The influence of diet on the distribution of nitrogen isotopes in animals was investigated by analyzing animals grown in the laboratory on diets of constant nitrogen isotopic composition. 
The isotopic composition of the nitrogen in an animal reflects the nitrogen isotopic composition of its diet. The δ^(15)N values of the whole bodies of animals are usually more positive than those of their diets. Different individuals of a species raised on the same diet can have significantly different δ^(15)N values. The variability of the relationship between the δ^(15)N values of animals and their diets is greater for different species raised on the same diet than for the same species raised on different diets. Different tissues of mice are also enriched in ^(15)N relative to the diet, with the difference between the δ^(15)N values of a tissue and the diet depending on both the kind of tissue and the diet involved. The δ^(15)N values of collagen and chitin, biochemical components that are often preserved in fossil animal remains, are also related to the δ^(15)N value of the diet. 
The dependence of the δ^(15)N values of whole animals and their tissues and biochemical components on the δ^(15)N value of diet indicates that the isotopic composition of animal nitrogen can be used to obtain information about an animal's diet if its potential food sources had different δ^(15)N values. The nitrogen isotopic method of dietary analysis probably can be used to estimate the relative use of legumes vs non-legumes or of aquatic vs terrestrial organisms as food sources for extant and fossil animals. However, the method probably will not be applicable in those modern ecosystems in which the use of chemical fertilizers has influenced the distribution of nitrogen isotopes in food sources. 
The isotopic method of dietary analysis was used to reconstruct changes in the diet of the human population that occupied the Tehuacan Valley of Mexico over a 7000 yr span. Variations in the δ^(15)C and δ^(15)N values of bone collagen suggest that C_4 and/or CAM plants (presumably mostly corn) and legumes (presumably mostly beans) were introduced into the diet much earlier than suggested by conventional archaeological analysis.

Influence of Diet On the Distribtion of Nitrogen Isotopes in Animals

The use of stable isotopes to solve biogeochemical problems in ecosystem analysis is increasing rapidly because stable isotope data can contribute both source-sink (tracer) and process information: The elements C, N, S, H, and all have more than one isotope, and isotopic compositions of natural materials can be measured with great precision with a mass spectrometer. Isotopic compositions change in predictable ways as elements cycle through the biosphere. These changes have been exploited by geochemists to understand the global elemental cycles. Ecologists have not until quite recently employed these techniques. The reasons for this are, first, that most ecologists do not have the background in chemistry and geochemistry to be fully aware of the possibilities for exploiting the natural variations in stable isotopic compositions, and second, that stable isotope ratio measurements require equipment not normally available to ecologists. This is unfortunate because some of the more intractable problems in ecology can be profitably addressed using stable isotope measurements. Stable isotopes are ideally suited to increase our understanding of element cycles in ecosystems. This review is written for ecologists who would like to learn more about how stable isotope analyses have been and can be used in ecosystem studies. We begin with an explanation of isotope terminology and fractionation, then summarize isotopic distributions in the C, N, and S biogeochemical cycles, and conclude with five case studies that show how stable isotope measurements can provide crucial information for ecosystem analysis. We restrict this review to studies of natural variations in C, N, and S isotopic abundances, cxcluding from consideration ~5N enrichment studies and hydrogen and oxygen isotope studies. Our focus on C, N, and S derives in part from our

Stable isotopes in ecosystem studies

Use of stable isotope ratios to trace pathways of organic matter among consumers requires knowledge of the isotopic shift between diet and consumer. Variation in trophic shift among consumers can be substantial. For data from the published literature and supplementary original data (excluding fluid-feeding consumers), the mean isotopic shift for C was + 0.5 + 0.13%o rather than 0.0%o, as commonly assumed. The shift for C was higher for consumers analyzed as muscle (+ 1.3 + 0.30%o) than for consumers analyzed whole (+ 0.3 +0.14%o). Among consumers analyzed whole, the trophic shift for C was lower for consumers acidified prior to analysis (-0.2 + 0.21%o) than for unacidified samples ( +0.5 + 0.17%o). For N, trophic shift was lower for consumers raised on invertebrate diets (+ 1.4 + 0.21%o) than for consumers raised on other high-protein diets (+3.3 +0.26%o) and was intermediate for consumers raised on plant and algal diets (+2.2 +0.30%o). The trophic shift for S differed between high-protein (+ 2.0 + 0.65%o) and low-protein diets (-0.5 + 0.56%o). Thus, methods of analysis and dietary differences can affect trophic shift for consumers; the utility of stable isotope methods can be improved if this information is incorporated into studies of trophic relationships. Although few studies of stable isotope ratios have considered variation in the trophic shift, such variation is important because small errors in estimates of trophic shift can result in large errors in estimates of the contribution of sources to consumers or in estimates of trophic position.

Variation in trophic shift for stable isotope ratios of carbon, nitrogen, and sulfur

Stable isotope ratios provide clues about the origins and transformations of organic matter. A few key reactions control the isotopic composition of most organic matter. Isotopic variations introduced by these reactions are often passed on with little change so that isotopic measurements can indicate natural pathways and flows “downstream” from these key reactions. When chemical and metabolic processes scramble the information content of molecules, isotopic compositions are often preserved. This realization has prompted increasing use of stable isotope analyses as a tool for understanding complex ecological processes.

δ13C Measurements as Indicators of Carbon Flow in Marine and Freshwater Ecosystems

To understand the ecology of migratory animals it is important to link geographic regions used by individuals including breeding, wintering, and intermediate stopover sites. Previous conventional approaches used to track animal movements have relied on extrinsic markers and typically the subsequent recovery of individuals. This approach has generally been inappropriate for most small, or non-game animals. The use of intrinsic markers such as fatty acid profiles, molecular DNA analyses, and the measurement of naturally occurring stable isotopes in animal tissues offer alternative approaches. This paper reviews the use of stable isotope analyses (primarily δ13C, δ15N, δ34S, δD, δ87Sr) to trace nutritional origin and migration in animals. This approach relies on the fact that foodweb isotopic signatures are reflected in the tissues of organisms and that such signatures can vary spatially based on a variety of biogeochemical processes. Organisms moving between isotopically distinct foodwebs can carry with them information on the location of previous feeding. Such an approach has been used to track animal use of inshore versus offshore, marine versus freshwater, terrestrial C3 versus marine, terrestrial mesic versus xeric, and C3 versus C4 or Crassulacean acid metabolism foodwebs. More recently, the use of stable hydrogen isotope analyses (δD) to link organisms to broad geographic origin in North America is based on large-scale isotopic contours of growing-season average δD values in precipitation. This technique, especially when combined with the assay of other stable isotopes, will be extremely useful in helping to track migration and movement of a wide range of animals from insects to birds and mammals. Future research to refine our understanding of natural and anthropogenic-induced isotopic gradients in nature, and to explore the use of stable isotopes of other elements, is recommended.

Tracing origins and migration of wildlife using stable isotopes: a review

The use of stable carbon isotopes as a means of studying energy flow is increasing in ecology and paleo- ecology. However, secondary fractionation and turnover of stable isotopes in animals are poorly understood pro- cesses. This study shows that tissues of the gerbil (Meriones unguienlatus) have different 613C values when equilibrated on corn (C4) or wheat (C~) diets with constant 13c/1ac contents. Lipids were depleted 3.0%o and hair was enriched 1.0%o relative to the C4 diet. Tissue 6t3C values were ranked hair > brain > muscle > liver > fat. After changing the gerbils to a wheat (C3) diet, isotope ratios of the tissues shifted in the direction of the 6a3C value of the new diet. The rate at which carbon derived from the corn diet was replaced by carbon derived from the wheat diet was ade- quately described by a negative exponential decay model for all tissues examined. More metabolically active tissues such as liver and fat had more rapid turnover rates than less metabolically active tissues such as hair. The half-life for carbon ranged from 6.4 days in liver to 47.5 days in hair. The results of this study have important implications for the use of 6~3C values as indicators of animal diet. Both fractionation and turnover of stable carbon isotopes in animal tissues may obscure the relative contributions of isotopically distinct dietary components (such as C3 vs. C,, or marine vs. terrestrial) if an animal's diet varies through time. These complications deserve attention in any study using stable isotope ratios of animal tissue as dietary indicators and might be minimized by analysis of several tissues or products covering a range of turnover times.

N. A. Slade

Papers

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

The alluvial deposits of lower wahweap creek: linking climate and incision records in the central colorado plateau