (75.1%), and uniformly absorbed soil water from different layers in July. These observations could be explained by periodic fluctuations in the groundwater level and the consequent decrease in soil water availability, as well as plant root adjustments. As soil water availability decreases, N. tangutorum adapts to water variation by increasing its water use efficiency (WUE) and reducing its leaf water potential (Ψ). With intensified water stress, N. tangutorum gradually shifted from adventurous anisohydric regulation to conservative isohydric regulation. Thus, N. tangutorum responds to diverse degrees of environmental changes by altering its water-use strategy. A better understanding of the adaptive water use strategies developed by desert plants under varying water availability conditions provides insight into the diversity of species' reactions to long-term drought and quantifies the hydrological cycle of desert ecosystems against the background of worldwide climate warming.

In situ characterisation of whole-plant stomatal responses to VPD using leaf optical dendrometry.

Vapour pressure deficit (VPD) plays a crucial role in regulating plant carbon and water fluxes due to its influence on stomatal behaviour and transpiration. Yet, characterising stomatal responses of the whole plant to VPD remains challenging due to methodological limitations. Here, we develop a novel method for in situ assessment of whole-plant stomatal responses (gc ) to VPD in the herbaceous plant Tanacetum cinerariifolium. To do this, we examine the relationship between daytime VPD and the corresponding soil-stem water potential gradient (ΔΨ) monitored using the optical dendrometry in well-hydrated plants under nonlimiting light in both glasshouse and field conditions. In glasshouse plants, ΔΨ increased proportionally with the VPD up to a threshold of 1.53 kPa, beyond which the slope decreased, suggesting a two-phase response in gc . This pattern aligned with corresponding gravimetrically measured gc behaviour, which also showed a decline when VPD exceeded a similar threshold. This response was then compared with that of field plants monitored using the optical dendrometry technique over a growing season under naturally variable VPD conditions and nonlimiting light and water supply. Field plants exhibited a similar threshold-type response to VPD but were more sensitive than glasshouse individuals with a VPD threshold of 0.74 kPa. The results showed that whole-plant gc responses to VPD can be characterised optically in T. cinerariifolium, introducing a new tool for the monitoring and characterisation of stomatal behaviour in situ.

Climate warming negatively affects plant water-use efficiency in a seasonal hydroperiod wetland.

Climate warming has substantial influences on plant water-use efficiency (PWUE), which is defined as the ratio of plant CO2 uptake to water loss and is central to the cycles of carbon and water in ecosystems. However, it remains uncertain how does climate warming affect PWUE in wetland ecosystems, especially those with seasonally alternating water availability during the growing season. In this study, we used a continuous 10-year (2011-2020) eddy covariance (EC) dataset from a seasonal hydroperiod wetland coupled with a 15-year (2003-2017) satellite-based dataset (called PML-V2) and an in situ warming experiment to examine the climate warming impacts on wetland PWUE. The 10-year EC observational results revealed that rising temperatures had significant negative impacts on the interannual variations in wetland PWUE, and increased transpiration (Et) rather than changes in gross primary productivity (GPP) dominated these negative impacts. Furthermore, the 15-year satellite-based evidence confirmed that, in the study region, climate warming had significant negative consequences for the interannual variations in wetland PWUE by enhancing wetland Et. Lastly, at the leaf-scale, the light response curves of leaf photosynthesis, leaf Et, and leaf-scale PWUE indicated that wetland plants need to consume more water during the photosynthesis process under warmer conditions. These findings provide a fresh perspective on how climate warming influences carbon and water cycles in wetland ecosystems.

The influences of stomatal size and density on rice abiotic stress resilience.

A warming climate coupled with reductions in water availability and rising salinity are increasingly affecting rice (Oryza sativa) yields. Elevated temperatures combined with vapour pressure deficit (VPD) rises are causing stomatal closure, further reducing plant productivity and cooling. It is unclear what stomatal size (SS) and stomatal density (SD) will best suit all these environmental extremes. To understand how stomatal differences contribute to rice abiotic stress resilience, we screened the stomatal characteristics of 72 traditionally bred varieties. We found significant variation in SS, SD and calculated anatomical maximal stomatal conductance (gsmax ) but did not identify any varieties with SD and gsmax as low as transgenic OsEPF1oe plants. Traditionally bred varieties with high SD and small SS (resulting in higher gsmax ) typically had lower biomasses, and these plants were more resilient to drought than low SD and large SS plants, which were physically larger. None of the varieties assessed were as resilient to drought or salinity as low SD OsEPF1oe transgenic plants. High SD and small SS rice displayed faster stomatal closure during increasing temperature and VPD, but photosynthesis and plant cooling were reduced. Compromises will be required when choosing rice SS and SD to tackle multiple future environmental stresses.

Consistent responses to moisture stress despite diverse growth forms within mountain fynbos communities.

Understanding climate change impacts on the Cape Floristic Region requires improved knowledge of plant physiological responses to the environment. Studies examining physiological responses of mountain fynbos have consisted of campaign-based measurements, capturing snapshots of plant water relations and photosynthesis. We examine conclusions drawn from prior studies by tracking in situ physiological responses of three species, representing dominant growth forms (proteoid, ericoid, restioid), over 2 years using miniature continuous sap flow technology, long-term observations of leaf/culm water potential and gas exchange, and xylem vulnerability to embolism. We observed considerable inter-specific variation in the timing and extent of seasonal declines in productivity. Shallow-rooted Erica monsoniana exhibited steep within-season declines in sap flow and water potentials, and pronounced inter-annual variability in total daily sap flux (Js). Protea repens showed steady reductions in Js across both years, despite deeper roots and less negative water potentials. Cannomois congesta-a shallow-rooted restioid-was least negatively impacted. Following rehydrating rain at the end of summer, gas exchange recovery was lower in the drier year compared with the normal year, but did not differ between species. Loss of function in the drier year was partially accounted for by loss of xylem transport capacity in Erica and Cannomois, but not Protea. Hitherto unseen water use patterns, including inter-annual variability of gas exchange associated with contrasting water uptake properties, reveal that species use different mechanisms to cope with summer dry periods. Revealing physiological responses of key growth forms enhances predictions of plant function within mountain fynbos under future conditions.

Stable isotope approaches and opportunities for improving plant conservation.

Successful conservation of threatened species and ecosystems in a rapidly changing world requires scientifically sound decision-making tools that are readily accessible to conservation practitioners. Physiological applications that examine how plants and animals interact with their environment are now widely used when planning, implementing and monitoring conservation. Among these tools, stable-isotope physiology is a potentially powerful, yet under-utilized cornerstone of current and future conservation efforts of threatened and endangered plants. We review the underlying concepts and theory of stable-isotope physiology and describe how stable-isotope applications can support plant conservation. We focus on stable isotopes of carbon, hydrogen, oxygen and nitrogen to address plant ecophysiological responses to changing environmental conditions across temporal scales from hours to centuries. We review examples from a broad range of plant taxa, life forms and habitats and provide specific examples where stable-isotope analysis can directly improve conservation, in part by helping identify resilient, locally adapted genotypes or populations. Our review aims to provide a guide for practitioners to easily access and evaluate the information that can be derived from stable-isotope signatures, their limitations and how stable isotopes can improve conservation efforts.

Linking tree water use efficiency with calcium and precipitation.

Water use efficiency (WUE) is a key physiological trait in studying plant carbon and water relations. However, the determinants of WUE across a large geographical scale are not always clear, limiting our capacity to predict WUE in response to future global climate change. We propose that tree WUE is influenced by calcium (Ca) availability and precipitation. In addition, although it is well-known that transpiration is the major driving force for passive nutrient uptake, the linkage between these two processes has not been well-established. Because Ca uptake is an apoplastic and passive process that purely relies on transpiration, and there is no translocation once assimilated, we further developed a theoretical model to quantify the relationship between tree Ca accumulation and WUE using soil-to-plant calcium ratio (SCa/BCa) and tree WUE derived from δ13C. We tested our theoretical model and predicted relationships using three common tree species across their native habitats in Northern China, spanning 2300 km and a controlled greenhouse experiment with soil Ca concentrations manipulated. We found that tree WUE was negatively related to precipitation of the growing season (GSP) and positively with soil Ca. A multiple regression model and a path analysis suggested a higher contribution of soil Ca to WUE than GSP. As predicted by our theoretical model, we found a positive relationship between WUE and SCa/BCa across their distribution ranges in all three tree species and in the controlled experiment for one selected species. This relationship suggests a tight coupling between water and Ca uptake and the potential use of SCa/BCa to indicate WUE. A negative relationship between SCa/BCa and GSP also suggests a possible decrease in tree Ca accumulation efficiency in a drier future in Northern China.

Higher leaf nitrogen content is linked to tighter stomatal regulation of transpiration and more efficient water use across dryland trees.

The least-cost economic theory of photosynthesis shows that water and nitrogen are mutually substitutable resources to achieve a given carbon gain. However, vegetation in the Sahel has to cope with the dual challenge imposed by drought and nutrient-poor soils. We addressed how variation in leaf nitrogen per area (Narea ) modulates leaf oxygen and carbon isotopic composition (δ18 O, δ13 C), as proxies of stomatal conductance and water-use efficiency, across 34 Sahelian woody species. Dryland species exhibited diverging leaf δ18 O and δ13 C values, indicating large interspecific variation in time-integrated stomatal conductance and water-use efficiency. Structural equation modeling revealed that leaf Narea is a pivotal trait linked to multiple water-use traits. Leaf Narea was positively linked to both δ18 O and δ13 C, suggesting higher carboxylation capacity and tighter stomatal regulation of transpiration in N-rich species, which allows them to achieve higher water-use efficiency and more conservative water use. These adaptations represent a key physiological advantage of N-rich species, such as legumes, that could contribute to their dominance across many dryland regions. This is the first report of a robust mechanistic link between leaf Narea and δ18 O in dryland vegetation that is consistent with core principles of plant physiology.

Hydraulic architecture with high root-resistance fraction contributes to efficient carbon gain of plants in temperate habitats.

PREMISE: The hydraulic architecture in the leaves, stems and roots of plants constrains water transport and carbon gain through stomatal limitation to CO2 absorption. Because roots are the main bottleneck in water transport for a range of plant species, we assessed the ecophysiological mechanism and importance of a high fraction of root hydraulic resistance in woody and herbaceous species. METHODS: Biomass partitioning and hydraulic conductance of leaves, stems, and roots of Japanese knotweed (Fallopia japonica, a perennial herb) and Japanese zelkova (Zelkova serrata, a deciduous tall tree) were measured. Theoretical analyses were used to examine whether the measured hydraulic architecture and biomass partitioning maximized the plant photosynthetic rate (the product of leaf area and photosynthetic rate per leaf area). RESULTS: Root hydraulic resistance accounted for 83% and 68% of the total plant resistance for Japanese knotweed and Japanese zelkova, respectively. Comparisons of hydraulic and biomass partitioning revealed that high root-resistance fractions were attributable to low biomass partitioning to root organs rather than high mass-specific root conductance. The measured partitioning of hydraulic resistance closely corresponded to the predicted optimal partitioning, maximizing the plant photosynthetic rate for the two species. The high fraction of root resistance was predicted to be optimal with variations in air humidity and soil water potential. CONCLUSIONS: These results suggest that the hydraulic architecture of plants growing in mesic and fertile habitats not only results in high root resistance due to small biomass partitioning to root organs, but contributes to efficient carbon gain.

[Distribution of hydrogen and oxygen stable isotope of water in soil-plant-atmosphere continuum (SPAC)system of a typical forest area]. / å ¸åž‹æž—åŒºæ°´åˆ†æ°¢æ°§ç¨³å®šåŒä½ç´ åœ¨åœŸå£¤-植物-大气连续体中的分布特征.

Water cycle in the soil-plant-atmosphere continuum (SPAC) is an important research topic in hydrology and ecology. The differences in the composition of hydrogen and oxygen stable isotopes in different water bodies can indicate water cycle process. Based on the measurements of isotopic compositions in precipitation, soil water, and plant water, we analyzed water isotope evolution in a SPAC system located in a subtropical evergreen broad-leaved forest in Chengdu Plain. The different interface processes of regional water cycle was revealed. The results showed that the local meteoric water line (LMWL) equation was δD=7.13 δ18O+2.35 (R2=0.99), and the soil evaporation line (SEL) equation was δD=6.98δ18O-0.32 (R2=0.92). In the water transportation process of precipitation-soil water-plant water, hydrogen and oxygen isotopes were gradually enriched. The δ18O in water of the surface soil layer (0-35 cm) was sensitive to precipitation input, as it was directly affected by precipitation. In contrast, the δ18O in water of the middle and deep layers (35-100 cm) was relatively stable. The isotopes of plant xylem water were slightly more enriched than those of soil water, indicating the possibility of slight evaporation or transpiration through phloem or bark in plant water transportation. The estimation of plant water intake from different soil layers was performed by direct correlation method. Cinnamomum camphora mainly used water from the middle layer, Broussonetia papyrifera mainly used that from the surface layer, and Parathelypteris glanduligera tended to use surface soil water and precipitation intercepted by plants because of the shallow root system. Compared with P. glanduligera, C. camphora and B. papyrifera experienced stronger water evaporation and the isotopes were influenced by more intense dynamic fractionation.

Water use efficiency, grain yield, and economic benefits of common beans (Phaseolus vulgaris L.) under four soil tillage systems in Mukono District, Uganda.

With the increasing climate change impacts and variabilities, water is becoming a limiting factor for rainfed crop production in Uganda. Conservation tillage practices could improve soil and water conservation in croplands. Field experiments were conducted for three consecutive seasons from April 2019 to June 2020. The experiments evaluated the effect of soil tillage treatments on soil water storage, water use efficiency, grain yield, and economic benefits of the common beans (Phaseolus vulgaris L.) in two sub-counties of Mukono District, central Uganda. The soil tillage treatments were: no-tillage, stubble-mulching, deep tillage, and conventional tillage. The no-tillage and stubble-mulching improved soil water storage by 46 and 45%, respectively, compared with the conventional tillage in the 0-100 cm soil depth over the 14 months. Soil tillage treatments significantly (p < 0.05) affected the water use efficiency, with water use efficiency values generally higher under no-tillage and stubble-mulching than under deep tillage and conventional tillage treatments. The grain yield was highest under no-tillage and stubble-mulching than deep tillage and conventional tillage treatments, with over 5, 38, and 43% higher grain yield under no-tillage than under stubble-mulching, deep tillage, and conventional tillage treatments, respectively. Although no-tillage and stubble-mulching improved soil water storage and grain yield, seasonal precipitation distribution had a greater influence on the final grain yield, soil water storage, and water use efficiency. The net profit was 3 and 5 times higher under no-tillage than under conventional tillage and deep tillage treatments, respectively. The overall results showed that no-tillage and stubble-mulching were the optimum tillage treatments for increasing soil water storage and common bean yield, enhancing water use efficiency, and improving economic returns in central Uganda.

Water uptake depth is coordinated with leaf water potential, water-use efficiency and drought vulnerability in karst vegetation.

Root access to bedrock water storage or groundwater is an important trait allowing plant survival in seasonally dry environments. However, the degree of coordination between water uptake depth, leaf-level water-use efficiency (WUEi) and water potential in drought-prone plant communities is not well understood. We conducted a 135-d rainfall exclusion experiment in a subtropical karst ecosystem with thin skeletal soils to evaluate the responses of 11 co-occurring woody species of contrasting life forms and leaf habits to a severe drought during the wet growing season. Marked differences in xylem water isotopic composition during drought revealed distinct ecohydrological niche separation among species. The contrasting behaviour of leaf water potential in coexisting species during drought was largely explained by differences in root access to deeper, temporally stable water sources. Smaller-diameter species with shallower water uptake, more negative water potentials and lower WUEi showed extensive drought-induced canopy defoliation and/or mortality. By contrast, larger-diameter species with deeper water uptake, higher leaf-level WUEi and more isohydric behaviour survived drought with only moderate canopy defoliation. Severe water limitation imposes strong environmental filtering and/or selective pressures resulting in tight coordination between tree diameter, water uptake depth, iso/anisohydric behaviour, WUEi and drought vulnerability in karst plant communities.

Mesophyll CO2 conductance and leakiness are not responsive to short- and long-term soil water limitations in the C4 plant Sorghum bicolor.

Breeding economically important C4 crops for enhanced whole-plant water-use efficiency (WUEplant ) is needed for sustainable agriculture. WUEplant is a complex trait and an efficient phenotyping method that reports on components of WUEplant , such as intrinsic water-use efficiency (WUEi , the rate of leaf CO2 assimilation relative to water loss via stomatal conductance), is needed. In C4 plants, theoretical models suggest that leaf carbon isotope composition (δ13 C), when the efficiency of the CO2 -concentrating mechanism (leakiness, ϕ) remains constant, can be used to screen for WUEi . The limited information about how ϕ responds to water limitations confines the application of δ13 C for WUEi screening of C4 crops. The current research aimed to test the response of ϕ to short- or long-term moderate water limitations, and the relationship of δ13 C with WUEi and WUEplant , by addressing potential mesophyll CO2 conductance (gm ) and biochemical limitations in the C4 plant Sorghum bicolor. We demonstrate that gm and ϕ are not responsive to short- or long-term water limitations. Additionally, δ13 C was not correlated with gas-exchange estimates of WUEi under short- and long-term water limitations, but showed a significant negative relationship with WUEplant . The observed association between the δ13 C and WUEplant suggests an intrinsic link of δ13 C with WUEi in this C4 plant, and can potentially be used as a screening tool for WUEplant in sorghum.

Global response patterns of plant photosynthesis to nitrogen addition: A meta-analysis.

A mechanistic understanding of plant photosynthetic response is needed to reliably predict changes in terrestrial carbon (C) gain under conditions of chronically elevated atmospheric nitrogen (N) deposition. Here, using 2,683 observations from 240 journal articles, we conducted a global meta-analysis to reveal effects of N addition on 14 photosynthesis-related traits and affecting moderators. We found that across 320 terrestrial plant species, leaf N was enhanced comparably on mass basis (Nmass , +18.4%) and area basis (Narea , +14.3%), with no changes in specific leaf area or leaf mass per area. Total leaf area (TLA) was increased significantly, as indicated by the increases in total leaf biomass (+46.5%), leaf area per plant (+29.7%), and leaf area index (LAI, +24.4%). To a lesser extent than for TLA, N addition significantly enhanced leaf photosynthetic rate per area (Aarea , +12.6%), stomatal conductance (gs , +7.5%), and transpiration rate (E, +10.5%). The responses of Aarea were positively related with that of gs , with no changes in instantaneous water-use efficiency and only slight increases in long-term water-use efficiency (+2.5%) inferred from 13 C composition. The responses of traits depended on biological, experimental, and environmental moderators. As experimental duration and N load increased, the responses of LAI and Aarea diminished while that of E increased significantly. The observed patterns of increases in both TLA and E indicate that N deposition will increase the amount of water used by plants. Taken together, N deposition will enhance gross photosynthetic C gain of the terrestrial plants while increasing their water loss to the atmosphere, but the effects on C gain might diminish over time and that on plant water use would be amplified if N deposition persists.