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.     

Isotope fractionation and 13C enrichment in soil profiles during the decomposition of soil organic matter

The mechanisms behind the ¹³C enrichment of organic matter with increasing soil depth in forests are unclear. To determine if ¹³C discrimination during respiration could contribute to this pattern, we compared δ¹³C signatures of respired CO₂ from sieved mineral soil, litter layer and litterfall with measurements of δ¹³C and δ¹⁵N of mineral soil, litter layer, litterfall, roots and fungal mycelia sampled from a 68-year-old Norway spruce forest stand planted on previously cultivated land. Because the land was subjected to ploughing before establishment of the forest stand, shifts in δ¹³C in the top 20 cm reflect processes that have been active since the beginning of the reforestation process. As ¹³C-depleted organic matter accumulated in the upper soil, a 1.0‰ δ¹³C gradient from −28.5‰ in the litter layer to −27.6‰ at a depth of 2–6 cm was formed. This can be explained by the 1‰ drop in δ¹³C of atmospheric CO₂ since the beginning of reforestation together with the mixing of new C (forest) and old C (farmland). However, the isotopic change of the atmospheric CO₂ explains only a portion of the additional 1.0‰ increase in δ¹³C below a depth of 20 cm. The δ¹³C of the respired CO₂ was similar to that of the organic matter in the upper soil layers but became increasingly ¹³C enriched with depth, up to 2.5‰ relative to the organic matter. We hypothesise that this ¹³C enrichment of the CO₂ as well as the residual increase in δ¹³C of the organic matter below a soil depth of 20 cm results from the increased contribution of ¹³C-enriched microbially derived C with depth. Our results suggest that ¹³C discrimination during microbial respiration does not contribute to the ¹³C enrichment of organic matter in soils. We therefore recommend that these results should be taken into consideration when natural variations in δ¹³C of respired CO₂ are used to separate different components of soil respiration or ecosystem respiration.     

Carbon nutrition of mature green orchid <Emphasis Type="Italic">Serapias strictiflora</Emphasis> and its mycorrhizal fungus <Emphasis Type="Italic">Epulorhiza</Emphasis> sp.

We studied the nutritional modes of the orchid Serapias strictiflora and its mycorrhizal fungus Epulorhiza sp. using the differences in carbon isotopic composition (δ¹³C) of C₃ orchid and C₄ maize tissues. We found that if cultivated in substrate lacking any organic compounds, the mycorrhizal extraradical mycelia (δ¹³C = −26.3 ± 0.2 ‰) developed well, despite being fully dependent on nutrition from orchid roots (δ¹³C = −28.6 ± 0.1 ‰). If the mycorrhizal fungus had additional access to and colonized decaying maize roots (δ¹³C = −14.6 ± 0.1 ‰), its isotopic composition (δ¹³C = −21.6 ± 0.4 ‰) reflected a mixture of biotrophy and saprotrophy. No statistically significant differences in δ¹³C of new storage tubers were found between Epulorhiza-associated orchids with (δ¹³C = -28.2 ± 0.1 ‰) and without access to maize roots (δ¹³C = −28.6 ± 0.2 ‰). We conclude that autotrophy is the predominant nutritional mode of mature S. strictiflora plants and that they supply their mycorrhizal fungus with substantial amount of carbon (69 ± 3 % of the fungus demand), even if the fungus feeds saprotrophically.     

Deep Autotrophic Soil Respiration in Shrubland and Woodland Ecosystems in Central New Mexico

Quantifying the controls on soil respiration is important for understanding ecosystem physiology and for predicting the response of soil carbon reservoirs to climate change. The majority of soil respiration is typically considered to occur in the top 20–30 cm of soils. In desert soils, where organic matter concentrations tend to be low and plants are deeply rooted, deeper respiration might be expected. However, little is known about the depth distribution of respiration in dryland soils. Here we show that the average depth of soil respiration between pulse precipitation events is almost always greater than 20 cm and is frequently greater than 50 cm in two central New Mexico desert shrublands. The average depth of soil respiration in a pi?on-juniper woodland was shallower, between 5 and 40 cm. In the shrublands, 8‰ seasonal variations in the carbon isotope composition of soil-respired CO₂ (δ¹³C_r-soil) that correlate with vapor pressure deficit support root/rhizosphere respiration as the dominant source of soil CO₂. Such deep autotrophic respiration indicates that shrubs preferentially allocate photosynthate to deep roots when conditions near the surface are unfavorable. Therefore, respiration rates in these soils are not necessarily correlated with root biomass. The δ¹³C_r-soil values provide no evidence for CO₂ evolved from soil inorganic carbon. Our results also suggest that organic carbon cycling is rapid and efficient in these soils and that the δ¹³C value of CO₂ respired from soils in much of the southwestern US, and perhaps in other semiarid regions, varies seasonally by at least 4‰.     

Changes in stable isotopic signatures of soil nitrogen and carbon during 40 years of forest development

Understanding what governs patterns of soil δ¹⁵N and δ¹³C is limited by the absence of these data assembled throughout the development of individual ecosystems. These patterns are important because stable isotopes of soil organic N and C are integrative indicators of biogeochemical processing of soil organic matter. We examined δ¹⁵N of soil organic matter (δ¹⁵N_SOM) and δ¹³C_SOM of archived soil samples across four decades from four depths of an aggrading forest in southeastern USA. The site supports an old-field pine forest in which the N cycle is affected by former agricultural fertilization, massive accumulation of soil N by aggrading trees over four decades, and small to insignificant fluxes of N via NH₃ volatilization, nitrification, and denitrification. We examine isotopic data and the N and C dynamics of this ecosystem to evaluate mechanisms driving isotopic shifts over time. With forest development, δ¹³C_SOM became depth-dependent. This trend resulted from a decline of ~2‰ in the surficial 15 cm of mineral soil to −26.0‰, due to organic matter inputs from forest vegetation. Deeper layers exhibited relatively little trend in δ¹³C_SOM with time. In contrast, δ¹⁵N_SOM was most dynamic in deeper layers. During the four decades of forest development, the deepest layer (35–60 cm) reached a maximum δ¹⁵N value of 9.1‰, increasing by 7.6‰. The transfer of >800 kg ha⁻¹ of soil organic N into aggrading vegetation and the forest floor and the apparent large proportion of ectomycorrhizal (ECM) fungi in these soils suggest that fractionation via microbial transformations must be the major process changing δ¹⁵N in these soils. Accretion of isotopically enriched compounds derived from microbial cells (i.e., ECM fungi) likely promote isotopic enrichment of soils over time. The work indicates the rapid rate at which ecosystem development can impart δ¹⁵N_SOM and δ¹³C_SOM signatures associated with undisturbed soil profiles.     

Nocturnal and seasonal patterns of carbon isotope composition of leaf dark-respired carbon dioxide differ among dominant species in a semiarid savanna

The C isotope composition of leaf dark-respired CO₂ (δ¹³C_l) integrates short-term metabolic responses to environmental change and is potentially recorded in the isotopic signature of ecosystem-level respiration. Species differences in photosynthetic pathway, resource acquisition and allocation patterns, and associated isotopic fractionations at metabolic branch points can influence δ¹³C_l, and differences are likely to be modified by seasonal variation in drought intensity. We measured δ¹³C_l in two deep-rooted C₃ trees (Prosopis velutina and Celtis reticulata), and two relatively shallow-rooted perennial herbs (a C₃ dicot Viguiera dentata and a C₄ grass Sporobolus wrightii) in a floodplain savanna ecosystem in southeastern Arizona, USA during the dry pre-monsoon and wet monsoon seasons. δ¹³C_l decreased during the nighttime and reached minimum values at pre-dawn in all species. The magnitude of nocturnal shift in δ¹³C_l differed among species and between pre-monsoon and monsoon seasons. During the pre-monsoon season, the magnitude of the nocturnal shift in δ¹³C_l in the deep-rooted C₃ trees P. velutina (2.8 ± 0.4‰) and C. reticulata (2.9 ± 0.2‰) was greater than in the C₃ herb V. dentata (1.8 ± 0.4‰) and C₄ grass S. wrightii (2.2 ± 0.4‰). The nocturnal shift in δ¹³C_l in V. dentata and S. wrightii increased to 3.2 ± 0.1‰ and 4.6 ± 0.6‰, respectively, during the monsoon season, but in C₃ trees did not change significantly from pre-monsoon values. Cumulative daytime net CO₂ uptake was positively correlated with the magnitude of the nocturnal decline in δ¹³C_l across all species, suggesting that nocturnal δ¹³C_l may be controlled by ¹³C/¹²C fractionations associated with C substrate availability and C metabolite partitioning. Nocturnal patterns of δ¹³C_l in dominant plant species in the semiarid savanna apparently have predictable responses to seasonal changes in water availability, which is important for interpreting and modeling the C isotope signature of ecosystem-respired CO₂.     

The carbon origin of arbuscular mycorrhizal fungi estimated from δ13C values of individual spores

 The origin of carbon in the spores of arbuscular mycorrhizal (AM) fungi was quantified based on their obligate symbiosis with C₃ and C₄ plants showing clearly different δ¹³C values. The δ¹³C values of individual spores of the AM fungus Gigaspora margarita were analyzed. In monoculture pots of a C₃ or a C₄ plant species, spore δ¹³C values were ca. 3.5‰ lower than those of host roots. In coculture pots of a C₃ and a C₄ plant species, spore δ¹³C values varied between those of the roots of C₃ and C₄ plants, and increased linearly from the C₃ to the proximity of the C₄ plant (P<0.01). This reflects the higher δ¹³C values in C₄ plants than in C₃ plants. Thus the carbon origin of G. margarita spores changed with growth state and combination of host plants. In the presence of fresh plant residue instead of living host plants, spore δ¹³C values did not vary with distance from the residue. This finding supports the current view that AM fungi are obligate symbionts. Accepted: 12 February 1999     

Interseasonal comparison of CO2 concentrations, isotopic composition, and carbon dynamics in an Amazonian rainforest (French Guiana)

Canopy CO₂ concentrations in a tropical rainforest in French Guiana were measured continuously for 5 days during the 1994 dry season and the 1995 wet season. Carbon dioxide concentrations ([CO₂]) throughout the canopy (0.02–38 m) showed a distinct daily pattern, were well-stratified and decreased with increasing height into the canopy. During both seasons, daytime [CO₂] in the upper and middle canopy decreased on average 7–10 μmol mol⁻¹ below tropospheric baseline values measured at Barbados. Within the main part of the canopy (≥ 0.7 m), [CO₂] did not differ between the wet and dry seasons. In contrast, [CO₂] below 0.7 m were generally higher during the dry season, resulting in larger [CO₂] gradients. Supporting this observation, soil CO₂ efflux was on average higher during the dry season than during the wet season, either due to diffusive limitations and/or to oxygen deficiency of root and microbial respiration. Soil respiration rates decreased by 40% after strong rain events, resulting in a rapid decrease in canopy [CO₂] immediately above the forest floor of about 50␣μmol mol⁻¹. Temporal and spatial variations in [CO₂]_canopy were reflected in changes of δ¹³C_canopy and δ¹⁸O_canopy values. Tight relationships were observed between δ¹³C and δ¹⁸O of canopy CO₂ during both seasons (r ² > 0.86). The most depleted δ¹³C_canopy and δ¹⁸O_canopy values were measured immediately above the forest floor (δ¹³C = −16.4‰; δ¹⁸O = 39.1‰ SMOW). Gradients in the isotope ratios of CO₂ between the top of the canopy and the forest floor ranged between 2.0‰ and 6.3‰ for δ¹³C, and between 1.0‰ and 3.5‰ for δ¹⁸O. The δ¹³C_leaf and calculated c _i/c _a of foliage at three different positions were similar for the dry and wet seasons indicating that the canopy maintained a constant ratio of photosynthesis to stomatal conductance. About 20% of the differences in δ¹³C_leaf within the canopy was accounted for by source air effects, the remaining 80% must be due to changes in c _i/c _a. Plotting 1/[CO₂] vs. the corresponding δ¹³C ratios resulted in very tight, linear relationships (r ² = 0.99), with no significant differences between the two seasons, suggesting negligible seasonal variability in turbulent mixing relative to ecosystem gas exchange. The intercepts of these relationships that should be indicative of the δ¹³C of respired sources were close to the measured δ¹³C of soil respired CO₂ and to the δ¹³C of litter and soil organic matter. Estimates of carbon isotope discrimination of the entire ecosystem, Δ_e, were calculated as 20.3‰ during the dry season and as 20.5‰ during the wet season. Received: 3 March 1996 / Accepted: 19 October 1996     

Effect of C3–C4 Vegetation Change on δ13C and δ15N Values of Soil Organic Matter Fractions Separated by Thermal Stability

Carbon isotopic composition of soils subjected to C₃–C₄ vegetation change can be used to estimate C turnover in bulk soil and in soil organic matter (SOM) pools with fast and intermediate turnover rates. We hypothesized that the biological availability of SOM pools is inversely proportional to their thermal stability, so that thermogravimetry can be used to separate SOM pools with contrasting turnover rates. Soil samples from a field plot cultivated for 10.5 years with the perennial C₄ plant Miscanthus×gigantheus were analyzed by thermogravimetry coupled with differential scanning calorimetry (DSC). Three SOM fractions were distinguished according to the differential weight losses and exothermic or endothermic reactions measured by DSC. The δ¹³C and δ¹⁵N values of these three fractions obtained by gradual soil heating were measured by IRMS. The weight losses up to 190 °C mainly reflected water evaporation because no significant C and N losses were detected and δ¹³C and δ¹⁵N values of the residual SOM remained unchanged. The δ¹³C values (−16.4‰) of SOM fraction decomposed between 190 and 390 °C (containing 79% of total soil C) were slightly closer to that of the Miscanthus plant tissues (δ¹³C = −11.8‰) compared to the δ¹³C values (−16.8‰) of SOM fraction decomposed above 390 °C containing the residual 21% of SOM. Thus, the C turnover in the thermally labile fraction was faster than that in thermally stable fractions, but the differences were not very strong. Therefore, in this first study combining TG-DSC with isotopic analysis, we conclude that the thermal stability of SOM was not very strongly related to biological availability of SOM fractions. In contrast to δ¹³C, the δ¹⁵N values strongly differed between SOM fractions, suggesting that N turnover in the soil was different from C turnover. More detailed fractionation of SOM by thermal analysis with subsequent isotopic analysis may improve the resolution for δ¹³C.     

Determinants of isotopic coupling of CO2 and water vapour within a Quercus petraea forest canopy

Concentration and isotopic composition (δ¹³C and δ¹⁸O) of ambient CO₂ and water vapour were determined within a Quercus petraea canopy, Northumberland, UK. From continuous measurements made across a 36-h period from three heights within the forest canopy, we generated mixing lines (Keeling plots) for δ_a ¹³CO₂, δ_a C¹⁸O¹⁶O and δ_a H₂ ¹⁸O, to derive the isotopic composition of the signal being released from forest to atmosphere. These were compared directly with measurements of different respective pools within the forest system, i.e. δ¹³C of organic matter input for δ_a ¹³CO₂, δ¹⁸O of exchangeable water for δ_a C¹⁸O¹⁶O and transpired water vapour for δ_a H₂ ¹⁸O. [CO₂] and δ_a ¹³CO₂ showed strong coupling, where the released CO₂ was, on average, 4 per mil enriched compared to the organic matter of plant material in the system, suggesting either fractionation of organic material before eventual release as soil-respired CO₂, or temporal differences in ecosystem discrimination. δ_a C¹⁸O¹⁶O was less well coupled to [CO₂], probably due to the heterogeneity and transient nature of water pools (soil, leaf and moss) within the forest. Similarly, δ_a H₂ ¹⁸O was less coupled to [H₂O], again reflecting the transient nature of water transpired to the forest, seen as uncoupling during times of large changes in vapour pressure deficit. The δ¹⁸O of transpired water vapour, inferred from both mixing lines at the canopy scale and direct measurement at the leaf level, approximated that of source water, confirming that an isotopic steady state held for the forest integrated over the daily cycle. This demonstrates that isotopic coupling of CO₂ and water vapour within a forest canopy will depend on absolute differences in the isotopic composition of the respective pools involved in exchange and on the stability of each of these pools with time. Received: 21 March 1998 / Accepted: 10 December 1998     

Biological and physical influences on the carbon isotope content of CO<Subscript>2</Subscript> in a subalpine forest snowpack,Niwot Ridge,Colorado

Considerable research has recently been devoted to understanding biogeochemical processes under winter snow cover, leading to enhanced appreciation of the importance of many winter ecological processes. In this study, a comprehensive investigation of the stable carbon isotope composition (δ¹³C) of CO₂ within a high-elevation subalpine forest snowpack was conducted. Our goals were to study the δ¹³C of biological soil respiration under snow in winter, and to assess the relative importance of diffusion and advection (ventilation by wind) for gas transport within snow. In agreement with other studies, we found evidence of an active microbial community under a roughly 1-m deep snowpack during winter and into spring as it melted. Under-snow CO₂ mole fractions were observed up to 3,500 μmol mol⁻¹, and δ¹³C of CO₂ varied from ~−22 to ~−8‰. The δ¹³C of soil respiration calculated from mixing relationships was −26 to −24‰, and although it varied in time, it was generally close to that of the bulk organic horizon (−26.0‰). Subnivean CO₂ and δ¹³C were quite dynamic in response to changes in soil temperature, liquid water availability, and wind events. No clear biologically-induced isotopic changes were observed during periods when microbial activity and root/rhizosphere activity were expected to vary, although such changes cannot be eliminated. There was clear evidence of isotopic enrichment associated with diffusive transport as predicted by theory, but simple diffusive enrichment (4.4‰) was not observed. Instead, ventilation of the snowpack by sustained wind events in the forest canopy led to changes in the diffusively-enriched gas profile. The isotopic influence of diffusion on gases in the snowpack and litter was greatest at greater depths, due to the decreased relative contribution of advection at depth. There were highly significant correlations between the apparent isotopic content of respiration from the soil with wind speed and pressure. In summary, physical factors influencing gas transport substantially modified and potentially obscured biological factors in their effects on δ¹³C of CO₂ within this subalpine forest snowpack.     

Controlling for anthropogenically induced atmospheric variation in stable carbon isotope studies

Increased use of stable isotope analysis to examine food-web dynamics, migration, transfer of nutrients, and behavior will likely result in expansion of stable isotope studies investigating human-induced global changes. Recent elevation of atmospheric CO₂ concentration, related primarily to fossil fuel combustion, has reduced atmospheric CO₂ δ¹³C (¹³C/¹²C), and this change in isotopic baseline has, in turn, reduced plant and animal tissue δ¹³C of terrestrial and aquatic organisms. Such depletion in CO₂ δ¹³C and its effects on tissue δ¹³C may introduce bias into δ¹³C investigations, and if this variation is not controlled, may confound interpretation of results obtained from tissue samples collected over a temporal span. To control for this source of variation, we used a high-precision record of atmospheric CO₂ δ¹³C from ice cores and direct atmospheric measurements to model modern change in CO₂ δ¹³C. From this model, we estimated a correction factor that controls for atmospheric change; this correction reduces bias associated with changes in atmospheric isotopic baseline and facilitates comparison of tissue δ¹³C collected over multiple years. To exemplify the importance of accounting for atmospheric CO₂ δ¹³C depletion, we applied the correction to a dataset of collagen δ¹³C obtained from mountain lion (Puma concolor) bone samples collected in California between 1893 and 1995. Before correction, in three of four ecoregions collagen δ¹³C decreased significantly concurrent with depletion of atmospheric CO₂ δ¹³C (n ≥ 32, P ≤ 0.01). Application of the correction to collagen δ¹³C data removed trends from regions demonstrating significant declines, and measurement error associated with the correction did not add substantial variation to adjusted estimates. Controlling for long-term atmospheric variation and correcting tissue samples for changes in isotopic baseline facilitate analysis of samples that span a large temporal range.     

Assimilate partitioning affects 13C fractionation of recently assimilated carbon in maize

Coupling ¹³C natural abundance and ¹⁴C pulse labelling enabled us to investigate the dependence of ¹³C fractionation on assimilate partitioning between shoots, roots, exudates, and CO₂ respired by maize roots. The amount of recently assimilated C in these four pools was controlled by three levels of nutrient supply: full nutrient supply (NS), 10 times diluted nutrient supply (DNS), and deionised water (DW). After pulse labelling of maize shoots in a ¹⁴CO₂ atmosphere, ¹⁴C was traced to determine the amounts of recently assimilated C in the four pools and the δ¹³C values of the four pools were measured. Increasing amounts of recently assimilated C in the roots (from 8% to 10% of recovered ¹⁴C in NS and DNS treatments) led to a 0.3‰ ¹³C enrichment from NS to DNS treatments. A further increase of C allocation in the roots (from 10% to 13% of recovered ¹⁴C in DNS and DW treatments) resulted in an additional enrichment of the roots from DNS to DW treatments by 0.3‰. These findings support the hypothesis that ¹³C enrichment in a pool increases with an increasing amount of C transferred into that pool. δ¹³C of CO₂ evolved by root respiration was similar to that of the roots in DNS and DW treatments. However, if the amount of recently assimilated C in root respiration was reduced (NS treatment), the respired CO₂ became 0.7‰ ¹³C depleted compared to roots. Increasing amounts of recently assimilated C in the CO₂ from NS via DNS to DW treatments resulted in a 1.6‰ δ¹³C increase of root respired CO₂ from NS to DW treatments. Thus, for both pools, i.e. roots and root respiration, increasing amounts of recently assimilated C in the pool led to a δ¹³C increase. In DW and DNS plants there was no ¹³C fractionation between roots and exudates. However, high nutrient supply decreased the amount of recently assimilated C in exudates compared to the other two treatments and led to a 5.3‰ ¹³C enrichment in exudates compared to roots. We conclude that ¹³C discrimination between plant pools and within processes such as exudation and root respiration is not constant but strongly depends on the amount of C in the respective pool and on partitioning of recently assimilated C between plant pools. Section Editor: H. Lambers     

Stable isotope characteristics across narrow savanna/woodland ecotones in Wolfe Creek Meteorite Crater,Western Australia

The stable isotopic composition (δ¹³C) of sediments from lakes are frequently analyzed to reconstruct the proportion of the regional vegetation that used either the C₃ or C₄ photosynthetic pathways, often without conducting a detailed survey of the current local vegetation. We performed a study on the modern vegetation composition within the Wolfe Creek Meteorite Crater to complement our future paleoecological investigation of the crater. A bull’s-eye pattern exists where C₄ grasses dominate an outer ring and salt tolerant species, including shrubs, herbs, chenopods, and halophytic algae, dominate the inner pan of the crater. The ecotone between the inner and outer zones is narrow and occupied by tall (>7 m) Acacia ampliceps, with some C₄ grasses in the understory. Along with the highest water table and most saline soils the center of the crater has C₃ plants present with the highest δ¹³C and δ¹⁵N values. The range of δ¹³C and δ¹⁵N values from the analysis of surface soil organic matter (OM) was much smaller compared with the range of values from plant materials implying that either: (1) the current plant OM has not yet been integrated into the soils, or (2) processes within the soil have acted to homogenize isotopic variability within the crater. The application of a two end member mixing model to calculate %C₄ and %C₃ biomass from the δ¹³C of surface soil OM was complicated by: (1) the crater containing both a dry habitat with C₄ grasses and a central pan with C₄ halophytic plants and, (2) the large variation in the δ¹³C of the plants and soil OM.     

Patterns and controls of seasonal variability of carbon stable isotopes of particulate organic matter in lakes

Carbon stable isotopes (δ¹³C) of particulate organic matter (POM) have been used as indicators for energy flow, primary productivity and carbon dioxide concentration in individual lakes. Here, we provide a synthesis of literature data from 32 freshwater lakes around the world to assess the variability of δ¹³C_POM along latitudinal, morphometric and biogeochemical gradients. Seasonal mean δ¹³C_POM, a temporally integrated measure of the δ¹³C_POM, displayed weak relationships with all trophic state indices [total phosphorus (TP), total nitrogen (TN), and chlorophyll a (Chl a)], but decreased significantly with the increase in latitude, presumably in response to the corresponding decrease in water temperature and increase in CO₂ concentration. The seasonal minimum δ¹³C_POM also correlated negatively with latitude while seasonal maximum δ¹³C_POM correlated positively with all trophic state indices, pH, and δ¹³C of dissolved inorganic carbon (DIC). Seasonal amplitude of δ¹³C_POM (the difference between seasonal maximum and minimum values) correlated significantly with pH, TP and Chl a concentrations and displayed small variations in oligotrophic, mesotrophic and low latitude eutrophic lakes, which is attributed to low primary productivity and abundant non-living POM in the low trophic state lakes and relatively stable environmental conditions in the subtropics. Seasonal amplitude of δ¹³C_POM was the greatest in high latitude eutrophic lakes. Greater seasonal changes in solar energy and light regime may be responsible for the large seasonal variability in high latitude productive lakes. This synthesis provides new insights on the factors controlling variations in stable carbon isotopes of POM among lakes on the global scale.     

Insights into nitrogen and carbon dynamics of ectomycorrhizal and saprotrophic fungi from isotopic evidence

The successful use of natural abundances of carbon (C) and nitrogen (N) isotopes in the study of ecosystem dynamics suggests that isotopic measurements could yield new insights into the role of fungi in nitrogen and carbon cycling. Sporocarps of mycorrhizal and saprotrophic fungi, vegetation, and soils were collected in young, deciduous-dominated sites and older, coniferous-dominated sites along a successional sequence at Glacier Bay National Park, Alaska. Mycorrhizal fungi had consistently higher δ¹⁵N and lower δ¹³C values than saprotrophic fungi. Foliar δ¹³C values were always isotopically depleted relative to both fungal types. Foliar δ¹⁵N values were usually, but not always, more depleted than those in saprotrophic fungi, and were consistently more depleted than in mycorrhizal fungi. We hypothesize that an apparent isotopic fractionation by mycorrhizal fungi during the transfer of nitrogen to plants may be attributed to enzymatic reactions within the fungi producing isotopically depleted amino acids, which are subsequently passed on to plant symbionts. An increasing difference between soil mineral nitrogen δ¹⁵N and foliar δ¹⁵N in later succession might therefore be a consequence of greater reliance on mycorrhizal symbionts for nitrogen supply under nitrogen-limited conditions. Carbon signatures of mycorrhizal fungi may be more enriched than those of foliage because the fungi use isotopically enriched photosynthate such as simple sugars, in contrast to the mixture of compounds present in leaves. In addition, some ¹³C fractionation may occur during transport processes from leaves to roots, and during fungal chitin biosynthesis. Stable isotopes have the potential to help clarify the role of fungi in ecosystem processes. Received: 7 January 1998 / Accepted: 9 November 1998     

Measurement of net ecosystem production and ecosystem respiration in a Zoysia japonica grassland,central Japan,by the chamber method

Measuring light, temperature, soil moisture, and growth provides a better understanding of net ecosystem production (NEP), ecosystem respiration (R _eco), and their response functions. Here, we studied the variations in NEP and R _eco in a grassland dominated by a perennial warm-season C₄ grass, Zoysia japonica. We used the chamber method to measure NEP and R _eco from August to September 2007. Biomass and leaf area index (LAI) were also measured to observe their effects on NEP and R _eco. Diurnal variations in NEP and R _eco were predicted well by light intensity (PPFD) and by soil temperature, respectively. Maximum NEP (NEP_max) values on days of year 221, 233, 247, and 262, were 2.44, 2.55, 3.90, and 4.17 μmol m⁻² s⁻¹, respectively. Throughout the growing period, the apparent quantum yield (α) increased with increasing NEP_max that ranged from 0.0154 to 0.0515, and NEP responded to the soil temperature changes by 44% and R _eco changes by 48%, and R _eco responded from 88 to 94% with the soil temperature diurnally. NEP’s light response and R _eco’s temperature response were affected by soil water content; more than 27% of the variation in NEP and 67% of the variation in R _eco could be explained by this parameter. NEP was strongly correlated with biomass and LAI, but R _eco was not, because environmental variables affected R _eco more strongly than growth parameters. Using the light response of NEP, the temperature response of R _eco, and meteorological data, daily NEP and R _eco were estimated at 0.67, 0.81, 1.17, and 1.56 g C m⁻², and at 2.88, 2.50, 3.51, and 3.04 g C m⁻², respectively, on days of year 221, 233, 247, and 262. The corresponding daily gross primary production (NEP + R _eco) was 3.5, 3.3, 4.6, and 4.6 g C m⁻².     

Short-term variations in δ<Superscript>13</Superscript>C of ecosystem respiration reveals link between assimilation and respiration in a deciduous forest

We present a comprehensive dataset of hourly, daily, and monthly measurements of carbon isotope measurements of CO₂ in canopy air from a temperate deciduous forest with the aim to identify the relevance of short-term variations in the isotopic signature of ecosystem respiration (¹³C_R) and to understand its underlying physiological processes. We show that during daytime low vertical mixing inside the canopy can lead to decoupling of the air in the lower and upper canopy layer resulting in large spatial variation of ¹³C in CO₂ of canopy air. Intercept of Keeling Plots also showed large temporal variation (3.8) over the course of the day demonstrating that intercepts can differ between day and night and suggesting that choosing the right time for sampling is essential to capture the isotopic signature of ecosystem respiration (¹³C_R). ¹³C_R as obtained from night-time measurements showed large variation of up to 2.65 on a day-to-day basis, which was similar to the observed variation of ¹³C_R over the seasonal cycle (3.08). This highlights the importance of short-term physiological processes within ecosystems for the isotopic composition of CO₂ in the atmosphere, not reflected by bulk plant and soil organic samples. At daily and monthly time scales, ¹³C_R increased with increasing ratio of vapour pressure deficit to photosynthetically active radiation, measured 4–5 days before. This suggests that ecosystem respiration was isotopically linked to assimilation. Furthermore, assimilates recently fixed in the canopy seem to form a labile carbon pool with a short mean residence time that is respired back to the atmosphere after 4–5 days.     

Comparisons of δ<Superscript>13</Superscript>C of photosynthetic products and ecosystem respiratory CO<Subscript>2</Subscript> and their responses to seasonal climate variability

This study investigated the relationship between ¹³C of ecosystem components, soluble plant carbohydrates and the isotopic signature of ecosystem respired CO₂ (¹³C_R) during seasonal changes in soil and atmospheric moisture in a beech (Fagus sylvatica L.) forest in the central Apennine mountains, Italy. Decrease in soil moisture and increase in air vapour pressure deficit during summer correlated with substantial increase in ¹³C of leaf and phloem sap soluble sugars. Increases in ¹³C of ecosystem respired CO₂ were linearly related to increases in phloem sugar ¹³C (r²=0.99, P0.001) and leaf sugar ¹³C (r²=0.981, P0.01), indicating that a major proportion of ecosystem respired CO₂ was derived from recent assimilates. The slopes of the best-fit lines differed significantly (P0.05), however, and were about 0.86 (SE=0.04) for phloem sugars and about 1.63 (SE=0.16) for leaf sugars. Hence, changes in isotopic signature in phloem sugars were transferred to ecosystem respiration in the beech forest, while leaf sugars, with relatively small seasonal changes in ¹³C, must have a slower turnover rate or a significant storage component. No significant variation in ¹³C was observed in bulk dry matter of various plant and ecosystem components (including leaves, bark, wood, litter and soil organics). The apparent coupling between the ¹³C of soluble sugars and ecosystem respiration was associated with large apparent isotopic disequilibria. Values of ¹³C_R were consistently more depleted by about 4 relative to phloem sugars, and by about 2 compared to leaf sugars. Since no combination of the measured pools could produce the observed ¹³C_R signal over the entire season, a significant isotopic discrimination against ¹³C might be associated with short-term ecosystem respiration. However, these differences might also be explained by substantial contributions of other not measured carbon pools (e.g., lipids) to ecosystem respiration or contributions linked to differences in footprint area between Keeling plots and carbohydrate sampling. Linking the seasonal and inter-annual variations in carbon isotope composition of carbohydrates and respiratory CO₂ should be applicable in carbon cycle models and help the understanding of inter-annual variation in biospheric sink strength.     

Effects of growth and tissue type on the kinetics of <Superscript>13</Superscript>C and <Superscript>15</Superscript>N incorporation in a rapidly growing ectotherm

The use of stable isotopes to investigate animal diets, habitat use, and trophic level requires understanding the rate at which animals incorporate the ¹³C and ¹⁵N from their diets and the factors that determine the magnitude of the difference in isotopic composition between the animal’s diet and that of its tissues. We determined the contribution of growth and catabolic turnover to the rate of ¹³C and ¹⁵N incorporation into several tissues that can be sampled non-invasively (skin, scute, whole blood, red blood cells, and plasma solutes) in two age classes of a rapidly growing ectotherm (loggerhead turtles, Caretta caretta). We found significant differences in C and N incorporation rates and isotopic discrimination factors (Δ¹³C = δ¹³C_tissues − δ¹³C_diet and Δ¹⁵N = δ¹⁵N_tissues − δ¹⁵N_diet) among tissues and between age classes. Growth explained from 26 to 100% of the total rate of incorporation in hatchling turtles and from 15 to 52% of the total rate of incorporation in juvenile turtles. Because growth contributed significantly to the rate of isotopic incorporation, variation in rates among tissues was lower than reported in previous studies. The contribution of growth can homogenize the rate of isotopic incorporation and limit the application of stable isotopes to identify dietary changes at contrasting time scales and to determine the timing of diet shifts. The isotopic discrimination factor of nitrogen ranged from −0.64 to 1.77‰ in the turtles’ tissues. These values are lower than the commonly assumed average 3.4‰ discrimination factors reported for whole body and muscle isotopic analyses. The increasing reliance on non-invasive and non-destructive sampling in animal isotopic ecology requires that we recognize and understand why different tissues differ in isotopic discrimination factors.