Stable isotopes of carbon and nitrogen as indicators of diet and trophic structure of the fish community in a shallow hypereutrophic lake

Stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes were employed to elucidate energy flows and trophic interactions in Lake Apopka, a hypereutrophic lake in central Florida, U.S.A. Isotope compositions of lake biota ranged from −27·1 to −3·0‰ for δ¹³C, and from 3·7 to 13·9‰ for δ¹⁵N. The food web was based primarily on plankton production with diatoms, Microcystis and zooplankton dominating the diet of fish. Carbon isotope evidence showed that pico- and nano-phytoplankton were not a direct carbon source for fish, but were important to zooplankton. δ¹⁵N mass balance estimates indicated that planktivorous fish obtained 48–85% of their diets from zooplankton. The ∼3‰ range of δ¹⁵N in gizzard shad reflected increasing dependence on zooplankton as fish grew whereas the positive relationship between total length and δ¹⁵N of largemouth bass reflected increasing predation on larger planktivorous fish with growth. The broad ranges of δ¹³C (−25·9 to −9·5‰) and δ¹⁵N (5·8 to 14·4‰) of blue tilapia were indicators of diet diversity. Two presumed omnivores (brown bullhead and white catfish) and piscivores (black crappie, largemouth bass and Florida gar) were found to depend on planktivorous fish. However, stable isotope data revealed no trophic links between blue tilapia, an abundant fish in the near-shore area, and piscivores.     

Starvation and low feeding levels result in an enrichment of 13C in lipids and 15N in protein of Nile tilapia Oreochromis niloticus L.

The influence of different feeding levels below and slightly above maintenance on whole body δ¹³C and δ¹⁵N values of Nile tilapia Oreochromis niloticus was examined. The energy budget of each fish was determined by indirect calorimetry. The δ¹³C values of the lipid-free material of Nile tilapia fed below and slightly above maintenance level did not differ between the feeding groups, but the δ¹³C values in the lipids and the δ¹⁵N values of the lipid-free material showed small but significant differences. Those fish with a negative lipid retention had significantly higher δ¹³C values in the lipid fraction compared to fish that synthesized fatty acids. There was a significant negative correlation between the amount of energy metabolized by the fish and both the δ¹³C values in the lipids and the δ¹⁵N values of the lipid-free material. Fasting and feeding below the maintenance level may influence the isotopic composition of animals and should therefore be considered in ecological and nutritional studies.     

A stable isotopic investigation into the causes of decline in a sub-Antarctic predator, the rockhopper penguin Eudyptes chrysocome

The rockhopper penguin (Eudyptes chrysocome) is a conspicuous apex marine predator that has experienced marked population declines throughout most of its circumpolar breeding distribution. The cause(s) for the declines remain elusive, but the relatively large spatio‐temporal scale over which population decreases have occurred implies that ecosystem‐scale, at‐sea factors are likely to be involved. We employ stable isotope analyses of carbon (¹³C/¹²C, expressed as δ¹³C) and nitrogen (¹⁵N/¹⁴N, δ¹⁵N) in time‐series of rockhopper penguin feather samples, dating back to 1861, in order to reconstruct the species' ecological history. Specifically, we examine whether rockhopper penguin population decline has been associated with a shift towards lower primary productivity in the ecosystem in which they feed, or with a shift to a diet of lower trophic status and lower quality, and we use long‐term temperature records to evaluate whether shifts in isotope ratios are associated with annual variations in sea surface temperature. Having controlled temporally for the Suess Effect and for increases in CO₂ concentrations in seawater, we found that overall, δ¹³C signatures decreased significantly over time in rockhopper penguins from seven breeding sites, supporting the hypothesis that decreases in primary productivity, and hence, carrying capacity, for which δ¹³C signature is a proxy, have been associated with the decline of penguin populations. There was some evidence of a long‐term decline in δ¹⁵N at some sites, and strong evidence that δ¹⁵N signatures were negatively related to sea surface temperatures across sites, indicative of a shift in diet to prey of lower trophic status over time and in warm years. However, a site‐by‐site analysis revealed divergent isotopic trends among sites: five of seven sites exhibited significant temporal or temperature‐related trends in isotope signatures. This study highlights the utility of stable isotope analyses when applied over relatively long timescales to apex predators.     

Effect of host shoot clipping on carbon and nitrogen sources for arbuscular mycorrhizal fungi

The effect of clipping of the host-plant shoot on the sources of carbon and nitrogen for the arbuscular mycorrhizal (AM) fungus Gigaspora margarita was determined by measuring ¹³C in spores and hyphae in cocultures of C₃ and C₄ plants and by differential ¹⁵N labeling. C₃ and C₄ plants, which have different δ¹³C values, were grown in the same container separated by a series of hyphal compartments. The C₃ and C₄ plants were applied with ¹⁴N- and ¹⁵N-urea, respectively. After clipping of the C₃ shoots, spore δ ¹³C gradually approached that of the C₄ roots. Hyphal δ ¹³C paralleled that of spores. Spore % ¹⁵N was similar to that of mineral N in the C₄ plant compartment. Thus C in G. margarita coming from the clipped plants decreased with time. This demonstrates that C in AM fungi comes from living plants, whilst the N in spores comes mostly from the soil. Accepted: 28 November 2000     

Stable isotope composition of organic compounds transported in the phloem of European beech--evaluation of different methods of phloem sap collection and assessment of gradients in carbon isotope composition during leaf-to-stem transport

The analysis of stable isotope composition (delta13C, delta15N, delta18O) of phloem-transported organic matter is a useful tool for assessing short-term carbon and water balance of trees. A major constraint of the general application of this method to trees at natural field sites is that the collection of phloem sap with the "phloem bleeding" technique is restricted to particular species and plant parts. To overcome this restriction, we compared the contents (amino compounds and sugars) and isotope signatures (delta13C, delta15N, delta18O) of phloem sap directly obtained from incisions in the bark (bleeding technique) with phloem exudates where bark pieces were incubated in aqueous solutions (phloem exudation technique with and without chelating agents [EDTA, polyphosphate] in the initial sampling solution, which prevent blocking of sieve tubes). A comparable spectrum of amino compounds and sugars was detected using the different techniques. O, C, or N compounds in the initial sampling solution originating from the chelating agents always decreased precision of determination of the respective isotopic signatures, as indicated by higher standard deviation, and/or led to a significant difference of mean delta as compared to the phloem bleeding technique. Hence, depending on the element from which the ratio of heavy to light isotope is determined, compounds lacking C, N, and/or O should be used as chelating agents in the exudation solution. In applying the different techniques, delta13C of organic compounds transported in the phloem of the twig (exudation technique with polyphosphate as chelating agent) were compared with those in the phloem of the main stem (phloem bleeding technique) in order to assess possible differences in carbon isotope composition of phloem carbohydrates along the tree axis. In July, organic compounds in the stem phloem were significantly enriched in 13C by > 1.3 per thousand as compared to the twig phloem, whereas this effect was not observed in September. Correlation analysis between delta13C and stomatal conductance (Gs) revealed the gradient from the twigs to the stem observed in July may be attributed to temporal differences rather than to spatial differences in carbon isotope composition of sugars. As various authors have produced conflicting results regarding the enrichment/depletion of 13C in organic compounds in the leaf-to-stem transition, the different techniques presented in this paper can be used to provide further insight into fractionation processes associated with transport of C compounds from leaves to branches and down the main stem.     

壳斗科三种植物种子大小对昆虫寄生及种子存活率的影响

种子内的寄生昆虫可以严重影响种子的发育、损害种子活力。种子足余策略理论认为大种子有利于抵御和适应昆虫寄生取食,但动物最优觅食理论推测,大种子更易遭受昆虫寄生。为对这两种对立观点进行验证,本实验以青冈、苦槠和麻栎各2个种群的种子为材料,对昆虫寄生与完好种子间的体积和萌发率进行比较,并对寄生种子萌发率与种子体积的关系进行了分析。结果显示:(1)在6个种群的种子中,只有松阳麻栎和青冈种群的寄生种子体积大于完好种子,其余4个种群的寄生种子体积小于完好种子,但这种差异不显著;(2)所有寄生种子的整体萌发率(18%)显著低于完好种子(45.66%)(P<0.001),在不同种群内,寄生种子的萌发率也分别显著低于完好种子。(3)比较同种植物体积差异显著的寄生种子的萌发率发现,大种子总比小种子具有更高的萌发率,但差异不显著;在不同植物的寄生种子间比较时,体积最大的麻栎种子萌发率显著高于体积较小的青冈和苦槠种子。研究结果表明,象虫在种子上产卵时对大种子没有选择偏好,在昆虫寄生取食严重损害种子活力的压力下,大种子比小种子具有更强的耐受力。     

中国东部南北样带森林优势植物叶片的水分利用效率和氮素利用效率

通过测定中国东部南北样带主要森林生态系统中10种优势植物(兴安落叶松、蒙古栎、水曲柳、紫椴、色木槭、红松、杉木、木荷、马尾松、锥栗)叶片的碳氮含量(Cmass、Nmass)、同位素丰度(δ13C、δ15N)以及光合响应曲线,分析了不同优势植物叶片的水分利用效率和氮素利用效率之间的差异及其相互关系.结果表明： 不同生活型植物叶片的Nmass和δ15N差异显著,表现为阔叶植物>针叶植物,落叶植物>常绿植物;最大光合速率(Pn max)表现为针叶植物>阔叶植物,落叶植物>常绿植物;植物叶片的瞬时水分利用效率(WUEi)和长期水分利用效率(WUE)均表现为阔叶植物>针叶植物,常绿植物>落叶植物;植物叶片的瞬时氮素利用效率(NUEi)和长期氮素利用效率(NUE)则表现出相反的规律,且常绿植物和落叶植物叶片的NUE差异显著;WUEi和WUE之间相关性不显著,而NUEi和NUE之间呈显著正相关.植物叶片的水分利用效率与氮素利用效率显著负相关.两种资源利用效率均受植物生活型的影响,并且存在一定的制约关系.     

Continuous variation in the pattern of marine v. freshwater foraging in brown trout Salmo trutta L. from Loch Lomond, Scotland

Carbon stable-isotope analysis showed that individual brown trout Salmo trutta in Loch Lomond adopted strategies intermediate to that of freshwater residency or anadromy, suggesting either repeated movement between freshwater and marine environments, or estuarine residency. Carbon stable-isotope (δ¹³C) values from Loch Lomond brown trout muscle tissue ranged from those indicative of assimilation of purely freshwater-derived carbon to those reflecting significant utilization of marine-derived carbon. A single isotope, two-source mixing model indicated that, on average, marine C made a 33% contribution to the muscle tissue C of Loch Lomond brown trout. Nitrogen stable isotope, δ¹⁵N, but not δ¹³C was correlated with fork length suggesting that larger fish were feeding at a higher trophic level but that marine feeding was not indicated by larger body size. These results are discussed with reference to migration patterns in other species.     

Using the δ13C and δ15N of whitefish scales for retrospective ecological studies: changes in isotope signatures during the restoration of Lake Geneva, 1980–2001

The scales of whitefish Coregonus lavaretus were used in place of dorsal muscle, which necessitates killing the fish, to study food webs from the δ¹³C and δ¹⁵N isotopic ratios in the organic fraction. As scales are composed of both organic and calcified fractions, a protocol for scale decalcification was first devised. The δ¹³C and δ¹⁵N values of the decalcified scales were then shown to be closely correlated to those of the dorsal muscle, demonstrating that scales could be used in place of muscle to study food webs. Changes in the δ¹³C of whitefish were determined from a scale collection that extended over the period during which the trophic state of Lake Geneva was recovering.     

Bulk leaf δ18O and δ13C reflect the intensity of intraspecific competition for water in a semi-arid tussock grassland

We investigated the extent to which plant water and nutrient status are affected by intraspecific competition intensity and microsite quality in a monodominant tussock grassland. Leaf gas exchange and stable isotope measurements were used to assess the water relations of Stipa tenacissima tussocks growing along a gradient of plant cover and soil depth in a semi-arid catchment of Southeast Spain. Stomatal conductance and photosynthetic rate decreased with increasing intensity of competition during the wet growing season, leading to foliar δ ¹⁸O and δ ¹³C enrichment. A high potential for runoff interception by upslope neighbours exerted strong detrimental effects on the water and phosphorus status of downslope S. tenacissima tussocks. Foliar δ ¹⁵N values became more enriched with increasing soil depth. Multiple stepwise regression showed that competition potential and/or rhizosphere soil depth accounted for large proportions of variance in foliar δ ¹³C, δ ¹⁸O and δ ¹⁵N among target tussocks (57, 37 and 64%, respectively). The results presented here highlight the key role that spatial redistribution of resources (water and nutrients) by runoff plays in semi-arid ecosystems. It is concluded that combined measurement of δ ¹³C, δ ¹⁸O and nutrient concentrations in bulk leaf tissue can provide insight into the intensity of competitive interactions occurring in natural plant communities.     

Altitudinal gradients of grassland carbon and nitrogen isotope composition are recorded in the hair of grazers

Aim The hair of grazers provides an isotopic record of environmental and nutritional signals. Here, we assess the effect of altitude on the carbon and nitrogen isotope composition of the hair of ruminant grazers and its relation to grassland vegetation, to evaluate the use of hair isotope data for ecosystem reconstruction, animal nutritional ecology and biogeochemical studies in montane environments. Location European Alps. Methods We sampled grassland vegetation (pure C₃) and the hair of ruminants along an altitudinal gradient (400–2500 m), and analysed their isotope composition (δ¹³C and δ¹⁵N). Results were compared with published effects of altitude on ¹³C in C₃ plants at the species level and on ¹⁵N at the community level. The study was complemented with a comparison of diet and hair isotope composition in ruminants held in confinement. Results δ¹³C of hair increased (c. 1.1‰ km⁻¹) and δ¹⁵N decreased (c. 1.1‰ km⁻¹) with altitude. The same changes occurred in local grassland vegetation, and in regional to global grassland data sets. Offsets between hair and vegetation ¹³C or ¹⁵N (‘diet–hair shift’) were independent of altitude. Sheep (Ovis aries) and cattle (Bos taurus) exhibited a ¹³C shift near +3‰, but that of goats (Capra hircus) was larger (+4.2‰) in alpine environments and in confinement. The diet–hair shift for ¹⁵N was more variable (+2.1 to +3.6‰). Main conclusions Grazer hair provides a faithful spatially and temporally integrated record of grassland isotope composition, useful for ecosystem and environment reconstruction. The effect of altitude on hair ¹⁵N is important for studies of trophic relationships: an altitude shift of 2000 m produced the same effect in hair ¹⁵N as would a shift from an animal tissue‐based to a plant‐based diet. The similarity of altitude effects on δ¹³C of individual plant species, vegetation and hair indicates that the effect of altitude on species‐level ‘intrinsic water use efficiency’ scales up linearly to the community and landscape level.     

Natural 15N abundance of epiphytes depends on the position within the forest canopy: source signals and isotope fractionation

The natural ¹⁵N abundance (δ¹⁵N) of epiphytes and its N sources were studied in the canopy of a lowland rainforest in Costa Rica. Vascular and non‐vascular epiphytes and canopy soils were collected from four canopy zones and analysed for N contents and δ¹⁵N signals. In addition, the N concentrations and δ¹⁵N signatures of bulk precipitation, throughfall and stemflow were measured during the wet and the dry season. The δ¹⁵N values of epiphyte leaves decreased significantly from the lower zones (means of ?3·9 and ?4·3‰) to the upper zones (means of ?5·4 and ?6·1‰) of the canopy. In contrast, δ¹⁵N signatures of canopy soils (average ?0·3‰) differed little between the zones. Bulk deposition was enriched in ¹⁵N (+4·3‰) compared to all other potential N sources and was higher than throughfall and stemflow (+0·5 to ?1·3‰). δ¹⁵N values of atmospheric deposition were inversely related to those of the epiphyte leaves, whereas N isotopic composition of canopy soils did not vary significantly. Consequently, it is concluded that the variations in foliar N isotope composition of epiphytes were not simply caused by utilization of isotopically different N sources, but by different ¹⁵N discrimination during N acquisition.     

Allometry, root/shoot ratio and root architecture in understory saplings of deciduous dicotyledonous trees in central Japan

Plant allometry that is related to plant architecture and biomass allocation strongly influences a plants ability to grow in shaded forest understory. Some allometric traits can change with plant size. The present study compared crown and trunk allometries, root/shoot biomass allometry, and root architecture among understory saplings (0.25--5m height, except for two trees > 5 < 7 m) of seven deciduous dicotyledonous species in central Japan. Associations of the crown and trunk allometries with several plant morphological attributes were analyzed. Branch morphology (plagiotropyvs orthotropy) and life size were correlated with sapling crown and trunk allometries. Both large leaves and orthotropic branches were associated with a narrow small crown and slender trunk. The root/shoot ratio decreased rapidly with increasing plant height for saplings shorter than about 1.5 m. Less shade-tolerant species tended to have smaller root/shoot ratios for saplings taller than 1.5 m. With an increase in plant height, the branch/trunk biomass ratio decreased for saplings with plagiotropic branches but increased for saplings with orthotropic branches. Four subcanopy species (Acer distylum, Carpinus cordata, Fraxinus lanuginosa and Acanthopanax sciadophylloides) had superficial root systems; a common understory species (Sapium japonica) had a deep tap root system; and a canopy species (Magnolia obovata) and a subcanopy species (Acer tenuifolium) had heart root systems of intermediate depth. The root depth was not related to shade tolerance. Among species of the same height, the difference in fine root length can be 30-fold.     

Carbon isotope discrimination and the ratio of carbon gained to water lost in barley cultivars

Abstract A negative correlation between water-use efficiency (W), defined as the ratio of moles of carbon in the plant to moles of water transpired, and carbon isotope discrimination (Δ) was established for barley in pot experiments using 12 cultivars. The correlation was strong in two independent experiments in four different controlled environment where ambient temperature and vapour pressure deficit were varied and plants were either well-watered or given limited amounts of water. Variation among cultivars was found in both Δ and W and rankings of both parameters, according to cultivar, were similar in different environments. Limiting water usually increased water-use efficiency of plants. Total dry matter can be substituted for moles of carbon when calculating water-use efficiency but the correlation between W and Δ were calculated using the carbon content of dry matter. There were differences varied significantly among cultivars. Despite these differences, correlations were also large between whole plant W and Δ of any of the plant parts. The amount of dry matter partitioned into reproductive growth varied genetically, as did the effect of stress on the partitioning. Growth, W and Δ of barley were compared with theory derived from gas exchange properties and with other literature. The effect on W of variation in vapour pressure deficit in these experiments was removed by multiplying W by vapour pressure deficit to derive the parameter, k(Pa mol C/mol H₂O). This allowed comparisons among experiments with different vapour pressure deficits. The mean k for these barley cultivars was similar to that calculated by others for grasses. However, variation was found, and, in contrast with previous work which treats k as a species constant, we conclude that there is promise in selecting for increased k.