Soil‐ and plant‐water dynamics in a C3/C4 grassland exposed to a subambient to superambient CO2 gradient

Plants may be more sensitive to carbon dioxide (CO₂) enrichment at subambient concentrations than at superambient concentrations, but field tests are lacking. We measured soil‐water content and determined xylem pressure potentials and δ¹³C values of leaves of abundant species in a C3/C4 grassland exposed during 1997–1999 to a continuous gradient in atmospheric CO₂ spanning subambient through superambient concentrations (200–560 µmol mol^2?1). We predicted that CO₂ enrichment would lessen soil‐water depletion and increase xylem potentials more over subambient concentrations than over superambient concentrations. Because water‐use efficiency of C3 species (net assimilation/leaf conductance; A/g) typically increases as soils dry, we hypothesized that improvements in plant‐water relations at higher CO₂ would lessen positive effects of CO₂ enrichment on A/g. Depletion of soil water to 1.35 m depth was greater at low CO₂ concentrations than at higher CO₂ concentrations during a mid‐season drought in 1998 and during late‐season droughts in 1997 and 1999. During droughts each year, mid‐day xylem potentials of the dominant C4 perennial grass (Bothriochloa ischaemum (L.) Keng) and the dominant C3 perennial forb (Solanum dimidiatum Raf.) became less negative as CO₂ increased from subambient to superambient concentrations. Leaf A/g—derived from leaf δ¹³C values—was insensitive to feedbacks from CO₂ effects on soil water and plant water. Among most C3 species sampled—including annual grasses, perennial grasses and perennial forbs—A/g increased linearly with CO₂ across subambient concentrations. Leaf and air δ¹³C values were too unstable at superambient CO₂ concentrations to reliably determine A/g. Significant changes in soil‐ and plant‐water relations over subambient to superambient concentrations and in leaf A/g over subambient concentrations generally were not greater over low CO₂ than over higher CO₂. The continuous response of these variables to CO₂ suggests that atmospheric change has already improved water relations of grassland species and that periodically water‐limited grasslands will remain sensitive to CO₂ enrichment.     

Assessing the effects of nitrogen deposition and climate on carbon isotope discrimination and intrinsic water‐use efficiency of angiosperm and conifer trees under rising CO2 conditions

The objective of this study is to globally assess the effects of atmospheric nitrogen deposition and climate, associated with rising levels of atmospheric CO₂, on the variability of carbon isotope discrimination (Δ¹³C), and intrinsic water‐use efficiency (iWUE) of angiosperm and conifer tree species. Eighty‐nine long‐term isotope tree‐ring chronologies, representing 23 conifer and 13 angiosperm species for 53 sites worldwide, were extracted from the literature, and used to obtain long‐term time series of Δ¹³C and iWUE. Δ¹³C and iWUE were related to the increasing concentration of atmospheric CO₂ over the industrial period (1850–2000) and to the variation of simulated atmospheric nitrogen deposition and climatic variables over the period 1950–2000. We applied generalized additive models and linear mixed‐effects models to predict the effects of climatic variables and nitrogen deposition on Δ¹³C and iWUE. Results showed a declining Δ¹³C trend in the angiosperm and conifer species over the industrial period and a 16.1% increase of iWUE between 1850 and 2000, with no evidence that the increased rate was reduced at higher ambient CO₂ values. The temporal variation in Δ¹³C supported the hypothesis of an active plant mechanism that maintains a constant ratio between intercellular and ambient CO₂ concentrations. We defined linear mixed‐effects models that were effective to describe the variation of Δ¹³C and iWUE as a function of a set of environmental predictors, alternatively including annual rate (N_rate) and long‐term cumulative (N_cum) nitrogen deposition. No single climatic or atmospheric variable had a clearly predominant effect, however, Δ¹³C and iWUE showed complex dependent interactions between different covariates. A significant association of N_rate with iWUE and Δ¹³C was observed in conifers and in the angiosperms, and N_cum was the only independent term with a significant positive association with iWUE, although a multi‐factorial control was evident in conifers.     

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.     

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 (WUE_plant) is needed for sustainable agriculture. WUE_plant is a complex trait and an efficient phenotyping method that reports on components of WUE_plant, such as intrinsic water‐use efficiency (WUE_i, 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 (δ¹³C), when the efficiency of the CO₂‐concentrating mechanism (leakiness, ?) remains constant, can be used to screen for WUE_i. The limited information about how ? responds to water limitations confines the application of δ¹³C for WUE_i screening of C₄ crops. The current research aimed to test the response of ? to short‐ or long‐term moderate water limitations, and the relationship of δ¹³C with WUE_i and WUE_plant, by addressing potential mesophyll CO₂ conductance (g_m) and biochemical limitations in the C₄ plant Sorghum bicolor. We demonstrate that g_m and ? are not responsive to short‐ or long‐term water limitations. Additionally, δ¹³C was not correlated with gas‐exchange estimates of WUE_i under short‐ and long‐term water limitations, but showed a significant negative relationship with WUE_plant. The observed association between the δ¹³C and WUE_plant suggests an intrinsic link of δ¹³C with WUE_i in this C₄ plant, and can potentially be used as a screening tool for WUE_plant in sorghum.     

Nitrogen fertilization enhances water‐use efficiency in a saline environment

Effects of salinity and nutrients on carbon gain in relation to water use were studied in the grey mangrove, Avicennia marina, growing along a natural salinity gradient in south‐eastern Australia. Tall trees characterized areas of seawater salinities (fringe zone) and stunted trees dominated landward hypersaline areas (scrub zone). Trees were fertilized with nitrogen (+N) or phosphorus (+P) or unfertilized. There was no significant effect of +P on shoot growth, whereas +N enhanced canopy development, particularly in scrub trees. Scrub trees maintained greater CO₂ assimilation per unit water transpired (water‐use efficiency, WUE) and had lower nitrogen‐use efficiency (NUE; CO₂ assimilation rate per unit leaf nitrogen) than fringe trees. The CO₂ assimilation rates of +N trees were similar to those in other treatments, but were achieved at lower transpiration rates, stomatal conductance and intercellular CO₂ concentrations. Maintaining comparable assimilation rates at lower stomatal conductance requires greater ribulose 1·5‐bisphosphate carboxylase/oxygenase activity, consistent with greater N content per unit leaf area in +N trees. Hence, +N enhanced WUE at the expense of NUE. Instantaneous WUE estimates were supported by less negative foliar δ¹³C values for +N trees and scrub control trees. Thus, nutrient enrichment may alter the structure and function of mangrove forests along salinity gradients.     

Nutrient supply enhanced the increase in intrinsic water‐use efficiency of a temperate seminatural grassland in the last century

Under the increase in atmospheric CO₂ during the last century, variable increases in the intrinsic water‐use efficiency (W_i), i.e., the ratio between carbon assimilation rate (A) and stomatal conductance (g_s), of C₃ vegetation have been observed. Here, we ask if long‐term nutrient status and especially nitrogen supply have an effect on the CO₂ response of W_i in a temperate seminatural C₃ grassland. This analysis draws on the long‐term trends (1915–2009) in W_i, derived from carbon isotope analysis, of archived hay and herbage from the Park Grass Experiment at Rothamsted (South‐East England). Plant samples came from five fertilizer treatments, each with different annual nitrogen (N; 0, 48 or 96 kg ha^?1), phosphorus (P; 0 or 35 kg ha^?1) and potassium (K; 0 or 225 kg ha^?1) applications, with lime as required to maintain soil pH near 7. Carbon isotope discrimination (¹³Δ) increased significantly (P < 0.001) on the Control (0.9‰ per 100 ppm CO₂ increase). This trend differed significantly (P < 0.01) from those observed on the fertilized treatments (PK only: 0.4‰ per 100 ppm CO₂ increase, P < 0.001; Low N only, Low N+PK, High N+PK: no significant increase). The ¹³Δ trends on fertilized treatments did not differ significantly from each other. However, N status, assessed as N fertilizer supply plus an estimate of biologically fixed N, was negatively related (r² = 0.88; P < 0.02) to the trend for ¹³Δ against CO₂. Other indices of N status exhibited similar relationships. Accordingly, the increase in W_i at High N+PK was twice that of the Control (+28% resp. +13% relative to 1915). In addition, the CO₂ responsiveness of ¹³Δ was related to the grass content of the plant community. This may have been due to the greater CO₂ responsiveness of g_s in grasses relative to forbs. Thus, the greater CO₂ response of grass‐rich fertilized swards may be related to effects of nutrient supply on botanical composition.     

Intrinsic water‐use efficiency of temperate seminatural grassland has increased since 1857: an analysis of carbon isotope discrimination of herbage from the Park Grass Experiment

A 150‐year‐long record of intrinsic water‐use efficiency (W_i) was derived from community‐level carbon isotope discrimination (¹³Δ) in the herbage of the unfertilized, unlimed control treatment (plot 3) of the Park Grass Experiment at Rothamsted (England) between 1857 and 2007. ¹³Δ during spring growth (first cut harvested in June) averaged 21.0‰ (±0.5‰ SD) and has not shown a long‐term trend (P=0.5) since 1857. ¹³Δ of summer/autumn growth (second cut harvested between September and November) increased from 21.3‰ to 22.0‰ (P < 0.001) between 1875 and 2007. W_i during spring growth has therefore increased by 33% since the beginning of the experiment, and W_i of summer/autumn growth has increased by 18%. The variation in ¹³Δ was mainly related to weather conditions. Plant available soil water explained 51% and 40% of the variation in spring growth ¹³Δ and summer/autumn growth ¹³Δ, respectively. In the 1857–2007 period yields have not increased, suggesting that community‐level photosynthesis has not increased either. Therefore, the increased W_i probably resulted from a decreased stomatal conductance. Vapour pressure deficit (VPD) during spring growth (March–June) has not changed since 1915, meaning that instantaneous water‐use efficiency (W_t) in spring time has increased and transpiration has probably decreased, provided that leaf temperature followed air temperature. Conversely, VPD in the months between the first and second cut has increased by 0.07 kPa since 1915, offsetting the effect of increased W_i on W_t during summer and early autumn. Our results suggest that vegetation has adjusted physiologically to elevated CO₂ by decreasing stomatal conductance in this nutrient‐limited grassland.     

Canopy radiation‐ and water‐use efficiencies as affected by elevated [CO2]

This study used an environmentally controlled plant growth facility, EcoCELLs, to measure canopy gas exchanges directly and to examine the effects of elevated [CO₂] on canopy radiation‐ and water‐use efficiencies. Sunflowers (Helianthus annus var. Mammoth) were grown at ambient (399 μmol mol^?1) and elevated [CO₂] (746 μmol mol^?1) for 53 days in EcoCELLs. Whole canopy carbon‐ and water‐fluxes were measured continuously during the period of the experiment. The results indicated that elevated [CO₂] enhanced daily total canopy carbon‐ and water‐fluxes by 53% and 11%, respectively, on a ground‐area basis, resulting in a 54% increase in radiation‐use efficiency (RUE) based on intercepted photosynthetic active radiation and a 26% increase in water‐use efficiency (WUE) by the end of the experiment. Canopy carbon‐ and water‐fluxes at both CO₂ treatments varied with canopy development. They were small at 22 days after planting (DAP) and gradually increased to the maxima at 46 DAP. When canopy carbon‐ and water‐fluxes were expressed on a leaf‐area basis, no effect of CO₂ was found for canopy water‐flux while elevated [CO₂] still enhanced canopy carbon‐flux by 29%, on average. Night‐time canopy carbon‐flux was 32% higher at elevated than at ambient [CO₂]. In addition, RUE and WUE displayed strong diurnal variations, high at noon and low in the morning or afternoon for WUE but opposite for RUE. This study provided direct evidence that plant canopy may consume more, instead of less, water but utilize both water and radiation more efficiently at elevated than at ambient [CO₂], at least during the exponential growth period as illustrated in this experiment.     

Variation in the carbon and oxygen isotope composition of plant biomass and its relationship to water‐use efficiency at the leaf‐ and ecosystem‐scales in a northern Great Plains grassland

Measurements of the carbon (δ¹³C_m) and oxygen (δ¹⁸O_m) isotope composition of C₃ plant tissue provide important insights into controls on water‐use efficiency. We investigated the causes of seasonal and inter‐annual variability in water‐use efficiency in a grassland near Lethbridge, Canada using stable isotope (leaf‐scale) and eddy covariance measurements (ecosystem‐scale). The positive relationship between δ¹³C_m and δ¹⁸O_m values for samples collected during 1998–2001 indicated that variation in stomatal conductance and water stress‐induced changes in the degree of stomatal limitation of net photosynthesis were the major controls on variation in δ¹³C_m and biomass production during this time. By comparison, the lack of a significant relationship between δ¹³C_m and δ¹⁸O_m values during 2002, 2003 and 2006 demonstrated that water stress was not a significant limitation on photosynthesis and biomass production in these years. Water‐use efficiency was higher in 2000 than 1999, consistent with expectations because of greater stomatal limitation of photosynthesis and lower leaf c_i/ca during the drier conditions of 2000. Calculated values of leaf‐scale water‐use efficiency were 2–3 times higher than ecosystem‐scale water‐use efficiency, a difference that was likely due to carbon lost in root respiration and water lost during soil evaporation that was not accounted for by the stable isotope measurements.     

From pattern to process: linking intrinsic water‐use efficiency to drought‐induced forest decline

The rise in atmospheric CO₂ concentrations (Ca) has been related to tree growth enhancement and increasing intrinsic water‐use efficiency (iWUE). However, the extent that rising Ca has led to increased long‐term iWUE and whether climate could explain deviations from expected Ca‐induced growth enhancement are still poorly understood. The aim of this research was to use Ca and local climatic variability to explain changes during the 20th century in growth and tree ring and needle δ¹³C in declining and nondeclining Abies alba stands from the Spanish Pyrenees, near the southern distribution limit of this species. The temporal trends of iWUE were calculated under three theoretical scenarios for the regulation of plant‐gas exchange at increasing Ca. We tested different linear mixed‐effects models by multimodel selection criteria to predict basal area increment (BAI), a proxy of tree radial growth, using these scenarios and local temperature together with precipitation data as predictors. The theoretical scenario assuming the strongest response to Ca explained 66–81% of the iWUE variance and 28–56% of the observed BAI variance, whereas local climatic variables together explained less than 11–21% of the BAI variance. Our results are consistent with a drought‐induced limitation of the tree growth response to rising CO₂ and a decreasing rate of iWUE improvement from the 1980s onward in declining A. alba stands subjected to lower water availability.     

Transpiration of a 64-year-old maritime pine stand in Portugal

Alpine plant species have been shown to exhibit a more pronounced increase in leaf photosynthesis under elevated CO₂ than lowland plants. In order to test whether this higher carbon fixation efficiency will translate into increased biomass production under CO₂ enrichment we exposed plots of narrow alpine grassland (Swiss Central Alps, 2470 m) to ambient (355 l l^-1) and elevated (680 l l^-1) CO₂ concentration using open top chambers. Part of the plost received moderate mineral nutrient additions (40 kg ha^-1 year^-1 of nitrogen in a complete fertilizer mix). Under natural nutrient supply CO₂ enrichment had no effect on biomass production per unit land area during any of the three seasons studied so far. Correspondingly, the dominant species Carex curvula and Leontodon helveticus as well as Trifolium alpinum did not show a growth response either at the population level or at the shoot level. However, the subdominant generalistic species Poa alpina strongly increased shoot growth (+47%). Annual root production (in ingrowth cores) was significantly enhanced in C. curvula in the 2nd and 3rd year of investigation (+43%) but was not altered in the bulk samples for all species. Fertilizer addition generally stimulated above-ground (+48%) and below-ground (+26%) biomass production right from the beginning. Annual variations in weather conditions during summer also strongly influenced above-ground biomass production (19–27% more biomass in warm seasons compared to cool seasons). However, neither nutrient availability nor climate had a significant effect on the CO₂ response of the plants. Our results do not support the hypothesis that alpine plants, due to their higher carbon uptake efficiency, will increase biomass production under future atmospheric CO₂ enrichment, at least not in such late successional communities. However, as indicated by the response of P. alpina, species-specific responses occur which may lead to altered community structure and perhaps ecosystem functioning in the long-term. Our findings further suggest that possible climatic changes are likely to have a greater impact on plant growth in alpine environments than the direct stimulation of photosynthesis by CO₂. Counter-intuitively, our results suggest that even under moderate climate warming or enhanced atmospheric nitrogen deposition positive biomass responses to CO₂ enrichment of the currently dominating species are unlikely.     

Maintenance of leaf N controls the photosynthetic CO2 response of grassland species exposed to 9 years of free‐air CO2 enrichment

Determining underlying physiological patterns governing plant productivity and diversity in grasslands are critical to evaluate species responses to future environmental conditions of elevated CO₂ and nitrogen (N) deposition. In a 9‐year experiment, N was added to monocultures of seven C₃ grassland species exposed to elevated atmospheric CO₂ (560 μmol CO₂ mol^?1) to evaluate how N addition affects CO₂ responsiveness in species of contrasting functional groups. Functional groups differed in their responses to elevated CO₂ and N treatments. Forb species exhibited strong down‐regulation of leaf N_mass concentrations (?26%) and photosynthetic capacity (?28%) in response to elevated CO₂, especially at high N supply, whereas C₃ grasses did not. Hence, achieved photosynthetic performance was markedly enhanced for C₃ grasses (+68%) in elevated CO₂, but not significantly for forbs. Differences in access to soil resources between forbs and grasses may distinguish their responses to elevated CO₂ and N addition. Forbs had lesser root biomass, a lower distribution of biomass to roots, and lower specific root length than grasses. Maintenance of leaf N, possibly through increased root foraging in this nutrient‐poor grassland, was necessary to sustain stimulation of photosynthesis under long‐term elevated CO₂. Dilution of leaf N and associated photosynthetic down‐regulation in forbs under elevated [CO₂], relative to the C₃ grasses, illustrates the potential for shifts in species composition and diversity in grassland ecosystems that have significant forb and grass components.     

Lack of photosynthetic or stomatal regulation after 9 years of elevated [CO2] and 4 years of soil warming in two conifer species at the alpine treeline

Alpine treelines are temperature‐limited vegetation boundaries. Understanding the effects of elevated [CO₂] and warming on CO₂ and H₂O gas exchange may help predict responses of treelines to global change. We measured needle gas exchange of Larix decidua Mill. and Pinus mugo ssp. uncinata DC trees after 9 years of free air CO₂ enrichment (575 µmol mol^?1) and 4 years of soil warming (+4 °C) and analysed δ¹³C and δ¹⁸O values of needles and tree rings. Tree needles under elevated [CO₂] showed neither nitrogen limitation nor end‐product inhibition, and no down‐regulation of maximal photosynthetic rate (A_max) was found. Both tree species showed increased net photosynthetic rates (A_n) under elevated [CO₂] (L. decidua: +39%; P. mugo: +35%). Stomatal conductance (g_H2O) was insensitive to changes in [CO₂], thus transpiration rates remained unchanged and intrinsic water‐use efficiency (iWUE) increased due to higher A_n. Soil warming affected neither A_n nor g_H2O. Unresponsiveness of g_H2O to [CO₂] and warming was confirmed by δ¹⁸O needle and tree ring values. Consequently, under sufficient water supply, elevated [CO₂] induced sustained enhancement in A_n and lead to increased C inputs into this ecosystem, while soil warming hardly affected gas exchange of L. decidua and P. mugo at the alpine treeline.     

Photosynthetic responses of 13 grassland species across 11 years of free‐air CO2 enrichment is modest,consistent and independent of N supply

If long‐term responses of photosynthesis and leaf diffusive conductance to rising atmospheric carbon dioxide (CO₂) levels are similar or predictably different among species, functional types, and ecosystem types, general global models of elevated CO₂ effects can effectively be developed. To address this issue we measured gas exchange rates of 13 perennial grassland species from four functional groups across 11 years of long‐term free‐air CO₂ enrichment (eCO₂, +180 ppm above ambient CO₂) in the BioCON experiment in Minnesota, USA. Eleven years of eCO₂ produced consistent but modest increases in leaf net photosynthetic rates of 10% on average compared with plants grown at ambient CO₂ concentrations across the 13 species. This eCO₂‐induced enhancement did not depend on soil N treatment, is much less than the average across other longer‐term studies, and represents strong acclimation (i.e. downregulation) as it is also much less than the instantaneous response to eCO₂. The legume and C3 nonlegume forb species were the most responsive among the functional groups (+13% in each), the C4 grasses the least responsive (+4%), and C3 grasses intermediate in their photosynthetic response to eCO₂ across years (+9%). Leaf stomatal conductance and nitrogen content declined comparably across species in eCO₂ compared with ambient CO₂ and to degrees corresponding to results from other studies. The significant acclimation of photosynthesis is explained in part by those eCO₂‐induced decreases in leaf N content and stomatal conductance that reduce leaf photosynthetic capacity in plants grown under elevated compared with ambient CO₂ concentrations. Results of this study, probably the longest‐term with the most species, suggest that carbon cycle models that assume and thereby simulate long‐lived strong eCO₂ stimulation of photosynthesis (e.g.> 25%) for all of Earth's terrestrial ecosystems should be viewed with a great deal of caution.     

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.     

Increasing water‐use efficiency and age‐specific growth responses of old‐growth ponderosa pine trees in the Northern Rockies

We examined radial growth responses of ponderosa pine (Pinus ponderosa var. ponderosa) between 1905–1954 and 1955–2004 to determine if the effects of increased intrinsic water‐use efficiencies (iWUE) caused by elevated atmospheric CO₂ concentrations were age‐specific. We collected 209 cores from five sites in the Northern Rockies and calculated iWUE using carbon isotope data from 1850 to 2004. Standardized radial growth responses were age dependent, with older trees exhibiting significantly higher values than younger trees during the later period at four sites and all sites combined. No significant differences in radial growth existed either for the individual sites or combined site during the earlier period. Increases in iWUE during 1955–2004 were 11% greater than during 1905–1954, and pentadal fluctuations in iWUE were significantly correlated with the radial growth of older trees from 1850 to 2004. Radial growth of younger trees and iWUE were not significantly correlated. Our results suggest that: (1) responses to elevated atmospheric CO₂ in old‐growth ponderosa forests are age‐specific; (2) radial growth increases in older trees coincided with increased iWUE; (3) ponderosa had increased growth rates in their third, fourth, and fifth centuries of life; and (4) age‐specific growth responses during 1955–2004 are unique since at least the mid‐16th century.     

Variations in nitrogen use efficiency reflect the biochemical subtype while variations in water use efficiency reflect the evolutionary lineage of C4 grasses at inter‐glacial CO2

C₄ photosynthesis evolved multiple times in diverse lineages. Most physiological studies comparing C₄ plants were not conducted at the low atmospheric CO₂ prevailing during their evolution. Here, 24 C₄ grasses belonging to three biochemical subtypes [nicotinamide adenine dinucleotide malic enzyme (NAD‐ME), phosphoenolpyruvate carboxykinase (PCK) and nicotinamide adenine dinucleotide phosphate malic enzyme (NADP‐ME)] and six major evolutionary lineages were grown under ambient (400 μL L^?1) and inter‐glacial (280 μL L^?1) CO₂. We hypothesized that nitrogen‐related and water‐related physiological traits are associated with subtypes and lineages, respectively. Photosynthetic rate and stomatal conductance were constrained by the shared lineage, while variation in leaf mass per area (LMA), leaf N per area, plant dry mass and plant water use efficiency were influenced by the subtype. Subtype and lineage were equally important for explaining variations in photosynthetic nitrogen use efficiency (PNUE) and photosynthetic water use efficiency (PWUE). CO₂ treatment impacted most parameters. Overall, higher LMA and leaf N distinguished the Chloridoideae/NAD‐ME group, while NADP‐ME and PCK grasses were distinguished by higher PNUE regardless of lineage. Plants were characterized by high photosynthesis and PWUE when grown at ambient CO₂ and by high conductance at inter‐glacial CO₂. In conclusion, the evolutionary and biochemical diversity among C₄ grasses was aligned with discernible leaf physiology, but it remains unknown whether these traits represent ecophysiological adaptation.     

Increased water‐use efficiency during the 20th century did not translate into enhanced tree growth

Aim The goals of this study are: (1) to determine whether increasing atmospheric CO₂ concentrations and changing climate increased intrinsic water use efficiency (iWUE, as detected by changes in Δ¹³C) over the last four decades; and if it did increase iWUE, whether it led to increased tree growth (as measured by tree‐ring growth); (2) to assess whether CO₂ responses are biome dependent due to different environmental conditions, including availability of nutrients and water; and (3) to discuss how the findings of this study can better inform assumptions of CO₂ fertilization and climate change effects in biospheric and climate models. Location A global range of sites covering all major forest biome types. Methods The analysis encompassed 47 study sites including boreal, wet temperate, mediterranean, semi‐arid and tropical biomes for which measurements of tree ring Δ¹³C and growth are available over multiple decades. Results The iWUE inferred from the Δ¹³C analyses of comparable mature trees increased 20.5% over the last 40 years with no significant differences between biomes. This increase in iWUE did not translate into a significant overall increase in tree growth. Half of the sites showed a positive trend in growth while the other half had a negative or no trend. There were no significant trends within biomes or among biomes. Main conclusions These results show that despite an increase in atmospheric CO₂ concentrations of over 50 p.p.m. and a 20.5% increase in iWUE during the last 40 years, tree growth has not increased as expected, suggesting that other factors have overridden the potential growth benefits of a CO₂‐rich world in many sites. Such factors could include climate change (particularly drought), nutrient limitation and/or physiological long‐term acclimation to elevated CO₂. Hence, the rate of biomass carbon sequestration in tropical, arid, mediterranean, wet temperate and boreal ecosystems may not increase with increasing atmospheric CO₂ concentrations as is often implied by biospheric models and short‐term elevated CO₂ experiments.     

Vapour pressure deficit during growth has little impact on genotypic differences of transpiration efficiency at leaf and whole‐plant level: an example from Populus nigra L.

Poplar genotypes differ in transpiration efficiency (TE) at leaf and whole‐plant level under similar conditions. We tested whether atmospheric vapour pressure deficit (VPD) affected TE to the same extent across genotypes. Six Populus nigra genotypes were grown under two VPD. We recorded (1) ¹³C content in soluble sugars; (2) ¹⁸O enrichment in leaf water; (3) leaf‐level gas exchange; and (4) whole‐plant biomass accumulation and water use. Whole‐plant and intrinsic leaf TE and ¹³C content in soluble sugars differed significantly among genotypes. Stomatal conductance contributed more to these differences than net CO₂ assimilation rate. VPD increased water use and reduced whole‐plant TE. It increased intrinsic leaf‐level TE due to a decline in stomatal conductance. It also promoted higher ¹⁸O enrichment in leaf water. VPD had no genotype‐specific effect. We detected a deviation in the relationship between ¹³C in leaf sugars and ¹³C predicted from gas exchange and the standard discrimination model. This may be partly due to genotypic differences in mesophyll conductance, and to its lack of sensitivity to VPD. Leaf‐level ¹³C discrimination was a powerful predictor of the genetic variability of whole‐plant TE irrespective of VPD during growth.     

Transpiration of a 64-year old maritime pine stand in Portugal

Alpine plant species have been shown to exhibit a more pronounced increase in leaf photosynthesis under elevated CO₂ than lowland plants. In order to test whether this higher carbon fixation efficiency will translate into increased biomass production under CO₂ enrichment we exposed plots of narrow alpine grassland (Swiss Central Alps, 2470 m) to ambient (355 μl l^-1) and elevated (680 μl l^-1) CO₂ concentration using open top chambers. Part of the plost received moderate mineral nutrient additions (40 kg ha^-1 year^-1 of nitrogen in a complete fertilizer mix). Under natural nutrient supply CO₂ enrichment had no effect on biomass production per unit land area during any of the three seasons studied so far. Correspondingly, the dominant species Carex curvula and Leontodon helveticus as well as Trifolium alpinum did not show a growth response either at the population level or at the shoot level. However, the subdominant generalistic species Poa alpina strongly increased shoot growth (+47%). Annual root production (in ingrowth cores) was significantly enhanced in C. curvula in the 2nd and 3rd year of investigation (+43%) but was not altered in the bulk samples for all species. Fertilizer addition generally stimulated above-ground (+48%) and below-ground (+26%) biomass production right from the beginning. Annual variations in weather conditions during summer also strongly influenced above-ground biomass production (19–27% more biomass in warm seasons compared to cool seasons). However, neither nutrient availability nor climate had a significant effect on the CO₂ response of the plants. Our results do not support the hypothesis that alpine plants, due to their higher carbon uptake efficiency, will increase biomass production under future atmospheric CO₂ enrichment, at least not in such late successional communities. However, as indicated by the response of P. alpina, species-specific responses occur which may lead to altered community structure and perhaps ecosystem functioning in the long-term. Our findings further suggest that possible climatic changes are likely to have a greater impact on plant growth in alpine environments than the direct stimulation of photosynthesis by CO₂. Counter-intuitively, our results suggest that even under moderate climate warming or enhanced atmospheric nitrogen deposition positive biomass responses to CO₂ enrichment of the currently dominating species are unlikely.