A genetic link between leaf carbon isotope composition and whole‐plant water use efficiency in the C4 grass Setaria

Genetic selection for whole‐plant water use efficiency (yield per transpiration; WUE_plant) in any crop‐breeding programme requires high‐throughput phenotyping of component traits of WUE_plant such as intrinsic water use efficiency (WUE_i; CO₂ assimilation rate per stomatal conductance). Measuring WUE_i by gas exchange measurements is laborious and time consuming and may not reflect an integrated WUE_i over the life of the leaf. Alternatively, leaf carbon stable isotope composition (δ¹³C_leaf) has been suggested as a potential time‐integrated proxy for WUE_i that may provide a tool to screen for WUE_plant. However, a genetic link between δ¹³C_leaf and WUE_plant in a C₄ species has not been well established. Therefore, to determine if there is a genetic relationship in a C₄ plant between δ¹³C_leaf and WUE_plant under well watered and water‐limited growth conditions, a high‐throughput phenotyping facility was used to measure WUE_plant in a recombinant inbred line (RIL) population created between the C₄ grasses Setaria viridis and S. italica. Three quantitative trait loci (QTL) for δ¹³C_leaf were found and co‐localized with transpiration, biomass accumulation, and WUE_plant. Additionally, WUE_plant for each of the δ¹³C_leaf QTL allele classes was negatively correlated with δ¹³C_leaf, as would be predicted when WUE_i influences WUE_plant. These results demonstrate that δ¹³C_leaf is genetically linked to WUE_plant, likely to be through their relationship with WUE_i, and can be used as a high‐throughput proxy to screen for WUE_plant in these C₄ species.     

Aerial decay influence on the stable oxygen and carbon isotope ratios in tree ring cellulose

Sub-fossil wood is often affected by the decaying process that introduces uncertainties in the measurement of oxygen and carbon stable isotope composition in cellulose. Although the cellulose stable isotopes are widely used as climatic proxies, our understanding of processes controlling their behavior is very limited. We present here a comparative study of stable oxygen and carbon isotope ratios in tree ring cellulose in decayed and non-decayed wood samples of Swiss stone pine (Pinus cembra) trees. The intra-ring stable isotope variability (around the circumference of a single ring) was between 0.1 and 0.5‰ for δ¹⁸O values and between 0.5 and 1.6‰ for δ¹³C values for both decayed and non-decayed wood. Observed intra-tree δ¹⁸O variability is less than that reported in the literature (0.5–1.5‰), however, for δ¹³C it is larger than the reported values (0.7–1.2‰). The inter-tree variability for non-decayed wood ranges between 1.1 and 2.3‰ for δ¹⁸O values, and between 2 and 4.7‰ for δ¹³C values. The inter-tree differences for δ¹⁸O values are similar to those reported in the literature (1–2‰ for oxygen and 1–3‰ for carbon) but are larger for δ¹³C values. We have found that the differences for δ¹⁸O and δ¹³C values between decayed and non-decayed wood are smaller than the variation among different trees from the same site, suggesting that the decayed wood can be used for isotopic paleoclimate research.     

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.     

Water-use efficiency of a mallee eucalypt growing naturally and in short-rotation coppice cultivation

This study compared mature Eucalyptus kochii subsp. plenissima trees in inner regions or edges of natural bushland to young trees belt-planted through cleared agricultural land as uncut saplings or regenerating coppice over 2.7 years at Kalannie, Western Australia (320 mm annual rainfall). We assessed the ability of the species to alter its gas exchange characteristics, leaf physical attributes, and water-use efficiency of foliar carbon assimilation (WUE _i) or of total dry matter production (WUE _DM). Stomatal conductance (g _s) varied five-fold between treatment means, with coppices exhibiting greatest values and mature bush least. Photosynthetic rates followed this trend. Leaf photosynthetic capacity estimated by chlorophyll content varied 1.3-fold parallel with variations in leaf thickness, with coppices rating lowest and mature edge trees most highly. WUE _i varied 1.5-fold between treatments and was greatest in mature inner-bush and edge trees. Leaf photosynthetic capacity and g _s were both correlated with WUE _i. Carbon isotope composition (δ¹³C values) of new shoot dry matter produced early in a seasonal flush were similar to those of root starch but when averaged over the whole season correlated well with WUE _i and gas exchange characteristics of trees of each treatment. Coppices showed poorest WUE _i and most negative shoot tip δ¹³C but their WUE _DM was high. This discrepancy was suggested to relate to carbon allocation strategies in coppices favouring fast growth of replacement shoots but not of roots. Physiology of coppice growth of E. kochii is usefully geared towards both rapid and water-use efficient production of woody biomass in water limited environments.     

Photosynthesis, water use efficiency and stable carbon isotope composition are associated with anatomical properties of leaf and xylem in six poplar species

Although fast‐growing Populus species consume a large amount of water for biomass production, there are considerable variations in water use efficiency (WUE) across different poplar species. To compare differences in growth, WUE and anatomical properties of leaf and xylem and to examine the relationship between photosynthesis/WUE and anatomical properties of leaf and xylem, cuttings of six poplar species were grown in a botanical garden. The growth performance, photosynthesis, intrinsic WUE (WUE_i), stable carbon isotope composition (δ¹³C) and anatomical properties of leaf and xylem were analysed in these poplar plants. Significant differences were found in growth, photosynthesis, WUE_i and anatomical properties among the examined species. Populus cathayana was the clone with the fastest growth and the lowest WUE_i/δ¹³C, whereas P. × euramericana had a considerable growth increment and the highest WUE_i/δ¹³C. Among the analysed poplar species, the highest total stomatal density in P. cathayana was correlated with its highest stomatal conductance (g_s) and lowest WUE_i/δ¹³C. Moreover, significant correlations were observed between WUE_i and abaxial stomatal density and stem vessel lumen area. These data suggest that photosynthesis, WUE_i and δ¹³C are associated with leaf and xylem anatomy and there are tradeoffs between growth and WUE_i. It is anticipated that some poplar species, e.g. P. × euramericana, are better candidates for water‐limited regions and others, e.g. P. cathayana, may be better for water‐abundant areas.     

Stable carbon isotopes in tree rings indicate improved water use efficiency and drought responses of a tropical dry forest tree species

Understanding the responses of tropical trees to increasing [CO₂] and climate change is important as tropical forests play an important role in carbon and hydrological cycles. We used stable carbon isotopes (δ¹³C) in tree rings to study the physiological responses of a tropical dry forest tree species in southern Mexico, Mimosa acantholoba to changes in atmospheric [CO₂] and variation in climate. Based on annual records of tree ring δ¹³C, we calculated intrinsic water use efficiency (W _i) and intercellular [CO₂] (c _i). Our results showed that trees responded strongly to the increase in atmospheric [CO₂] over the last four decades; W _i increased dramatically by 40%, while c _i remained largely constant. The maintenance of a constant c _i indicates that photosynthetic rates are unlikely to have increased in response to higher [CO₂], and that improvements in W _i are probably due to a reduction in stomatal conductance. This may have large consequences for the hydrological cycle. Inter-annual variation in c _i was strongly correlated with total annual rainfall (r = 0.70), and not influenced by temperature, solar radiation or cloud cover. Our results show that δ¹³C in tree rings of tropical dry forest trees may be a powerful tool to evaluate long-term responses of trees to increasing [CO₂] and to variation in climate.     

More than climate: Hydrogen isotope ratios in tree rings as novel plant physiological indicator for stress conditions

The analysis of stable carbon and oxygen isotopes in tree-rings is a widely applied tool which allows to retrieve information about past climatic conditions, as well as tree physiological responses to environmental changes. This is based on well-established mechanistic models and firm statistical relationships with climate variables. In contrast, the hydrogen isotopic signature (δ²H) of tree-rings has been reported to be poorly correlated to climate or difficult to explain, and as a consequence, hydrogen isotopes are far less utilized. However, recent plant-physiological experiments have highlighted the role of autotrophic versus heterotrophic processes affecting δ²H values, i.e. use of fresh assimilates versus stored carbohydrates, and have much improved our understanding of the role of post-photosynthetic ²H-fractionation. Using unpublished and literature δ²H data of tree-ring cellulose (δ²H_C) of 5 study sites in Europe and Asia, we systematically investigated the relationships between δ²H_C and tree-ring width (TRW), which, in contrast to previous research, could now be explained through post-photosynthetic ²H-fractionation. In most cases, these relationships were found to be negative (r² = 0.23 to 0.51, all P < 0.05) when the main growth limiting factors are precipitation and light, while in temperature-limited sites we observed a positive trend (r² = 0.14, P < 0.05). Our results suggest that, under stress conditions, trees use a surplus of carbon from reserves for wood formation. Therefore, in combination with TRW chronologies, δ²H_C may allow to infer about physiological information on stressful time periods independently of biotic and abiotic origin. Here, we discuss implications of these findings for tree-ring research, summarize them in a conceptual framework and suggest future research directions.     

Compiled records of atmospheric CO2 concentrations and stable carbon isotopes to reconstruct climate and derive plant ecophysiological indices from tree rings

The stable carbon isotopic composition (δ¹³C) measured in tree rings is a standard proxy for paleoclimate reconstructions and is increasingly being used as a paleophysiological proxy. To fully exploit the potential of tree ring δ¹³C proxy, atmospheric CO₂ concentration and δ¹³C (δ¹³CO₂) data are required to correct tree ring δ¹³C from the declining trend of δ¹³CO₂ due to fossil fuel burning since 1850 CE, and to derive physiological parameters using biochemical models that link photosynthesis to δ¹³C. These atmospheric data are available from direct measurements or can be inferred from indirect proxies such as ice cores covering the Common Era (CE) at variable temporal resolutions. For almost two decades, tree-ring researchers have relied on a dataset derived from fitted linear regressions of ice core measurements available through the seminal McCarroll and Loader (2004) article for the 1850−2003 CE period. However, new calibrations and compilations of ice core data and direct measurements are now available as part of Earth System Modelling efforts which remain overlooked by the tree ring research community.Here, we present an overview of the new and freely available datasets and provide recommendations for their use in ecophysiology and paleoclimate research, that we expect will stimulate cross-disciplinary collaborations.     

Dendroecological and genetic insights for future management of an old-planted forest of the endangered Mediterranean fir Abies pinsapo

Pinsapo fir (Abies pinsapo Boiss.) is an endangered Mediterranean conifer that has raised strong conservation interest as a paradigmatic example of species characterized by small and fragmented populations. We studied an old reforestation stand composed of A. pinsapo, Pinus nigra and Pinus sylvestris established in the 1910s in central-eastern Spain (about 500 km north of the species native distribution range), with the aim of evaluating the stand’s suitability as an ex situ conservation unit for the fir. To this end, we investigated the long-term performance of the stand and assessed genetic diversity of A. pinsapo. Tree-ring width (TRW) and carbon isotope discrimination (Δ¹³C) were used to characterise growth dynamics and intrinsic water-use efficiency (WUE_i), respectively. Furthermore, 42 pinsapo firs were genotyped at five microsatellite loci to compare their genetic variation with published data on natural populations. A. pinsapo showed ca. two-fold higher radial growth than pines in the last 80 years; however, a growth decrease was observed for all species from the 1990s onwards. Indexed TRW was positively associated with Δ¹³C at the species level, denoting inter-annual growth dependence on water availability. Overall, Δ¹³C was higher for A. pinsapo compared to pines, indicating lower WUE_i, but Δ¹³C significantly decreased over the last 50 years for all species, likely as the result of tighter stomatal regulation of water loss, resulting in WUE_i increases of about 25 %. Recently, however, A. pinsapo showed reduced WUE_i increase in concord with growth slowdown, suggesting a threshold response for stomatal regulation. Although genetic diversity of A. pinsapo was about half of natural populations, the old-planted stand could be important for the conservation of this endemic species considering its good long-term growth and physiology. The latest decrease in performance of A. pinsapo, however, asks for urgent management measures aimed at reducing the competition for water and promoting growth and natural regeneration. This study illustrates the potential of combining tree-ring-based long-term physiological information with genetic data to ascertain the prospects of artificial stands for conservation purposes.     

Temporal instability of isotopes–climate statistical relationships – A study of black spruce trees in northeastern Canada

Climate reconstructions using stable isotopes (δ¹⁸O and δ¹³C values) in tree rings are based on relationships between present climatic conditions and isotopic series. This widely used approach relies on the assumption that correlations between stable isotopes and climatic conditions are steady over time. In this paper, we evaluate the strength of the correlations between δ¹⁸O and δ¹³C series with several climatic parameters on fourteen black spruce trees coming from three different sites, in northeastern Canada. We applied a 21-year moving window on the r Pearson calculated between stable isotopes and March–May and June–August precipitation, June–August and April–June maximal temperatures. Our results indicate that despite the large distance and differences in stand conditions between the sites, the three sites responded in the same way over time. We show that because the climatic ambiance has changed during the 1980–1990 period due to a positive North Atlantic Oscillation index the δ¹³C values are not controlled anymore by spring precipitation or summer maximal temperature in the following two decades. As opposed to δ¹³C series, the relationship between summer maximal temperature and δ¹⁸O values was stable over time, and decreased only in the last decade. All these results attest of a “divergence problem” in the last decades which is most pronounced for δ¹³C series. We conclude that the spruce δ¹⁸O series appears to be the most appropriate indicator for reconstructing June–August maximal temperature in the studied area despite the divergence issue, given that the calibration–validation tests and reconstruction can exclude the divergent last decade.     

The legacy of enhanced N and S deposition as revealed by the combined analysis of δ13C, δ18O and δ15N in tree rings

This study aimed to evaluate the effects of long‐term repeated aerial nitrogen (N) and sulphur (S) misting over tree canopies of a Sitka spruce plantation in Scotland. We combined δ¹³C and δ¹⁸O in tree rings to evaluate the changes in CO₂ assimilation (A) and stomatal conductance (g_s) and to assess their contribution to variations in the intrinsic water‐use efficiency (WUE_i, i.e., the A/g_s ratio). Measurements of δ¹⁵N enabled shifts in the ecosystem N cycling following misting to be assessed. We found that: (i) N applications, with or without S, increased the ratio between A and g_s in favour of A, thus supporting a fertilizer effect of added N. (ii) After the treatments ceased, the trees quickly adjusted to the reductions of N deposition, but not to the reduction in S deposition, which had a negative effect on WUE_i by reducing A. This indicates that the beneficial role of N deposition may be negated in forests that previously received a high load of acid rain. (iii) δ¹⁵N in tree rings reflected the N dynamics caused by canopy retention, with the fingerprint also present in the litter, after the experiment stopped. (iv) Both our results (obtained using canopy applications) and a collection of published data (obtained using soil applications) showed that generally WUE_i increased in response to an increase of N applications, with the magnitude of the changes related to soil conditions and the availability of other nutrients. The shifts observed in δ¹⁵N in tree rings also suggest that both the quantity of the applied N and its quality, mediated by processes occurring during canopy N retention, are important determinants of the interactions between N and C cycles. Stable isotopes are useful probes to understand these processes and to put the results of short‐term experiments into context.     

Water-use responses of ��living fossil�� conifers to CO2 enrichment in a simulated Cretaceous polar environment

Background and Aims

During the Mesozoic, the polar regions supported coniferous forests that experienced warm climates, a CO₂-rich atmosphere and extreme seasonal variations in daylight. How the interaction between the last two factors might have influenced water use of these conifers was investigated. An experimental approach was used to test the following hypotheses: (1) the expected beneficial effects of elevated [CO₂] on water-use efficiency (WUE) are reduced or lost during the 24-h light of the high-latitude summer; and (2) elevated [CO₂] reduces plant water use over the growing season.

Methods

Measurements of leaf and whole-plant gas exchange, and leaf-stable carbon isotope composition were made on one evergreen (Sequoia sempervirens) and two deciduous (Metasequoia glyptostroboides and Taxodium distichum) ‘living fossil’ coniferous species after 3 years'' growth in controlled-environment simulated Cretaceous Arctic (69°N) conditions at either ambient (400 µmol mol⁻¹) or elevated (800 µmol mol⁻¹) [CO₂].

Key Results

Stimulation of whole-plant WUE (WUE_P) by CO₂ enrichment was maintained over the growing season for the three studied species but this pattern was not reflected in patterns of WUE inferred from leaf-scale gas exchange measurements (iWUE_L) and δ¹³C of foliage (tWUE_L). This response was driven largely by increased rates of carbon uptake, because there was no overall CO₂ effect on daily whole-plant transpiration or whole-plant water loss integrated over the study period. Seasonal patterns of tWUE_L differed from those measured for iWUE_L. The results suggest caution against over simplistic interpretations of WUE_P based on leaf isotopic composition.

Conclusions

The data suggest that the efficiency of whole-tree water use may be improved by CO₂ enrichment in a simulated high-latitude environment, but that transpiration is relatively insensitive to atmospheric CO₂ in the living fossil species investigated.Key words: Water-use efficiency, elevated CO₂, living fossil plants, conifers, paleoecology, ancient polar forests, stable carbon isotopes, stomatal conductance, canopy transpiration     

Isotopic and anatomical signals for interpreting fire-related responses in Pinus halepensis

Key message

Our study aims to define isotopic and anatomical responses to fires of P. halepensis . Main results: decrease in tree growth and relative conductivity and increase in water use efficiency.

Abstract

We investigated the ecophysiological responses of a Pinus halepensis Mill. stand surviving two wildfires in southern France. Basal area, isotope composition and anatomical traits were analysed before and after fires, using tree rings to assess the ecological responses of trees to heat-related damage. The years were determined based on the presence of fire scars. Stable isotopes (δ¹³C and δ¹⁸O) were measured in tree rings before and after the “fire years”. Anatomical observations allowed qualitative analysis of the scar region and quantification of tracheid size in tree rings before and after the fire years. Relative and percentage conductivity of earlywood and latewood far from the woundwood were estimated. Results showed a decrease in tree growth after the fire events accompanied by an increase in ¹³C-derived water use efficiency (WUE_i) and a decrease in relative conductivity. The positive relationship between δ¹³C and δ¹⁸O suggested that both isotopic variations are mostly driven by changes in stomatal conductance following fire events. P. halepensis proved to be a strong isohydric species, able to survive frequent fires with temporary ecophysiological modifications and anatomical adaptations. Our findings afford new insights into post-fire survival strategies of this species in an environment where fires are predicted to increase in frequency during the twenty-first century.     

Bimodality in stable isotope composition facilitates the tracing of carbon transfer from macrophytes to higher trophic levels

Even though the suitability of macrophytes to act as a carbon source to food webs has been questioned by some studies, some others indicate that macrophyte-derived carbon may play an important role in the trophic transfer of organic matter in the food web of shallow lakes. To evaluate the importance of macrophytes to food webs, we collected primary producers—macrophytes and periphyton—and consumers from 19 South American shallow lakes and analyzed their carbon stable isotopes composition (δ¹³C). Despite the diversity of inorganic carbon sources available in our study lakes, the macrophytes’ δ¹³C signatures showed a clear bimodal distribution: ¹³C-depleted and ¹³C-enriched, averaging at ?27.2 and ?13.5‰, respectively. We argue that the use of either CO₂ or HCO₃ ^? by the macrophytes largely caused the bimodal pattern in δ¹³C signals. The contribution of carbon from macrophytes to the lake’s food webs was not straightforward in most of the lakes because the macrophytes’ isotopic composition was quite similar to the isotopic composition of periphyton, phytoplankton, and terrestrial carbon. However, in some lakes where the macrophytes had a distinct isotopic signature, our data suggest that macrophytes can represent an important carbon source to shallow lake food webs.     

Fingerprinting environmental conditions and related stress using stable isotopic composition of rice (Oryza sativa L.) grain organic matter

Sea level rise (SLR) is a primary factor responsible for inundation of low-lying coastal regions across the world, which in turn governs the agricultural productivity. In this study, rice (Oryza sativa L.) cultivated seasonally in the Kuttanad Wetland, a SLR prone region on the southwest coast of India, were analysed for oxygen, hydrogen and carbon isotopic ratios (δ¹⁸O, δ²H and δ¹³C) to distinguish the seasonal environmental conditions prevalent during rice cultivation. The region receives high rainfall during the wet season which promotes large supply of fresh water to the local water bodies via the rivers. In contrast, during the dry season reduced river discharge favours sea water incursion which adversely affects the rice cultivation. The water for rice cultivation is derived from regional water bodies that are characterised by seasonal salinity variation which co-varies with the δ¹⁸O and δ²H values. Rice cultivated during the wet and the dry season bears the isotopic imprints of this water. We explored the utility of a mechanistic model to quantify the contribution of two prominent factors, namely relative humidity and source water composition in governing the seasonal variation in oxygen isotopic composition of rice grain OM. δ¹³C values of rice grain OM were used to deduce the stress level by estimating the intrinsic water use efficiency (WUE_i) of the crop during the two seasons. 1.3 times higher WUE_i was exhibited by the same genotype during the dry season. The approach can be extended to other low lying coastal agro-ecosystems to infer the growth conditions of cultivated crops and can further be utilised for retrieving paleo-environmental information from well preserved archaeological plant remains.     

Carbon isotope compositions (δ13C) of leaf,wood and holocellulose differ among genotypes of poplar and between previous land uses in a short‐rotation biomass plantation

The efficiency of water use to produce biomass is a key trait in designing sustainable bioenergy‐devoted systems. We characterized variations in the carbon isotope composition (δ¹³C) of leaves, current year wood and holocellulose (as proxies for water use efficiency, WUE) among six poplar genotypes in a short‐rotation plantation. Values of δ¹³C_wood and δ¹³C_{holocellulose} were tightly and positively correlated, but the offset varied significantly among genotypes (0.79–1.01‰). Leaf phenology was strongly correlated with δ¹³C, and genotypes with a longer growing season showed a higher WUE. In contrast, traits related to growth and carbon uptake were poorly linked to δ¹³C. Trees growing on former pasture with higher N‐availability displayed higher δ¹³C as compared with trees growing on former cropland. The positive relationships between δ¹³C_leaf and leaf N suggested that spatial variations in WUE over the plantation were mainly driven by an N‐related effect on photosynthetic capacities. The very coherent genotype ranking obtained with δ¹³C in the different tree compartments has some practical outreach. Because WUE remains largely uncoupled from growth in poplar plantations, there is potential to identify genotypes with satisfactory growth and higher WUE.     

Species-dependent responses of juniper and spruce to increasing CO<Subscript>2</Subscript> concentration and to climate in semi-arid and arid areas of northwestern China

Qilian juniper (Sabina przewalskii Kom.) and Qinghai spruce (Picea crassifolia Kom.) represent different tree functional types, which can be found extensively in northwestern China. The former is drought-tolerant, whereas the latter is hygrophilous and shade-tolerant. We compared their intrinsic water-use efficiency (iWUE, inferred from carbon isotopic discrimination, δ¹³C, in their wood) as a function of atmospheric CO₂ concentration, [CO₂], and climate. δ¹³C of spruce was consistently about higher than that of juniper in semi-arid areas but was lower in arid areas. This difference was stable over time and demonstrated strong cross-correlations between species, although some subtle high-frequency (2 or 3 years) variations existed in both species, suggesting that regional climate may control carbon isotope discrimination. The ratio (the [CO₂] values in leaf intercellular and the atmosphere, respectively) of the juniper increased steadily over time, whereas that of the spruce showed a long-term downward trend. IWUE increased at all sites over the 150-year study period, mainly caused by increasing [CO₂]. The relationship between iWUE and [CO₂] reveals that the spruce was more sensitive than the juniper to increasing [CO₂], suggesting a species-specific adaptation to long-term environmental changes. Correlations between the high-frequency variations in stable carbon discrimination (Δ) and climate indicate similar intra-site responses to climate in both species, but different response strengths. Overall, complex interactions of temperature and moisture on stable carbon discrimination during current growth seasons of both species were environmental-determined. Regulation of gas exchange and reduced transpiration may influence water and energy budgets directly; therefore species-dependent responses of trees to elevated CO₂ should be considered in future research on global plant physiological ecology.     

Comparing the performance of different stomatal conductance models using modelled and measured plant carbon isotope ratios (δ13C): implications for assessing physiological forcing

Accurate modelling of long‐term changes in plant stomatal functioning is vital to global climate change studies because changes in evapotranspiration influence temperature via physiological forcing of the climate. Various stomatal models are included in land surface schemes, but their robustness over longer timescales is difficult to validate. We compare the performance of three stomatal models, varying in their degree of complexity, and coupled to a land surface model. This is carried out by simulating the carbon isotope ratio of tree leaves (δ¹³C_leaf) over a period of 53 years, and comparing the results with carbon isotope ratios obtained from tree rings (δ¹³C_stem) measured at six sites in northern Europe. All three stomatal models fail to capture the observed interannual variability in the measured δ¹³C_stem time series. However, the Soil‐Plant‐Atmosphere (SPA) model performs significantly better than the Ball‐Berry (BB) or COX models when tested for goodness‐of‐fit against measured δ¹³C_stem. The δ¹³C_leaf time series simulated using the SPA model are significantly positively correlated (P < 0.05) with measured results over the full time period tested, at all six sites. The SPA model underestimates interannual variability measured in δ¹³C_stem, but is no worse than the BB model and significantly better than the COX model. The inability of current models to adequately replicate changes in stomatal response to rising levels of CO₂ concentrations, and thus to quantify the associated physiological forcing, warrants further investigation.     

Climatic information recorded in stable carbon isotopes in tree rings of Cryptomeria fortunei,Tianmu Mountain,China

The Cryptomeria fortunei (CF) tree-ring δ¹³C_p series, which was collected from the West Tianmu Mountain forestland (30°20′ N, 119°26′ E), located in the north-west of Zhejiang Province, China, belonging to the northern margin of the mid-subtropical region of Eastern China, were determined based on cross-dated tree-ring age. There was a significant decline in the δ¹³C_p series occurring from 1685 to 1985, more especially from 1835 to 1985 in response to increasing atmospheric CO₂ concentrations and decreasing atmospheric δ¹³C_a. To reduce the noise and enhance the climatic signals, we compared the polynomial function with the correction method developed by McCarroll and Loader (2004) to remove the low-frequency variation in the raw tree ring δ¹³C_p series (defined as the δ¹³C_poly series, δ¹³C_cor series, respectively), and found the most suited correction method was the correction method developed by McCarroll and Loader (2004) in our study area. High-frequency correlation analysis between the δ¹³C_cor series and many meteorological parameters recorded by Xian Rending weather station revealed that the current August–September mean maximum temperature and previous year mean minimum and mean maximum temperature (P < 0.005) most strongly influenced tree ring δ¹³C_p discrimination from 1956 to 1985, and the strongest temperature signal captured was the current August–September mean maximum temperature (r = 0.54, P < 0.005). Mainly on this basis, the varied history of current August–September mean maximum temperatures in the West Tianmu Mountain area were reconstructed from 1685 to 1985. The reconstructed maximum temperatures revealed a slight warming trend and showed close correlation with the climatic fluctuations of the Little Ice Age cold period before 1900 as well as the 20th century warm period after 1900. It also better corresponded with some climate events recorded in historical records. Spectrum analysis showed that in the reconstructed series there was quasi-periodicity of 66.7 yr, 21.1 yr, 3.2 yr, 2.3 yr and 2.0 yr. These cycles coincided with the “torque effect” variation of planets and the geocentric convergence, and changes in solar activity and irradiance, as well as the “Quasi-biennial oscillation” (QBO). This indicated that the δ¹³C_p chronology of tree rings in West Tianmu Mountain showed a good record of the sun's activities, the change in the sun radiation and ENSO events.     

Do centennial tree-ring and stable isotope trends of Larix gmelinii (Rupr.) Rupr. indicate increasing water shortage in the Siberian north?

Tree-ring width of Larix gmelinii (Rupr.) Rupr., ratios of stable isotopes of C (δ¹³C) and O (δ¹⁸O) of whole wood and cellulose chronologies were obtained for the northern part of central Siberia (Tura, Russia) for the period 1864–2006. A strong decrease in the isotope ratios of O and C (after atmospheric δ¹³C corrections) and tree-ring width was observed for the period 1967–2005, while weather station data show a decrease in July precipitation, along with increasing July air temperature and vapor pressure deficit (VPD). Temperature at the end of May and the whole month of June mainly determines tree radial growth and marks the beginning of the vegetation period in this region. A positive correlation between tree-ring width and July precipitation was found for the calibration period 1929–2005. Positive significant correlations between C isotope chronologies and temperatures of June and July were found for whole wood and cellulose and negative relationships with July precipitation. These relationships are strengthened when the likely physiological response of trees to increased CO₂ is taken into account (by applying a recently developed δ¹³C correction). For the O isotope ratios, positive relationships with annual temperature, VPD of July and a negative correlation with annual precipitation were observed. The δ¹⁸O in tree rings may reflect annual rather than summer temperatures, due to the late melting of the winter snow and its contribution to the tree water supply in summer. We observed a clear change in the isotope and climate trends after the 1960s, resulting in a drastic change in the relationship between C and O isotope ratios from a negative to a positive correlation. According to isotope fractionation models, this indicates reduced stomatal conductance at a relatively constant photosynthetic rate, as a response of trees to water deficit for the last half century in this permafrost region.