Differential C isotope discrimination by fungi during decomposition of C(3)- and C(4)-derived sucrose

Stable isotope analysis is a major tool used in ecosystem studies to establish pathways and rates of C exchange between various ecosystem components. Little is known about isotopic effects of many such components, especially microbes. Here we report on the discovery of an unexpected pattern of C isotopic discrimination by basidiomycete fungi with far-reaching consequences for our understanding of isotopic processing in ecosystems where these microbes mediate material transfers across trophic levels. We measured fractionation effects on three ecologically relevant basidiomycete species under controlled laboratory conditions. Sucrose derived from C(3) and C(4) plants is fractionated differentially by these microbes in a taxon-specific manner. The differentiation between mycorrhizal and saprotrophic fungi observed in the field by others is not explained by intrinsic discrimination patterns. Fractionation occurs during sugar uptake and is sensitive to the nonrandom distribution of stable isotopes in the sucrose molecule. The balance between respiratory physiology and fermentative physiology modulates the degree of fractionation. These discoveries disprove the assumption that fungal C processing does not significantly alter the distribution of stable C isotopes and provide the basis for a reevaluation of ecosystem models based on isotopic evidence that involve C transfer across microbial interfaces. We provide a mechanism to account for the observed differential discrimination effects.     

Disentangling drought-induced variation in ecosystem and soil respiration using stable carbon isotopes

Combining C flux measurements with information on their isotopic composition can yield a process-based understanding of ecosystem C dynamics. We studied the variations in both respiratory fluxes and their stable C isotopic compositions (δ¹³C) for all major components (trees, understory, roots and soil microorganisms) in a Mediterranean oak savannah during a period with increasing drought. We found large drought-induced and diurnal dynamics in isotopic compositions of soil, root and foliage respiration (δ¹³C_res). Soil respiration was the largest contributor to ecosystem respiration (R _eco), exhibiting a depleted isotopic signature and no marked variations with increasing drought, similar to ecosystem respired δ¹³CO₂, providing evidence for a stable C-source and minor influence of recent photosynthate from plants. Short-term and diurnal variations in δ¹³C_res of foliage and roots (up to 8 and 4‰, respectively) were in agreement with: (1) recent hypotheses on post-photosynthetic fractionation processes, (2) substrate changes with decreasing assimilation rates in combination with increased respiratory demand, and (3) decreased phosphoenolpyruvate carboxylase activity in drying roots, while altered photosynthetic discrimination was not responsible for the observed changes in δ¹³C_res. We applied a flux-based and an isotopic flux-based mass balance, yielding good agreement at the soil scale, while the isotopic mass balance at the ecosystem scale was not conserved. This was mainly caused by uncertainties in Keeling plot intercepts at the ecosystem scale due to small CO₂ gradients and large differences in δ¹³C_res of the different component fluxes. Overall, stable isotopes provided valuable new insights into the drought-related variations of ecosystem C dynamics, encouraging future studies but also highlighting the need of improved methodology to disentangle short-term dynamics of isotopic composition of R _eco.     

The use of stable isotopes to study ecosystem gas exchange

Stable isotopes are a powerful research tool in environmental sciences and their use in ecosystem research is increasing. In this review we introduce and discuss the relevant details underlying the use of carbon and oxygen isotopic compositions in ecosystem gas exchange research. The current use and potential developments of stable isotope measurements together with concentration and flux measurements of CO₂ and water vapor are emphasized. For these applications it is critical to know the isotopic identity of specific ecosystem components such as the isotopic composition of CO₂, organic matter, liquid water, and water vapor, as well as the associated isotopic fractionations, in the soil-plant- atmosphere system. Combining stable isotopes and concentration measurements is very effective through the use of ”Keeling plots.” This approach allows the identification of the isotopic composition and the contribution of ecosystem, or ecosystem components, to the exchange fluxes with the atmosphere. It also allows the estimation of net ecosystem discrimination and soil disequilibrium effects. Recent modifications of the Keeling plot approach permit examination of CO₂ recycling in ecosystems. Combining stable isotopes with dynamic flux measurements requires precision in isotopic sampling and analysis, which is currently at the limit of detection. Combined with the micrometeorological gradient approach (applicable mostly in grasslands and crop fields), stable isotope measurements allow separation of net CO₂ exchange into photosynthetic and soil respiration components, and the evapotranspiration flux into soil evaporation and leaf transpiration. Similar applications in conjunction with eddy correlation techniques (applicable to forests, in addition to grasslands and crop fields) are more demanding, but can potentially be applied in combination with the Keeling plot relationship. The advance and potential in using stable isotope measurements should make their use a standard component in the limited arsenal of ecosystem-scale research tools. Received: 8 July 1999 / Accepted: 10 January 2000     

Stable isotopes of carbon and nitrogen in soil ecological studies

The development of stable isotope techniques is one of the main methodological advances in ecology of the last decades of the 20th century. Many biogeochemical processes are accompanied by changes in the ratio between stable isotopes of carbon and nitrogen (¹²C/¹³C and ¹⁴N/¹⁵N), which allows different ecosystem components and different ecosystems to be distinguished by their isotopic composition. Analysis of isotopic composition makes it possible to trace matter and energy flows through biological systems and to evaluate the rate of many ecological processes. The main concepts and methods of stable isotope ecology and patterns of stable isotope fractionation during organic matter decomposition are considered with special emphasis on the fractionation of isotopes in food chains and the use of stable isotope studies of trophic relationships between soil animals in the field.     

Ecophysiology of 13C and 15N isotopic fractionation in forest fungi and the roots of the saprotrophic-mycorrhizal divide

To quantify and characterize N and C isotopic fractionation effects due to fungal transformation of organic substrates in forest ecosystems, we performed a field study in California and a meta-analysis of three additional studies conducted by others across the Northern Hemisphere. Basidiomycete fungal biomass was consistently enriched for the heavier isotope for C relative to substrate and either enriched or depleted for N relative to atmospheric N. Extent and pattern of fractionation was very variable, but the distinction between ectomycorrhizal and saprotrophic basidiomycetes was strongly supported, particularly when dual isotope analyses were performed. This differentiation, which we call the "EM-SAP Divide" holds for studies within a restricted ecosystem, but becomes less distinct over larger geographical regions, removing the rationale for using direct isotopic values from single specimens as diagnostic of ecophysiological role. For C, the EM-SAP Divide seems to reflect substrate effects, potentially due to differential access to recently synthesized versus recycled organic compounds, rather than distinct physiological pathways. Once substrate and ecophysiological role effects are removed, our meta-analysis suggests the existence of more than one mechanism causing C fractionations in fungi which is found equally in ectomycorrhizal and saprotrophic fungi. Similarly, a multimodal distribution of '¹⁵N values suggests that physiological effects may play a much stronger influence on N natural isotopic distributions in fungi. Our meta-analysis provides a firm statistical base to evaluate fungal ecological statements based on natural isotopic distributions of C and N. We call into question the current practice of using direct isotopic measurements to make statements about trophic relationships of fungi in the absence of other supporting evidence.     

The contribution of C<Subscript>3</Subscript> and C<Subscript>4</Subscript> plants to the carbon cycle of a tallgrass prairie: an isotopic approach

The photosynthetic pathway composition (C₃:C₄ mixture) of an ecosystem is an important controller of carbon exchanges and surface energy flux partitioning, and therefore represents a fundamental ecophysiological distinction. To assess photosynthetic mixtures at a tallgrass prairie pasture in Oklahoma, we collected nighttime above-canopy air samples along concentration and isotopic gradients throughout the 1999 and 2000 growing seasons. We analyzed these samples for their CO₂ concentration and carbon isotopic composition and calculated C₃:C₄ proportions with a two-source mixing model. In 1999, the C₄ percentage increased from 38% in spring (late April) to 86% in early fall (mid-September). The C₄ percentages inferred from ecosystem respiration measurements in 2000 indicate a smaller shift, from 67% in spring (early May) to 77% in mid-summer (late July). We also sampled daytime CO₂ concentration and carbon isotope gradients above the canopy to determine ecosystem discrimination against ¹³CO₂ during net uptake. These discrimination values were always lower than corresponding nighttime ecosystem respiration isotopic signatures would suggest. After accounting for the isotopic disequilibria between respiration and photosynthesis resulting from seasonal variations in the C₃:C₄ mixture, we estimated canopy photosynthetic discrimination. The C₄ percentage calculated from this approach agrees with the percentage determined from nighttime respiration for sampling periods in both growing seasons. Isotopic imbalances between photosynthesis and respiration are likely to be common in mixed C₃:C₄ ecosystems and must be considered when using daytime isotopic measurements to constrain ecosystem physiology. Given the global extent of such ecosystems, isotopic imbalances likely contribute to global variations in the carbon isotopic composition of atmospheric CO₂.     

Use of stable carbon and nitrogen isotopes in insect trophic ecology

Insects are the most diverse organisms and often the most abundant animals in some ecosystems. Despite the importance of their functional roles and of the knowledge for conservation, the trophic ecology of many insect species is not fully understood. In this review, I present how stable carbon (C) and nitrogen (N) isotopes have been used to reveal the trophic ecology of insects over the last 30 years. The isotopic studies on insects have used differences in C isotope ratios between C₃ and C₄ plants, along vertical profiles of temperate and tropical forest stands, and between terrestrial and aquatic resources. These differences enable exploration of the relative importance of the food resources, as well as movement and dispersal of insects across habitats. The ¹³C‐enrichment (approximately 3‰) caused by saprotrophic fungi can allow the estimation of the importance of fungi in insect diets. Stable N isotopes have revealed food resource partitioning across diverse insect species above and belowground. Detritivorous insects often show a large trophic enrichment in ¹³C (up to 3‰) and ¹⁵N (up to 10‰) relative to the food substrates, soil organic matter. These values are greater than those commonly used for estimation of trophic level. This enrichment likely reflects the prevalence of soil microbial processes, such as fungal development and humification, influencing the isotopic signatures of diets in detritivores. Isotope analysis can become an essential tool in the exploration of insect trophic ecology in terms of biogeochemical C and N cycles, including trophic interactions, plant physiological and soil microbial processes.     

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.     

Interpretation of nitrogen isotope signatures using the NIFTE model

Nitrogen cycling in forest soils has been intensively studied for many years because nitrogen is often the limiting nutrient for forest growth. Complex interactions between soil, microbes, and plants and the consequent inability to correlate δ¹⁵N changes with biologic processes have limited the use of natural abundances of nitrogen isotopes to study nitrogen (N) dynamics. During an investigation of N dynamics along the 250-year-old successional sequence in Glacier Bay, Alaska, United States, we observed several puzzling isotopic patterns, including a consistent decline in δ¹⁵N of the late successional dominant Picea at older sites, a lack of agreement between mineral N δ¹⁵N and foliar δ¹⁵N, and high isotopic signatures for mycorrhizal fungi. In order to understand the mechanisms creating these patterns, we developed a model of N dynamics and N isotopes (Nitrogen Isotope Fluxes in Terrestrial Ecosystems, NIFTE), which simulated the major transformations of the N cycle and predicted isotopic signatures of different plant species and soil pools. Comparisons with field data from five sites along the successional sequence indicated that NIFTE can duplicate observed patterns in δ¹⁵N of soil, foliage, and mineral N over time. Different scenarios that could account for the observed isotopic patterns were tested in model simulations. Possible mechanisms included increased isotopic fractionation on mineralization, fractionation during the transfer of nitrogen from mycorrhizal fungi to plants, variable fractionation on uptake by mycorrhizal fungi compared to plants, no fractionation on mycorrhizal transfer, and elimination of mycorrhizal fungi as a pool in the model. The model results suggest that fractionation during mineralization must be small (˜2‰), and that no fractionation occurs during plant or mycorrhizal uptake. A net fractionation during mycorrhizal transfer of nitrogen to vegetation provided the best fit to isotopic data on mineral N, plants, soils, and mycorrhizal fungi. The model and field results indicate that the importance of mycorrhizal fungi to N uptake is probably less under conditions of high N availability. Use of this model should encourage a more rigorous assessment of isotopic signatures in ecosystem studies and provide insights into the biologic transformations which affect those signatures. This should lead to an enhanced understanding of some of the fundamental controls on nitrogen dynamics. Received: 1 July 1998 / Accepted: 23 December 1998     

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.     

Isotopic Fractionation of Hydrogen in Plants

The naturally-occurring stable isotopes deuterium and hydrogen are fractionated by a number of physical and biological processes. Deuterium has a tendency to precipitate out first from a moist air mass. Thus ground water will become isotopically lighter with an increase in latitude, altitude, or distance inland. Water taken up by the plant from the soil undergoes little change until evapotranspiration results in leaf water becoming isotopically heavier. Thus hydrogen isotopes in plants can reveal something of geography (groundwater) and climate. Hydrogen isotopes undergo little fractionation by passage through the food chain, although plant parasites tend to be enriched in D as compared to their hosts, possibly due to higher rates of transpiration in the parasitic plants. The splitting of water in photosynthesis results in the lighter isotope being incorporated into organic matter. An even larger isotopic fractionation results during lipid synthesis and other processes involving the pyruvate dehydrogenase complex. Differences in metabolic pathway between species can be detected by D/H ratios. Hydrogen isotopic differences can be detected between CAM, C₄, and C₃ species. Within C₄ plants, the NADP-ME plants are isotopically distinguishable from NAD-ME and PEP-CK plants.     

稳定性同位素技术与Keeling曲线法在陆地生态系统碳/水交换研究中的应用

稳定性同位素技术和Keeling曲线法是现代生态学研究的重要手段和方法之一。稳定性同位素能够整合生态系统复杂的生物学、生态学和生物地球化学过程在时间和空间尺度上对环境变化的响应。Keeling曲线法是以生物过程前后物质平衡理论为基础,将CO₂或H₂O的同位素组成(δD、δ¹³C或δ¹⁸O)与其对应浓度测量结合起来,将生态系统净碳通量区分为光合固定和呼吸释放通量,或将整个生态系统水分蒸散区分为植物蒸腾和土壤蒸发。在全球尺度上,稳定性同位素技术、Keeling曲线法与全球尺度陆地生态系统模型相结合,还可区分陆地生态系统和海洋生态系统对全球碳通量的贡献以及不同植被类型(C₃或C₄)在全球CO₂同化量中所占的比例。然而,生态系统的异质性使得稳定性同位素技术和Keeling曲线法从冠层尺度外推到生态系统、区域或全球尺度时存在有一定程度的不确定性。此外,取样时间、地点的选取也会影响最终的研究结果。尽管如此,随着分析手段的不断精确和研究方法的日趋完善,稳定性同位素技术和Keeling曲线法与其它测量方法(如微气象法)的有机结合将成为未来陆地生态系统碳/水交换研究的重要手段和方法之一。     

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     

Fractionation and turnover of stable carbon isotopes in animal tissues: Implications for δ13C analysis of diet

The use of stable carbon isotopes as a means of studying energy flow is increasing in ecology and paleoecology. However, secondary fractionation and turnover of stable isotopes in animals are poorly understood processes. This study shows that tissues of the gerbil (Meriones unguienlatus) have different δ¹³C values when equilibrated on corn (C₄) or wheat (C₃) diets with constant ¹³C/¹²C contents. Lipids were depleted 3.0‰ and hair was enriched 1.0‰ relative to the C₄ diet. Tissue δ¹³C values were ranked hair>brain>muscle>liver>fat. After changing the gerbils to a wheat (C₃) diet, isotope ratios of the tissues shifted in the direction of the δ¹³C value of the new diet. The rate at which carbon derived from the corn diet was replaced by carbon derived from the wheat diet was adequately described by a negative exponential decay model for all tissues examined. More metabolically active tissues such as liver and fat had more rapid turnover rates than less metabolically active tissues such as hair. The half-life for carbon ranged from 6.4 days in liver to 47.5 days in hair. The results of this study have important implications for the use of δ¹³C values as indicators of animal diet. Both fractionation and turnover of stable carbon isotopes in animal tissues may obscure the relative contributions of isotopically distinct dietary components (such as C₃ vs. C₄, or marine vs. terrestrial) if an animal's diet varies through time. These complications deserve attention in any study using stable isotope ratios of animal tissue as dietary indicators and might be minimized by analysis of several tissues or products covering a range of turnover times.     

Trophic level delineation and resource partitioning in a South African afromontane forest bird community using carbon and nitrogen stable isotopes

Southern African forests are naturally fragmented yet hold a disproportionately high number of bird species. Carbon and nitrogen stable isotopes were measured in feathers from birds captured at Woodbush (n = 27 species), a large afromontane forest in the eastern escarpment of Limpopo province, South Africa. The δ¹³C signatures of a range of forest plants were measured to categorise the food base. Most plants sampled, including two of five grass species, had δ¹³C signatures typical of a C₃ photosynthetic pathway (?29.5 ± 1.9‰). Three grass species had a C₄ signature (?12.0 ± 0.6‰). Most bird species had δ¹³C values representing a predominantly C₃‐based diet (?24.8‰ to ?20.7‰). δ¹⁵N values were as expected, with higher levels of enrichment associated with a greater proportion of dietary animal matter. The cohesive isotopic niche defining most species (n = 22), where the ranges for δ¹³C and δ¹⁵N were 2.4‰ and 3.4‰, respectively, highlight the difficulties in understanding diets of birds in a predominantly C₃‐based ecosystem using carbon and nitrogen stable isotopes. However, variation in isotopic values between and within species provides insight into possible niche width and the use of resources by different birds within a forest environment.     

13C and 15N in microarthropods reveal little response of Douglas-fir ecosystems to climate change

Understanding ecosystem carbon (C) and nitrogen (N) cycling under global change requires experiments maintaining natural interactions among soil structure, soil communities, nutrient availability, and plant growth. In model Douglas-fir ecosystems maintained for five growing seasons, elevated temperature and carbon dioxide (CO₂) increased photosynthesis and increased C storage belowground but not aboveground. We hypothesized that interactions between N cycling and C fluxes through two main groups of microbes, mycorrhizal fungi (symbiotic with plants) and saprotrophic fungi (free-living), mediated ecosystem C storage. To quantify proportions of mycorrhizal and saprotrophic fungi, we measured stable isotopes in fungivorous microarthropods that efficiently censused the fungal community. Fungivorous microarthropods consumed on average 35% mycorrhizal fungi and 65% saprotrophic fungi. Elevated temperature decreased C flux through mycorrhizal fungi by 7%, whereas elevated CO₂ increased it by 4%. The dietary proportion of mycorrhizal fungi correlated across treatments with total plant biomass (n= 4, r²= 0.96, P= 0.021), but not with root biomass. This suggests that belowground allocation increased with increasing plant biomass, but that mycorrhizal fungi were stronger sinks for recent photosynthate than roots. Low N content of needles (0.8–1.1%) and A horizon soil (0.11%) coupled with high C : N ratios of A horizon soil (25–26) and litter (36–48) indicated severe N limitation. Elevated temperature treatments increased the saprotrophic decomposition of litter and lowered litter C : N ratios. Because of low N availability of this litter, its decomposition presumably increased N immobilization belowground, thereby restricting soil N availability for both mycorrhizal fungi and plant growth. Although increased photosynthesis with elevated CO₂ increased allocation of C to ectomycorrhizal fungi, it did not benefit plant N status. Most N for plants and soil storage was derived from litter decomposition. N sequestration by mycorrhizal fungi and limited N release during litter decomposition by saprotrophic fungi restricted N supply to plants, thereby constraining plant growth response to the different treatments.     

C4-derived soil organic carbon decomposes faster than its C3 counterpart in mixed C3/C4 soils

The large difference in the degree of discrimination of stable carbon isotopes between C3 and C4 plants is widely exploited in global change and carbon cycle research, often with the assumption that carbon retains the carbon isotopic signature of its photosynthetic pathway during later stages of decomposition in soil and sediments. We applied long-term incubation experiments and natural ¹³C-labelling of C3 and C4-derived soil organic carbon (SOC) collected from across major environmental gradients in Australia to elucidate a significant difference in the rate of decomposition of C3- and C4-derived SOC. We find that the active pool of SOC (ASOC) derived from C4 plants decomposes at over twice the rate of the total pool of ASOC. As a result, the proportion of C4 photosynthesis represented in the heterotrophic CO₂ flux from soil must be over twice the proportional representation of C4-derived biomass in SOC. This observation has significant implications for much carbon cycle research that exploits the carbon isotopic difference in these two photosynthetic pathways.     

Carbon isotope fractionation in plants

Plants with the C₃, C₄, and crassulacean acid metabolism (CAM) photosynthetic pathways show characteristically different discriminations against ¹³C during photosynthesis. For each photosynthetic type, no more than slight variations are observed within or among species. CAM plants show large variations in isotope fractionation with temperature, but other plants do not. Different plant organs, subcellular fractions and metabolises can show widely varying isotopic compositions. The isotopic composition of respired carbon is often different from that of plant carbon, but it is not currently possible to describe this effect in detail. The principal components which will affect the overall isotope discrimination during photosynthesis are diffusion of CO₂, interconversion of CO₂ and HCO^?₃, incorporation of CO₂ by phosphoenolpyruvate carboxylase or ribulose bisphosphate carboxylase, and respiration. Theisotope fractionations associated with these processes are summarized. Mathematical models are presented which permit prediction of the overall isotope discrimination in terms of these components. These models also permit a correlation of isotope fractionations with internal CO₂ concentrations. Analysis of existing data in terms of these models reveals that CO₂ incorporation in C₃ plants is limited principally by ribulose bisphosphate carboxylase, but CO₂ diffusion also contributes. In C₄ plants, carbon fixation is principally limited by the rate of CO₂ diffusion into the leaf. There is probably a small fractionation in C₄ plants due to ribulose bisphosphate carboxylase.     

Natural abundance of 13C and 15N in C3 and C4 vegetation of southern Africa: patterns and implications

The mean annual rainfall in southern Africa is found to explain over half of the observed variance in the stable nitrogen (N) isotopic signatures of C₃ vegetation in southern Africa (r²=0.54, P<0.01). The inverse relationship between the stable N isotopic signatures of foliar samples from C₃ vegetation and long‐term southern African rainfall is found on a scale larger than previously observed. A modest relationship is found between stable carbon (C) isotopic signatures of C₃ vegetation and rainfall across the region (r²=0.20, P<0.01). No such relationship is found between stable C and N isotopic signatures of C₄ vegetation and rainfall. The explanation of the relationship between ¹⁵N in C₃ vegetation and the mean annual rainfall presented here is that nutrient availability varies inversely with water availability. This suggests that water‐limited systems in southern Africa are more open in terms of nutrient cycling and therefore the resulting natural abundance of foliar ¹⁵N in these systems is enriched. The use of this relationship may be of value to those researchers modeling both the dynamics of vegetation and biogeochemistry across this region. The use of the isotopic enrichment in C₃ vegetation as a function of rainfall may provide an insight into nutrient cycling across the semi‐arid and arid regions of southern Africa. This finding has implications for the study of global change, especially as it relates to vegetation responses to changing regional rainfall regimes over time.     

Factors controlling the stable isotope composition and C:N ratio of seston and periphyton in shallow lake mesocosms with contrasting nutrient loadings and temperatures

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