Associations between carbon isotope ratios of ecosystem respiration, water availability and canopy conductance

We tested the hypothesis that the stable carbon isotope signature of ecosystem respiration (δ¹³C_R) was regulated by canopy conductance (G_c) using weekly Keeling plots (n=51) from a semiarid old‐growth ponderosa pine (Pinus ponderosa) forest in Oregon, USA. For a comparison of forests in two contrasting climates we also evaluated trends in δ¹³C_R from a wet 20‐year‐old Douglas‐fir (Pseudotsuga menziesii) plantation located near the Pacific Ocean. Intraannual variability in δ¹³C_R was greater than 8.0‰ at both sites, was highest during autumn, winter, and spring when rainfall was abundant, and lowest during summer drought. The δ¹³C_R of the dry pine forest was consistently more positive than the wetter Douglas‐fir forest (mean annual δ¹³C_R: ?25.41‰ vs. ?26.23‰, respectively, P=0.07). At the Douglas‐fir forest, δ¹³C_R–climate relationships were consistent with predictions based on stomatal regulation of carbon isotope discrimination (Δ). Soil water content (SWC) and vapor pressure deficit (vpd) were the most important factors governing δ¹³C_R in this forest throughout the year. In contrast, δ¹³C_R at the pine forest was relatively insensitive to SWC or vpd, and exhibited a smaller drought‐related enrichment (～2‰) than the enrichment observed during drought at the Douglas‐fir forest (～5‰). Groundwater access at the pine forest may buffer canopy–gas exchange from drought. Despite this potential buffering, δ¹³C_R at the pine forest was significantly but weakly related to canopy conductance (G_c), suggesting that δ¹³C_R remains coupled to canopy–gas exchange despite groundwater access. During drought, δ¹³C_R was strongly correlated with soil temperature at both forests. The hypothesis that canopy‐level physiology is a critical regulator of δ¹³C_R was supported; however, belowground respiration may become more important during rain‐free periods.     

Canopy-scale δ13C of photosynthetic and respiratory CO2 fluxes: observations in forest biomes across the United States

The δ¹³C values of atmospheric carbon dioxide (CO₂) can be used to partition global patterns of CO₂ source/sink relationships among terrestrial and oceanic ecosystems using the inversion technique. This approach is very sensitive to estimates of photosynthetic ¹³C discrimination by terrestrial vegetation (Δ_A), and depends on δ¹³C values of respired CO₂ fluxes (δ¹³C_R). Here we show that by combining two independent data streams – the stable isotope ratios of atmospheric CO₂ and eddy‐covariance CO₂ flux measurements – canopy scale estimates of Δ_A can be successfully derived in terrestrial ecosystems. We also present the first weekly dataset of seasonal variations in δ¹³C_R from dominant forest ecosystems in the United States between 2001 and 2003. Our observations indicate considerable summer‐time variation in the weekly value of δ¹³C_R within coniferous forests (4.0‰ and 5.4‰ at Wind River Canopy Crane Research Facility and Howland Forest, respectively, between May and September). The monthly mean values of δ¹³C_R showed a smaller range (2–3‰), which appeared to significantly correlate with soil water availability. Values of δ¹³C_R were less variable during the growing season at the deciduous forest (Harvard Forest). We suggest that the negative correlation between δ¹³C_R and soil moisture content observed in the two coniferous forests should represent a general ecosystem response to the changes in the distribution of water resources because of climate change. Shifts in δ¹³C_R and Δ_A could be of sufficient magnitude globally to impact partitioning calculations of CO₂ sinks between oceanic and terrestrial compartments.     

18O composition of CO2 and H2O ecosystem pools and fluxes in a tallgrass prairie: Simulations and comparisons to measurements

In this paper we describe measurements and modeling of ¹⁸O in CO₂ and H₂O pools and fluxes at a tallgrass prairie site in Oklahoma. We present measurements of the δ¹⁸O value of leaf water, depth‐resolved soil water, atmospheric water vapor, and Keeling plot δ¹⁸O intercepts for net soil‐surface CO₂ and ecosystem CO₂ and H₂O fluxes during three periods of the 2000 growing season. Daytime discrimination against C¹⁸OO, as calculated from measured above‐canopy CO₂ and δ¹⁸O gradients, is also presented. To interpret the isotope measurements, we applied an integrated land‐surface and isotope model (ISOLSM) that simulates ecosystem H₂¹⁸O and C¹⁸OO stocks and fluxes. ISOLSM accurately predicted the measured isotopic composition of ecosystem water pools and the δ¹⁸O value of net ecosystem CO₂ and H₂O fluxes. Simulations indicate that incomplete equilibration between CO₂ and H₂O within C₄ plant leaves can have a substantial impact on ecosystem discrimination. Diurnal variations in the δ¹⁸O value of above‐canopy vapor had a small impact on the predicted δ¹⁸O value of ecosystem water pools, although sustained differences had a large impact. Diurnal variations in the δ¹⁸O value of above‐canopy CO₂ substantially affected the predicted ecosystem discrimination. Leaves dominate the ecosystem ¹⁸O‐isoflux in CO₂ during the growing season, while the soil contribution is relatively small and less variable. However, interpreting daytime measurements of ecosystem C¹⁸OO fluxes requires accurate predictions of both soil and leaf ¹⁸O‐isofluxes.     

Temperature and precipitation controls over leaf‐ and ecosystem‐level CO2 flux along a woody plant encroachment gradient

Conversion of grasslands to woodlands may alter the sensitivity of CO₂ exchange of individual plants and entire ecosystems to air temperature and precipitation. We combined leaf‐level gas exchange and ecosystem‐level eddy covariance measurements to quantify the effects of plant temperature sensitivity and ecosystem temperature responses within a grassland and mesquite woodland across seasonal precipitation periods. In so doing, we were able to estimate the role of moisture availability on ecosystem temperature sensitivity under large‐scale vegetative shifts. Optimum temperatures (T_opt) for net photosynthetic assimilation (A) and net ecosystem productivity (NEP) were estimated from a function fitted to A and NEP plotted against air temperature. The convexities of these temperature responses were quantified by the range of temperatures over which a leaf or an ecosystem assimilated 50% of maximum NEP (Ω₅₀). Under dry pre‐ and postmonsoon conditions, leaf‐level Ω₅₀ in C₃ shrubs were two‐to‐three times that of C₄ grasses, but under moist monsoon conditions, leaf‐level Ω₅₀ was similar between growth forms. At the ecosystems‐scale, grassland NEP was more sensitive to precipitation, as evidenced by a 104% increase in maximum NEP at monsoon onset, compared to a 57% increase in the woodland. Also, woodland NEP was greater across all temperatures experienced by both ecosystems in all seasons. By maintaining physiological function across a wider temperature range during water‐limited periods, woody plants assimilated larger amounts of carbon. This higher carbon‐assimilation capacity may have significant implications for ecosystem responses to projected climate change scenarios of higher temperatures and more variable precipitation, particularly as semiarid regions experience conversions from C₄ grasses to C₃ shrubs. As regional carbon models, CLM 4.0, are now able to incorporate functional type and photosynthetic pathway differences, this work highlights the need for a better integration of the interactive effects of growth form/functional type and photosynthetic pathway on water resource acquisition and temperature sensitivity.     

Combining meteorology, eddy fluxes, isotope measurements, and modeling to understand environmental controls of carbon isotope discrimination at the canopy scale

Estimates of terrestrial carbon isotope discrimination are useful to quantify the terrestrial carbon sink. Carbon isotope discrimination by terrestrial ecosystems may vary on seasonal and interannual time frames, because it is affected by processes (e.g. photosynthesis, stomatal conductance, and respiration) that respond to variable environmental conditions (e.g. air humidity, temperature, light). In this study, we report simulations of the temporal variability of canopy‐scale C₃ photosynthetic carbon isotope discrimination obtained with an ecophysiologically based model (ISOLSM) designed for inclusion in global models. ISOLSM was driven by half‐hourly meteorology, and parameterized with eddy covariance measurements of carbon and energy fluxes and foliar carbon isotope ratios from a pine forest in Metolius (OR). Comparing simulated carbon and energy fluxes with observations provided a range of parameter values that optimized the simulated fluxes. We found that the sensitivity of photosynthetic carbon isotope discrimination to the slope of the stomatal conductance equation (m, Ball–Berry constant) provided an additional constraint to the model, reducing the wide parameter space obtained from the fluxes alone. We selected values of m that resulted in similar simulated long‐term discrimination as foliar isotope ratios measured at the site. The model was tested with ¹³C measurements of ecosystem (δ_R) and foliar (δ_f) respiration. The daily variability of simulated ¹³C values of assimilated carbon (δ_A) was similar to that of observed δ_f, and higher than that of observed and simulated δ_R. We also found similar relationships between environmental factors (i.e. vapor pressure deficit) and simulated δ_R as measured in ecosystem surveys of δ_R. Therefore, ISOLSM reasonably simulated the short‐term variability of δ_A controlled by atmospheric conditions at the canopy scale, which can be useful to estimate the variability of terrestrial isotope discrimination. Our study also shows that including the capacity to simulate carbon isotope discrimination, together with simple ecosystem isotope measurements, can provide a useful constraint to land surface and carbon balance models.     

Environmental and physiological controls on the carbon isotope composition of CO2 respired by leaves and roots of a C3 woody legume (Prosopis velutina) and a C4 perennial grass (Sporobolus wrightii)

Accurate estimates of the δ¹³C value of CO₂ respired from roots (δ¹³C_{R_root}) and leaves (δ¹³C_{R_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 δ¹³C_{R_leaf}, δ¹³C_{R_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 δ¹³C_{R_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 δ¹³C_{R_root} and root glucose δ¹³C values in P. velutina suggest large diel changes in δ¹³C_{R_root} (were up to 3.9‰) influenced by short‐term changes in δ¹³C of leaf‐derived phloem C. No diel variation in δ¹³C_{R_root} was observed in S. wrightii. Our findings show that short‐term changes in δ¹³C_{R_leaf} and δ¹³C_{R_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.     

Effect of precipitation variability on net primary production and soil respiration in a Chihuahuan Desert grassland

Precipitation regimes are predicted to become more variable with more extreme rainfall events punctuated by longer intervening dry periods. Water‐limited ecosystems are likely to be highly responsive to altered precipitation regimes. The bucket model predicts that increased precipitation variability will reduce soil moisture stress and increase primary productivity and soil respiration in aridland ecosystems. To test this hypothesis, we experimentally altered the size and frequency of precipitation events during the summer monsoon (July through September) in 2007 and 2008 in a northern Chihuahuan Desert grassland in central New Mexico, USA. Treatments included (1) ambient rain, (2) ambient rain plus one 20 mm rain event each month, and (3) ambient rain plus four 5 mm rain events each month. Throughout two monsoon seasons, we measured soil temperature, soil moisture content (θ), soil respiration (R_s), along with leaf‐level photosynthesis (A_net), predawn leaf water potential (Ψ_pd), and seasonal aboveground net primary productivity (ANPP) of the dominant C₄ grass, Bouteloua eriopoda. Treatment plots receiving a single large rainfall event each month maintained significantly higher seasonal soil θ which corresponded with a significant increase in R_s and ANPP of B. eriopoda when compared with plots receiving multiple small events. Because the strength of these patterns differed between years, we propose a modification of the bucket model in which both the mean and variance of soil water change as a consequence of interannual variability from 1 year to the next. Our results demonstrate that aridland ecosystems are highly sensitive to increased precipitation variability, and that more extreme precipitation events will likely have a positive impact on some aridland ecosystem processes important for the carbon cycle.     

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.     

Does Drought Influence the Relationship Between Biodiversity and Ecosystem Functioning in Boreal Forests?

In mixed forests, interactions among species influence ecosystem functioning but environmental conditions also play an important role in shaping relationships between biodiversity and ecosystem functioning. In the context of climate change, the carbon and water balance in pure versus mixed forest stands may be differentially influenced by changing soil water availability. To test this hypothesis, we compared the influence of biodiversity on stand water use efficiency (WUE_S) in boreal forests between wet and dry years. We assessed the carbon isotope composition (δ ¹³C) of tree rings in Betula pendula, Pinus sylvestris, and Picea abies growing in pure versus mixed stands. In addition, we tested whether differences in WUE_S affected patterns of stand basal area increment (BAI_S). No biodiversity effect was found for stand δ ¹³C (δ ¹³C_S) during the wet year. However, there was a significant increase in δ ¹³C_S between the wet and the dry year and a significant effect of biodiversity on δ ¹³C_S in the dry year. The increase in δ ¹³C_S in mixed stands was associated with both selection and complementarity effects. Although BAI_S decreased significantly in the dry year, changes in δ ¹³C_S did not translate into variations in BAI_S along the biodiversity gradient. Our results confirmed that the physiological response of boreal forest ecosystems to changing soil water conditions is influenced by species interactions and that during dry growing seasons, species interactions in mixed stands can lead to lower soil moisture availability. This illustrates that biodiversity effects can also be negative in mixed stands in the sense that soil resources can be more intensively exhausted. Overall, our results confirm that in boreal forests, the biodiversity–ecosystem functioning relationship depends on local environmental conditions.     

Carbon cycling in temperate grassland under elevated temperature

An increase in mean soil surface temperature has been observed over the last century, and it is predicted to further increase in the future. The effect of increased temperature on ecosystem carbon fluxes in a permanent temperate grassland was studied in a long‐term (6 years) field experiment, using multiple temperature increments induced by IR lamps. Ecosystem respiration (R‐eco) and net ecosystem exchange (NEE) were measured and modeled by a modified Lloyd and Taylor model including a soil moisture component for R‐eco (average R² of 0.78) and inclusion of a photosynthetic component based on temperature and radiation for NEE (R² = 0.65). Modeled NEE values ranged between 2.3 and 5.3 kg CO₂ m^?2 year^?1, depending on treatment. An increase of 2 or 3°C led to increased carbon losses, lowering the carbon storage potential by around 4 tonnes of C ha^?1 year^?1. The majority of significant NEE differences were found during night‐time compared to daytime. This suggests that during daytime the increased respiration could be offset by an increase in photosynthetic uptake. This was also supported by differences in δ¹³C and δ¹⁸O, indicating prolonged increased photosynthetic activity associated with the higher temperature treatments. However, this increase in photosynthesis was insufficient to counteract the 24 h increase in respiration, explaining the higher CO₂ emissions due to elevated temperature.     

Comparison of ecosystem water-use efficiency among Douglas-fir forest, aspen forest and grassland using eddy covariance and carbon isotope techniques

Comparisons were made among Douglas‐fir forest, aspen (broad leaf deciduous) forest and wheatgrass (C₃) grassland for ecosystem‐level water‐use efficiency (WUE). WUE was defined as the ratio of photosynthetic CO₂ assimilation rate and evapotranspiration (ET) rate. The ET data measured by eddy covariance were screened so that they overwhelmingly represented transpiration. The three sites used in this comparison spanned a range of vegetation (plant functional) types and environmental conditions within western Canada. When compared in the relative order Douglas‐fir (located on Vancouver Island, BC), aspen (northern Saskatchewan), grassland (southern Alberta), the sites demonstrated a progressive decline in precipitation and a general increase in maximum air temperature and atmospheric saturation deficit (D_max) during the mid‐summer. The average (±SD) WUE at the grassland site was 2.6±0.7 mmol mol^?1, which was much lower than the average values observed for the two other sites (aspen: 5.4±2.3, Douglas‐fir: 8.1±2.4). The differences in WUE among sites were primarily because of variation in ET. The highest maximum ET rates were approximately 5, 3.2 and 2.7 mm day^?1 for the grassland, aspen and Douglas‐fir sites, respectively. There was a strong negative correlation between WUE and D_max for all sites. We also made seasonal measurements of the carbon isotope ratio of ecosystem respired CO₂ (δ_R) in order to test for the expected correlation between shifts in environmental conditions and changes to the ecosystem‐integrated ratio of leaf intercellular to ambient CO₂ concentration (c_i/c_a). There was a consistent increase in δ_R values in the grassland, aspen forest and Douglas‐fir forest associated with a seasonal reduction in soil moisture. Comparisons were made between WUE measured using eddy covariance with that calculated based on D and δ_R measurements. There was excellent agreement between WUE values calculated using the two techniques. Our δ_R measurements indicated that c_i/c_a values were quite similar among the Douglas‐fir, aspen and grassland sites, despite large variation in environmental conditions among sites. This implied that the shorter‐lived grass species had relatively high c_i/c_a values for the D of their habitat. By contrast, the longer‐lived Douglas‐fir trees were more conservative in water‐use with lower c_i/c_a values relative to their habitat D. This illustrates the interaction between biological and environmental characteristics influencing ecosystem‐level WUE. The strong correlation we observed between the two independent measurements of WUE, indicates that the stable isotope composition of respired CO₂ is a useful ecosystem‐scale tool to help study constraints to photosynthesis and acclimation of ecosystems to environmental stress.     

Carbon and oxygen isotope composition of organic compounds in the phloem sap provides a short-term measure for stomatal conductance of European beech (Fagus sylvatica L.)

At eight different dates during the 2000 growing season, δ¹³C and δ¹⁸O were determined in the phloem of adult beech trees growing in natural beech stands in south‐west Germany differing in stand density and local climate. In addition, stand transpiration, precipitation, photosynthetic active radiation, relative air humidity, water pressure deficit of the air, air and soil temperature, soil water potential, and sugar concentration of the phloem sap were determined directly and evaporation and canopy stomatal conductance were modelled. All parameters were related to δ¹³C. The study aimed to identify the time integral within which the δ¹³C of organic compounds transported in the phloem is an indicative measure of these environmental influences. δ¹³C of soluble carbon transported in the phloem was well correlated with mean stomatal conductance in a two‐day integral prior to phloem sampling but did not depend on either light intensity or soil water availability. A strong positive relationship between δ¹³C and δ¹⁸O pointed to observed variation in δ¹³C of phloem sap being a result of variation in stomatal conductance. Bulk leaf δ¹³C was a poor indicator of changes in environmental conditions during the growing season. From these results we conclude that the analysis of δ¹³C in soluble carbon transported in the phloem is a reliable indicator of short‐term changes in C_i/C_a. In contrast, the δ¹³C of structural carbon in beech foliage represents an integration of a range of factors that mask short‐term influences responsible for C_i/C_a.     

Diel variations in carbon isotopic composition and concentration of organic acids and their impact on plant dark respiration in different species

Leaf respiration in the dark and its C isotopic composition (δ¹³C_R) contain information about internal metabolic processes and respiratory substrates. δ¹³C_R is known to be less negative compared to potential respiratory substrates, in particular shortly after darkening during light enhanced dark respiration (LEDR). This phenomenon might be driven by respiration of accumulated ¹³C‐enriched organic acids, however, studies simultaneously measuring δ¹³C_R during LEDR and potential respiratory substrates are rare. We determined δ¹³C_R and respiration rates (R) during LEDR, as well as δ¹³C and concentrations of potential respiratory substrates using compound‐specific isotope analyses. The measurements were conducted throughout the diel cycle in several plant species under different environmental conditions. δ¹³C_R and R patterns during LEDR were strongly species‐specific and showed an initial peak, which was followed by a progressive decrease in both values. The species‐specific differences in δ¹³C_R and R during LEDR may be partially explained by the isotopic composition of organic acids (e.g., oxalate, isocitrate, quinate, shikimate, malate), which were ¹³C‐enriched compared to other respiratory substrates (e.g., sugars and amino acids). However, the diel variations in both δ¹³C and concentrations of the organic acids were generally low. Thus, additional factors such as the heterogeneous isotope distribution in organic acids and the relative contribution of the organic acids to respiration are required to explain the strong ¹³C enrichment in leaf dark‐respired CO₂.     

Land use and season affect fluxes of CO2, CH4, CO,N2O,H2 and isotopic source signatures in Panama: evidence from nocturnal boundary layer profiles

Conversion of tropical rainforests to pastures and plantations is associated with changes in soil properties and biogeochemical cycling, with implications for carbon cycling and trace gas fluxes. The stable isotopic composition of ecosystem respiration (δ¹³C_R and δ¹⁸O_R) is used in inversion models to quantify regional patterns of CO₂ sources and sinks, but models are limited by sparse measurements in tropical regions. We measured soil respiration rates, concentrations of CO₂, CH₄, CO, N₂O and H₂ and the isotopic composition of CO₂, CH₄ and H₂ at four heights in the nocturnal boundary layer (NBL) above three common land‐use types in central Panama, during dry and rainy seasons. Soil respiration rates were lowest in Plantation (average 3.4 μmol m^?2 s^?1), highest in Pasture (8.3 μmol m^?2 s^?1) and intermediate in Rainforest (5.2 μmol m^?2 s^?1). δ¹³C_R closely reflected land use and increased during the dry season where C₃ vegetation was present. δ¹⁸O_R did not differ by land use but was lower during the rainy than the dry season. CO₂ was correlated with other species in approximately half of the NBL profiles, allowing us to estimate trace gas fluxes that were generally within the range of literature values. The Rainforest soil was a sink for CH₄ but emissions were observed in Pasture and Plantation, especially during the wet season. N₂O emissions were higher in Pasture and Plantation than Rainforest, contrary to expectations. Soil H₂ uptake was highest in Rainforest and was not observable in Pasture and Plantation during the wet season. We observed soil CO uptake during the dry season and emissions during the wet season across land‐use types. This study demonstrated that strong impacts of land‐use change on soil–atmosphere trace gas exchange can be detected in the NBL, and provides useful observational constraints for top‐down and bottom‐up biogeochemistry models.     

Seasonal variation in δ13C and δ18O of cellulose from growth rings of Pinus radiata

Seasonal variation in δ¹³C and δ¹⁸O of cellulose (δ¹³C_c and δ¹⁸O_c) was measured within two annual rings of Pinus radiata growing at three sites in New Zealand. In general, both δ¹³C_c and δ¹⁸O_c increased to a peak over summer. The three sites differed markedly in annual water balance, and these differences were reflected in δ¹³C_c and δ¹⁸O_c. Average δ¹³C_c and δ¹⁸O_c from each site were positively related, so that the driest site had the most enriched cellulose. δ¹³C_c and δ¹⁸O_c were also related within each site, although both the slope and the closeness of fit of the relationship varied between sites. Supporting the theory, the site with the lowest average relative humidity also had the greatest change in δ¹⁸O_c‰ change in δ¹³C_c. Specific climatic events, such as drought or high rainfall, were recorded as a peak or a trough in enrichment, respectively. These results suggest that seasonal and between‐site variation in δ¹³C_c and δ¹⁸O_c are driven by the interaction between variation in climatic conditions and soil water availability, and plant response to this variation.     

Carbon and nitrogen isotope fractionation of amino acids in an avian marine predator,the gentoo penguin (Pygoscelis papua)

Compound‐specific stable isotope analysis (CSIA) of amino acids (AA) has rapidly become a powerful tool in studies of food web architecture, resource use, and biogeochemical cycling. However, applications to avian ecology have been limited because no controlled studies have examined the patterns in AA isotope fractionation in birds. We conducted a controlled CSIA feeding experiment on an avian species, the gentoo penguin (Pygoscelis papua), to examine patterns in individual AA carbon and nitrogen stable isotope fractionation between diet (D) and consumer (C) (Δ¹³C_C‐D and Δ¹⁵N_C‐D, respectively). We found that essential AA δ¹³C values and source AA δ¹⁵N values in feathers showed minimal trophic fractionation between diet and consumer, providing independent but complimentary archival proxies for primary producers and nitrogen sources respectively, at the base of food webs supporting penguins. Variations in nonessential AA Δ¹³C_C‐D values reflected differences in macromolecule sources used for biosynthesis (e.g., protein vs. lipids) and provided a metric to assess resource utilization. The avian‐specific nitrogen trophic discrimination factor (TDF_Glu‐Phe = 3.5 ± 0.4‰) that we calculated from the difference in trophic fractionation (Δ¹⁵N_C‐D) of glutamic acid and phenylalanine was significantly lower than the conventional literature value of 7.6‰. Trophic positions of five species of wild penguins calculated using a multi‐TDF_Glu‐Phe equation with the avian‐specific TDF_Glu‐Phe value from our experiment provided estimates that were more ecologically realistic than estimates using a single TDF_Glu‐Phe of 7.6‰ from the previous literature. Our results provide a quantitative, mechanistic framework for the use of CSIA in nonlethal, archival feathers to study the movement and foraging ecology of avian consumers.     

Root exclusion through trenching does not affect the isotopic composition of soil CO2 efflux

Disentangling the autotrophic and heterotrophic components of soil CO₂ efflux is critical to understanding the role of soil system in terrestrial carbon (C) cycling. In this study, we combined a stable C-isotope natural abundance approach with the trenched plot method to determine if root exclusion significantly affected the isotopic composition (δ¹³C) of soil CO₂ efflux (R_S). This study was performed in different forest ecosystems: a tropical rainforest and two temperate broadleaved forests, where trenched plots had previously been installed. At each site, R_S and its δ¹³C (δ¹³C_Rs) tended to be lower in trenched plots than in control plots. Contrary to R_S, δ¹³C_Rs differences were not significant. This observation is consistent with the small differences in δ¹³C measured on organic matter from root, litter and soil. The lack of an effect on δ¹³C_Rs by root exclusion could be from the small difference in δ¹³C between autotrophic and heterotrophic soil respirations, but further investigations are needed because of potential artefacts associated with the root exclusion technique.     

Soil carbon inventories and δ13C along a moisture gradient in Botswana

We present a study of soil organic carbon (SOC) inventories and δ¹³C values for 625 soil cores collected from well‐drained, coarse‐textured soils in eight areas along a 1000 km moisture gradient from Southern Botswana, north into southern Zambia. The spatial distribution of trees and grass in the desert, savannah and woodland ecosystems along the transect control large systematic local variations in both SOC inventories and δ¹³C values. A stratified sampling approach was used to smooth this variability and obtain robust weighted‐mean estimates for both parameters. Weighted SOC inventories in the 0–5 cm interval of the soils range from 7 mg cm^?2 in the driest area (mean annual precipitation, MAP=225 mm) to 41±12 mg cm^?2 in the wettest area (MAP=910 mm). For the 0–30 cm interval, the inventories are 37.8 mg cm^?2 for the driest region and 157±33 mg cm^?2 for the wettest region. SOC inventories at intermediate sites increase as MAP increases to approximately 400–500 mm, but remain approximately constant thereafter. This plateau may be the result of feedbacks between MAP, fuel load and fire frequency. Weighted δ ¹³C values decrease linearly in both the 0–5 and 0–30 cm depth intervals as MAP increases. A value of –17.5±1.0‰ characterizes the driest areas, while a value of ?25±0.7‰ characterizes the wettest area. The decrease in δ ¹³C value with increasing MAP reflects an increasing dominance of C₃ vegetation as MAP increases. SOC in the deeper soil (5–30 cm depth) is, on average, 0.4±0.3‰ enriched in ¹³C relative to SOC in the 0–5 cm interval.     

Soil respiration is stimulated by elevated CO2 and reduced by summer drought: three years of measurements in a multifactor ecosystem manipulation experiment in a temperate heathland (CLIMAITE)

This study investigated the impact of predicted future climatic and atmospheric conditions on soil respiration (R_S) in a Danish Calluna‐Deschampsia‐heathland. A fully factorial in situ experiment with treatments of elevated atmospheric CO₂ (+130 ppm), raised soil temperature (+0.4 °C) and extended summer drought (5–8% precipitation exclusion) was established in 2005. The average R_S, observed in the control over 3 years of measurements (1.7 μmol CO₂ m^?2 sec^?1), increased 38% under elevated CO₂, irrespective of combination with the drought or temperature treatments. In contrast, extended summer drought decreased R_S by 14%, while elevated soil temperature did not affect R_S overall. A significant interaction between elevated temperature and drought resulted in further reduction of R_S when these treatments were combined. A detailed analysis of short‐term R_S dynamics associated with drought periods showed that R_S was reduced by ~50% and was strongly correlated with soil moisture during these events. Recovery of R_S to pre‐drought levels occurred within 2 weeks of rewetting; however, unexpected drought effects were observed several months after summer drought treatment in 2 of the 3 years, possibly due to reduced plant growth or changes in soil water holding capacity. An empirical model that predicts R_S from soil temperature, soil moisture and plant biomass was developed and accounted for 55% of the observed variability in R_S. The model predicted annual sums of R_S in 2006 and 2007, in the control, were 672 and 719 g C m^?2 y^?1, respectively. For the full treatment combination, i.e. the future climate scenario, the model predicted that soil respiratory C losses would increase by ~21% (140–150 g C m^?2 y^?1). Therefore, in the future climate, stimulation of C storage in plant biomass and litter must be in excess of 21% for this ecosystem to not suffer a reduction in net ecosystem exchange.     

Wetting and drying cycles drive variations in the stable carbon isotope ratio of respired carbon dioxide in semi-arid grassland

In semi-arid regions, where plants using both C₃ and C₄ photosynthetic pathways are common, the stable C isotope ratio (δ¹³C) of ecosystem respiration (δ¹³C_R) is strongly variable seasonally and inter-annually. Improved understanding of physiological and environmental controls over these variations will improve C cycle models that rely on the isotopic composition of atmospheric CO₂. We hypothesized that timing of precipitation events and antecedent moisture interact with activity of C₃ and C₄ grasses to determine net ecosystem CO₂ exchange (NEE) and δ¹³C_R. Field measurements included CO₂ and δ¹³C fluxes from the whole ecosystem and from patches of different plant communities, biomass and δ¹³C of plants and soils over the 2000 and 2001 growing seasons. NEE shifted from C source to sink in response to rainfall events, but this shift occurred after a time lag of up to 2 weeks if a dry period preceded the rainfall. The seasonal average of δ¹³C_R was higher in 2000 (−16‰) than 2001 (20‰), probably due to drier conditions during the 2000 growing season (79.7 mm of precipitation from April up to and including July) than in 2001 (189 mm). During moist conditions, δ¹³C averaged −22‰ from C₃ patches, −16‰ from C₄ patches, and −19‰ from mixed C₃ and C₄ patches. However, during dry conditions the apparent spatial differences were not obvious, suggesting reduced autotrophic activity in C₄ grasses with shallow rooting depth, soon after the onset of dry conditions. Air and soil temperatures were negatively correlated with δ¹³C_R; vapor pressure deficit was a poor predictor of δ¹³C_R, in contrast to more mesic ecosystems. Responses of respiration components to precipitation pulses were explained by differences in soil moisture thresholds between C₃ and C₄ species. Stable isotopic composition of respiration in semi-arid ecosystems is more temporally and spatially variable than in mesic ecosystems owing to dynamic aspects of pulse precipitation episodes and biological drivers.