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Aim: Within C3 plants, photosynthesis is a balance between CO2 supply from the atmosphere via stomata and demand by enzymes within chloroplasts. This process is dynamic and a complex but crucial aspect of photosynthesis. We sought to understand the spatial pattern in CO2 supply–demand balance on a global scale, via analysis of stable isotopes of carbon within leaves (Δ13C), which provide an integrative record of CO2 drawdown during photosynthesis. LocationGlobal. Time period1951–2011. Major taxa studiedVascular plants. Methods: We assembled a database of leaf carbon isotope ratios containing 3,979 species–site combinations from across the globe, including 3,645 for C3 species. We examined a wide array of potential climate and soil drivers of variation in Δ13C. Results: The strongest drivers of carbon isotope discrimination at the global scale included atmospheric pressure, potential evapotranspiration and soil pH, which explained 44% of the variation in Δ13C. Addition of eight more climate and soil variables (each explaining small but highly significant amounts of variation) increased the explained variation to 60%. On top of this, the largest plant trait effect was leaf nitrogen per area, which explained 11% of Δ13C variation. Main conclusions: By considering variation in Δ13C at a considerably larger scale than previously, we were able to identify and quantify key drivers in CO2 supply–demand balance previously unacknowledged. Of special note is the key role of soil properties, with greater discrimination on low-pH and high-silt soils. Unlike other plant traits, which show typically wide variation within sets of coexisting species, the global pattern in carbon stable isotope ratios is much more conservative; there is relatively narrow variation in time-integrated CO2 concentrations at the site of carboxylation among plants in a given soil and climate.

Terrestrial paleoclimate records are rare in the South Atlantic, limiting opportunities to provide a prehistoric context for current global changes. The tussock grass, Poa flabellata , grows abundantly along the coasts of the Falkland Islands and other subantarctic islands. It forms extensive peat records, providing a promising opportunity to reconstruct high-resolution regional climate records. The isotopic composition of leaf and root tissues deposited in these peats has the potential to record variation in precipitation, temperature, and relative humidity over time, but these relationships are unknown for P. flabellata. Here, we measured the isotopic composition of P. flabellata and precipitation and explore relationships with seasonal temperature and humidity variations across four study locations in the Falkland Islands. We reveal that inter-seasonal differences in carbon and oxygen stable isotopes of leaf α -cellulose of living P. flabellata correlated with monthly mean temperature and relative humidity. The carbon isotope composition of leaf α -cellulose (δ13C leaf) records the balance of CO2 supply through stomata and the demand by photosynthesis. The positive correlation between δ13C leaf and temperature and negative correlation between δ13 C leaf and relative humidity suggest that photosynthetic demand for CO2 relative to stomatal supply is enhanced when conditions are warm and dry. Further, the positive correlation between δ13 C leaf and δ18O leaf (r=0.88 ; p<0.001 ; n=24) indicates that stomatal closure during warm dry periods explains seasonal variation in δ13C leaf. We observed significant differences between winter and summer seasons for both δ18 O leaf and δ13C leaf and among study locations for δ18O leaf but not δ13C leaf. δ18 O values of monthly composite precipitation were similar between seasons and among study locations, yet characteristic of the latitudinal origin of storm tracks and seasonal winds. The weak correlation between δ18O in monthly composite precipitation and δ18 O leaf further suggests that relative humidity is the main driver of the δ18O leaf. The oxygen isotopes in root α -cellulose did not reflect, or only partially reflected (at one study location), the δ18O in precipitation. Overall, this study supports the use of peat records formed by P. flabellata to fill a significant gap in our knowledge of the long-term trends in Southern Hemisphere climate dynamics. [ABSTRACT FROM AUTHOR]

Aim: Within C3 plants, photosynthesis is a balance between CO2 supply from the atmosphere via stomata and demand by enzymes within chloroplasts. This process is dynamic and a complex but crucial aspect of photosynthesis. We sought to understand the spatial pattern in CO2 supply–demand balance on a global scale, via analysis of stable isotopes of carbon within leaves (Δ13C), which provide an integrative record of CO2 drawdown during photosynthesis. Location: Global Time period: 1951–2011. Major taxa studied: Vascular plants. Methods: We assembled a database of leaf carbon isotope ratios containing 3,979 species–site combinations from across the globe, including 3,645 for C3 species. We examined a wide array of potential climate and soil drivers of variation in Δ13C. Results: The strongest drivers of carbon isotope discrimination at the global scale included atmospheric pressure, potential evapotranspiration and soil pH, which explained 44% of the variation in Δ13C. Addition of eight more climate and soil variables (each explaining small but highly significant amounts of variation) increased the explained variation to 60%. On top of this, the largest plant trait effect was leaf nitrogen per area, which explained 11% of Δ13C variation. Main conclusions: By considering variation in Δ13C at a considerably larger scale than previously, we were able to identify and quantify key drivers in CO2 supply–demand balance previously unacknowledged. Of special note is the key role of soil properties, with greater discrimination on low‐pH and high‐silt soils. Unlike other plant traits, which show typically wide variation within sets of coexisting species, the global pattern in carbon stable isotope ratios is much more conservative; there is relatively narrow variation in time‐integrated CO2 concentrations at the site of carboxylation among plants in a given soil and climate. [ABSTRACT FROM AUTHOR]

ABSTRACT Accurate estimates of the δ13C value of CO2 respired from roots ( δ13CR_root) and leaves ( δ13CR_leaf) are important for tracing and understanding changes in C fluxes at the ecosystem scale. Yet the mechanisms underlying temporal variation in these isotopic signals are not fully resolved. We measured δ13CR_leaf, δ13CR_root, and the δ13C values and concentrations of glucose and sucrose in leaves and roots in the C4 grass Sporobolus wrightii and the C3 tree Prosopis velutina in a savanna ecosystem in southeastern Arizona, USA. Night-time variation in δ13CR_leaf of up to 4.6 ± 0.6‰ in S. wrightii and 3.0 ± 0.6‰ in P. velutina were correlated with shifts in leaf sucrose concentration, but not with changes in δ13C values of these respiratory substrates. Strong positive correlations between δ13CR_root and root glucose δ13C values in P. velutina suggest large diel changes in δ13CR_root (were up to 3.9‰) influenced by short-term changes in δ13C of leaf-derived phloem C. No diel variation in δ13CR_root was observed in S. wrightii. Our findings show that short-term changes in δ13CR_leaf and δ13CR_root were both related to substrate isotope composition and concentration. Changes in substrate limitation or demand for biosynthesis may largely control short-term variation in the δ13C of respired CO2 in these species. [ABSTRACT FROM AUTHOR]

We review the relevance and use of stable isotopes for the study of plant community succession. Stable isotope measurements provide information on the origin of resources acquired by plants, the processes governing resource uptake and transformation, and the physiological and environmental conditions of plant growth. When combined with measurements of the stable isotope ratio values of soil microbial biomass, soil organic matter and plant litter, isotope measurements of plants can indicate effects of successional changes on ecosystem processes. However, their application to questions of plant succession and ecosystem change is limited by the degree to which the underlying assumptions are met in each study, and complementary measures may be required, depending upon the question of interest. First, we discuss the changes that occur in the stable isotope composition of plants and ecosystems with ontogeny and species replacements, as well as their potential evolutionary significance. Second, we discuss the imprints of plant competition and facilitation on leaf and wood tissue, as well as how stable isotopes can provide novel insights on the mechanisms underlying plant interactions. Finally, we discuss the capacity for stable isotope measurements to serve as a proxy record for past disturbances such as fire, logging and cyclones. [ABSTRACT FROM PUBLISHER]

We evaluated diurnal and seasonal patterns of carbon isotope composition of leaf dark-respired CO2 ( δ13Cl) in the C3 perennial shrub velvet mesquite ( Prosopis velutina) across flood plain and upland savanna ecosystems in the south-western USA. δ13Cl of darkened leaves increased to maximum values late during daytime periods and declined gradually over night-time periods to minimum values at pre-dawn. The magnitude of the diurnal shift in δ13Cl was strongly influenced by seasonal and habitat-related differences in soil water availability and leaf surface vapour pressure deficit. δ13Cl and the cumulative flux-weighted δ13C value of photosynthates were positively correlated, suggesting that progressive 13C enrichment of the CO2 evolved by darkened leaves during the daytime mainly resulted from short-term changes in photosynthetic 13C discrimination and associated shifts in the δ13C signature of primary respiratory substrates. The 13C enrichment of dark-respired CO2 relative to photosynthates across habitats and seasons was 4 to 6‰ at the end of the daytime period (1800 h), but progressively declined to 0‰ by pre-dawn (0300 h). The origin of night-time and daytime variations in δ13Cl is discussed in terms of the carbon source(s) feeding respiration and the drought-induced changes in carbon metabolism. [ABSTRACT FROM AUTHOR]

As atmospheric carbon dioxide concentrations, [CO2Air], continue their uncontrolled rise, the capacity of soils to accumulate or retain carbon is uncertain. Free-air CO2 enrichment (FACE) experiments have been conducted to better understand the plant, soil and ecosystem response to elevated [CO2], frequently employing commercial CO2 that imparts a distinct isotopic signal to the system for tracing carbon. We conducted a FACE experiment in 1998 and 1999, whereby sorghum (C4 photosynthetic pathway) was grown in four replicates of four treatments using a split-strip plot design: (i) ambient CO2/ample water (365 μmol mol−1, "Control–Wet"), (ii) ambient CO2/water stress ("Control–Dry"), (iii) CO2-enriched (560 μmol mol−1, "FACE–Wet"), and (iv) CO2-enriched/water stressed ("FACE–Dry"). The stable-carbon isotope composition of the added CO2 (in FACE treatments) was close to that of free atmosphere background values, so the subsequent similar 13C-enriched carbon signal photosynthetically fixed by C4 sorghum plants could be used to trace the fate of carbon in both FACE and control treatments. Measurement of soil organic carbon content (SOC (%) = gC/gdry soil × 100%) and δ13C at three depths (0–15, 15–30, and 30–60 cm) were made on soils from the beginning and end of the two experimental growing seasons. A progressive ca. 0.5‰–1.0‰ δ13C increase in the upper soil SOC in all treatments over the course of the experiment indicated common entry of new sorghum carbon into the SOC pools. The 0–15 cm SOC in FACE treatments was 13C-enriched relative to the Control by ca. 1‰, and according to isotopic mass balance, the fraction of the new sorghum-derived SOC in the Control–Wet treatment at the end of the second season was 8.4%, 14.2% in FACE–Wet, 6.5% in Control–Dry, and 14.2% in FACE–Dry. The net SOC enhancement resulting from CO2 enrichment was therefore 5.8% (or 2.9% y−1 of experiment) under ample water and 7.7% (3.8% y−1 of experiment) under limited water, which matches the pattern of greater aboveground biomass increase with elevated [CO2Air] under the Dry treatment, but no parallel isotopic shifts were found in deeper soils. However, these increased fractions of new carbon in SOC at the end of the experiment do not necessarily mean an increase in total SOC content, because gravimetric measurements of SOC did not reveal a significant increase under elevated [CO2Air], at least within the limits of SOC-content error bars. Thus, new carbon gains might be offset by pre-experiment carbon losses. The results demonstrate successful isotopic tracing of carbon from plants to soils in this sorghum FACE experiment showing differences between FACE and Control treatments, which suggest more dynamic cycling of SOC under elevated [CO2Air] than in the Control treatment. [ABSTRACT FROM AUTHOR]

Abstract: Seasonal and annual variations of rainfall and humidity are recorded in the carbon and oxygen stable isotope ratios of sequentially grown spines found on the columnar cactus, Carnegiea gigantea. A 26-year long composite δ 18O and δ 13C isotope record from the spines of five saguaro cacti was created using bomb radiocarbon and semi-annual variations in δ 13C. Once dating errors in the composite record are corrected, mean annual spine δ 18O is negatively correlated (P <0.001) with total annual precipitation (TAP) from November through October and positively correlated (P <0.01) with mean annual nighttime vapor pressure deficit (VPD). Year-to-year decreases (>2‰) in the maximum annual spine δ 18O are positively correlated (P <0.01) with the Southern Oscillation Index (SOI). We attribute these decreases to enhanced winter rainfall associated with the El Niño phase of the El Niño-Southern Oscillation. Minimum annual δ 13C is negatively correlated with TAP (P <0.05) and mean nighttime VPD (P <0.05). These results bolster proposed mechanistic models of isotopic variation in the spines of columnar cactus and demonstrate how isotopic spine series may be used as climate proxies in regions of the Americas where trees suitable for traditional or isotopic dendrochonology are absent. [Copyright &y& Elsevier]