Stable carbon and nitrogen incorporation in blood and fin tissue of the catfish <Emphasis Type="Italic">Pterygoplichthys disjunctivus</Emphasis> (Siluriformes,Loricariidae)

A feeding trial was performed in the laboratory with the catfish species Pterygoplichthys disjunctivus to determine stable carbon (¹³C) and nitrogen (¹⁵ N) turnover rates and discrimination factors in non-lethally sampled tissues (red blood cells, plasma solutes, and fin). A second feeding trial was conducted to determine what P. disjunctivus could assimilate from low-quality wood-detritus—refractory polysaccharides (e.g., cellulose), or soluble wood-degradation products inherent in wood-detritus. This was performed by feeding the fish an artificial wood-detritus diet with fibrous (δ¹³C = −26.36‰; δ¹⁵  N = 2.13‰) and soluble portions (δ¹³C = −11.82‰; δ¹⁵  N = 3.39‰) that had different isotopic signatures and monitoring the dynamics of isotopic incorporation in the different tissues over time. Plasma solutes turned over more quickly than red blood cells for ¹³C and ¹⁵ N. However, in contrast to previous studies of juvenile fishes, C and N incorporation was primarily driven by catabolic tissue turnover as opposed to growth rate. Tissue-diet discrimination factors for ¹⁵ N varied from 4.08 to 5.17‰, whereas they were <2‰ for ¹³C (and less than 0.3‰ for plasma and red blood cells). The results of trial two suggested that P. disjunctivus could not assimilate refractory polysaccharides. Moreover, the δ¹³C and δ¹⁵ N signatures of wild-caught P. disjunctivus from Florida confirmed their detrital trophic standing in Floridian aquatic ecosystems.     

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     

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

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

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.     

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.     

Carbon sources of Antarctic nematodes as revealed by natural carbon isotope ratios and a pulse-chase experiment

δ¹³C of nematode communities in 27 sites was analyzed, spanning a large depth range (from 130 to 2,021 m) in five Antarctic regions, and compared to isotopic signatures of sediment organic matter. Sediment organic matter δ¹³C ranged from −24.4 to −21.9‰ without significant differences between regions, substrate types or depths. Nematode δ¹³C showed a larger range, from −34.6 to −19.3‰, and was more depleted than sediment organic matter typically by 1‰ and by up to 3‰ in silty substrata. These, and the isotopically heavy meiofauna at some stations, suggest substantial selectivity of some meiofauna for specific components of the sedimenting plankton. However, ¹³C-depletion in lipids and a potential contribution of chemoautotrophic carbon in the diet of the abundant genus Sabatieria may confound this interpretation. Carbon sources for Antarctic nematodes were also explored by means of an experiment in which the fate of a fresh pulse of labile carbon to the benthos was followed. This organic carbon was remineralized at a rate (11–20 mg C m⁻² day⁻¹) comparable to mineralization rates in continental slope sediments. There was no lag between sedimentation and mineralization; uptake by nematodes, however, did show such a lag. Nematodes contributed negligibly to benthic carbon mineralization.     

Contributions of Different Organic Carbon Sources to <Emphasis Type="Italic">Daphnia</Emphasis> in the Pelagic Foodweb of a Small Polyhumic Lake: Results from Mesocosm DI<Superscript>13</Superscript>C-Additions

Abstract Freshwater ecosystems derive organic carbon from both allochthonous and autochthonous sources. We studied the relative contributions of different carbon sources to zooplankton in a small, polyhumic, steeply stratified lake, using six replicate surface-to-sediment enclosures established during summer and autumn 2004. We added ¹³C-enriched bicarbonate to the epilimnion of half the enclosures for three weeks during each season and monitored carbon stable isotope ratios of DIC, DOC, POC and Daphnia, along with physical, chemical and biological variables. During summer, ¹³C-enriched DIC (δ¹³C up to 44 ± 7.2‰) was soon taken up by phytoplankton (δ¹³C up to −5.1 ± 13.6‰) and was transmitted to Daphnia (δ¹³C up to −1.7 ± 7.2‰), demonstrating consumption of phytoplankton. In contrast, during autumn, ¹³C-enriched DIC (δ¹³C up to 56.3 ± 9.8‰) was not transmitted to Daphnia, whose δ¹³C became progressively lower (δ¹³C down to −45.6 ± 3.3‰) concomitant with decreasing methane concentration. Outputs from a model suggested phytoplankton contributed 64–84% of Daphnia diet during summer, whereas a calculated pelagic carbon mass balance indicated only 30–40% could have come from phytoplankton. Although autumn primary production was negligible, zooplankton biomass persisted at the summer level. The model suggested methanotrophic bacteria contributed 64–87% of Daphnia diet during autumn, although the calculated carbon mass balance indicated a contribution of 37–112%. Thus methanotrophic bacteria could supply virtually all the carbon requirement of Daphnia during autumn in this lake. The strongly ¹³C-depleted Daphnia values, together with the outputs from the models and the calculated carbon mass balance showed that methanotrophic bacteria can be a greater carbon source for Daphnia in lakes than previously suspected.     

Variations in leaf β13C along a vertical profile of irradiance in a temperate Japanese forest

The vertical profile of stable carbon isotope ratios (δ¹³C) of leaves was analyzed for 13 tree species in a cool-temperate deciduous forest in Japan. The vertical distribution of long-term averaged δ¹³C in atmospheric CO₂ (δ_a) was estimated from δ¹³C of dry matter from NADP-malic enzyme type C₄ plant (Zea mays L. var. saccharata Sturt.) grown at a tower in the forest for 32␣days, assuming constant Δ value (3.3‰) in Z. mays against height. The δ_a value obtained from δ¹³C in Z.␣mays was lowest at the forest floor (−9.30 ± 0.03‰), increased with height, and was almost constant above 10␣m (−7.14 ± 0.14‰). Then leaf Δ values for the tree species were calculated from tree leaf δ¹³ C andδ_a. Mean leaf Δ values for the three tall deciduous species (Fraxinus mandshurica, Ulmus davidiana, and Alnus hirsuta) were significantly different among three height levels in the forest: 23.1 ± 0.7‰ at the forest floor (understory), 21.4 ± 0.5‰ in lower canopy, and 20.5 ± 0.3‰ in upper canopy. The true difference in tree leaf Δ among the forest height levels might be even greater, because Δ in Z. mays probably increased with shading by up to ∼‰. The difference in tree leaf Δ among the forest height levels would be mainly due to decreasing intercellular CO₂ (C _i) with the increase in irradiance. Potential assimilation rate for the three tree species probably increased with height, since leaf nitrogen content on an area basis for these species also increased with height. However, the increase in stomatal conductance for these tree species would fail to meet the increase in potential assimilation rate, which might lead to increasing the degree of stomatal limitation in photosynthesis with height. Received: 30 September 1995 / Accepted: 25 October 1996     

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

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

Unusually low carbon isotope ratios in plants from hanging gardens in southern Utah

Leaf carbon isotope ratios (δ¹³C) and photosynthetic gas exchange were measured on plants growing in hanging garden communities in southern Utah, USA. Hanging gardens are unusual, mesic cliff communities occurring where water seeps from the sandstone bedrock in an otherwise extremely arid region; there is very limited overlap in species distributions inside and outside these gardens. Solar exposure in hanging gardens varied with orientation and one of the gardens (Ribbon Garden) was shaded throughout the day. The leaf δ¹³C values of plants in hanging gardens were significantly more negative than for plants from either nearby ephemeral wash or riparian communities. In Ribbon Garden, the observed δ¹³C values were as low as −34.8‰, placing them among the most negative values reported for any terrestrial plant species growing in a natural environment. Hanging garden plants were exposed to normal atmospheric CO₂ with an average δ¹³C value of −7.9‰ and so the low leaf δ¹³C values could not be attributed to exposure to a CO₂ source with low ¹³C content. There was a seasonal change toward more negative leaf δ¹³C values at the end of the growing season. The observed leaf δ¹³C values were consistent with photosynthetic gas exchange measurements that indicated unusually high leaf intercellular CO₂ concentrations associated with the relatively low light levels in hanging gardens. Thus, extremely negative leaf δ¹³C values would be expected if significant amounts of the seasonal carbon gain occur at light levels low enough to be near the light compensation point. Maximum observed photosynthetic rates varied with light levels at each of the gardens, with maximum rates averaging 20.3, 14.6, and 3.1 μmol m⁻²s⁻¹ at Double Garden, Lost Garden, and Ribbon Garden, respectively. Leaf nitrogen contents averaged 18.5 mg g⁻¹ in species from the more shaded hanging gardens (Lost and Ribbon). When expressed on a leaf area basis, nitrogen contents averaged 117 mmol N m⁻² at Lost Garden and 65 mmol N m⁻² at Ribbon Garden (shadiest of the two gardens). Leaf nitrogen isotope ratios averaged −2.3‰ (range of −0.7 to −6.1‰), suggesting that most of the nitrogen was derived from a biological fixation source which is most likely the Nostoc growing on the sandstone walls at the seep. These values contrast with leaf nitrogen isotope ratios of 5–9‰ which have been previously reported for arid zone plants in nearby ecosystems. Received: 19 January 1997 / Accepted: 19 April 1997     

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.     

Contributions of C3 and C4 plants to higher trophic levels in an Amazonian savanna

We studied the energy flow from C₃ and C₄ plants to higher trophic levels in a central Amazonian savanna by comparing the carbon stable-isotope ratios of potential food plants to the isotope ratios of species of different consumer groups. All C₄ plants encountered in our study area were grasses and all C₃ plants were bushes, shrubs or vines. Differences in δ¹³C ratios among bushes (xˉ = −30.8, SD = 1.2), vines (xˉ = −30.7, SD = 0.46) and trees (xˉ = −29.7, SD = 1.5) were small. However the mean δ¹³C ratio of dicotyledonous plants (xˉ = −30.4, SD = 1.3) was much more negative than that of the most common grasses (xˉ = −13.4, SD = 0.27). The insect primary consumers had δ¹³C ratios which ranged from a mean of −29.5 (SD = 0.47) for the grasshopper Tropidacris collaris to a mean of −14.7 (SD = 0.56) for a termite (Nasutitermes sp.), a range similar to that of the vegetation. However, the common insectivorous and omnivorous vertebrates had intermediate values for δ¹³C, indicating that carbon from different autotrophic sources mixes rapidly as it moves up the food chain. Despite this mixing, the frogs and lizards generally had higher values of δ¹³C (xˉ = −21.7, SD = 1.6; xˉ = −21.9, SD = 1.8, respectively) than the birds (xˉ = −24.8, SD = 1.8) and the only species of mammal resident in the savanna (xˉ = −25.4), indicating that they are generally more dependent on, or more able to utilise, food chains based on C₄ grasses. Received: 7 May 1998 / Accepted: 30 November 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     

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.     

Nitrogen and carbon isotope responses of Chinese cabbage and chrysanthemum to the application of liquid pig manure

The effects of the liquid pig manure (LM) used in organic farming on the natural abundance of ¹⁵N and ¹³C signatures in plant tissues have not been studied. We hypothesized that application of LM will (1) increase δ¹⁵N of plant tissues due to the high δ¹⁵N of N in LM as compared with soil N or inorganic fertilizer N, and (2) increase δ¹³C of plant tissues as a result of high salt concentration in LM that decreases stomatal conductance of plants. To test these hypotheses, variations in the δ¹⁵N and δ¹³C of Chinese cabbage (Brassica campestris L.) and chrysanthemum (Chrysanthemum morifolium Ramatuelle) with two different LMs (with δ¹⁵N of +15.6 and +18.2‰) applied at two rates (323 and 646 kg N ha^-1 for cabbage and 150 and 300 kg N ha^-1 for chrysanthemum), or urea (δ¹⁵N = -2.7‰) applied at the lower rate above for the respective species, in addition to the control (no N input) were investigated through a 60-day pot experiment. Application of LM significantly increased plant tissue δ¹⁵N (range +9.4 to +14.9‰) over the urea (+3.2 to +3.3‰) or control (+6.8 to 7.7‰) treatments regardless of plant species, strongly reflecting the δ¹⁵N of the N source. Plant tissue δ¹³C were not affected by the treatments for cabbage (range −30.8 to −30.2‰) or chrysanthemum (−27.3 to −26.8‰). However, cabbage dry matter production decreased while its δ¹³C increased with increasing rate of LM application or increasing soil salinity (P < 0.05), suggesting that salinity stress caused by high rate of LM application likely decreased stomatal conductance and limited growth of cabbage. Our study expanded the use of the δ¹⁵N technique in N source (organic vs. synthetic fertilizer) identification and suggested that plant tissue δ¹³C maybe a sensitive indicator of plant response to salinity stress caused by high LM application rates.     

Functional diversity in an Amazonian rainforest of French Guyana: a dual isotope approach (δ15N and δ13C)

Functional aspects of biodiversity were investigated in a lowland tropical rainforest in French Guyana (5°2′N, annual precipitation 2200 mm). We assessed leaf δ¹⁵N as a presumptive indicator of symbiotic N₂ fixation, and leaf and wood cellulose δ¹³C as an indicator of leaf intrinsic water-use efficiency (CO₂ assimilation rate/leaf conductance for water vapour) in dominant trees of 21 species selected for their representativeness in the forest cover, their ecological strategy (pioneers or late successional stage species, shade tolerance) or their potential ability for N₂ fixation. Similar measurements were made in trees of native species growing in a nearby plantation after severe perturbation (clear cutting, mechanical soil disturbance). Bulk soil δ¹⁵N was spatially quite uniform in the forest (range 3–5‰), whereas average leaf δ¹⁵N ranged from −0.3‰ to 3.5‰ in the different species. Three species only, Diplotropis purpurea, Recordoxylon speciosum (Fabaceae), and Sclerolobium melinonii (Caesalpiniaceae), had root bacterial nodules, which was also associated with leaf N concentrations higher than 20 mg g⁻¹. Although nodulated trees displayed significantly lower leaf δ¹⁵N values than non-nodulated trees, leaf δ¹⁵N did not prove a straightforward indicator of symbiotic fixation, since there was a clear overlap of δ¹⁵N values for nodulated and non-nodulated species at the lower end of the δ¹⁵N range. Perturbation did not markedly affect the difference δ¹⁵N_soil−δ¹⁵N_leaf, and thus the isotopic data provide no evidence of an alteration in the different N acquisition patterns. Extremely large interspecific differences in sunlit leaf δ¹³C were observed in the forest (average values from −31.4 to −26.7‰), corresponding to intrinsic water-use efficiencies (ratio CO₂ assimilation rate/leaf conductance for water vapour) varying over a threefold range. Wood cellulose δ¹³C was positively related to total leaf δ¹³C, the former values being 2–3‰ higher than the latter ones. Leaf δ¹³C was not related to leaf δ¹⁵N at either intraspecific or interspecific levels. δ¹³C of sunlit leaves was highest in shade hemitolerant emergent species and was lower in heliophilic, but also in shade-tolerant species. For a given species, leaf δ¹³C did not differ between the pristine forest and the disturbed plantation conditions. Our results are not in accord with the concept of existence of functional types of species characterized by common suites of traits underlying niche differentiation; rather, they support the hypothesis that each trait leads to a separate grouping of species. Received: 18 August 1997 / Accepted: 14 April 1998     

Ontogenetic dietary shifts in European common frog (Rana temporaria) revealed by stable isotopes

During migrations and ontogeny amphibians change their habitat and feeding, and thus are important in linking terrestrial and aquatic ecosystems. We measured δ ¹³C and δ ¹⁵N values of early stages (egg, embryo, tadpole) and toes of adult frogs Rana temporaria, collected from a small wetland in Lithuania. We compared the isotopic composition of these tissues with potential food sources, excrements of tadpoles, and filled intestinal tracts. We found that δ ¹³C values in R. temporaria tadpoles were markedly depleted in comparison to adults, eggs or embryos, demonstrating a terrestrial to aquatic shift in energy sources. After the onset of feeding, tadpoles approached isotopic equilibrium with available food (algae and litter). Tadpoles had higher δ ¹⁵N than both algae and litter, differing by 3.6 and 2.4‰, respectively, and similar δ ¹³C to these sources. However, tadpole excrements and body tissue diverged, with mean δ ¹³C values of excrements (−30.3 ± 1.6‰ SD) more similar to litter (−31.7 ± 1.2‰ SD) and body tissue δ ¹³C (−34.8 ± 0.7‰ SD) more similar to algae (−34.2 ± 4.1‰ SD). This suggests that algal resources are critical in early life stages of this anuran, particularly at stages characterized by high growth and low development (stages: 25–35).     

Isotopic signature of nitrate in two contrasting watersheds of Brush Brook,Vermont, USA

We used the dual isotope method to study differences in nitrate export in two subwatersheds in Vermont, USA. Precipitation, soil water and streamwater samples were collected from two watersheds in Camels Hump State Forest, located within the Green Mountains of Vermont. These samples were analyzed for the δ¹⁵N and δ¹⁸O of NO₃⁻. The range of δ¹⁵N–NO₃⁻ values overlapped, with precipitation −4.5‰ to +2.0‰ (n = 14), soil solution −10.3‰ to +6.2‰ (n = 12) and streamwater +0.3‰ to +3.1‰ (n = 69). The δ¹⁸O of precipitation NO₃⁻ (mean 46.8 ± 11.5‰) was significantly different (P < 0.001) from that of the stream (mean 13.2 ± 4.3‰) and soil waters (mean 14.5 ± 4.2‰) even during snowmelt periods. Extracted soil solution and streamwater δ¹⁸O of NO₃⁻ were similar and within the established range of microbially produced NO₃⁻, demonstrating that NO₃⁻ was formed by microbial processes. The δ¹⁵N and δ¹⁸O of NO₃⁻ suggests that although the two tributaries have different seasonal NO₃⁻ concentrations, they have a similar NO₃⁻ source.     

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.     

Variation in carbon isotope composition among years in the riparian tree Populus fremontii

Water availability is an important factor limiting the productivity of desert plants but little is known about the impact of water-limiting conditions on the physiology of plants in mesic environments. Riparian ecosystems of the western US receive significantly more water than the surrounding desert environments but experience dramatic interannual fluctuations in water availability because both stream flow and precipitation are highly variable over time. This variability results in different growing conditions each year which may influence the physiology of riparian species such as Fremont cottonwood (Populus fremontii), the dominant, native canopy tree species in lowland southwestern US river systems. We wished to determine if the physiology of this species varies among years, what climatic parameters are related to any observed physiological variation and if individuals within a P. fremontii population differ in their physiological response to variation through time. We collected tree ring cores from a central New Mexico cottonwood population and analyzed carbon isotope composition (δ¹³C) in each year from 1981 to 1995. We used δ¹³C analysis in this study because it allowed us to obtain multi-year estimates of physiological activity. During these years, mean stream flow at our study site ranged over two orders of magnitude from 0.82 to 80.94 m³ s⁻¹, precipitation ranged fourfold from 49 to 215 ccmm and mean temperature ranged from 20.5 to 22.6°C during the growing season. δ¹³C varied from a low of −26.7‰ in 1984 to a high of −24.7‰ in 1981. Low δ¹³C values were associated with years in which stream flow and/or precipitation were high and temperature was low. The opposite was true of years with high δ¹³C values. We observed a strong linear relationship between δ¹³C and stream flow during years when stream flow was <25 m³ s⁻¹ but no significant relationship between these variables when stream flow was >25 m³ s⁻¹. Additionally, there was a linear relationship between δ¹³C and precipitation during years when stream flow was <25 m³ s⁻¹ but not in years when stream flow was >25 m³ s⁻¹. These data suggest that above a threshold of total stream flow, increased flow does not influence physiology. Below this threshold, precipitation can be an important water source. The ten individuals within our study population varied significantly in mean δ¹³C values but responded to interannual variation in a similar manner (i.e., all individuals had low δ¹³C values when water was abundant). These results suggest that precipitation as well as stream flow are important factors influencing the physiology of this riparian tree. Received: 25 November 1998 / Accepted: 9 February 1999