Restricted internal diffusion weakens transpiration-photosynthesis coupling during heatwaves: Evidence from leaf carbonyl sulphide exchange.

Increasingly frequent and intense heatwaves threaten ecosystem health in a warming climate. However, plant responses to heatwaves are poorly understood. A key uncertainty concerns the intensification of transpiration when heatwaves suppress photosynthesis, known as transpiration-photosynthesis decoupling. Field observations of such decoupling are scarce, and the underlying physiological mechanisms remain elusive. Here, we use carbonyl sulphide (COS) as a leaf gas exchange tracer to examine potential mechanisms leading to transpiration-photosynthesis decoupling on a coast live oak in a southern California woodland in spring 2013. We found that heatwaves suppressed both photosynthesis and leaf COS uptake but increased transpiration or sustained it at non-heatwave levels throughout the day. Despite statistically significant decoupling between transpiration and photosynthesis, stomatal sensitivity to environmental factors did not change during heatwaves. Instead, midday photosynthesis during heatwaves was restricted by internal diffusion, as indicated by the lower internal conductance to COS. Thus, increased evaporative demand and nonstomatal limitation to photosynthesis act jointly to decouple transpiration from photosynthesis without altering stomatal sensitivity. Decoupling offered limited potential cooling benefits, questioning its effectiveness for leaf thermoregulation in xeric ecosystems. We suggest that adding COS to leaf and ecosystem flux measurements helps elucidate diverse physiological mechanisms underlying transpiration-photosynthesis decoupling.

Influences of light and humidity on carbonyl sulfide-based estimates of photosynthesis.

Understanding climate controls on gross primary productivity (GPP) is crucial for accurate projections of the future land carbon cycle. Major uncertainties exist due to the challenge in separating GPP and respiration from observations of the carbon dioxide (CO2) flux. Carbonyl sulfide (COS) has a dominant vegetative sink, and plant COS uptake is used to infer GPP through the leaf relative uptake (LRU) ratio of COS to CO2 fluxes. However, little is known about variations of LRU under changing environmental conditions and in different phenological stages. We present COS and CO2 fluxes and LRU of Scots pine branches measured in a boreal forest in Finland during the spring recovery and summer. We find that the diurnal dynamics of COS uptake is mainly controlled by stomatal conductance, but the leaf internal conductance could significantly limit the COS uptake during the daytime and early in the season. LRU varies with light due to the differential light responses of COS and CO2 uptake, and with vapor pressure deficit (VPD) in the peak growing season, indicating a humidity-induced stomatal control. Our COS-based GPP estimates show that it is essential to incorporate the variability of LRU with environmental variables for accurate estimation of GPP on ecosystem, regional, and global scales.

COSORE: A community database for continuous soil respiration and other soil-atmosphere greenhouse gas flux data.

Globally, soils store two to three times as much carbon as currently resides in the atmosphere, and it is critical to understand how soil greenhouse gas (GHG) emissions and uptake will respond to ongoing climate change. In particular, the soil-to-atmosphere CO2 flux, commonly though imprecisely termed soil respiration (RS ), is one of the largest carbon fluxes in the Earth system. An increasing number of high-frequency RS measurements (typically, from an automated system with hourly sampling) have been made over the last two decades; an increasing number of methane measurements are being made with such systems as well. Such high frequency data are an invaluable resource for understanding GHG fluxes, but lack a central database or repository. Here we describe the lightweight, open-source COSORE (COntinuous SOil REspiration) database and software, that focuses on automated, continuous and long-term GHG flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation. Contributed datasets are mapped to a single, consistent standard, with metadata on contributors, geographic location, measurement conditions and ancillary data. The design emphasizes the importance of reproducibility, scientific transparency and open access to data. While being oriented towards continuously measured RS , the database design accommodates other soil-atmosphere measurements (e.g. ecosystem respiration, chamber-measured net ecosystem exchange, methane fluxes) as well as experimental treatments (heterotrophic only, etc.). We give brief examples of the types of analyses possible using this new community resource and describe its accompanying R software package.

Sources and sinks of carbonyl sulfide in an agricultural field in the Southern Great Plains.

Net photosynthesis is the largest single flux in the global carbon cycle, but controls over its variability are poorly understood because there is no direct way of measuring it at the ecosystem scale. We report observations of ecosystem carbonyl sulfide (COS) and CO2 fluxes that resolve key gaps in an emerging framework for using concurrent COS and CO2 measurements to quantify terrestrial gross primary productivity. At a wheat field in Oklahoma we found that in the peak growing season the flux-weighted leaf relative uptake of COS and CO2 during photosynthesis was 1.3, at the lower end of values from laboratory studies, and varied systematically with light. Due to nocturnal stomatal conductance, COS uptake by vegetation continued at night, contributing a large fraction (29%) of daily net ecosystem COS fluxes. In comparison, the contribution of soil fluxes was small (1-6%) during the peak growing season. Upland soils are usually considered sinks of COS. In contrast, the well-aerated soil at the site switched from COS uptake to emissions at a soil temperature of around 15 °C. We observed COS production from the roots of wheat and other species and COS uptake by root-free soil up to a soil temperature of around 25 °C. Our dataset demonstrates that vegetation uptake is the dominant ecosystem COS flux in the peak growing season, providing support of COS as an independent tracer of terrestrial photosynthesis. However, the observation that ecosystems may become a COS source at high temperature needs to be considered in global modeling studies.

Resilience to seasonal heat wave episodes in a Mediterranean pine forest.

Short-term, intense heat waves (hamsins) are common in the eastern Mediterranean region and provide an opportunity to study the resilience of forests to such events that are predicted to increase in frequency and intensity. The response of a 50-yr-old Aleppo pine (Pinus halepensis) forest to hamsin events lasting 1-7 d was studied using 10 yr of eddy covariance and sap flow measurements. The highest frequency of heat waves was c. four per month, coinciding with the peak productivity period (March-April). During these events, net ecosystem carbon exchange (NEE) and canopy conductance (gc ) decreased by c. 60%, but evapotranspiration (ET) showed little change. Fast recovery was also observed with fluxes reaching pre-stress values within a day following the event. NEE and gc showed a strong response to vapor pressure deficit that weakened as soil moisture decreased, while sap flow was primarily responding to changes in soil moisture. On an annual scale, heat waves reduced NEE and gross primary productivity by c. 15% and 4%, respectively. Forest resilience to short-term extreme events such as heat waves is probably a key to its survival and must be accounted for to better predict the increasing impact on productivity and survival of such events in future climates.

Farmer preference for macadamia varieties and constraints to production in Malawi.

Macadamia nuts constitute a vital component of both nutrition and livelihoods for smallholder producers in Malawi. We conducted a comprehensive mixed-methods study, combining qualitative and quantitative analyses, to explore varietal preferences and production challenges among these farmers. Leveraging cross-sectional data from 144 members of the Highlands Macadamia Cooperative Union Limited, our study underscores several significant findings. Our findings reveal that the majority of smallholder macadamia farmers (62%) are aged over 50, with farming as their primary occupation. Varied preferences are driven by yield-related traits, including high yield potential (38%), nut quality (29%), and extended flowering patterns (15%). Among the macadamia varieties, the top five choices, grown by over half of the farmers, include HAES 660 (18%), 800 (10%), 791 (9%), 816 (8%), and 246 (7%). Additionally, our study identifies five primary constraints faced by smallholder macadamia farmers: insect pests (81%), diseases (34%), limited market access (33%), wind damage (25%), and inadequate agricultural advisory services (17%). Based on these findings, we propose two policy recommendations to enhance smallholder macadamia production and productivity in Malawi and other regions. Specifically, we advocate for informed breeding programs that align with farmer preferences to promote greater adoption of macadamia varieties. Additionally, we emphasize the crucial role of the Malawian government in the macadamia value chain, suggesting active participation in providing extension services and marketing support, akin to its support for other cash crops.

Variability of 13C-labeling in plant leaves.

RATIONALE: Plant tissues artificially labeled with (13)C are increasingly used in environmental studies to unravel biogeochemical and ecophysiological processes. However, the variability of (13)C-content in labeled tissues has never been carefully investigated. Hence, this study aimed at documenting the variability of (13)C-content in artificially labeled leaves. METHODS: European beech and Italian ryegrass were subjected to long-term (13)C-labeling in a controlled-environment growth chamber. The (13)C-content of the leaves obtained after several months labeling was determined by isotope ratio mass spectrometry. RESULTS: The (13)C-content of the labeled leaves exhibited inter- and intra-leaf variability much higher than those naturally occurring in unlabeled plants, which do not exceed a few per mil. This variability was correlated with labeling intensity: the isotope composition of leaves varied in ranges of ca 60‰ and 90‰ for experiments that led to average leaf (13)C-content of ca +15‰ and +450‰, respectively. CONCLUSIONS: The reported variability of isotope composition in (13)C-enriched leaves is critical, and should be taken into account in subsequent experimental investigations of environmental processes using (13)C-labeled plant tissues.

Reducing and correcting for contamination of ecosystem water stable isotopes measured by isotope ratio infrared spectroscopy.

Concern exists about the suitability of laser spectroscopic instruments for the measurement of the (18)O/(16)O and (2)H/(1)H values of liquid samples other than pure water. It is possible to derive erroneous isotope values due to optical interference by certain organic compounds, including some commonly present in ecosystem-derived samples such as leaf or soil waters. Here we investigated the reliability of wavelength-scanned cavity ring-down spectroscopy (CRDS) (18)O/(16)O and (2)H/(1)H measurements from a range of ecosystem-derived waters, through comparison with isotope ratio mass spectrometry (IRMS). We tested the residual of the spectral fit S(r) calculated by the CRDS instrument as a means to quantify the difference between the CRDS and IRMS δ-values. There was very good overall agreement between the CRDS and IRMS values for both isotopes, but differences of up to 2.3‰ (δ(18)O values) and 23‰ (δ(2)H values) were observed in leaf water extracts from Citrus limon and Alnus cordata. The S(r) statistic successfully detected contaminated samples. Treatment of Citrus leaf water with activated charcoal reduced, but did not eliminate, δ(2)H(CRDS) - δ(2)H(IRMS) linearly for the tested range of 0-20% charcoal. The effect of distillation temperature on the degree of contamination was large, particularly for δ(2)H values but variable, resulting in positive, negative or no correlation with distillation temperature. S(r) and δ(CRDS) - δ(IRMS) were highly correlated, in particular for δ(2)H values, across the range of samples that we tested, indicating the potential to use this relationship to correct the δ-values of contaminated plant water extracts. We also examined the sensitivity of the CRDS system to changes in the temperature of its operating environment. We found that temperature changes ≥4 °C for δ(18)O values and ≥10 °C for δ(2)H values resulted in errors larger than the CRDS precision for the respective isotopes and advise the use of such instruments only in sufficiently temperature-stabilised environments.

The impact of soil microorganisms on the global budget of delta18O in atmospheric CO2.

Improved global estimates of terrestrial photosynthesis and respiration are critical for predicting the rate of change in atmospheric CO(2). The oxygen isotopic composition of atmospheric CO(2) can be used to estimate these fluxes because oxygen isotopic exchange between CO(2) and water creates distinct isotopic flux signatures. The enzyme carbonic anhydrase (CA) is known to accelerate this exchange in leaves, but the possibility of CA activity in soils is commonly neglected. Here, we report widespread accelerated soil CO(2) hydration. Exchange was 10-300 times faster than the uncatalyzed rate, consistent with typical population sizes for CA-containing soil microorganisms. Including accelerated soil hydration in global model simulations modifies contributions from soil and foliage to the global CO(18)O budget and eliminates persistent discrepancies existing between model and atmospheric observations. This enhanced soil hydration also increases the differences between the isotopic signatures of photosynthesis and respiration, particularly in the tropics, increasing the precision of CO(2) gross fluxes obtained by using the delta(18)O of atmospheric CO(2) by 50%.

Short-range multispectral imaging is an inexpensive, fast, and accurate approach to estimate biodiversity in a temperate calcareous grassland.

Image sensing technologies are rapidly increasing the cost-effectiveness of biodiversity monitoring efforts. Species differences in the reflectance of electromagnetic radiation can be used as a surrogate estimate plant biodiversity using multispectral image data. However, these efforts are often hampered by logistical difficulties in broad-scale implementation. Here, we investigate the utility of multispectral imaging technology from commercially available unmanned aerial vehicles (UAVs, or drones) in estimating biodiversity metrics at a fine spatial resolution (0.1-0.5 cm pixel resolution) in a temperate calcareous grassland in Oxfordshire, UK. We calculate a suite of moments (coefficient of variation, standard deviation, skewness, and kurtosis) for the distribution of radiance from multispectral images at five wavelength bands (Blue 450 ± 16 nm; Green 560 ± 16 nm; Red 650 ± 16 nm; Red Edge 730 ± 16 nm; Near Infrared 840 ± 16 nm) and test their effectiveness at estimating ground-truthed biodiversity metrics from in situ botanical surveys for 37-1 × 1 m quadrats. We find positive associations between the average coefficient of variation in spectral radiance and both the Shannon-Weiner and Simpson's biodiversity indices. Furthermore, the average coefficient of variation in spectral radiance is consistent and highly repeatable across sampling days and recording heights. Positive associations with biodiversity indices hold irrespective of the image recording height (2-8 m), but we report reductions in estimates of spectral diversity with increases to UAV recording height. UAV imaging reduced sampling time by a factor of 16 relative to in situ botanical surveys. We demonstrate the utility of multispectral radiance moments as an indicator of biodiversity in this temperate calcareous grassland at a fine spatial resolution using a widely available UAV monitoring system with a coarse spectral resolution. The use of UAV technology with multispectral sensors has far-reaching potential to provide cost-effective and high-resolution monitoring of biodiversity.

Increase in water-use efficiency and underlying processes in pine forests across a precipitation gradient in the dry Mediterranean region over the past 30 years.

Motivated by persistent predictions of warming and drying in the entire Mediterranean and other regions, we have examined the interactions of intrinsic water-use efficiency (W(i)) with environmental conditions in Pinus halepensis. We used 30-year (1974-2003) tree-ring records of basal area increment (BAI) and cellulose (13)C and (18)O composition, complemented by short-term physiological measurements, from three sites across a precipitation (P) gradient (280-700 mm) in Israel. The results show a clear trend of increasing W(i) in both the earlywood (EW) and latewood (LW) that varied in magnitude depending on site and season, with the increase ranging from ca. 5 to 20% over the study period. These W(i) trends were better correlated with the increase in atmospheric CO(2) concentration, C(a), than with the local increase in temperature (~0.04°C year(-1)), whereas age, height and density variations had minor effects on the long-term isotope record. There were no trends in P over time, but W(i) from EW and BAI were dependent on the interannual variations in P. From reconstructed C(i) values, we demonstrate that contrasting gas-exchange responses at opposing ends of the hydrologic gradient underlie the variation in W(i) sensitivity to C(a) between sites and seasons. Under the mild water limitations typical of the main seasonal growth period, regulation was directed at increasing C(i)/C(a) towards a homeostatic set-point observed at the most mesic site, with a decrease in the W(i) response to C(i) with increasing aridity. With more extreme drought stress, as seen in the late season at the drier sites, the response was W(i) driven, and there was an increase in the W(i) sensitivity to C(a) with aridity and a decreasing sensitivity of C(i) to C(a). The apparent C(a)-driven increases in W(i) can help to identify the adjustments to drying conditions that forest ecosystems can make in the face of predicted atmospheric change.

Climate suitability predictions for the cultivation of macadamia (Macadamia integrifolia) in Malawi using climate change scenarios.

Climate change is altering suitable areas of crop species worldwide, with cascading effects on people reliant upon those crop species as food sources and for income generation. Macadamia is one of Malawi's most important and profitable crop species; however, climate change threatens its production. Thus, this study's objective is to quantitatively examine the potential impacts of climate change on the climate suitability for macadamia in Malawi. We utilized an ensemble model approach to predict the current and future (2050s) suitability of macadamia under two Representative Concentration Pathways (RCPs). We achieved a good model fit in determining suitability classes for macadamia (AUC = 0.9). The climatic variables that strongly influence macadamia's climatic suitability in Malawi are suggested to be the precipitation of the driest month (29.1%) and isothermality (17.3%). Under current climatic conditions, 57% (53,925 km2) of Malawi is climatically suitable for macadamia. Future projections suggest that climate change will decrease the suitable areas for macadamia by 18% (17,015 km2) and 21.6% (20,414 km2) based on RCP 4.5 and RCP 8.5, respectively, with the distribution of suitability shifting northwards in the 2050s. The southern and central regions of the country will suffer the greatest losses (≥ 8%), while the northern region will be the least impacted (4%). We conclude that our study provides critical evidence that climate change will reduce the suitable areas for macadamia production in Malawi, depending on climate drivers. Therefore area-specific adaptation strategies are required to build resilience among producers.

Chlorophyll a fluorescence illuminates a path connecting plant molecular biology to Earth-system science.

For decades, the dynamic nature of chlorophyll a fluorescence (ChlaF) has provided insight into the biophysics and ecophysiology of the light reactions of photosynthesis from the subcellular to leaf scales. Recent advances in remote sensing methods enable detection of ChlaF induced by sunlight across a range of larger scales, from using instruments mounted on towers above plant canopies to Earth-orbiting satellites. This signal is referred to as solar-induced fluorescence (SIF) and its application promises to overcome spatial constraints on studies of photosynthesis, opening new research directions and opportunities in ecology, ecophysiology, biogeochemistry, agriculture and forestry. However, to unleash the full potential of SIF, intensive cross-disciplinary work is required to harmonize these new advances with the rich history of biophysical and ecophysiological studies of ChlaF, fostering the development of next-generation plant physiological and Earth-system models. Here, we introduce the scale-dependent link between SIF and photosynthesis, with an emphasis on seven remaining scientific challenges, and present a roadmap to facilitate future collaborative research towards new applications of SIF.

Concentration effects on laser-based δ18 O and δ2 H measurements and implications for the calibration of vapour measurements with liquid standards.

Recently available isotope ratio infrared spectroscopy can directly measure the isotopic composition of atmospheric water vapour (δ(18) O, δ(2) H), overcoming one of the main limitations of isotope ratio mass spectrometry (IRMS) methods. Calibrating these gas-phase instruments requires the vapourisation of liquid standards since primary standards in principle are liquids. Here we test the viability of calibrating a wavelength-scanned cavity ring-down spectroscopy (CRDS) instrument with vapourised liquid standards. We also quantify the dependency of the measured isotope values on the water concentration for a range of isotopic compositions. In both liquid and vapour samples, we found an increase in δ(18) O and δ(2) H with water vapour concentration. For δ(18) O, the slope of this increase was similar for liquid and vapour, with a slight positive relationship with sample δ-value. For δ(2) H, we found diverging patterns for liquid and vapour samples, with no dependence on δ-value for vapour, but a decreasing slope for liquid samples. We also quantified tubing memory effects to step changes in isotopic composition, avoiding concurrent changes in the water vapour concentration. Dekabon tubing exhibited much stronger, concentration-dependent, memory effects for δ(2) H than stainless steel or perfluoroalkoxy (PFA) tubing. Direct vapour measurements with CRDS in a controlled experimental chamber agreed well with results obtained from vapour simultaneously collected in cold traps analysed by CRDS and IRMS. We conclude that vapour measurements can be calibrated reliably with liquid standards. We demonstrate how to take the concentration dependencies of the δ-values into account. Copyright © 2010 John Wiley & Sons, Ltd.

Quantification of leaf-scale light energy allocation and photoprotection processes in a Mediterranean pine forest under extensive seasonal drought.

Photoprotection strategies in a Pinus halepensis Mill. forest at the dry timberline that shows sustained photosynthetic activity during 6-7 month summer drought were characterized and quantified under field conditions. Measurements of chlorophyll fluorescence, leaf-level gas exchange and pigment concentrations were made in both control and summer-irrigated plots, providing the opportunity to separate the effects of atmospheric from soil water stress on the photoprotection responses. The proportion of light energy incident on the leaf surface ultimately being used for carbon assimilation was 18% under stress-free conditions (irrigated, winter), declining to 4% under maximal stress (control, summer). Allocation of absorbed light energy to photochemistry decreased from 25 to 15% (control) and from 50% to 30% (irrigated) between winter and summer, highlighting the important role of pigment-mediated energy dissipation processes. Photorespiration or other non-assimilatory electron flow accounted for 15-20% and ~10% of incident light energy during periods of high and low carbon fixation, respectively, representing a proportional increase in photochemical energy going to photorespiration in summer but a decrease in the absolute amount of photorespiratory CO2 loss. Resilience of the leaf photochemical apparatus was expressed in the complete recovery of photosystem II (PSII) efficiency (ΦPSII) and relaxation of the xanthophyll de-epoxidation state on the diurnal cycle throughout the year, and no seasonal decrease in pre-dawn maximal PSII efficiency (Fv/Fm). The response of CO2 assimilation and photoprotection strategies to stomatal conductance and leaf water potential appeared independent of whether stress was due to atmospheric or soil water deficits across seasons and treatments. The range of protection characteristics identified provides insights into the relatively high carbon economy under these dry conditions, conditions that are predicted for extended areas in the Mediterranean and other regions due to global climate change.

Physiology-phenology interactions in a productive semi-arid pine forest.

This study explored possible advantages conferred by the phase shift between leaf phenology and photosynthesis seasonality in a semi-arid Pinus halepensis forest system, not seen in temperate sites. Leaf-scale measurements of gas exchange, nitrogen and phenology were used on daily, seasonal and annual time-scales. Peak photosynthesis was in late winter, when high soil moisture, mild temperatures and low leaf vapour pressure deficit (D(L)) allowed high rates associated with high water- and nitrogen-use efficiencies. Self-sustained new needle growth through the dry and hot summer maximized photosynthesis in the following wet season, without straining carbon storage. Low rates of water loss were associated with increasing sensitivity of stomatal conductance (g(s)) to soil moisture below a relative extractable water (REW) of 0.4, and decreased g(s )sensitivity to D(L) below REW of approx. 0.2. This response was captured by the modified Ball-Berry (Leuning) model. While most physiological parameters and responses measured were typical of temperate pines, the photosynthesis-phenological phasing contributed to high productivity under warm-dry conditions. This contrasts with reported effects of short-term periodical droughts and could lead to different predictions of the effect of warming and drying climate on pine forest productivity.

Biotic and abiotic factors affecting the delta(13)C of soil respired CO(2) in a Mediterranean oak woodland.

The flux (R(s)) and carbon isotopic composition (delta(13)C (Rs)) of soil respired CO (2) was measured every 2 h over the course of three diel cycles in a Mediterranean oak woodland, together with measurements of the delta(13)C composition of leaf, root and soil organic matter (delta(13)C (SOM)) and metabolites. Simulations of R(s) and delta(13)C (Rs) were also made using a numerical model parameterised with the SOM data and assuming short-term production rates were driven mainly by temperature. Average values of delta(13)C (Rs) over the study period were within the range of root metabolite and average delta(13)C (SOM) values, but enriched in (13)C relative to the bulk delta(13)C of leaf, litter, and roots and the upper soil organic layers. There was good agreement between model output and observed CO (2) fluxes and the underlying features of delta(13)C (Rs). Observed diel variations of 0.5 per thousand in delta(13)C (Rs) were predicted by the model in response to temperature-related shifts in production rates along a approximately 3 per thousand gradient observed in the profile of delta(13)C (SOM). However, observed delta(13)C (Rs) varied by over 2 per thousand, indicating that both dynamics in soil respiratory metabolism and physical processes can influence short-term variability of delta(13)C (Rs).

Large daily variation in 13C-enrichment of leaf-respired CO2 in two Quercus forest canopies.

The use of the 13C : 12C isotopic ratio (delta13C) of leaf-respired CO2 to trace carbon fluxes in plants and ecosystems is limited by little information on temporal variations in delta13C of leaf dark-respired CO2 (delta13Cr) under field conditions. Here, we explored variability in delta13Cr and its relationship to key respiratory substrates from collections of leaf dark-respired CO2, carbohydrate extractions and gas exchange measurements over 24-h periods in two Quercus canopies. Throughout both canopies, delta13Cr became progressively 13C-enriched during the photoperiod, by up to 7%, then 13C-depleted at night relative to the photoperiod. This cycle could not be reconciled with delta13C of soluble sugars (delta13Css), starch (delta13Cst), lipids (delta13Cl), cellulose (delta13Cc) or with calculated photosynthetic discrimination (Delta). However, photoperiod progressive enrichment in delta13Cr was correlated with cumulative carbon assimilation (r2 = 0.91). We concluded that there is considerable short-term variation in delta13Cr in forest canopies, that it is consistent with current hypotheses for 13C fractionation during leaf respiration, that leaf carbohydrates cannot be used as surrogates for delta13Cr, and that diel changes in leaf carbohydrate status could be used to predict changes in delta13Cr empirically.

Association between tree-ring and needle delta13C and leaf gas exchange in Pinus halepensis under semi-arid conditions.

Associations between delta13C values and leaf gas exchanges and tree-ring or needle growth, used in ecophysiological compositions, can be complex depending on the relative timing of CO2 uptake and subsequent redistribution and allocation of carbon to needle and stem components. For palaeoenvironmental and dendroecological studies it is often interpreted in terms of a simple model of delta13C fractionation in C3 plants. However, in spite of potential complicating factors, few studies have actually examined these relationships in mature trees over inter- and intra-annual time-scales. Here, we present results from a 4 years study that investigated the links between variations in leaf gas-exchange properties, growth, and dated delta13C values along the needles and across tree rings of Aleppo pine trees growing in a semi-arid region under natural conditions or with supplemental summer irrigation. Sub-sections of tissue across annual rings and along needles, for which time of formation was resolved from growth rate analyses, showed rapid growth and delta13C responses to changing environmental conditions. Seasonal cycles of growth and delta13C (up to approximately 4 per thousand) significantly correlated (P < 0.01) with photosynthetically active radiation, vapour pressure deficit, air temperature, and soil water content. The irrigation significantly increased leaf net assimilation, stomatal conductance and needle and tree-ring growth rate, and markedly decreased needle and tree-ring delta13C values and its sensitivity to environmental parameters. The delta13C estimates derived from gas-exchange parameters, and weighted by assimilation, compared closely with seasonal and inter-annual delta13C values of needle- and tree-ring tissue. Higher stomatal conductances of the irrigated trees (0.22 vs. 0.08 mol m(-2) s(-1) on average) corresponded with approximately 2.0 per thousand lower average delta13C values, both measured and derived. Derived and measured delta13C values also indicated that needle growth, which occurs throughout the stressful summer was supported by carbon from concurrent, low rate assimilation. For Aleppo pine under semi-arid and irrigated conditions, the delta13C of tree-ring and needle material proved, in general, to be a reasonable indicator of integrated leaf gas-exchange properties.