Research article: small-scale spatial variation of δ13C and δ15N isotopes in Antarctic carbon sources and consumers

Regional food web studies that fail to account for small-scale isotopic variability can lead to a mismatch between an organism’s inferred and true trophic position. Misinterpretation of trophic status may result, substantially limiting spatial and temporal comparability of food web studies. We sampled several carbon sources and consumers in a nested design to assess the variability of food web members across small spatial scales (100 s of m to several km) in regions around the Windmill Islands and Vestfold Hills in East Antarctica. For carbon sources, δ¹³C in sea ice POM was particularly variable between locations (km apart) and between sites (100 s of m apart) with replicate samples varying by up to 16‰. Macroalgae δ¹³C was less variable (replicate samples ranging up to 6.9‰ for the red alga Iridaea cordata), yet still differed between locations. Sediment POM and pelagic POM were the least variable, displaying minimal differences between locations or sites for δ¹³C and δ¹⁵N. Three out of eight consumers were significantly different between locations for δ¹³C, and five out of eight for δ¹⁵N, with the fish Trematomus bernacchii the most variable for both δ¹³C and δ¹⁵N. At smaller scales, the amphipod Paramorea walkeri showed significant variation between sites in δ¹³C but not in δ¹⁵N. We attribute small-scale variability to the dynamic physical environment for carbon sources in coastal systems and a close coupling of diet to habitat for consumers. We highlight the need to account for small-scale spatial variation in sampling designs for regional food web studies.     

Diel variation in lacunal CH4 and CO2 concentration and δ13C in Phragmites australis

We tested the hypothesis that the diurnal patterns of variationin lacunal gas concentrations and isotopic fractionationpreviously reported in a single plant genera (Typha)typified the patterns of all through-flow convective plantsby extending our observations to Phragmites australisCav. In daylight, Phragmites CH₄ transport isdriven by internal pressurization which results in gas flowdown young green culms and its exit from one year old deadbrown culms. Flow rates of 10.4 ± 4.0 mL min^–1 weremeasured in this study. At night, CH₄ is transportedfrom the sediments to the atmosphere via the lacunal plantspaces by molecular diffusion. Within green culms, lacunalCH₄ concentrations varied by a factor of 1000, from 3%(parts by volume) pre-dawn to lows of 25 ppmv during midday.Methane in brown culms varied by a factor of 10 diurnally,from 5% pre-dawn to 0.3% at midday. Lacunal CO₂concentrations varied similarly.Concentrations of both gases varied inversely with lacunalpressure. In green culms, large isotopic fractionations wereobserved in CH₄ and CO₂ in the morning and eveningduring transitions in gas transport mode and were associatedwith slight downward flows counter to the upward diffusionof these gases. Methane ¹³C as depletedas –100 was observed. In daylight, lacunal CH₄ wassimilar to or ¹³C depleted relative to sedimentary andemitted CH₄ isotopic values, but at night lacunalCH₄ was ¹³C enriched relative to sedimentarymethane. Overall, the diurnal variations of CH₄concentration and ¹³C value inPhragmiteswere similar to those observed in Typha andindicate that these patterns should be consistent in otherconvective-flow plants. Furthermore, our results demonstratethat the large isotopic fractionations found in aquaticplants can result solely from isotopic fractionationassociated with gas transport.     

Foliar δ13C within a temperate deciduous forest: spatial,temporal, and species sources of variation

Summary Foliar ¹³C-abundance (¹³C) was analyzed in the dominant trees of a temperate deciduous forest in east Tennessee (Walker Branch Watershed) to investigate the variation in foliar ¹³C as a function of time (within-year and between years), space (canopy height, watershed topography and habitat) and species (deciduous and coniferous taxa). Various hypotheses were tested by analyzing (i) samples collected from the field during the growing season and (ii) foliar tissues maintained in an archived collection. The ¹³C-value for leaves from the tops of trees was 2 to 3%. more positive than for leaves sampled at lower heights in the canopy. Quercus prinus leaves sampled just prior to autumn leaf fall had significantly more negative ¹³C-values than those sampled during midsummer. On the more xeric ridges, needles of Pinus spp. had more positive ¹³C-values than leaves from deciduous species. Foliar ¹³C-values differed significantly as a function of topography. Deciduous leaves from xeric sites (ridges and slopes) had more positive ¹³C-values than those from mesic (riparian and cove) environments. On the more xeric sites, foliar ¹³C was significantly more positive in 1988 (a dry year) relative to that in 1989 (a year with above-normal precipitation). In contrast, leaf ¹³C in trees from mesic valley bottoms did not differ significantly among years with disparate precipitation. Patterns in foliar ¹³C indicated a higher ratio of net CO₂ assimilation to transpiration (A/E) for trees in more xeric versus mesic habitats, and for trees in xeric habitats during years of drought versus years of normal precipitation. However, A/E (units of mmol CO₂ fixed/mol H₂O transpired) calculated on the basis of ¹³C-values for leaves from the more xeric sites was higher in a wet year (6.6±1.2) versus a dry year (3.4±0.4). This difference was attributed to higher transpiration (and therefore lower A/E) in the year with lower relative humidity and higher average daily temperature. The calculated A/E values for the forest in 1988–89, based on ¹³C, were within ±55% of estimates made over a 17 day period at this site in 1984 using micrometeorological methods.     

Does the 13C of foliage‐respired CO2 and biochemical pools reflect the 13C of recently assimilated carbon?

Isotopic labelling experiments were conducted to assess relationships among ¹³C of recently assimilated carbon ( δC _A), foliage respiration ( δC _F), soluble carbohydrate ( δC _SC), leaf waxes ( δC _LW) and bulk organic matter ( δC _OM). Slash pine, sweetgum and maize were grown under ¹³C depleted CO₂ to label biomass and then placed under ambient conditions to monitor the loss of label. In pine and sweetgum, δC _F of labelled plants (∼−44 and −35‰, respectively) rapidly approached control values but remained depleted by ∼4–6‰ after 3–4 months. For these tree species, no or minimal label was lost from δC _SC, δC _LW and δC _OM during the observation periods. δC _F and δC _SC of labelled maize plants rapidly changed and were indistinguishable from controls after 1 month, while δC _LW and δC _OM more slowly approached control values and remained depleted by 2–6‰. Changes in δC _F in slash pine and sweetgum fit a two-pool exponential model, with the fast turnover metabolic pool (∼3–4 d half-life) constituting only 1–2% of the total. In maize, change in δC _F fits a single pool model with a half-life of 6.4 d. The ¹³C of foliage respiration and biochemical pools reflect temporally integrated values of δC _A, with change in isotopic composition dampened by the size of metabolic carbon reserves and turnover rates.     

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     

Responses of pigmentation and δ13C in Qilian juniper to cold and drought stresses under natural conditions in the Qilian Mountains,China

The seasonal changes of pigments and stable carbon isotope composition (δ¹³C values) were investigated in needles of Qilian juniper (Sabina przewalskii Kom.) from two typical sites, one drier and the other wetter, in the Qilian Mountains, China. The anthocyanins and rhodoxanthin content from both sites were much higher in winter than in summer. Plant needles contained more carotenoids and xanthophylls in winter at the wetter site, while no seasonal difference appeared at the drier site. However, lower chlorophyll content and higher proline and δ¹³C were observed at the drier site. Average tree height was shorter at the drier site trees than at the wetter site. Our results suggested that under natural conditions, pigmentation in S. przewalskii may serve to improve the energy balance of the photosynthetic apparatus under cold and drought stresses. Proline and δ¹³C could be used as drought indicators for S. przewalskii.     

Fate and dynamics of recently fixed C in pasture plant–soil system under field conditions

The flow of photosynthetically fixed C from plants to selected soil C pools was studied after ¹³CO₂ pulse labeling of pasture plants under field conditions, dynamics of root-derived C in soil was assessed and turnover times of the soil C pools were estimated. The transport of the fixed C from shoots to the roots and into the soil was very fast. During 27 h, net C belowground allocation reached more than 10% of the fixed C and most of the C was already found in soil. Soil microbial biomass (C_MIC) was the major sink of the fixed C within soil C pools (ca 40–70% of soil ¹³C depending on sampling time). Significant amounts of ¹³C were also found in other labile soil C pools connected with microbial activity, in soluble organic C and C associated with microbial biomass (hot-water extract from the soil residue after chloroform fumigation-extraction) and the ¹³C dynamics of all these pools followed that of the shoots. When the labelling (2 h) finished, the fixed ¹³C was exponentially lost from the plant–soil system. The loss had two phases; the first rapid phase corresponded to the immediate respiration of ¹³C during the first 24 h and the second slower loss was attributable to the turnover of ¹³C assimilated in C_MIC. The corresponding turnover times for C_MIC were 1.1 days and 3.4 days respectively. Such short turnover times are comparable to those measured by growth kinetics after the substrate amendment in other studies, which indicates that microbial growth in the rhizosphere is probably not limited by substrate availability. Our results further confirmed the main role of the soil microbial community in the transformation of recently fixed C, short turnover time of the easily degradable C in the rhizosphere, and its negligible contribution to more stable soil C storage.     

Effect of salinity and humidity on δ13C value of halophytes—Evidence for diffusional isotope fractionation determined by the ratio of intercellular/atmospheric partial pressure of CO2 under different environmental conditions

Summary Seedlings of two mangrove species, Avicennia marina and Aegiceras corniculatum, were grown in a range of salinities and humidities in controlled environment chambers, and Phaseolus vulgaris plants were grown in the glasshouse. The fractionation of carbon isotopes in the three species was correlated with the ratio of intercellular and ambient partial pressures of CO₂. The results are consistent with fractionation being due both to diffusion in air and to carboxylation in the leaf. It was concluded that the latter process discriminates against ¹³CO₂ relative to ¹²CO₂ by about 27.     

Regional variation in soil carbon and δ<Superscript>13</Superscript>C in forests and pastures of northeastern Costa Rica

Recent studies suggest that the direction and magnitude of changes in soil organic carbon (soil C) pools following forest-to-pasture conversion in the tropics are dependent upon initial soil conditions and local factors (e.g. pre-conversion soil C content, soil texture, vegetation productivity, and management practices). The goal of this study was to understand how landscape-scale variation in soil-forming factors influenced the response of soil C pools to forest clearing and pasture establishment in northeastern Costa Rica. We measured soil C and its stable isotopic composition in 24 paired pasture and reference forest sites distributed over large gradients of edaphic characteristics and slope throughout a 1400 km² region. We used the large difference in stable C isotopic signatures of C3 vegetation (rain forest) versus C4 vegetation (pasture grasses) as a tracer of soil C dynamics. Soil C pools to 30 cm depth ranged from 26% lower to 23% higher in pastures compared to paired forests. The presence of non-crystalline clays and percent slope explained between 27 and 37% of the variation in the direction and magnitude of the changes in soil C storage following pasture establishment. Stable carbon isotopes (¹³C) in the top soil (0–10 cm) showed a rapid incorporation of pasture-derived C following pasture establishment, but the vegetation in these pastures never became pure C4 communities. The amount of forest-derived soil C in pasture topsoils (0–10 cm) was negatively correlated to both pasture age and the concentrations of non-crystalline iron oxides. Together these results imply that site factors such as soil mineralogy are an important control over soil C storage and turnover in this region.     

Temporal records of δ13C and δ15N in North Pacific pinnipeds: inferences regarding environmental change and diet

Sea lion and seal populations in Alaskan waters underwent various degrees of decline during the latter half of the twentieth century and the cause(s) for the declines remain uncertain. The stable carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) isotope ratios in bone collagen from wild Steller sea lions (Eumetopias jubatus), northern fur seals (Callorhinus ursinus) and harbor seals (Phoca vitulina) from the Bering Sea and Gulf of Alaska were measured for the period 1951-1997 to test the hypothesis that a change in trophic level may have occurred during this interval and contributed to the population declines. A significant change in '¹⁵N in pinniped tissues over time would imply a marked change in trophic level. No significant change in bone collagen '¹⁵N was found for any of the three species during the past 47 years in either the Bering Sea or the Gulf of Alaska. However, the ¹⁵N in the Steller sea lion collagen was significantly higher than both northern fur seals and harbor seals. A significant decline in '¹³C (almost 2 ‰ over the 47 years) was evident in Steller sea lions, while a declining trend, though not significant, was evident in harbor seals and northern fur seals. Changes in foraging location, in combination with a trophic shift, may offer one possible explanation. Nevertheless, a decrease in '¹³C over time with no accompanying change in '¹⁵N suggests an environmental change affecting the base of the foodweb rather than a trophic level change due to prey switching. A decline in the seasonal primary production in the region, possibly resulting from decreased phytoplankton growth rates, would exhibit itself as a decline in '¹³C. Declining production could be an indication of a reduced carrying capacity in the North Pacific Ocean. Sufficient quantities of optimal prey species may have fallen below threshold sustaining densities for these pinnipeds, particularly for yearlings and subadults who have not yet developed adequate foraging skills.     

The diel imprint of leaf metabolism on the δ13 C signal of soil respiration under control and drought conditions

Recent (13) CO(2) canopy pulse chase labeling studies revealed that photosynthesis influences the carbon isotopic composition of soil respired CO(2) (δ(13) C(SR)) even on a diel timescale. However, the driving mechanisms underlying these short-term responses remain unclear, in particular under drought conditions. The gas exchange of CO(2) isotopes of canopy and soil was monitored in drought/nondrought-stressed beech (Fagus sylvatica) saplings after (13) CO(2) canopy pulse labeling. A combined canopy/soil chamber system with gas-tight separated soil and canopy compartments was coupled to a laser spectrometer measuring mixing ratios and isotopic composition of CO(2) in air at high temporal resolution. The measured δ(13) C(SR) signal was then explained and substantiated by a mechanistic carbon allocation model. Leaf metabolism had a strong imprint on diel cycles in control plants, as a result of an alternating substrate supply switching between sugar and transient starch. By contrast, diel cycles in drought-stressed plants were determined by the relative contributions of autotrophic and heterotrophic respiration throughout the day. Drought reduced the speed of the link between photosynthesis and soil respiration by a factor of c. 2.5, depending on the photosynthetic rate. Drought slows the coupling between photosynthesis and soil respiration and alters the underlying mechanism causing diel variations of δ(13) C(SR).     

Effects of Ontogeny on δ13C of Plant- and Soil-Respired CO2 and on Respiratory Carbon Fractionation in C3 Herbaceous Species

Knowledge gaps regarding potential ontogeny and plant species identity effects on carbon isotope fractionation might lead to misinterpretations of carbon isotope composition (δ¹³C) of respired CO₂, a widely-used integrator of environmental conditions. In monospecific mesocosms grown under controlled conditions, the δ¹³C of C pools and fluxes and leaf ecophysiological parameters of seven herbaceous species belonging to three functional groups (crops, forage grasses and legumes) were investigated at three ontogenetic stages of their vegetative cycle (young foliage, maximum growth rate, early senescence). Ontogeny-related changes in δ¹³C of leaf- and soil-respired CO₂ and ¹³C/¹²C fractionation in respiration (Δ_R) were species-dependent and up to 7‰, a magnitude similar to that commonly measured in response to environmental factors. At plant and soil levels, changes in δ¹³C of respired CO₂ and Δ_R with ontogeny were related to changes in plant physiological status, likely through ontogeny-driven changes in the C sink to source strength ratio in the aboveground plant compartment. Our data further showed that lower Δ_R values (i.e. respired CO₂ relatively less depleted in ¹³C) were observed with decreasing net assimilation. Our findings highlight the importance of accounting for ontogenetic stage and plant community composition in ecological studies using stable carbon isotopes.     

Origin of carbon in agricultural soil profiles deduced from depth gradients of C:N ratios,carbon fractions, δ13C and δ15N values

Plant and Soil - Agricultural soils in Germany store 2.54 Pg of organic carbon (C). However, information about how and when this C entered the soils is limited. This study illustrates how depth...     

Shrub (Prosopis velutina) encroachment in a semidesert grassland: spatial–temporal changes in soil organic carbon and nitrogen pools

Recent trends of increasing woody vegetation in arid and semiarid ecosystems may contribute substantially to the North American C sink. There is considerable uncertainty, however, in the extent to which woody encroachment alters dryland soil organic carbon (SOC) and total nitrogen (TN) pools. To date, studies assessing SOC and TN response to woody plant proliferation have not explicitly assessed the variability caused by shrub age or size and subcanopy spatial gradients. These factors were quantified via spatially intensive soil sampling around Prosopis velutina shrubs in a semidesert grassland, using shrub size as a proxy for age. We found that bulk density increased with distance from the bole (P < 0.005) and decreased with increasing shrub size (P= 0.056), while both SOC and TN increased with shrub size and decreased with distance from the bole (P < 0.001 for both). Significant (and predictable) spatial variation in bulk density suggests that use of generic values would generate unreliable estimates of SOC and TN mass, and subcanopy SOC pools could be overestimated by nearly 30% if intercanopy bulk density values were applied to subcanopy sites. Predictive models based on field-documented spatial patterns were used to generate integrated estimates of under-shrub SOC and TN pools, and these were compared with results obtained by typical area-weighting protocols based on point samples obtained next to the bole or at a specified distance from the bole. Values obtained using traditional area-weighting approaches generally overestimated SOC pools relative to those obtained using the spatially integrated approach, the discrepancy increasing with increasing shrub size and proximity of the point sample to the bole. These discrepancies were observed at the individual plant scale and for landscapes populated by various shrub size classes. Results suggest that sampling aimed at quantifying shrub encroachment impacts on SOC and TN pools will require area-weighting algorithms that simultaneously account for shrub size (age) and subcanopy spatial patterns.     

δ13C values of an herbivore and the ratio of C3 to C4 plant carbon in its diet

Summary The ¹³C values of whole body samples of the beetle Tribolium castaneum are closely correlated with the ¹³C values of the plant carbon in its diet. The correlation is always high for diets ranging from 100% C₄ to 100% C₃ plant material. The degree of correlation is independent of the growth rate of the animals.     

Temporal–spatial variation and source analysis of carbon and nitrogen in a tidal wetland of Luoyuan Bay

Carbon and nitrogen are important elements in biogeochemical studies of tidal wetlands. Three wetland zones in Luoyuan Bay in the Fujian province were chosen for this study; the Spartina alterniflora flat zone with Spartina alterniflora growing, the silt zone with no Spartina alterniflora growing and the Spartina alterniflora-silt flat zone – a transition zone between the two. The spatial and seasonal variations of total organic carbon (TOC), total nitrogen (TN), stable isotopes of organic material (δ¹³C, δ¹⁵N), C/N ratio, average particle size and sediment composition in surface and vertical sediments of different ecological zones were analyzed. Carbon and nitrogen accumulation and particle size effects in the different ecological zones were discussed and the indicators of δ¹³C and C/N ratios were also compared. TOC, TN, δ¹³C contents, C/N ratios, and average particle size varied within the ranges of 0.611–1.133%, 0.053–0.090%, ?22.60 to ?18.92‰, 12.3–15.7, and 6.4–8.7 μm, respectively. Sediments were mainly silt-sized. Besides δ¹⁵N values, the other parameters, such as TOC, TN, δ¹³C contents, C/N ratios, and average particle size showed an obvious zonal distribution in surface sediments. The distribution of TOC and TN contents reflected the distribution of Spartina alterniflora within the bay. The profile and seasonal variations of these parameters in different ecological zones indicated that variations in the Spartina alterniflora flat and transition zones were complex because of the effect of Spartina alterniflora. Vertical and seasonal variations were sampled in the silt flat area. The profile and seasonal variations of TOC, TN and δ¹³C were similar in the transition zone and the Spartina alterniflora flat zone. Seasonal concentrations of TOC, TN and δ¹³C decreased from autumn > spring > winter > summer. The seasonal variation of carbon and nitrogen in the sediments may be influenced by temperature, particle size, plankton and benthos. The particle size effect was significant in the surface sediments and profile sediments of the transition zone. However, other factors had a greater effect on the distributions of TOC and TN in the Spartina alterniflora flat and silt flat zones. C/N ratios in sediments of the Spartina alterniflora flat, transition zone and silt flat were close to or > 12, indicating that the organic material source was dominated by terrestrial inputs. However, δ¹³C values decreased from the Spartina alterniflora flat zone > transition zone > silt flat zone indicating that the organic material source was predominantly from marine inputs. Thus the indications from C/N ratios and δ¹³C were different. There was no clear relationship between C/N ratios and δ¹³C values and a better relationship between δ¹³C values and TOC concentrations suggested that δ¹³C values provided a better indication of the organic source. Limited amounts of organic material came from Spartina alterniflora. This study has provided basic data for researching biogeochemical processes of biogenic elements in tidal wetlands and vegetation restoration, and has also provided a reference for assessing and protecting the environment and ecological systems in wetlands.     

Estimation of growth and food consumption in juvenile Japanese flounder Paralichthys olivaceus using carbon stable isotope ratio δC under laboratory conditions

To estimate the accumulated food consumption and growth of juvenile Japanese flounder Paralichthys olivaceus, we investigated the relationships between individual food consumption and growth, and the change in the stable carbon isotope ratio (δ¹³C). Japanese flounder juveniles were individually reared and their diet was switched from one formulated feed EP1 (δ¹³C = − 19.47‰) to another EP3 (δ¹³C = − 17.21‰) and fed at different feeding regimes. After the switch, the δ¹³C content of the dorsal muscle was exponentially shifted to a different level in proportion to the feeding and growth rates. Therefore, measuring the carbon stable isotope ratio is a useful tool for estimating the food consumption and growth rate of juveniles. In addition, since the velocity of change and the asymptotic value of the carbon stable isotope ratio varied in muscle, caudal fin and liver tissue, different tissues can be used for different time scale estimations.     

Diversity,metabolic types and δ13C carbon isotope ratios in the grass flora of Namibia in relation to growth form,precipitation and habitat conditions

The grass flora of Namibia (374 species in 110 genera) shows surprisingly little variation in ¹³C values along a rainfall gradient (50–600 mm) and in different habitat conditions. However, there are significant differences in the ¹³C values between the metabolic types of the C4 photosynthetic pathway. NADP-ME-type C4 species exhibit the highest ¹³C values (–11.7 ) and occur mainly in regions with high rainfall. NAD-ME-type C4 species have significantly lower ¹³C values (–13.4 ) and dominate in the most arid part of the precipitation regime. PCK-type C4 species play an intermediate role (–12.5 ) and reach a maximum abundance in areas of intermediate precipitation. This pattern is also evident in genera containing species of different metabolic types. Within the same genus NAD species reach more negative ¹³C values than PCK species and ¹³C values decreased with rainfall. Also in Aristida, with NADP-ME-type photosynthesis, ¹³C values decreased from –11 in the inland region (600 mm precipitation) to –15 near the coast (150 mm precipitation), which is a change in discrimination which is otherwise associated by a change in metabolism. The exceptional C3 species Eragrostis walteri and Panicum heterostachyum are coastal species experiencing 50 mm precipitation only. Many of the rare species and monotypic genera grow in moist habitats rather than in the desert, and they are not different in their carbon isotope ratios from the more common flora. The role of species diversity with respect to habitat occupation and carbon metabolism is discussed.     

The intramolecular ¹³C-distribution in ethanol reveals the influence of the CO₂ -fixation pathway and environmental conditions on the site-specific ¹³C variation in glucose

Efforts to understand the cause of ¹²C versus ¹³C isotope fractionation in plants during photosynthesis and post‐photosynthetic metabolism are frustrated by the lack of data on the intramolecular ¹³C‐distribution in metabolites and its variation with environmental conditions. We have exploited isotopic carbon‐13 nuclear magnetic resonance (¹³C NMR) spectrometry to measure the positional isotope composition (δ¹³C_i, ‰) in ethanol samples from different origins: European wines, liquors and sugars from C₃, C₄ and crassulacean acid metabolism (CAM) plants. In C₃‐ethanol samples, the methylene group was always ¹³C‐enriched (～2‰) relative to the methyl group. In wines, this pattern was correlated with both air temperature and δ¹⁸O of wine water, indicating that water vapour deficit may be a critical defining factor. Furthermore, in C₄‐ethanol, the reverse relationship was observed (methylene‐C relatively ¹³C‐depleted), supporting the concept that photorespiration is the key metabolic process leading to the ¹³C distribution in C₃‐ethanol. By contrast, in CAM‐ethanol, the isotopic pattern was similar to but stronger than C₃‐ethanol, with a relative ¹³C‐enrichment in the methylene‐C of up to 13‰. Plausible causes of this ¹³C‐pattern are briefly discussed. As the intramolecular δ¹³C_i‐values in ethanol reflect that in source glucose, our data point out the crucial impact on the ratio of metabolic pathways sustaining glucose synthesis.