Ecological Studies Analysis and Synthesis Edited by W.D. Billings, Durham (USA) F. Golley, Athens (USA) O.L. Lange, Wilrzburg (FRG) 1.S. Olson, Oak Ridge (USA) H. Remmert, Marburg (FRG) Volume 68 Ecological Studies Volume 56 Volume 62 Resources and Society Frost Survival of Plants A Systems Ecology Study of the Island By A. Sakai and W. Larcher of Gotland, Sweden 1987. XII, 321p., 200 figures, 78 tables. cloth By James J. Zucchetto ISBN 0-387-17332-3 and Ann-Mari Jansson 1985. X, 248p., 70 figures. cloth Volume 63 Long-Term Forest Dynamics Volume 57 of the Temperate Zone Forest Ecosystems in the Alaskan Taiga By Paul A. Delcourt and Hazel R Delcourt A Synthesis of Structure and Function 1987. XIV, 45Op., 90 figures, 333 maps. cloth Edited by K. Van aeve, F.S. Chapin III, ISBN 0-387-96495-9 LA. Viereck, C.T. Dymess and P.W. Flanagan Volume 64 1986. X, 240p., 81 figures. cloth Land~ape Heterogeneity and Disturbance ISBN 0-387-96251-4 Edited by Monica Goigel Turner 1987. XII, 241p., 56 figures. cloth Volume 58 ISBN 0-387-96497-5 Ecology of Biological Invasions of North America and Hawaii Volume 65 Edited by HA. Mooney and JA. Drake Community Ecology of Sea Otters 1986. X, 32Op., 25 figures. cloth Edited by G.R van Blaricom and JA. Estes ISBN 0-387-96289-1 1987. X, 280p., 71 figures. cloth ISBN 3-540-18090-7 Volume 59 Acid Deposition and the Acidification Volume 66 of Soils and Waters Forest Hydrology and Ecology at Coweeta An Analysis Edited by W.T. Swank and DA. Crossley, Jr. By J.O. Reuss and D.W. Johnson 1987. XIV, 512p., 151 figures. cloth 1986. VIII, 12Op., 37 figures. cloth ISBN 0-387-96547-5 ISBN 0-387-96290-5 Volume 67 Volume 60 Concepts of Ecosystem Ecology Amazonian Rain Forests A Comparative View Ecosystem Disturbance and Recovery Edited by L.R Pomeroy Edited by Carl F. Jordan and JJ. Alberts 1987. X, 133p., 55 figures. cloth 1988. XII, 384p., 93 figures. cloth ISBN 0-387-96397-9 ISBN 0-387-96686-2 Volume 61 Volume 68 Potentials and Limitations Stable Isotopes in Ecological Research of Ecosystem Analysis Edited by P.W. Rundel, J.R Ehleringer Edited by E.-D. Schulze and H. Zwolfer and KA. Nagy 1987. XII, 435p., 141 figures. cloth 1989. XVI, 544p., 164 figures. cloth ISBN 0-387-17138-X ISBN 0-387-96712-5 P. W. Rundel J .R. Ehleringer K. A. Nagy Editors Stable Isotopes in Ecological Research With 164 Illustrations Springer-Verlag New York Berlin Heidelberg London Paris Tokyo P.W. Rundel l.R. Ehleringer K.A. Nagy Laboratory of Biomedical Department of Biology Laboratory of Biomedical and Environmental Sciences University of Utah and Environmental Sciences University of California Salt Lake City, UT 84112 University of California Los Angeles, CA 90024 USA Los Angeles, CA 90024 USA USA Library of Congress Cataloging-in-Publication Data Stable isotopes in ecological research / P.W. Rundel, J.R. Ehleringer, K.A. Nagy, editors. p. cm.- (Ecological studies; v. 68) Bibliography: p. Includes index. \. Stable isotopes in ecological research. I. Rundel, Philip W. (Philip Wilson) II. Ehleringer, J.R. III. Nagy, Kenneth A. IV. Series. QH541.15.S68S72 1988 574.5'072-dcl9 88-20171 Printed on acid-free paper. "'1989 by Springer-Verlag New York Inc. Softcover reprint of the hardcover 1st edition 1989 All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag, 175 Fifth Avenue, New York, NY 10010, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use of general descriptive names, trade names, trademarks, etc. in this publication, even if the former are not especially identified, is not to be taken as a sign that such names, as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Typeset by David Seham Associates, Metuchen, New Jersey. 9 8 7 6 5 4 3 2 I ISBN-13: 978-1-4612-8127-6 e-ISBN-13: 978-1-4612-3498-2 DOl: 10.1007/978-1-4612-3498-2 Preface In past years, ecologists and environmental biologists have largely equated the word "isotopes" with short-lived radioactive isotopes useful in tracer studies. However, virtually all of the elements of biological importance occur naturally with two or more stable isotopes. There are two stable isotopes of hydrogen, IH and 0 or deuterium. Similarly, there are two stable isotopes of carbon, two of nitrogen, three of oxygen, and four of sulfur. These stable isotopes, which vary in their ratios of natural abundance, offer tremendous potentials for new approaches to research on a wide range of ecological processes. Natural differences in the stable isotope composition of biological and abiotic compounds of ecological interest result from differences in a variety of pre dictable factors which influence fractionation. These include source effects, diffusional constraints, enzyme selectivity, and/or interactions between com pounds. Stable isotope investigations can thus provide new insights into flux rates among organisms, between organisms and their abiotic environment, and between compartments of the abiotic environment. Much of the early research on stable isotope ratios in biological tissues came from the geological sciences. This work was a logical outgrowth of pioneering applications of stable isotope analyses to research in geochemistry, sedimen tology, and oceanography. Applications of stable isotope analyses in environ mental biology were slow to develop, largely due to a combination of ignorance in the field and the difficulty of access to isotope ratio mass spectrometers. Until recently, applications of stable isotope ratios to ecological research were largely in the novelty or descriptive stage. In the last few years, however, in- vi Preface novative applications of stable isotope ratios to physiological and process-level studies have been expanding rapidly. This increased pace of research has re sulted from improved access by biologists to the necessary instrumentation, and reduced costs of analyses. As a result, there is every indication that stable isotope approaches to ecological research will become increasingly common and will lead to significant new levels of understanding of physiological processes and elemental fluxes through biological and abiotic systems. Our objective in this volume has been to provide both general background information and illustrative case studies to demonstrate how differences in stable isotope composition can be used as powerful tools for measuring integrated physiological responses and elemental fluxes through both abiotic and biotic compartments of natural ecosystems. Following an introductory chapter of background material on the history, units, and instrumentation used in stable isotope research, we have divided our book into three parts. The first of these deals with ecophysiological studies in plants, focusing on approaches to utilizing isotopes of carbon, hydrogen, and oxygen in research on physiological processes in plants. In the second section on animal food webs and feeding ecology, seven chap ters describe ways in which stable isotope ratios can be used to study food web dynamics. These contributions add to our understanding of the principles that "we are what we eat, plus or minus a few parts per million" as originally dem onstrated by Michael DeNiro and Samuel Epstein. Our focus in this section is on natural food chains, with only a single chapter on human diet analysis. An thropologists and archaeologists, nevertheless, should find this section, as well as other parts of the volume, highly relevant to the rapidly expanding interest in stable isotope approaches to their fields. The final section of eleven chapters broadly treats ecosystem process studies utilizing stable isotope ratios. Aspects of pedogenic processes, nitrogen fixation, paleoclimate, atmospheric fluxes of gases and particulates, and pollutant trans fers are all included. While these chapters only highlight the range of potential applications of stable isotope ratios to investigations of ecosystem-level fluxes, they should provide a good appreciation of the scope of the promise that such applications offer to researchers. Stable Isotopes in Ecological Research is an outgrowth of a workshop on this subject held at the Lake Arrowhead Conference of the University of Cal ifornia, Los Angeles, in April 1986. We are indebted to Drs. Helen McCammon and Janet Dorigan of the Ecological Research Division of the Office of Health and Environmental Research, U. S. Department of Energy, for sponsoring this workshop. Cosponsorship was provided by the National Center for Intermedia Transport Research at UCLA. P.W. Rundel J .R. Ehleringer K.A. Nagy Contents Preface V Contributors XI 1. Stable Isotopes: History, Units, and Instrumentation l.R. EHLERINGER and P.W. RUNDEL Section I Ecophysiological Studies in Plants 17 2. Carbon Isotope Fractionation and Plant Water-Use Efficiency 21 G.D. FARQUHAR, K.T. HUBICK, A.G. CONDON, and R.A. RICHARDS 3. Carbon Isotope Ratios and Physiological Processes in Aridland Plants 41 l.R. EHLERINGER 4. Stable Carbon Isotope Ratio as an Index of Water-Use Efficiency in C Halophytes-Possible Relationship to Strategies 3 for Osmotic Adjustment 55 R.D. GUY, P.G. WARNE, and D.M. REID viii Contents 5. Stable Carbon Isotopes in Vernal Pool Aquatics of Differing Photosynthetic Pathways 76 J.E. KEELEY 6. Studies of Mechanisms Affecting the Fractionation of Carbon Isotopes in Photosynthesis 82 J.A. BERRY c 7. Intertree Variability on,13 in Tree Rings 95 S.W. LEAVITT and A. LONG 8. Hydrogen Isotope Fractionation in Plant Tissues 105 H. ZIEGLER 9. Oxygen and Hydrogen Isotope Ratios in Plant Cellulose: Mechanisms and Applications 124 L. DA SILVEIRA LOBO STERNBERG 10. Stable Hydrogen Isotope Ratios in Plants: A Review of Current Theory and Some Potential Applications 142 J.W.c. WHITE Section II Animal Food Webs and Feeding Ecology 163 11. Stable Carbon Isotopes in Terrestrial Ecosystem Research 167 L.L. TIEszEN and T. W. BOUTTON 12. Ol3C Measurements as Indicators of Carbon Flow in Marine and Freshwater Ecosystems 196 B. FRY and E.B. SHERR 13. Natural Carbon Isotope Tracers in Arctic Aquatic Food Webs 230 D.M. SCHELL and P.J. ZIEMANN 14. Some Problems and Potentials of Strontium Isotope Analysis for Human and Animal Ecology 252 J.E. ERICSON 15. Natural Isotope Abundances in Bowhead Whale (Balaena mysticetus) Baleen: Markers of Aging and Habitat Usage 260 D.M. SCHELL, S.M. SAUPE, and N. HAUBENSTOCK 16. Doubly-Labeled Water Studies of Vertebrate Physiological Ecology 270 K.A. NAGY Contents ix 17. A Ol3C and o15N Tracer Study of Nutrition in Aquaculture: Penaeus vannamei in a Pond Growout System 288 P.L. PARKER, R.K. ANDERSON, and A. LAWRENCE Section III Ecosystem Process Studies 305 18. Stable Isotope Ratios and the Dynamics of Caliche in Desert Soils 309 W.H. SCHLESINGER, G.M. MARION, and P.J. FONTEYN 19. The Use of Stable Isotopes in Assessing the Effect of Agriculture on Arid and Semi-Arid Soils 318 R. AMUNDSON 20. Estimates of N2 Fixation in Ecosystems: The Need for and Basis of the 15N Natural Abundance Method 342 G. SHEARER and D.H. KOHL 21. The Use of Variation in the Natural Abundance of 15N to Assess Symbiotic Nitrogen Fixation by Woody Plants 375 R.A. VIRGINIA, W.M. JARRELL, P.W. RUNDEL, G. SHEARER, and D.H. KOHL 22. l3C/l2C Ratios in Atmospheric Methane and Some of Its Sources 395 S.c. TYLER 23. Temperature-Dependent Hydrogen Isotope Fractionation in Cyanobacterial Sheaths: Applications to Studies of Modern and Precambrian Stromatolites 410 G.E. STRATHEARN 24. Sulfur Isotope Studies of the Pedosphere and Biosphere 424 H.R. KROUSE 25. Sulfate Fertilization and Changes in Stable Sulfur Isotopic Compositions of Lake Sediments 445 B. FRY 26. The Use of Stable Sulfur and Nitrogen Isotopes in Studies of Plant Responses to Air Pollution 454 W.E. WINNER, V.S. BERG, and P.J. LANGSTON-UNKEFER 27. The Use of Stable Sulfur Isotope Ratios in Air Pollution Studies: An Ecosystem Approach in South Florida 471 L.L. JACKSON and L.P. GOUGH x Contents 28. 87Srf"Sr Ratios Measure the Sources and Flow of Strontium in Terrestrial Ecosystems 491 w.e. GRAUSTEIN Index 513