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

Phytoplankton blooms are prominent features of biological variability in shallow coastal ecosystems such as estuaries, lagoons, bays, and tidal rivers. Long-term observation and research in San Francisco Bay illustrates some patterns of phytoplankton spatial and temporal variability and the underlying mechanisms of this variability. Blooms are events of rapid production and accumulation of phytoplankton biomass that are usually responses to changing physical forcings originating in the coastal ocean (e.g., tides), the atmosphere (wind), or on the land surface (precipitation and river runoff). These physical forcings have different timescales of variability, so algal blooms can be short-term episodic events, recurrent seasonal phenomena, or rare events associated with exceptional climatic or hydrologic conditions. The biogeochemical role of phytoplankton primary production is to transform and incorporate reactive inorganic elements into organic forms, and these transformations are rapid and lead to measurable geochemical change during blooms. Examples include the depletion of inorganic nutrients (N, P, Si), supersaturation of oxygen and removal of carbon dioxide, shifts in the isotopic composition of reactive elements (C, N), production of climatically active trace gases (methyl bromide, dimethylsulfide), changes in the chemical form and toxicity of trace metals (As, Cd, Ni, Zn), changes in the biochemical composition and reactivity of the suspended particulate matter, and synthesis of organic matter required for the reproduction and growth of heterotrophs, including bacteria, zooplankton, and benthic consumer animals. Some classes of phytoplankton play special roles in the cycling of elements or synthesis of specific organic molecules, but we have only rudimentary understanding of the forces that select for and promote blooms of these species. Mounting evidence suggests that the natural cycles of bloom variability are being altered on a global scale by human activities including the input of toxic contaminants and nutrients, manipulation of river flows, and translocation of species. This hypothesis will be a key component of our effort to understand global change at the land-sea interface. Pursuit of this hypothesis will require creative approaches for distinguishing natural and anthropogenic sources of phytoplankton population variability, as well as recognition that the modes of human disturbance of coastal bloom cycles operate interactively and cannot be studied as isolated processes.

Phytoplankton bloom dynamics in coastal ecosystems: A review with some general lessons from sustained investigation of San Francisco Bay, California

We report measurements of seasonal variability in the C-N stable isotope ratios of plants collected across the habitat mosaic of San Francisco Bay, its marshes, and its tributary river system. Analyses of 868 plant samples were binned into 10 groups (e.g., terrestrial riparian, freshwater phytoplankton, salt marsh) to determine whether C-N isotopes can be used as biomarkers for tracing the origins of organic matter in this river‐marsh‐estuary complex. Variability of d 13 C and d 15 N was high (;5‐10‰) within each plant group, and we identified three modes of variability: (1) between species and their microhabitats, (2) over annual cycles of plant growth and senescence, and (3) between living and decomposing biomass. These modes of within-group variability obscure any sourcespecific isotopic signatures, confounding the application of C-N isotopes for identifying the origins of organic matter. A second confounding factor was large dissimilarity between the d 13 C-d 15 N of primary producers and the organicmatter pools in the seston and sediments. Both confounding factors impede the application of C-N isotopes to reveal the food supply to primary consumers in ecosystems supporting diverse autotrophs and where the isotopic composition of organic matter has been transformed and become distinct from that of its parent plant sources. Our results support the advice of others: variability of C-N stable isotopes within all organic-matter pools is high and must be considered in applications of these isotopes to trace trophic linkages from primary producers to primary consumers. Isotope-based approaches are perhaps most powerful when used to complement other tools, such as molecular biomarkers, bioassays, direct measures of production, and compilations of organic-matter budgets.

/pdf/stable-carbon-and-nitrogen-isotope-composition-of-aquatic-39b36tyw9n.pdf

Stable carbon and nitrogen isotope composition of aquatic and terrestrial plants of the San Francisco Bay estuarine system

Fatty acid (FA) analysis is a well- established tool for studying trophic interactions in marine habitats. However, its application to benthic food webs poses 2 particular challenges. First, unlike pelagic zooplankton, benthic con- sumers have access to different sizes and func- tional groups of primary producers and may con- sume a highly mixed diet. Classes of benthic primary producers are distinct in their overall FA composition, but most do not possess unique marker FAs that can be used to identify their con- tribution to higher trophic levels. Second, unlike mammalian predators, benthic invertebrates have the capacity to significantly modify their dietary FAs and thereby obscure markers for food sources. Controlled feeding studies have been used to dis- tinguish dietary tracer FAs from those that are modified by the consumer in several benthic invertebrates, but more such studies are needed. Despite these challenges, FAs have been used to study trophic structure in a variety of benthic habitats including the deep sea, polar regions, estuaries, and the rocky subtidal zone. However, the complexity of benthic food webs and lack of unique markers impose uncertainties in the inter- pretation of FA data from field studies. Multivari- ate analyses are necessary for analyzing FA datasets, although univariate tests can be useful for comparing levels of informative FAs among food sources and consumers. Combining FA analysis with other lines of evidence, such as sta- ble isotope analysis, offers a more reliable approach to examining trophic interactions in benthic systems.

/pdf/fatty-acids-as-dietary-tracers-in-benthic-food-webs-1jo4oyc6rt.pdf

Fatty acids as dietary tracers in benthic food webs

Use of 14C and 13C natural abundances for evaluating riverine, estuarine, and coastal DOC and POC sources and cycling: a review and synthesis

Multiple indicators (Chl a, C : N ratios, [6L3C]POC, and two classes of lipid biomarker compounds- sterols and phospholipid ester-linked fatty acids) were used to evaluate spatial and temporal variations in the origin of particulate organic matter (POM) in the San Francisco Bay (SFB) estuary. Comparisons were made between the northern and southern subestuaries of SFB, as well as along the salinity gradient of northern SFB. Two sample types were collected-seston, which was used to characterize the bulk POM, and tissues of the suspension-feeding bivalve Potamocorbula amurensis -in order to evaluate the assimilable portion of the POM. Samples were collected around biological and physical events (phytoplankton blooms and freshwater inflow) thought to be the primary mechanisms controlling temporal variability in organic matter sources. Seston samples indicate that phytoplankton sources of POM are important throughout the entire SFB system, with additional inputs of organic matter from bacterial and terrestrial vascular plant sources delivered to the northern region. Analysis of biomarker compounds in- P. amurensis tissues indicates that phytoplankton supply a large fraction of the assimilable carbon to clams throughout SFB, although isotopic analysis of clam tissues suggests that the origin of this reactive carbon varies spatially and that freshwater algae are an important source of reactive organic matter to clams living in northern SFB.

https://aslopubs.onlinelibrary.wiley.com/doi/pdf/10.4319/lo.1995.40.1.0067

James B. Guckert

Papers

Molecular and isotopic tracers used to examine sources of organic matter and its incorporation into the food webs of San Francisco Bay