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

Sea temperatures in many tropical regions have increased by almost 1 degrees C over the past 100 years, and are currently increasing at similar to 1-2 degrees C per century. Coral bleaching occurs when the thermal tolerance of corals and their photosynthetic symbionts (zooxanthellae) is exceeded. Mass coral bleaching has occurred in association with episodes of elevated sea temperatures over the past 20 years and involves the loss of the zooxanthellae following chronic photoinhibition. Mass bleaching has resulted in significant losses of live coral in many parts of the world. This paper considers the biochemical, physiological and ecological perspectives of coral bleaching. It also uses the outputs of four runs from three models of global climate change which simulate changes in sea temperature and hence how the frequency and intensity of bleaching events will change over the next 100 years. The results suggest that the thermal tolerances of reef-building corals are likely to be exceeded every year within the next few decades. Events as severe as the 1998 event, the worst on record, are likely to become commonplace within 20 years. Most information suggests that the capacity for acclimation by corals has already been exceeded, and that adaptation will be too slow to avert a decline in the quality of the world's reefs. The rapidity of the changes that are predicted indicates a major problem for tropical marine ecosystems and suggests that unrestrained warming cannot occur without the loss and degradation of coral reefs on a global scale.

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Climate change, coral bleaching and the future of the world's coral reefs

Rising atmospheric carbon dioxide (CO2), primarily from human fossil fuel combustion, reduces ocean pH and causes wholesale shifts in seawater carbonate chemistry. The process of ocean acidification is well documented in field data, and the rate will accelerate over this century unless future CO2 emissions are curbed dramatically. Acidification alters seawater chemical speciation and biogeochemical cycles of many elements and compounds. One well-known effect is the lowering of calcium carbonate saturation states, which impacts shell-forming marine organisms from plankton to benthic molluscs, echinoderms, and corals. Many calcifying species exhibit reduced calcification and growth rates in laboratory experiments under high-CO2 conditions. Ocean acidification also causes an increase in carbon fixation rates in some photosynthetic organisms (both calcifying and noncalcifying). The potential for marine organisms to adapt to increasing CO2 and broader implications for ocean ecosystems are not well known; both are high priorities for future research. Although ocean pH has varied in the geological past, paleo-events may be only imperfect analogs to current conditions.

/pdf/ocean-acidification-the-other-co-2-problem-4h18mw2cpj.pdf

Ocean Acidification: The Other CO 2 Problem

Couplings Between Changes in the Climate System and Biogeochemistry Coordinating Lead Authors: Kenneth L. Denman (Canada), Guy Brasseur (USA, Germany) Lead Authors: Amnat Chidthaisong (Thailand), Philippe Ciais (France), Peter M. Cox (UK), Robert E. Dickinson (USA), Didier Hauglustaine (France), Christoph Heinze (Norway, Germany), Elisabeth Holland (USA), Daniel Jacob (USA, France), Ulrike Lohmann (Switzerland), Srikanthan Ramachandran (India), Pedro Leite da Silva Dias (Brazil), Steven C. Wofsy (USA), Xiaoye Zhang (China) Contributing Authors: D. Archer (USA), V. Arora (Canada), J. Austin (USA), D. Baker (USA), J.A. Berry (USA), R. Betts (UK), G. Bonan (USA), P. Bousquet (France), J. Canadell (Australia), J. Christian (Canada), D.A. Clark (USA), M. Dameris (Germany), F. Dentener (EU), D. Easterling (USA), V. Eyring (Germany), J. Feichter (Germany), P. Friedlingstein (France, Belgium), I. Fung (USA), S. Fuzzi (Italy), S. Gong (Canada), N. Gruber (USA, Switzerland), A. Guenther (USA), K. Gurney (USA), A. Henderson-Sellers (Switzerland), J. House (UK), A. Jones (UK), C. Jones (UK), B. Karcher (Germany), M. Kawamiya (Japan), K. Lassey (New Zealand), C. Le Quere (UK, France, Canada), C. Leck (Sweden), J. Lee-Taylor (USA, UK), Y. Malhi (UK), K. Masarie (USA), G. McFiggans (UK), S. Menon (USA), J.B. Miller (USA), P. Peylin (France), A. Pitman (Australia), J. Quaas (Germany), M. Raupach (Australia), P. Rayner (France), G. Rehder (Germany), U. Riebesell (Germany), C. Rodenbeck (Germany), L. Rotstayn (Australia), N. Roulet (Canada), C. Sabine (USA), M.G. Schultz (Germany), M. Schulz (France, Germany), S.E. Schwartz (USA), W. Steffen (Australia), D. Stevenson (UK), Y. Tian (USA, China), K.E. Trenberth (USA), T. Van Noije (Netherlands), O. Wild (Japan, UK), T. Zhang (USA, China), L. Zhou (USA, China) Review Editors: Kansri Boonpragob (Thailand), Martin Heimann (Germany, Switzerland), Mario Molina (USA, Mexico) This chapter should be cited as: Denman, K.L., G. Brasseur, A. Chidthaisong, P. Ciais, P.M. Cox, R.E. Dickinson, D. Hauglustaine, C. Heinze, E. Holland, D. Jacob, U. Lohmann, S Ramachandran, P.L. da Silva Dias, S.C. Wofsy and X. Zhang, 2007: Couplings Between Changes in the Climate System and Biogeochemistry. In: Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change [Solomon, S., D. Qin, M. Manning, Z. Chen, M. Marquis, K.B. Averyt, M.Tignor and H.L. Miller (eds.)]. Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA.

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Couplings between changes in the climate system and biogeochemistry

In marine ecosystems, rising atmospheric CO2 and climate change are associated with concurrent shifts in temperature, circulation, stratification, nutrient input, oxygen content, and ocean acidification, with potentially wideranging biological effects. Population-level shifts are occurring because of physiological intolerance to new environments, altered dispersal patterns, and changes in species interactions. Together with local climate-driven invasion and extinction, these processes result in altered community structure and diversity, including possible emergence of novel ecosystems. Impacts are particularly striking for the poles and the tropics, because of the sensitivity of polar ecosystems to sea-ice retreat and poleward species migrations as well as the sensitivity of coral-algal symbiosis to minor increases in temperature. Midlatitude upwelling systems, like the California Current, exhibit strong linkages between climate and species distributions, phenology, and demography. Aggregated effects may modify energy and material flows as well as biogeochemical cycles, eventually impacting the overall ecosystem functioning and services upon which people and societies depend.

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Climate Change Impacts on Marine Ecosystems

SYNOPSIS. Photosynthesis and calcification in zooxanthellate scleractinian corals and coral reefs are reviewed at several scales: cellular (pathways and transport mechanisms of inorganic carbon and calcium), organismal (interaction between photosynthesis and calcification, effect of light) and ecosystemic (community primary production and calcification, and air-sea CO2 exchanges).

The coral host plays a major role in supplying carbon for the photosynthesis by the algal symbionts through a system similar to the carbon-concentrating mechanism described in free living algal cells. The details of carbon supply to the calcification process are almost unknown, but metabolic CO2 seems to be a significant source. Calcium supply for calcification is diffusional through oral layers, and active membrane transport only occurs between the calicoblastic cells and the site of calcification. Photosynthesis and calcification are tightly coupled in zooxanthellate scleractinian corals and coral reef communities. Calcification is, on average, three times higher in light than in darkness. The recent suggestion that calcification is dark-repressed rather than light-enhanced is not supported by the literature. There is a very strong correlation between photosynthesis and calcification at both the organism and community levels, but the ratios of calcification to gross photosynthesis (0.6 in corals and 0.2 in reef communities) differ from unity, and from each other as a function of level.

The potential effect of global climatic changes ( p CO2 and temperature) on the rate of calcification is also reviewed. In various calcifying photosynthetic organisms and communities, the rate of calcification decreases as a function of increasing p CO2 and decreasing calcium carbonate saturation state. The calculated decrease in CaCO3, production, estimated using the scenarios considered by the International Panel on Climate Change (IPCC), is 10% between 1880 and 1990, and 9–30% (mid estimate: 22%) from 1990 to 2100. Inadequate understanding of the mechanism of calcification and its interaction with photosynthesis severely limits the ability to provide an accurate prediction of future changes in the rate of calcification.

/pdf/photosynthesis-and-calcification-at-cellular-organismal-and-45k4v0r6ef.pdf

Photosynthesis and Calcification at Cellular, Organismal and Community Levels in Coral Reefs: A Review on Interactions and Control by Carbonate Chemistry

SUMMARY The symbiosis between cnidarians (e.g., corals or sea anemones) and intracellular dinoflagellate algae of the genus Symbiodinium is of immense ecological importance. In particular, this symbiosis promotes the growth and survival of reef corals in nutrient-poor tropical waters; indeed, coral reefs could not exist without this symbiosis. However, our fundamental understanding of the cnidarian-dinoflagellate symbiosis and of its links to coral calcification remains poor. Here we review what we currently know about the cell biology of cnidarian-dinoflagellate symbiosis. In doing so, we aim to refocus attention on fundamental cellular aspects that have been somewhat neglected since the early to mid-1980s, when a more ecological approach began to dominate. We review the four major processes that we believe underlie the various phases of establishment and persistence in the cnidarian/coral-dinoflagellate symbiosis: (i) recognition and phagocytosis, (ii) regulation of host-symbiont biomass, (iii) metabolic exchange and nutrient trafficking, and (iv) calcification. Where appropriate, we draw upon examples from a range of cnidarian-alga symbioses, including the symbiosis between green Hydra and its intracellular chlorophyte symbiont, which has considerable potential to inform our understanding of the cnidarian-dinoflagellate symbiosis. Ultimately, we provide a comprehensive overview of the history of the field, its current status, and where it should be going in the future.

https://ir.library.oregonstate.edu/downloads/2z10wr382

Cell Biology of Cnidarian-Dinoflagellate Symbiosis

NOTICE from NSF Any opinions, findings, and conclusions or recommendations expressed in this material are those of the authors and do not necessarily reflect the views of the National Science Foundation. NOTICE from NOAA Mention of a commercial company or product does not constitute an endorsement by NOAA/OAR. Use of information from this publication concerning proprietary products or the tests of such products for publicity or advertising purposes is not authorized. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the authors and do not necessarily reflect the views of the National Oceanic and Atmospheric Administration. NOTICE from USGS Reference herein to any specific commercial product, process, or service by trade name, trademark, manufacturer , or otherwise does not necessarily constitute or imply its endorsement, recommendation, or favoring by the United States Government or any agency thereof. Any views and opinions of authors expressed herein do not necessarily state or reflect those of the United States Government or any agency thereof. Cover design by Cheryl Markel (NCAR). Photo credits: clockwise form upper left: reef scene from NOAA's photo library (credit: Florida Keys National Marine Sanctuary); upper right: euthecosomatous pteropod Cavolinia tridentata (V. Fabry); lower right: coral Eusmilia fastigata from NOAA's photo library (credit: Florida Keys National Marine Sanctuary); coccol-ithophore Emiliania huxleyi (V. Fabry). Studies, who hosted the meeting. Special thanks goes to Ryan Layne Whitney of NOAA/PMEL for his excellent editing and formatting skills, to Janice Lough for her diligent attention to detail, and to Rebecca Haacker-Santos and Sandra Coffman for their hard work to ensure a well-executed meeting. We also acknowledge all of the workshop participants, who canceled classes, cooperated and cajoled, and creatively carried on with the many questions put to them in tackling the important questions about how the marine calcium carbonate system, and marine calcifiers, will respond to increasing concentrations of carbon dioxide.

Impacts of ocean acidification on coral reefs and other marine calcifiers : a guide for future research

In spite of more than one century and half of studies, mechanisms of coral biomineralization, leading to coral growth and reef formation, still remain poorly known, although major global threats to coral reefs, such as ocean acidification, primarily affect this process. Coral skeletons are used as environmental archives but the vital processes that govern incorporation of trace elements and stable isotope are still unknown. Our knowledge on coral physiology is restricted to the organismal level due to the lack of appropriate cell model, however the advent of new approaches, such as coral genomic, is changing drastically our knowledge on these animals even if only a few data are available concerning the field of biomineralization. This chapter reviews our present knowledge and discusses the different theories on coral calcification, from the molecular to the reef level. Conclusion is presented in a list of key issues to be resolved in order to understand the intimate mechanisms of calcification of corals, essential to determine the origin of the sensitivity of corals to ocean acidification, to improve paleoceanographic reconstructions or coral reef management, or “just” to understand how genes of a soft organism control the formation of an extracellular 3D-skeleton.

Denis Allemand

Papers

Photosynthesis and Calcification at Cellular, Organismal and Community Levels in Coral Reefs: A Review on Interactions and Control by Carbonate Chemistry

Cell Biology of Cnidarian-Dinoflagellate Symbiosis

Impacts of ocean acidification on coral reefs and other marine calcifiers : a guide for future research

Coral Calcification, Cells to Reefs

Coral biomineralization: From the gene to the environment