Precipitation‐use efficiency along a 4500‐km grassland transect

Aims Clarifying the spatiotemporal variations in precipitation‐use efficiency (PUE), the ratio of vegetation above‐ground productivity to annual precipitation, will advance our understanding of how ecosystems' carbon and water cycles respond to climate change. Our goal is to investigate the variations in PUE at both regional and site scales along a 4500‐km climate‐related grassland transect. Location The Inner Mongolian Plateau in northern China and the Qinghai‐Tibetan Plateau. Methods We collected data on 580 sites from four data sources. The data were acquired through field surveys and long‐term in situ observations. We investigated the relationships between precipitation and PUE at both regional and site scales, and we evaluated the effects of the main biotic and climatic factors on PUE at both spatial scales. Results PUE decreased with decreasing mean annual precipitation (MAP), except for a slight rise toward the dry end of the gradient. The maximum PUE showed large site‐to‐site variation along the transect. Vegetation cover significantly affected the spatial variations in PUE, and this probably accounts for the positive relationship between PUE and MAP. However, there was no significant relationship between inter‐annual variations in precipitation or vegetation cover and PUE within given ecosystems along the transect. Conclusions The findings of this research contradict the prevailing view that a convergent maximum PUE exists among diverse ecosystems, as presented in previous reports. Our findings also suggest the action of distinct mechanisms in controlling PUE at different spatial scales. We propose the use of a conceptual model for predicting vegetation productivity at continental and global scales with a sigmoid function, which illustrates an increasing PUE with MAP in arid regions. Our approach may represent an improvement over use of the popular Miami model.     

Radial growth response of four dominant boreal tree species to climate along a latitudinal gradient in the eastern Canadian boreal forest

To address the central question of how climate change influences tree growth within the context of global warming, we used dendroclimatological analysis to understand the reactions of four major boreal tree species –Populus tremuloides, Betula papyrifera, Picea mariana, and Pinus banksiana– to climatic variations along a broad latitudinal gradient from 46 to 54°N in the eastern Canadian boreal forest. Tree‐ring chronologies from 34 forested stands distributed at a 1° interval were built, transformed into principal components (PCs), and analyzed through bootstrapped correlation analysis over the period 1950–2003 to identify climate factors limiting the radial growth and the detailed radial growth–climate association along the gradient. All species taken together, previous summer temperature (negative influences), and current January and March–April temperatures (positive influences) showed the most consistent relationships with radial growth across the gradient. Combined with the identified species/site‐specific climate factors, our study suggested that moisture conditions during the year before radial growth played a dominant role in positively regulating P. tremuloides growth, whereas January temperature and growing season moisture conditions positively impacted growth of B. papyrifera. Both P. mariana and P. banksiana were positively affected by the current‐year winter and spring or whole growing season temperatures over the entire range of our corridor. Owing to the impacts of different climate factors on growth, these boreal species showed inconsistent responsiveness to recent warming at the transition zone, where B. papyrifera, P. mariana, and P. banksiana would be the most responsive species, whereas P. tremuloides might be the least. Under continued warming, B. papyrifera stands located north of 49°N, P. tremuloides at northern latitudes, and P. mariana and P. banksiana stands located north of 47°N might benefit from warming winter and spring temperatures to enhance their radial growth in the coming decades, whereas other southern stands might be decreasing in radial growth.     

Differences in leaf gas exchange strategies explain Quercus rubra and Liriodendron tulipifera intrinsic water use efficiency responses to air pollution and climate change

Trees continuously regulate leaf physiology to acquire CO₂ while simultaneously avoiding excessive water loss. The balance between these two processes, or water use efficiency (WUE), is fundamentally important to understanding changes in carbon uptake and transpiration from the leaf to the globe under environmental change. While increasing atmospheric CO₂ (iCO₂) is known to increase tree intrinsic water use efficiency (iWUE), less clear are the additional impacts of climate and acidic air pollution and how they vary by tree species. Here, we couple annually resolved long-term records of tree-ring carbon isotope signatures with leaf physiological measurements of Quercus rubra (Quru) and Liriodendron tulipifera (Litu) at four study locations spanning nearly 100 km in the eastern United States to reconstruct historical iWUE, net photosynthesis (A_net), and stomatal conductance to water (g_s) since 1940. We first show 16%–25% increases in tree iWUE since the mid-20th century, primarily driven by iCO₂, but also document the individual and interactive effects of nitrogen (NO_x) and sulfur (SO₂) air pollution overwhelming climate. We find evidence for Quru leaf gas exchange being less tightly regulated than Litu through an analysis of isotope-derived leaf internal CO₂ (C_i), particularly in wetter, recent years. Modeled estimates of seasonally integrated A_net and g_s revealed a 43%–50% stimulation of A_net was responsible for increasing iWUE in both tree species throughout 79%–86% of the chronologies with reductions in g_s attributable to the remaining 14%–21%, building upon a growing body of literature documenting stimulated A_net overwhelming reductions in g_s as a primary mechanism of increasing iWUE of trees. Finally, our results underscore the importance of considering air pollution, which remains a major environmental issue in many areas of the world, alongside climate in the interpretation of leaf physiology derived from tree rings.     

Interactions between CO2 enhancement and N addition on net primary productivity and water‐use efficiency in a mesocosm with multiple subtropical tree species

Carbon dioxide (CO₂) enhancement (eCO₂) and N addition (aN) have been shown to increase net primary production (NPP) and to affect water‐use efficiency (WUE) for many temperate ecosystems, but few studies have been made on subtropical tree species. This study compared the responses of NPP and WUE from a mesocosm composing five subtropical tree species to eCO₂ (700 ppm), aN (10 g N m^?2 yr^?1) and eCO₂ × aN using open‐top chambers. Our results showed that mean annual ecosystem NPP did not changed significantly under eCO₂, increased by 56% under aN and 64% under eCO₂ × aN. Ecosystem WUE increased by 14%, 55%, and 61% under eCO₂, aN and eCO₂ × aN, respectively. We found that the observed responses of ecosystem WUE were largely driven by the responses of ecosystem NPP. Statistical analysis showed that there was no significant interactions between eCO₂ and aN on ecosystem NPP (P = 0.731) or WUE (P = 0.442). Our results showed that increasing N deposition was likely to have much stronger effects on ecosystem NPP and WUE than increasing CO₂ concentration for the subtropical forests. However, different tree species responded quite differently. aN significantly increased annual NPP of the fast‐growing species (Schima superba). Nitrogen‐fixing species (Ormosia pinnata) grew significantly faster only under eCO₂ × aN. eCO₂ had no effects on annual NPP of those two species but significantly increased annual NPP of other two species (Castanopsis hystrix and Acmena acuminatissima). Differential responses of the NPP among different tree species to eCO₂ and aN will likely have significant implications on the species composition of subtropical forests under future global change.     

Seasonal and within‐canopy variation in shoot‐scale resource‐use efficiency trade‐offs in a Norway spruce stand

Previous leaf‐scale studies of carbon assimilation describe short‐term resource‐use efficiency (RUE) trade‐offs where high use efficiency of one resource requires low RUE of another. However, varying resource availabilities may cause long‐term RUE trade‐offs to differ from the short‐term patterns. This may have important implications for understanding canopy‐scale resource use and allocation. We used continuous gas exchange measurements collected at five levels within a Norway spruce, Picea abies (L.) karst., canopy over 3 years to assess seasonal differences in the interactions between shoot‐scale resource availability (light, water and nitrogen), net photosynthesis (A_n) and the use efficiencies of light (LUE), water (WUE) and nitrogen (NUE) for carbon assimilation. The continuous data set was used to develop and evaluate multiple regression models for predicting monthly shoot‐scale A_n. These models showed that shoot‐scale A_n was strongly dependent on light availability and was generally well described with simple one‐ or two‐parameter models. WUE peaked in spring, NUE in summer and LUE in autumn. However, the relative importance of LUE for carbon assimilation increased with canopy depth at all times. Our results suggest that accounting for seasonal and within‐canopy trade‐offs may be important for RUE‐based modelling of canopy carbon uptake.     

Long‐term increases in intrinsic water‐use efficiency do not lead to increased stem growth in a tropical monsoon forest in western Thailand

Rising atmospheric carbon dioxide [CO₂] can accelerate tree growth by stimulating photosynthesis and increasing intrinsic water‐use efficiency (iWUE). Little evidence exists, however, for the long‐term growth and gas‐exchange responses of mature trees in tropical forests to the combined effects of rising [CO₂] and other global changes such as warming. Using tree rings and stable isotopes of carbon and oxygen, we investigated long‐term trends in the iWUE and stem growth (basal area increment, BAI) of three canopy tree species in a tropical monsoon forest in western Thailand (Chukrasia tabularis, Melia azedarach, and Toona ciliata). To do this, we modelled the contribution of ontogenetic effects (tree diameter or age) and calendar year to variation in iWUE, oxygen isotopes, and BAI using mixed‐effects models. Although iWUE increased significantly with both tree diameter and calendar year in all species, BAI at a given tree diameter was lower in more recent years. For one species, C. tabularis, differences in crown dominance significantly influence stable isotopes and growth. Tree ring Δ¹⁸O increased with calendar year in all species, suggesting that increasing iWUE may have been driven by relatively greater reductions in stomatal conductance – leading to enrichment in Δ¹⁸O – than increases in photosynthetic capacity. Plausible explanations for the observed declines in growth include water stress resulting from rising temperatures and El Niño events, increased respiration, changes in allocation, or more likely, a combination of these factors.     

Relationship between photosynthetic phosphorus‐use efficiency and foliar phosphorus fractions in tropical tree species

How plants develop adaptive strategies to efficiently use nutrients on infertile soils is an important topic in plant ecology. It has been suggested that, with decreasing phosphorus (P) availability, plants increase photosynthetic P‐use efficiency (PPUE) (i.e., the ratio of instantaneous photosynthetic carbon assimilation rate per unit foliar P). However, the mechanism to increase PPUE remains unclear. In this study, we tested whether high PPUE is explained by an optimized allocation of P in cells among P‐containing biochemical compounds (i.e., foliar P fractions). We investigated the relationships among mass‐based photosynthetic carbon assimilation rate (A_mass), PPUE, total foliar P concentration, and foliar P fractions in 10 tree species in two tropical montane rain forests with differing soil P availability (five species on sedimentary soils and five species on P‐poorer ultrabasic serpentine soils) on Mount Kinabalu, Borneo. We chemically fractionated foliar P into the following four fractions: metabolic P, lipid P, nucleic acid P, and residual P. A_mass was positively correlated with the concentrations of total foliar P and of metabolic P across 10 tree species. Mean A_mass and mean concentrations of total foliar P and of each foliar P fraction were lower on the P‐poorer ultrabasic serpentine soils than on the sedimentary soils. There was a negative relationship between the proportion of metabolic P per total P and the proportion of lipid P per total P. PPUE was positively correlated with the ratio of metabolic P to lipid P. High PPUE is explained by the net effect of a relatively greater investment of P into P‐containing metabolites and a relatively lesser investment into phospholipids in addition to generally reduced concentrations of all P fractions. We conclude that plants optimize the allocation of P among foliar P fractions for maintaining their productivity and growth and for reducing demand for P as their adaptation to P‐poor soils.     

Arbuscular mycorrhiza influences carbon‐use efficiency and grain yield of wheat grown under pre‐ and post‐anthesis salinity stress

• Soil salinity severely affects and constrains crop production worldwide. Salinity causes osmotic and ionic stress, inhibiting gas exchange and photosynthesis, ultimately impairing plant growth and development. Arbuscular mycorrhiza (AM) have been shown to maintain light and carbon use efficiency under stress, possibly providing a tool to improve salinity tolerance of the host plants. Thus, it was hypothesized that AM will contribute to improved growth and yield under stress conditions.
• Wheat plants (Triticum aestivum L.) were grown with (AMF+) or without (AMF?) arbuscular mycorrhizal fungi (AMF) inoculation. Plants were subjected to salinity stress (200 mm NaCl) either at pre‐ or post‐anthesis or at both stages. Growth and yield components, leaf chlorophyll content as well as gas exchange parameters and AMF colonization were analysed.
• AM plants exhibited a higher rate of net photosynthesis and stomatal conductance and lower intrinsic water use efficiency. Furthermore, AM wheat plants subjected to salinity stress at both pre‐anthesis and post‐anthesis maintained higher grain yield than non‐AM salinity‐stressed plants.
• These results suggest that AMF inoculation mitigates the negative effects of salinity stress by influencing carbon use efficiency and maintaining higher grain yield under stress.

     

Differences in the soil microbial community and carbon‐use efficiency following development of Vochysia guatemalensis tree plantations in unproductive pastures in Costa Rica

This study shows that Vochysia guatemalensis tree plantations were associated with enhanced soil biotic and abiotic characteristics in previously cleared forests in the northern zone of Costa Rica, suggesting the possible use of this practice as a restoration strategy for local land owners. Soil samples from a primary forest, secondary forest, and a 13‐year‐old plantation of V. guatemalensis had greater relative abundances of DNA sequences of microbial genera critical for carbon‐use (C‐use) efficiency (i.e. the saprobe, complex C and wood rot/lignin decomposer fungi, and bacterial lignin and other complex C degraders), and greater levels of total organic carbon, C‐biomass, and microbial quotients as indicators of enhanced C‐use efficiency, than found in soils of adjacent 5‐year‐old V. guatemalensis plantations and abandoned non‐productive pasture/grasslands (GRs). The major research conclusions were that (1) conversion of forested land into abandoned pasture/GRs decreased the C‐use efficiency in the soils and the microbial groups associated with C‐use efficiency; (2) soils in plantations of V. guatemalensis were associated with increased abundances of the DNA of these same microbial groups and enhanced C‐use efficiency; (3) DNA‐based taxonomic analysis of microbes and analysis of the microbial quotient values can be used to monitor soil ecosystems for assessment of the efficacy of restoration activities. Thus, planting V. guatemalensis on damaged lands in the Maquenque National Wildlife Refuge should be encouraged to provide a sustainable forestry crop that can be harvested rotationally, while improving soil ecosystem health and reducing the pressure to harvest other forest sites.     

Divergent long‐term trends and interannual variation in ecosystem resource use efficiencies of a southern boreal old black spruce forest 1999–2017

Long‐term trends in ecosystem resource use efficiencies (RUEs) and their controlling factors are key pieces of information for understanding how an ecosystem responds to climate change. We used continuous eddy covariance and microclimate data over the period 1999–2017 from a 120‐year‐old black spruce stand in central Saskatchewan, Canada, to assess interannual variability, long‐term trends, and key controlling factors of gross ecosystem production (GEP) and the RUEs of carbon (CUE = net primary production [NPP]/GEP), light (LUE = GEP/absorbed photosynthetic radiation [APAR]), and water (WUE = GEP/evapotranspiration [E]). At this site, annual GEP has shown an increasing trend over the 19 years (p < 0.01), which may be attributed to rising atmospheric CO₂ concentration. Interannual variability in GEP, aside from its increasing trend, was most strongly related to spring temperatures. Associated with the significant increase in annual GEP were relatively small changes in NPP, APAR, and E, so that annual CUE showed a decreasing trend and annual LUE and WUE showed increasing trends over the 19 years. The long‐term trends in the RUEs were related to the increasing CO₂ concentration. Further analysis of detrended RUEs showed that their interannual variation was impacted most strongly by air temperature. Two‐factor linear models combining CO₂ concentration and air temperature performed well (R²~0.60) in simulating annual RUEs. LUE and WUE were positively correlated both annually and seasonally, while LUE and CUE were mostly negatively correlated. Our results showed divergent long‐term trends among CUE, LUE, and WUE and highlighted the need to account for the combined effects of climatic controls and the ‘CO₂ fertilization effect’ on long‐term variations in RUEs. Since most RUE‐based models rely primarily on one resource limitation, the observed patterns of relative change among the three RUEs may have important implications for RUE‐based modeling of C fluxes.     

Carbon and nitrogen economy of four Triticum aestivum cultivars differing in relative growth rate and water use efficiency

We investigated physiological and morphological traits underlying variation in relative growth rate (KGR) among wheat cultivars. Subsequently, we determined whether higher RGR is correlated with higher water demand and lower plant water use efficiency (WUE_p). Further, the correlation between water use efficiency and leaf nitrogen concentration was examined. For this purpose we chose lour cultivars contrasting in RGR or WUE_p. Gas exchange of shoots and respiration of roots were measured on intact plants over a 24 h period, and total carbon and nitrogen concentrations of all plant parts were determined. The highest RGR was achieved by the cultivars with the highest leaf area ratio. WUE_p was strongly dependent on photosynthetic water use efficiency and was highest for the cultivars with the highest rate of photosynthesis, which achieved higher rates of photosynthesis per unit leaf nitrogen. We found no evidence for a functional or genetic link between the physiological traits underlying differences in RGR (specific leaf area and leaf area ratio) and those causing variation in water use efficiency (photosynthetic rate and transpiration rate). These results indicate that, in wheat, it may be possible to select simultaneously for traits associated with a high WUE_p and a high RGR.     

Seed reserve composition in 19 tree species of a tropical deciduous forest in Mexico and its relationship to seed germination and seedling growth

Background and Aims

The size and composition of seed reserves may reflect the ecological strategy and evolutionary history of a species and also temporal variation in resource availability. The seed mass and composition of seed reserves of 19 co-existing tree species were studied, and we examined how they varied among species in relation to germination and seedling growth rates, as well as between two years with contrasting precipitation (652 and 384 mm).

Methods

Seeds were collected from a tropical deciduous forest in the northwest of Mexico (Chamela Biological Station). The seed dry mass, with and without the seed coat, and the concentrations of lipids, nitrogen and non-structural carbohydrates for the seed minus seed coat were determined. The anatomical localization of these reserves was examined using histochemical analysis. The germination capacity, rate and lag time were determined. The correlations among these variables, and their relationship to previously reported seedling relative growth rates, were evaluated with and without phylogenetic consideration.

Key Results

There were interannual differences in seed mass and reserve composition. Seed was significantly heavier after the drier year in five species. Nitrogen concentration was positively correlated with seed coat fraction, and was significantly higher after the drier year in 12 species. The rate and lag time of germination were negatively correlated with each other. These trait correlations were also supported for phylogenetic independent contrasts. Principal component analysis supported these correlations, and indicated a negative association of seedling relative growth rate with seed size, and a positive association of germination rate with nitrogen and lipid concentrations.

Conclusions

Nitrogen concentration tended to be higher after the drier year and, while interannual variations in seed size and reserve composition were not sufficient to affect interspecific correlations among seed and seedling traits, some of the reserves were related to germination variables and seedling relative growth rate.     

Temperature‐induced water stress in high‐latitude forests in response to natural and anthropogenic warming

The Arctic is particularly sensitive to climate change, but the independent effects of increasing atmospheric CO₂ concentration (pCO₂) and temperature on high‐latitude forests are poorly understood. Here, we present a new, annually resolved record of stable carbon isotope (δ¹³C) data determined from Larix cajanderi tree cores collected from far northeastern Siberia in order to investigate the physiological response of these trees to regional warming. The tree‐ring record, which extends from 1912 through 1961 (50 years), targets early twentieth‐century warming (ETCW), a natural warming event in the 1920s to 1940s that was limited to Northern hemisphere high latitudes. Our data show that net carbon isotope fractionation (Δ¹³C), decreased by 1.7‰ across the ETCW, which is consistent with increased water stress in response to climate warming and dryer soils. To investigate whether this signal is present across the northern boreal forest, we compiled published carbon isotope data from 14 high‐latitude sites within Europe, Asia, and North America. The resulting dataset covered the entire twentieth century and spanned both natural ETCW and anthropogenic Late Twentieth‐Century Warming (~0.7 °C per decade). After correcting for a ~1‰ increase in Δ¹³C in response to twentieth century pCO₂ rise, a significant negative relationship (r = ?0.53, P < 0.0001) between the average, annual Δ¹³C values and regional annual temperature anomalies is observed, suggesting a strong control of temperature on the Δ¹³C value of trees growing at high latitudes. We calculate a 17% increase in intrinsic water‐use efficiency within these forests across the twentieth century, of which approximately half is attributed to a decrease in stomatal conductance in order to conserve water in response to drying conditions, with the other half being attributed to increasing pCO₂. We conclude that annual tree‐ring records from northern high‐latitude forests record the effects of climate warming and pCO₂ rise across the twentieth century.     

Regional‐scale directional changes in abundance of tree species along a temperature gradient in Japan

Climate changes are assumed to shift the ranges of tree species and forest biomes. Such range shifts result from changes in abundances of tree species or functional types. Owing to global warming, the abundance of a tree species or functional type is expected to increase near the colder edge of its range and decrease near the warmer edge. This study examined directional changes in abundance and demographic parameters of forest trees along a temperature gradient, as well as a successional gradient, in Japan. Changes in the relative abundance of each of four functional types (evergreen broad‐leaved, deciduous broad‐leaved, evergreen temperate conifer, and evergreen boreal conifer) and the demography of each species (recruitment rate, mortality, and population growth rate) were analyzed in 39 permanent forest plots across the Japanese archipelago. Directional changes in the relative abundance of functional types were detected along the temperature gradient. Relative abundance of evergreen broad‐leaved trees increased near their colder range boundaries, especially in secondary forests, coinciding with the decrease in deciduous broad‐leaved trees. Similarly, relative abundance of deciduous broad‐leaved trees increased near their colder range boundaries, coinciding with the decrease in boreal conifers. These functional‐type‐level changes were mainly due to higher recruitment rates and partly to the lower mortality of individual species at colder sites. This is the first report to show that tree species abundances in temperate forests are changing directionally along a temperature gradient, which might be due to current or past climate changes as well as recovery from past disturbances.     

Does declining carbon‐use efficiency explain thermal acclimation of soil respiration with warming?

Enhanced soil respiration in response to global warming may substantially increase atmospheric CO₂ concentrations above the anthropogenic contribution, depending on the mechanisms underlying the temperature sensitivity of soil respiration. Here, we compared short‐term and seasonal responses of soil respiration to a shifting thermal environment and variable substrate availability via laboratory incubations. To analyze the data from incubations, we implemented a novel process‐based model of soil respiration in a hierarchical Bayesian framework. Our process model combined a Michaelis–Menten‐type equation of substrate availability and microbial biomass with an Arrhenius‐type nonlinear temperature response function. We tested the competing hypotheses that apparent thermal acclimation of soil respiration can be explained by depletion of labile substrates in warmed soils, or that physiological acclimation reduces respiration rates. We demonstrated that short‐term apparent acclimation can be induced by substrate depletion, but that decreasing microbial biomass carbon (MBC) is also important, and lower MBC at warmer temperatures is likely due to decreased carbon‐use efficiency (CUE). Observed seasonal acclimation of soil respiration was associated with higher CUE and lower basal respiration for summer‐ vs. winter‐collected soils. Whether the observed short‐term decrease in CUE or the seasonal acclimation of CUE with increased temperatures dominates the response to long‐term warming will have important consequences for soil organic carbon storage.     

Photosynthesis and water‐use efficiency: A comparison between invasive (exotic) and non‐invasive (native) species

Abstract Invasive species have been hypothesized to out‐compete natives though either a Jack‐of‐all‐trades strategy, where they are able to utilize resources effectively in unfavourable environments, a master‐of‐some, where resource utilization is greater than its competitors in favourable environments, or a combination of the two (Jack‐and‐master). We examined the invasive strategy of Berberis darwinii in New Zealand compared with four co‐occurring native species by examining germination, seedling survival, photosynthetic characteristics and water‐use efficiency of adult plants, in sun and shade environments. Berberis darwinii seeds germinated more in shady sites than the other natives, but survival was low. In contrast, while germination of B. darwinii was the same as the native species in sunny sites, seedling survival after 18 months was nearly twice that of the all native species. The maximum photosynthetic rate of B. darwinii was nearly double that of all native species in the sun, but was similar among all species in the shade. Other photosynthetic traits (quantum yield and stomatal conductance) did not generally differ between B. darwinii and the native species, regardless of light environment. Berberis darwinii had more positive values of δ¹³C than the four native species, suggesting that it gains more carbon per unit water transpired than the competing native species. These results suggest that the invasion success of B. darwinii may be partially explained by combination of a Jack‐of‐all‐trades scenario of widespread germination with a master‐of‐some scenario through its ability to photosynthesize at higher rates in the sun and, hence, gain a rapid height and biomass advantage over native species in favourable environments.     

Gas exchange and water‐use efficiency in plant canopies

In this review, I first address the basics of gas exchange, water‐use efficiency and carbon isotope discrimination in C₃ plant canopies. I then present a case study of water‐use efficiency in northern Australian tree species. In general, C₃ plants face a trade‐off whereby increasing stomatal conductance for a given set of conditions will result in a higher CO₂ assimilation rate, but a lower photosynthetic water‐use efficiency. A common garden experiment suggested that tree species which are able to establish and grow in drier parts of northern Australia have a capacity to use water rapidly when it is available through high stomatal conductance, but that they do so at the expense of low water‐use efficiency. This may explain why community‐level carbon isotope discrimination does not decrease as steeply with decreasing rainfall on the North Australian Tropical Transect as has been observed on some other precipitation gradients. Next, I discuss changes in water‐use efficiency that take place during leaf expansion in C₃ plant leaves. Leaf phenology has recently been recognised as a significant driver of canopy gas exchange in evergreen forest canopies, and leaf expansion involves changes in both photosynthetic capacity and water‐use efficiency. Following this, I discuss the role of woody tissue respiration in canopy gas exchange and how photosynthetic refixation of respired CO₂ can increase whole‐plant water‐use efficiency. Finally, I discuss the role of water‐use efficiency in driving terrestrial plant responses to global change, especially the rising concentration of atmospheric CO₂. In coming decades, increases in plant water‐use efficiency caused by rising CO₂ are likely to partially mitigate impacts on plants of drought stress caused by global warming.     

Water‐use efficiency in response to climate change: from leaf to ecosystem in a temperate steppe

Water‐use efficiency (WUE) has been recognized as an important characteristic of ecosystem productivity, which links carbon (C) and water cycling. However, little is known about how WUE responds to climate change at different scales. Here, we investigated WUE at leaf, canopy, and ecosystem levels under increased precipitation and warming from 2005 to 2008 in a temperate steppe in Northern China. We measured gross ecosystem productivity (GEP), net ecosystem CO₂ exchange (NEE), evapotranspiration (ET), evaporation (E), canopy transpiration (T_c), as well as leaf photosynthesis (P_max) and transpiration (T_l) of a dominant species to calculate canopy WUE (WUE_c=GEP/T), ecosystem WUE (WUE_gep=GEP/ET or WUE_nee=NEE/ET) and leaf WUE (WUE_l=P_max/T_l). The results showed that increased precipitation stimulated WUE_c, WUE_gep and WUE_nee by 17.1%, 10.2% and 12.6%, respectively, but decreased WUE_l by 27.4%. Climate warming reduced canopy and ecosystem WUE over the 4 years but did not affect leaf level WUE. Across the 4 years and the measured plots, canopy and ecosystem WUE linearly increased, but leaf level WUE of the dominant species linearly decreased with increasing precipitation. The differential responses of canopy/ecosystem WUE and leaf WUE to climate change suggest that caution should be taken when upscaling WUE from leaf to larger scales. Our findings will also facilitate mechanistic understanding of the C–water relationships across different organism levels and in projecting the effects of climate warming and shifting precipitation regimes on productivity in arid and semiarid ecosystems.     

Evidence of changing intrinsic water‐use efficiency under rising atmospheric CO2 concentrations in Boreal Fennoscandia from subfossil leaves and tree ring δ13C ratios

Investigating the many internal feedbacks within the climate system is a vital component of the effort to quantify the full effects of future anthropogenic climate change. The stomatal apertures of plants tend to close and decrease in number under elevated CO₂ concentrations, increasing water‐use efficiency (WUE) and reducing canopy evapotranspiration. Experimental and modelling studies reveal huge variations in these changes such that the warming associated with reduced evapotranspiration (known as physiological forcing) is neither well understood or constrained. Palaeo‐observations of changes in stomatal response and plant WUE under rising CO₂ might be used to better understand the processes underlying the physiological forcing feedback and to link measured changes in plant WUE to a specific physiological change in stomata. Here we use time series of tree ring (Pinus sylvestris L.) δ¹³C and subfossil leaf (Betula nana L.) measurements of stomatal density and geometry to derive records of changes in intrinsic water‐use efficiency (iWUE) and maximum stomatal conductance in the Boreal zone of northern Finland and Sweden. We investigate the rate of change in both proxies, over the recent past. The independent lines of evidence from these two different Boreal species indicate increased iWUE and reduced maximum stomatal conductance of similar magnitude from preindustrial times (ca. ad 1850) to around ad 1970. After this maximum stomatal conductance continues to decrease to ad 2000 in B. nana but iWUE in P. sylvestris reaches a plateau. We suggest that northern boreal P. sylvestris might have reached a threshold in its ability to increase WUE as CO₂ rises.     

A common thermal niche among geographically diverse populations of the widely distributed tree species Eucalyptus tereticornis: No evidence for adaptation to climate‐of‐origin

Impacts of climate warming depend on the degree to which plants are constrained by adaptation to their climate‐of‐origin or exhibit broad climatic suitability. We grew cool‐origin, central and warm‐origin provenances of Eucalyptus tereticornis in an array of common temperature environments from 18 to 35.5°C to determine if this widely distributed tree species consists of geographically contrasting provenances with differentiated and narrow thermal niches, or if provenances share a common thermal niche. The temperature responses of photosynthesis, respiration, and growth were equivalent across the three provenances, reflecting a common thermal niche despite a 2,200 km geographic distance and 13°C difference in mean annual temperature at seed origin. The temperature dependence of growth was primarily mediated by changes in leaf area per unit plant mass, photosynthesis, and whole‐plant respiration. Thermal acclimation of leaf, stem, and root respiration moderated the increase in respiration with temperature, but acclimation was constrained at high temperatures. We conclude that this species consists of provenances that are not differentiated in their thermal responses, thus rejecting our hypothesis of adaptation to climate‐of‐origin and suggesting a shared thermal niche. In addition, growth declines with warming above the temperature optima were driven by reductions in whole‐plant leaf area and increased respiratory carbon losses. The impacts of climate warming will nonetheless vary across the geographic range of this and other such species, depending primarily on each provenance's climate position on the temperature response curves for photosynthesis, respiration, and growth.