Interspecific variability of δ13C among trees in rainforests of French Guiana: functional groups and canopy integration

The interspecific variability of sunlit leaf carbon isotope composition (δ¹³C), an indicator of leaf intrinsic water-use efficiency (WUE, CO₂ assimilation rate/leaf conductance for water vapour), was investigated in canopy trees of three lowland rainforest stands in French Guiana, differing in floristic composition and in soil drainage characteristics, but subjected to similar climatic conditions. We sampled leaves with a rifle from 406 trees in total, representing 102 species. Eighteen species were common to the three stands. Mean species δ¹³C varied over a 6.0‰ range within each stand, corresponding to WUE varying over about a threefold range. Species occurring in at least two stands displayed remarkably stable δ¹³C values, suggesting a close genetic control of species δ¹³C. Marked differences in species δ¹³C values were found with respect to: (1) the leaf phenology pattern (average δ¹³C=–29.7‰ and –31.0‰ in deciduous-leaved and evergreen-leaved species, respectively), and (2) different types of shade tolerance defined by features reflecting the plasticity of growth dynamics with respect to contrasting light conditions. Heliophilic species exhibited more negative δ¹³C values (average δ¹³C=–30.5‰) (i.e. lower WUE) than hemitolerant species (–29.3‰). However, tolerant species (–31.4‰) displayed even more negative δ¹³C values than heliophilic ones. We could not provide a straightforward ecophysiological interpretation of this result. The negative relationship found between species δ¹³C and midday leaf water potential (Ψ_wm) suggests that low δ¹³C is associated with high whole tree leaf specific hydraulic conductance. Canopy carbon isotope discrimination (Δ _A ) calculated from the basal area-weighed integral of the species δ¹³C values was similar in the three stands (average Δ _A =23.1‰), despite differences in stand species composition and soil drainage type, reflecting the similar proportions of the three different shade-tolerance types among stands. Received: 30 November 1999 / Accepted: 23 March 2000     

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     

Spatial patterns of foliar stable carbon isotope compositions of C<Subscript>3</Subscript> plant species in the Loess Plateau of China

The spatial pattern of foliar stable carbon isotope compositions (δ¹³C) of dominant species and their relationships with environmental factors in seven sites, Yangling, Yongshou, Tongchuan, Fuxian, Ansai, Mizhi and Shenmu, standing from south to north in the Loess Plateau of China, was studied. The results showed that in the 121 C₃ plant samples collected from the Loess Plateau, the foliar δ¹³C value ranged from −22.66‰ to −30.70‰, averaging −27.04‰. The foliar δ¹³C value varied significantly (P<0.01) among the seven sites, and the average δ¹³C value increased by about 1.69‰ from Yangling in the south to Shenmu in the north as climatic drought increased. There was a significant difference in foliar δ¹³C value among three life-forms categorized from all the plant samples in the Loess Plateau (P<0.001). The trees (−26.74‰) and shrubs (−26.68‰) had similar mean δ¹³C values, both significantly (P<0.05) higher than the mean δ¹³C value of herbages (−27.69‰). It was shown that the trees and shrubs had higher WUEs and employed more conservative water-use patterns to survive drier habitats in the Loess Plateau. Of all the C₃ species in the Loess Plateau, the foliar δ¹³C values were significantly and negatively correlated with the mean annual rainfall (P<0.001) and mean annual temperature (P<0.05), while being significantly and positively correlated with the latitude (P<0.001) and the annual solar radiation (P<0.01). In general, the foliar δ¹³C values increased as the latitude and solar radiation increased and the rainfall and temperature decreased. The annual rainfall as the main influencing factor could explain 13.3% of the spatial variations in foliar δ¹³C value. A 100 mm increment in annual rainfall would result in a decrease by 0.88‰ in foliar δ¹³C values.     

Discrimination between12C and13C by marine plants

Summary The natural abundance¹³C/¹²C ratios (as δ¹³C) of organic matter of marine macroalgae from Fife and Angus (East Scotland) were measured for comparison with the species' ability to use CO₂ and HCO ₃ ^- for photosynthesis, as deduced from previously published pH-drift measurements. There was a clear difference in δ¹³C values for species able or unable to use HCO ₃ ^- . Six species of Chlorophyta, 12 species of Phaeophyta and 8 species of Rhodophyta that the pH-drift data suggested could use HCO ₃ ^- had δ¹³C values in the range -8.81‰ to -22.55‰. A further 6 species of Rhodophyta which the pH-drift data suggested could only use CO₂ had δ¹³C values in the range -29.90‰ to-34.51‰. One of these six species (Lomentaria articulata) is intertidal; the other five are subtidal and so have no access to atmospheric CO₂ to complicate the analysis. For these species, calculations based on the measured δ¹³C of the algae, the δ¹³C of CO₂ in seawater, and the known¹³C/¹²C discrimination of CO₂ diffusion and RUBISCO carboxylation suggest that only 15–21% of the limitation to photosynthesisin situ results from CO₂ diffusion from the bulk medium to the plastids; the remaining 79–85% is associated with carboxylation reactions (and, via feedback effects, down-stream processes). This analysis has been extended for one of these five species,Delesseria sanguinea, by incorporating data onin situ specific growth rates, respiratory rates measured in the laboratory, and applying Fick's law of diffusion to calculate a boundary layer thickness of 17–24 μm. This value is reasonable for aDelesseria sanguinea frondin situ. For HCO ₃ ^- -using marine macroalgae the range of δ¹³C values measured can be accommodated by a CO₂ efflux from algal cells which range from 0.306 of the gross HCO ₃ ^- influx forEnteromorpha intestinalis (δ¹³C=-8.81‰) in a rockpool to 0.787 forChondrus crispus (δ¹³C=-22.55‰). The relatively high computed CO₂ efflux for those HCO ₃ ^- -users with the more negative δ¹³C values implies a relatively high photon cost of C assimilation; the observed photon costs can be accommodated by assuming coupled, energy-independent inorganic carbon influx and efflux. The observed δ¹³C values are also interpreted in terms of water movement regimes and obtaining CO₂ from the atmosphere. Published δ¹³C values for freshwater macrophytes were compared with the ability of the species to use CO₂ and HCO ₃ ^- and again there was an apparent separation in δ¹³C values for these two groups. δ¹³C values obtained for marine macroalgae for which no pH-drift data are available permit predictions, as yet untested, as to whether they use predominantly CO₂ or HCO ₃ ^-     

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.     

Bat breath reveals metabolic substrate use in free-ranging vampires

We analysed the stable carbon isotope ratio in exhaled CO₂ (δ¹³C_breath) of free-ranging vampires to assess the type of metabolized substrate (endogenous or exogenous substrate) and its origin, i.e. whether the carbon atoms came from a C₄ food web (grass and cattle) or the C₃ food web in which they were captured (a rainforest remnant and its mammals). For an improved understanding of factors influencing the δ¹³C_breath of vampires, we conducted feeding experiments with captive animals. The mean δ¹³C_breath of starved bats was depleted in ¹³C in relation to the diet by 4.6‰ (n = 10). Once fed with blood, δ¹³C_breath levelled off within a short time approximately 2.2‰ above the stable carbon isotope signature of the diet. The median time required to exchange 50% of the carbon atoms in exhaled CO₂ with carbon atoms from the ingested blood was 18.6 min (mean 29.5 ± 19.0 min, n = 5). The average δ¹³C of wing membrane and fur in free-ranging vampire bats suggested that bats almost exclusively foraged for cattle blood during the past weeks. The δ¹³C_breath of the same bats averaged −19.1‰. Given that all free-ranging vampires were starving and that the δ¹³C of cattle was more in enriched in ¹³C by 5–6‰ than the δ¹³C_breath of vampires, we conclude that the vampire bats of our study metabolised fat that was predominantly built from carbon atoms originating from cattle blood. Since δ¹³C of wing membrane and fur integrates over weeks and months respectively and δ¹³C_breath over hours and days, we also conclude that vampire bats of the studied population consistently ignored rainforest mammals and chose cattle as their prey during and prior to our study.     

Isotopic enrichment in herbivorous insects: a comparative field-based study of variation

Researchers will be able to use stable isotope analysis to study community structure in an efficient way, without a need for extensive calibrations, if isotopic enrichment values are consistent, or if variation in enrichment values can be predicted. In this study, we generated an experimental data set of δ¹⁵N and δ¹³C enrichment means for 22 terrestrial herbivorous arthropods feeding on 18 different host plants. Mean enrichments observed across a single trophic transfer (plants to herbivores) were −0.53±0.26‰ for δ¹³C (range: −3.47‰ to 1.89‰) and 1.88±0.37‰ for δ¹⁵N (range: −0.20‰ to 6.59‰). The mean δ¹³C enrichment was significantly lower than that reported in recent literature surveys, whereas the mean δ¹⁵N enrichment was not significantly different. The experimental data set provided no support for recent hypotheses advanced to explain variation in enrichment values, including the proposed roles for consumer feeding mode, development type, and diet C:N ratio. A larger data set, formed by combining our experimental data with data from the literature, did suggest possible roles for feeding mode, nitrogen recycling, herbivore life stage, and host plant type. Our results indicate that species enrichment values are variable even in this relatively narrow defined group of organisms and that our ability to predict enrichment values of terrestrial herbivorous arthropods based on physiological, ecological, or taxonomic traits is low. The primary implications are that (1) mean enrichment may have to be measured empirically for each trophic link of interest, rather than relying on estimates from a broad survey of animal taxa and (2) the advantage of using stable isotope analysis to probe animal communities that are recalcitrant to other modes of study will be somewhat diminished as a consequence.Electronic Supplementary Material Supplementary material is available for this article at     

Spatial variability in Florida Bay particulate organic matter composition: combining flow cytometry with stable isotope analyses

Long-term management plans for restoration of natural flow conditions through the Everglades increase the importance of understanding potential nutrient impacts of increased freshwater delivery on Florida Bay biogeochemistry. Planktonic communities respond quickly to changes in water quality, thus spatial variability in community composition and relationships to nutrient parameters must be understood in order to evaluate future downstream impacts of modifications to Everglades hydrology. Here we present initial results combining flow cytometry analyses of phytoplankton and bacterial populations (0.1–50 μm size fraction) with measurements of δ¹³C and δ¹⁵N composition and dissolved inorganic nutrient concentrations to explore proxies for planktonic species assemblage compositions and nutrient cycling. Particulate organic material in the 0.1–50 μm size fraction was collected from five stations in Northeastern and Western Florida Bay to characterize spatial variability in species assemblage and stable isotopic composition. A dense bloom of the picocyanobacterium, Synechococcus elongatus, was observed at Western Florida Bay sites. Smaller Synechococcus sp. were present at Northeast sites in much lower abundance. Bacteria and detrital particles were also more abundant at Western Florida Bay stations than in the northeast region. The highest abundance of detritus occurred at Trout Creek, which receives freshwater discharge from the Everglades through Taylor Slough. In terms of nutrient availability and stable isotopic values, the S. elongatus population in the Western bay corresponded to low DIN (0.5 μM NH ₄ ⁺ ; 0.2 μM NO ₃ ⁻ ) concentrations and depleted δ¹⁵N signatures ranging from +0.3 to +0.8‰, suggesting that the bloom supported high productivity levels through N₂-fixation. δ¹⁵N values from the Northeast bay were more enriched (+2.0 to +3.0‰), characteristic of N-recycling. δ¹³C values were similar for all marine Florida Bay stations, ranging from −17.6 to −14.4‰, however were more depleted at the mangrove ecotone station (−25.5 to −22.3‰). The difference in the isotopic values reflects differences in carbon sources. These findings imply that variations in resource availability and nutrient sources exert significant control over planktonic community composition, which is reflected by stable isotopic signatures.     

Carbon isotopic fractionation in subtropical brazilian grassland soils. Comparison with tropical forest soils

The natural relationship¹³C/¹²C determined in three soil profiles under grass vegetation indicated a depletion in organic¹³C at depth: theδ ¹³C was between −18‰ and −15‰ in the A horizons and ranged from −18 to −22‰ at depth. Previous work showed that in forest soils, whereδ ¹³C was near −28‰ in the upper horizon, there was, on the contrary, a relative enrichment of the lower strata. This meant thatδ ¹³C, initially different in the various topsoils, became more equal at depth. Comparison between dark, deep horizons (sombric horizons), which are certainly of illuvial origine, would confirm this:δ ¹³C of grassland and a forest sombric horizon were almost equal at around −22‰. These results might mean that, in natural ecosystems, the isotopic carbon composition of the soil underlying humus would be independent of the vegetation type. This would have practical implications for the use of¹³C as a tracer for soil organic matter studies.     

Mechanism for the small-scale movement of carbon among estuarine habitats: organic matter transfer not crab movement

In theory, carbon is highly mobile in aquatic systems. Recent evidence from carbon stable isotopes of crabs (Parasesarma erythrodactyla and Australoplax tridentata), however, shows that in subtropical Australian waters, measurable carbon movement between adjacent mangrove and saltmarsh habitats is limited to no more than a few metres. We tested whether the pattern in crab δ¹³C values across mangrove and saltmarsh habitats was explained by crab movement, or the movement of particulate organic matter. We estimated crab movement in a mark–recapture program using an array of pitfall traps on 13 transects (a total of 65 traps) covering an area of 600 m² across the interface of these two habitats. Over a 19-day period, the majority of crabs (91% for P. erythrodactyla, 93% for A. tridentata) moved <2 m from the place of initial capture. Crab movement cannot, therefore, explain the patterns in δ¹³C values of crabs. δ¹³C values of detritus collected at 2-m intervals across the same habitat interface fitted a sigmoidal curve of a similar form to that fitting the δ¹³C values of crabs. δ¹³C values of detritus were 2–4‰ more depleted in saltmarsh (−18.5±0.6‰), and 4–7‰ more depleted in mangroves (−25.9±0.1‰) than δ¹³C values of crabs recorded previously in each habitat. Assimilation by crabs of very small detrital fragments or microphytobenthos, more enriched in ¹³C, may explain the disparity in δ¹³C values. Nevertheless, the pattern in δ¹³C values of detritus suggests that crabs obtain their carbon from up to several metres away, but without themselves foraging more then a metre or so from their burrow. Such detailed measurements of carbon movement in estuaries provide a spatially explicit understanding of the functioning of food webs in saltmarsh and mangrove habitats.     

Carbon isotope ratio and photosynthetic activity of phytoplankton in the eutrophic Mikawa Bay,Japan

Primary production by phytoplankton in the eutrophic Mikawa Bay, Japan, was studied by simultaneous measurements of natural carbon isotope ratio (δ ¹³C) and short-term carbon uptake rates (¹³C tracer study) of size-fractionated nannoplankton (<10 μm) and net plankton (>10 μm) samples. Short-term photosynthetic rates, which represent the physiological state of algae, were variable regardless of standing stock sizes. Theδ ¹³C values of particulate organic carbon (POC) in June and July displayed horizontal variations for both the net plankton fraction (−19.8 to −12.7‰) and the nannoplankton fraction (−22.0 to −12.8‰). For both fractions, low concentrations of POC had more negativeδ ¹³C values (−22 to −18‰). Highδ ¹³C values for the net plankton were found when POC concentrations were much higher, due to red tide. This suggests that the increase in algal standing crop for the net plankton fraction resulted from accelerated photosynthetic activity. However the nannoplankton fractions with higher POC values have relatively lowδ ¹³C values.     

Stable carbon isotope characteristics of desert plants in the Junggar Basin, China

Desert plants have unique strategies for survival and growth to cope with the limited water availability in arid regions. The stable carbon isotope (δ ¹³C) provides an integrated measurement of internal plant physiological and external environmental properties affecting photosynthetic gas exchange and water use efficiency. The δ ¹³C values of 84 species in the Junggar Basin were categorized into two groups (ranged from −30.1 to −23.3‰ for C₃ and −14.9 to −9.9‰ for C₄ species, respectively). No life form differences in δ ¹³C values were detected in C₃ (p = 0.78) and C₄ plants (p = 0.63). Small differences among life forms were observed in δ ¹³C values in C₄ species with shrubs slightly depleted (−13.3‰) relative to perennials (−13.1‰) and annuals (−12.5‰). These differences suggested that δ ¹³C value could not represent a plant functional group classification based on life forms in C₄ plants in extremely arid regions. Ephemerals are all using C₃ photosynthetic pathway and no significant differences (p = 0.92) in δ ¹³C values were observed between annuals (−26.5‰) and perennials (−26.4‰). The δ ¹³C values of Tulipa iliensis (an important ephemeral species distributed widely in the Junggar Basin) among nine natural populations were positively correlated with leaf (r ² = 0.46, p = 0.046) and soil (r ² = 0.67, p = 0.007) total nitrogen content, and negatively correlated with leaf (r ² = 0.48, p = 0.039) and soil (r ² = 0.79, p = 0.001) water content. This indicated that the variation in δ ¹³C values of T. iliensis was probably caused by both water availability associated stomatal openness and nitrogen availability associated photosynthetic capacity. T. iliensis is very sensitive to water and nitrogen availability in soil.     

Subantarctic Macquarie Island – a model ecosystem for studying animal-derived nitrogen sources using 15N natural abundance

Plants collected from diverse sites on subantarctic Macquarie Island varied by up to 30‰ in their leaf δ¹⁵N values. ¹⁵N natural abundance of plants, soils, animal excrement and atmospheric ammonia suggest that the majority of nitrogen utilised by plants growing in the vicinity of animal colonies or burrows is animal-derived. Plants growing near scavengers and animal higher in the food chain had highly enriched δ¹⁵N values (mean = 12.9‰), reflecting the highly enriched signature of these animals' excrement, while plants growing near nesting penguins and albatross, which have an intermediate food chain position, had less enriched δ¹⁵N values (>6‰). Vegetation in areas affected by rabbits had lower δ¹⁵N values (mean = 1.2‰), while the highly depleted δ¹⁵N values (below −5‰) of plants at upland plateau sites inland of penguin colonies, suggested that a portion of their nitrogen is derived from ammonia (mean ¹⁵N =−10‰) lost during the degradation of penguin guano. Vegetation in a remote area had δ¹⁵N values near −2‰. These results contrast with arctic and subarctic studies that attribute large variations in plant ¹⁵N values to nitrogen partitioning in nitrogen-limited environments. Here, plant ¹⁵N reflects the ¹⁵N of the likely nitrogen sources utilised by plants. Received: 18 December 1997 / Accepted: 13 June 1998     

Influence of stand structure on carbon-13 of vegetation, soils, and canopy air within deciduous and evergreen forests in Utah, United States

Carbon isotope ratios (δ¹³C) were studied in evergreen and deciduous forest ecosystems in semi-arid Utah (Pinus contorta, Populus tremuloides, Acer negundo and Acer grandidentatum). Measurements were taken in four to five stands of each forest ecosystem differing in overstory leaf area index (LAI) during two consecutive growing seasons. The δ¹³C_leaf (and carbon isotope discrimination) of understory vegetation in the evergreen stands (LAI 1.5–2.2) did not differ among canopies with increasing LAI, whereas understory in the deciduous stands (LAI 1.5–4.5) exhibited strongly decreasing δ¹³C_leaf values (increasing carbon isotope discrimination) with increasing LAI. The δ¹³C values of needles and leaves at the top of the canopy were relatively constant over the entire LAI range, indicating no change in intrinsic water-use efficiency with overstory LAI. In all canopies, δ¹³C_leaf decreased with decreasing height above the forest floor, primarily due to physiological changes affecting c _i/c _a (> 60%) and to a minor extent due to δ¹³C of canopy air (< 40%). This intra-canopy depletion of δ¹³C_leaf was lowest in the open stand (1‰) and greatest in the denser stands (4.5‰). Although overstory δ¹³C_leaf did not change with canopy LAI, δ¹³C of soil organic carbon increased with increasing LAI in Pinus contorta and Populus tremuloides ecosystems. In addition, δ¹³C of decomposing organic carbon became increasingly enriched over time (by 1.7–2.9‰) for all deciduous and evergreen dry temperate forests. The δ¹³C_canopy of CO₂ in canopy air varied temporally and spatially in all forest stands. Vertical canopy gradients of δ¹³C_canopy, and [CO₂]_canopy were larger in the deciduous Populus tremuloides than in the evergreen Pinu contorta stands of similar LAI. In a very wet and cool year, ecosystem discrimination (Δ_e) was similar for both deciduous Populus tremulodies (18.0 ± 0.7‰) and evergreen Pinus contorta (18.3 ± 0.9‰) stands. Gradients of δ¹³C_canopy and [CO₂]_canopy were larger in denser Acer spp. stands than those in the open stand. However, ¹³C enrichment above and photosynthetic draw-down of [CO₂]_canopy below tropospheric baseline values were larger in the open than in the dense stands, due to the presence of a vigorous understory vegetation. Seasonal patterns of the relationship δ¹³C_canopy versus 1/[CO₂]_canopy were strongly influenced by precipitation and air temperature during the growing season. Estimates of Δ_e for Acer spp. did not show a significant effect of stand structure, and averaged 16.8 ± 0.5‰ in 1933 and 17.4 ± 0.7‰ in 1994. However, Δ_e varied seasonally with small fluctuations for the open stand (2‰), but more pronounced changes for the dense stand (5‰). Received: 15 April 1996 / Accepted: 19 October 1996     

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     

Food web structure of a shallow eutrophic lake (Lake Taihu,China) assessed by stable isotope analysis

Lake Taihu is a large, shallow, and eutrophic lake in China. It has provided local communities with valuable fisheries for centuries, but little is known of the trophodynamics, or of its faunal communities. Carbon and nitrogen isotopic composition was used to assess its trophic pathways and the food web structure [food sources and trophic levels (TL)]. Basal food sources were distinguishable based on their δ¹³C values, ranging from −27.2 to −15.2‰. Consumers were also well separated in δ¹³C (−26.9 to −17.9‰ for invertebrates and −25.7 to −18.1‰ for fishes), which allowed for an effective discrimination of carbon sources between these fauna. An average trophic enrichment factor of 3.4‰ was used to calculate the TLs based on δ¹⁵N of zooplankton, with results indicating a food web having four TLs. Although δ¹⁵N values overlap and cover a large range within trophic compartments, the isotopic signatures of the species assessed revealed a general trend of ¹⁵N enrichment with increasing TL. Stable isotope signatures were also used to establish a general food web scheme in which five main trophic pathways were analyzed.     

Palaeozoic cold seep carbonates from Europe and North Africa—an integrated isotopic and geochemical approach

In this paper, we report the highest and lowest carbon isotope values known from Palaeozoic carbonate rocks. These unusual δ¹³C values (−50 to +23.5‰) are due to microbial methanogenesis and methanotrophy in Silurian to Carboniferous carbonates. Trace elements were used to decipher the primary mineralogy of the carbonate cements. Very high Sr values and low amounts of Mg, Fe and Mn point toward aragonite precursors, whereas high Fe and Mn values are indicative of primary calcites and allow reconstruction of the redox conditions. Four carbonate deposits are described from the Meseta and the Antiatlas of Morocco, the Pyrenees (France) and the Harz mountains (Germany). The highest δ¹³C values in concretion below the uppermost Silurian Spinatrypa Mound (Moroccan Meseta) give evidence, that CO₂ was produced during methanogenesis. δ¹³C values between −10 and −32‰indicate that the formation of microbial carbonates and cements in the Middle Devonian Hollard Mound (Antiatlas) and in the Lower Carboniferous sediments of the Iberg (Harz) formed at thermogenetic methane or petroleum seeps. The Late Bashkirian carbonate mound of the High Pyrenees (Tantes Mound) is the first Palaeozoic carbonate with seepage fluids being dominated by biogenic methane. Matrix carbonates exhibit δ¹³C values as low as −34‰. In some parts, voids make up more than 50 vol% of the mound. They are filled with several generations of cement. The earliest void filling is isopachous fibrous cement, which represents former aragonite. Most negative δ¹³C values of −50‰were measured in these isopachous fibrous cements. The difference of 55‰in δ¹³C values between normal sediments and early aragonite cements can only be explained by the contribution of CO₂ from anaerobic oxidation of biogenic methane in a cold seep setting.     

Leaf δ13C in Pinus resinosa trees and understory plants: variation associated with light and CO2 gradients

 Our objective was to evaluate the relative importance of gradients in light intensity and the isotopic composition of atmospheric CO₂ for variation in leaf carbon isotope ratios within a Pinus resinosa forest. In addition, we measured photosynthetic gas exchange and leaf carbon isotope ratios on four understory species (Dryopteris carthusiana, Epipactus helleborine, Hieracium floribundum, Rhamnus frangula), in order to estimate the consequence of the variation in the understory light microclimate for carbon gain in these plants. During midday, CO₂ concentration was relatively constant at vertical positions ranging from 15 m to 3 m above ground. Only at positions below 3 m was CO₂ concentration significantly elevated above that measured at 15 m. Based on the strong linear relationship between changes in CO₂ concentration and δ¹³C values for air samples collected during a diurnal cycle, we calculated the expected vertical profile for the carbon isotope ratio of atmospheric CO₂ within the forest. These calculations indicated that leaves at 3 m height and above were exposed to CO₂ of approximately the same isotopic composition during daylight periods. There was no significant difference between the daily mean δ¹³C values at 15 m (–7.77‰) and 3 m (–7.89‰), but atmospheric CO₂ was significantly depleted in ¹³C closer to the ground surface, with daily average δ¹³C values of –8.85‰ at 5 cm above ground. The light intensity gradient in the forest was substantial, with average photosynthetically active radiation (PAR) on the forest floor approximately 6% of that received at the top of the canopy. In contrast, there were only minor changes in air temperature, and so it is likely that the leaf-air vapour pressure difference was relatively constant from the top of the canopy to the forest floor. For red pine and elm tree samples, there was a significant correlation between leaf δ¹³C value and the height at which the leaf sample was collected. Leaf tissue sampled near the forest floor, on average, had lower δ¹³C values than samples collected near the top of the canopy. We suggest that the average light intensity gradient through the canopy was the major factor influencing vertical changes in tree leaf δ¹³C values. In addition, there was a wide range of variation (greater than 4‰) among the four understory plant species for average leaf δ¹³C values. Measurements of leaf gas exchange, under natural light conditions and with supplemental light, were used to estimate the influence of the light microclimate on the observed variation in leaf carbon isotope ratios in the understory plants. Our data suggest that one species, Epipactus helleborine, gained a substantial fraction of carbon during sunflecks. Received: 21 March 1996 / Accepted: 13 August 1996     

Sulfur,carbon, and nitrogen stable isotope ratios in soft tissues and trophic relationships of fish from the near-shore waters of the peter the great bay in the Sea of Japan

Stable isotope ratios of sulfur (³⁴S/³²S), carbon (¹³C/¹²C), and nitrogen (¹⁵N/¹⁴N) were analyzed in the soft tissues of 12 common species of fish from the near-shore waters of the Peter the Great Bay in the Sea of Japan. The average δ¹³C values of individual species varied from −20.7‰ for planktivorous fish to −16.8‰ for benthivorous fish, reflecting the growing relative contribution of benthic primary producers to fish nutrition. The majority of the various species representatives studied can be assigned to one trophic level, as indicated by their narrow range of δ¹⁵N values (9.9 to 12.6‰). Large interspecific variations were found in the sulfur stable isotope ratios of fish (the mean δ³⁴S values ranged from 11.2 to 19.5‰). This is the result of the different contributions to fish nutrition of infaunal invertebrates that are depleted in ³⁴S due to the microbial food chain of the bottom sediments.     

The δ13C changes in four plant species of the Loess Plateau over the last 70 years

The relative abundance of carbon isotope (δ¹³C) was measured in four C₃ species (Sophora viccifolia, Quercus liaotungensis, Ostryopsis davidiana and Zizyphus jujuba var. spinosa) of the Loess Plateau in China from the 1930’s to 2002. The results showed that the δ¹³C values in the four species varied from −25.05‰ to −29.75‰ with their a average at −27.04‰. A decrease in the δ¹³C value with time was found in all the four species, which indicating that the water use efficiencies (WUEs) of all the measured species declined during 70 years. However, the decrease in δ¹³C value differed among the four species with its significant decreases measured in two of the species, Sophora viciifolia and Quercus liaotungensis, its relatively significant decrease found in Ostryopsis davidiana, and its slight decrease appearing in Zizyphus jujuba var. spinosa. in the δ¹³C values in the four species decreased by 14.65 ‰, 14.46‰, 11.99‰ and 2.44‰, respectively. The different species were shown to have different sensitivities to climatic change, and Zizyphus jujuba var. spinosa was found to be the most drought-tolerant species of the four, which had a high WUE.