Robert W. Sterner is Professor and Head of the Department of Ecology, Evolution, and Behavior at the University of Minnesota. James J. Elser is Professor of Biology at Arizona State University. Peter Vitousek, one of the world's leading ecosystem ecologists, is Morrison Professor of Population and Resources at Stanford University.

"Few, if any, details of stoichiometry seem to have been overlooked by Sterner and Elser, and their book will be a useful reference to me for many years to come... The hundreds of references in the bibliography are worth the price of the book alone."--David W. Schindler, Nature "Robert Sterner and James Elser take a giant stride in knitting together perspectives across scales, biomes, and disciplines to craft an integrative and predictive vision of the topic... It knits together such a broad range of relevant topics that anyone interested in the connections between biology and elemental cycles should give it a try."--Anthony F. Michaels, Science "I believe that his is one of the most important books written in ecology in the last 10 years... I predict that in a few years it will be inconceivable to ignore the stoichiometric perspective when tackling an ecological problem... Ecological Stoichiometry is well written in colloquial and friendly prose. The authors strived to explain their arguments clearly and in detail. The many mathematical models are explained with laudable lucidity and the figures that illustrate them are consistently good."--Carlos Martinez Del Rio, Ecology "How often do you read a book that has a large number of 'aha!' moments in every chapter? This is a significant piece of synthesis and scholarship that brings together a very large number of disciplines and disparate chunks of data into a very satisfying whole... Never before have I seen a book which spans the scales from molecular biology to ecosystems so effectively... It is sure to become a classic."--Graham Harris, Journal of Plankton Research "Sterner and Elser adopt a simple, incremental logic in the book and follow it with fidelity... The book is an excellent introduction to ecological stoichiometry for graduate students; and for those already in the field, it is a thorough review of the complexities and nuances of stoichiometric ecology by two of its best practitioners."--Robert E. Hecky, Limnology and Oceanography Bulletin "A fresh and stimulating perspective on this area of ecology. The clear, readable style renders this text highly accessible."--Bulletin of the British Ecological Society "[This] book is seminal and presents a clear, well-reasoned overview of ES, thereby providing the biological community with an essential piece to the puzzle of integrating micro- and macro-level biology."--Van Savage, Complexity

• List of Figures ix List of Tables xiii Foreword xv Preface xvii 1. Stoichiometry and Homeostasis 1 Scope 3 Stoichiometry and Homeostasis 8 Yield 25 The Redfield Ratio 27 Conventions and Concerns about Element Ratios 31 Some Conventions about Growth Rate 34 A Logical Framework 35 The Structure of This Book 40 Summary and Synthesis 41 Key Definitions 42 2. Biological Chemistry: Building Cells from Elements 44 The Basis for Selection of Carbon, Nitrogen, and Phosphorus in Biochemical Evolution 45 The Elemental Composition of Major Biochemicals 51 Cell Components: The Elemental Composition of Cellular Structures 66 Summary and Synthesis 78 3. The Stoichiometry of Autotroph Growth: Variation at the Base of Food Webs 80 Cellular and Physiological Bases 81 C:N:P Stoichiometry of Entire Higher Plants 87 Autotrophs in Captivity 89 Theories of Autotroph Stoichiometry 107 Autotrophs in the Wild: Oceans, Lakes, and Land 120 Causes of Variation in Autotroph C:N:P in Nature 127 Catalysts for Ecological Stoichiometry 132 Summary and Synthesis 133 4. How to Build an Animal: The Stoichiometry of Metazoans 135 Biochemical and Biological Determinants of Body Elemental Composition 136 Invertebrate Stoichiometry: C:N:P in Zooplankton and Insects 138 Determinants of C:N:P in Invertebrates: The Growth Rate Hypothesis 142 Molecular Biology and the C:N:P Stoichiometry of Growth, or Ecosystem Scientists Go Astray 150 A Simple Molecular-Kinetic Model of the Growth Rate-C:N:P Connection 160 Structural Investment and the Stoichiometry of Vertebrates 168 Elemental Composition and Body Size 171 Catalysts for Ecological Stoichiometry 175 Summary and Synthesis 178 5. Imbalanced Resources and Animal Growth 179 Mass Balance in Growth Processes 180 Maximizing Yield in Chemistry and in Ecology 185 Limiting Factors for Heterotroph Growth: Development of Threshold Element Ratio Theory 189 A New Minimal Model of the Stoichiometry of Secondary Production 197 Some Real World Problems in Stoichiometric Balance 205 Growth Efficiency 222 Catalysts for Ecological Stoichiometry 227 Summary and Synthesis 229 6. The Stoichiometry of Consumer-Driven Nutrient Recycling 231 A Brief History of Studies of Consumer-Driven Nutrient Recycling 232 Stoichiometric Theories of Consumer-Driven Nutrient Recycling 235 Evidence That Consumers Differentially Recycle Nitrogen and Phosphorus 245 Microbial Mineralization 249 The Stoichiometry of Consumer-Driven Nutrient Recycling by Vertebrates 252 Catalysts for Ecological Stoichiometry 259 Summary and Synthesis 260 7. Stoichiometry in Communities: Dynamics and Interactions 262 Species Interactions 264 Positive Feedbacks and Multiple Stable States 277 Trophic Cascades 291 Light: Nutrient Effects at the Community Level 298 Feedbacks Owing to the "Constraints of Stuff": C:N Ratios in Tall-Grass Prairie 307 Catalysts for Ecological Stoichiometry 308 Summary and Synthesis 310 8. Big-Scale Stoichiometry: Ecosystems in Space and Time 313 Empirical Patterns in Ecosystem Stoichiometry 315 Linkages in the Stoichiometry of Biomass Yield: Using One Substance to Obtain Another 336 Nutrient Use Efficiency at the Ecosystem Level 341 The Stoichiometry of Food-Chain Production: A New Term, Carbon Use Efficiency 348 The Fate of Primary Production 350 Global Change 354 Catalysts for Ecological Stoichiometry 364 Summary and Synthesis 366 9. Recapitulation and Integration 370 Recapitulation 370 Integration: Toward a Biological Stoichiometry of Living Systems 376 Appendix 382 Literature Cited 385 Index 431

Robert W. Sterner is Professor and Head of the Department of Ecology, Evolution, and Behavior at the University of Minnesota. James J. Elser is Professor of Biology at Arizona State University. Peter Vitousek, one of the world's leading ecosystem ecologists, is Morrison Professor of Population and Resources at Stanford University.

"Few, if any, details of stoichiometry seem to have been overlooked by Sterner and Elser, and their book will be a useful reference to me for many years to come... The hundreds of references in the bibliography are worth the price of the book alone."--David W. Schindler, Nature "Robert Sterner and James Elser take a giant stride in knitting together perspectives across scales, biomes, and disciplines to craft an integrative and predictive vision of the topic... It knits together such a broad range of relevant topics that anyone interested in the connections between biology and elemental cycles should give it a try."--Anthony F. Michaels, Science "I believe that his is one of the most important books written in ecology in the last 10 years... I predict that in a few years it will be inconceivable to ignore the stoichiometric perspective when tackling an ecological problem... Ecological Stoichiometry is well written in colloquial and friendly prose. The authors strived to explain their arguments clearly and in detail. The many mathematical models are explained with laudable lucidity and the figures that illustrate them are consistently good."--Carlos Martinez Del Rio, Ecology "How often do you read a book that has a large number of 'aha!' moments in every chapter? This is a significant piece of synthesis and scholarship that brings together a very large number of disciplines and disparate chunks of data into a very satisfying whole... Never before have I seen a book which spans the scales from molecular biology to ecosystems so effectively... It is sure to become a classic."--Graham Harris, Journal of Plankton Research "Sterner and Elser adopt a simple, incremental logic in the book and follow it with fidelity... The book is an excellent introduction to ecological stoichiometry for graduate students; and for those already in the field, it is a thorough review of the complexities and nuances of stoichiometric ecology by two of its best practitioners."--Robert E. Hecky, Limnology and Oceanography Bulletin "A fresh and stimulating perspective on this area of ecology. The clear, readable style renders this text highly accessible."--Bulletin of the British Ecological Society "[This] book is seminal and presents a clear, well-reasoned overview of ES, thereby providing the biological community with an essential piece to the puzzle of integrating micro- and macro-level biology."--Van Savage, Complexity

• List of Figures ix List of Tables xiii Foreword xv Preface xvii 1. Stoichiometry and Homeostasis 1 Scope 3 Stoichiometry and Homeostasis 8 Yield 25 The Redfield Ratio 27 Conventions and Concerns about Element Ratios 31 Some Conventions about Growth Rate 34 A Logical Framework 35 The Structure of This Book 40 Summary and Synthesis 41 Key Definitions 42 2. Biological Chemistry: Building Cells from Elements 44 The Basis for Selection of Carbon, Nitrogen, and Phosphorus in Biochemical Evolution 45 The Elemental Composition of Major Biochemicals 51 Cell Components: The Elemental Composition of Cellular Structures 66 Summary and Synthesis 78 3. The Stoichiometry of Autotroph Growth: Variation at the Base of Food Webs 80 Cellular and Physiological Bases 81 C:N:P Stoichiometry of Entire Higher Plants 87 Autotrophs in Captivity 89 Theories of Autotroph Stoichiometry 107 Autotrophs in the Wild: Oceans, Lakes, and Land 120 Causes of Variation in Autotroph C:N:P in Nature 127 Catalysts for Ecological Stoichiometry 132 Summary and Synthesis 133 4. How to Build an Animal: The Stoichiometry of Metazoans 135 Biochemical and Biological Determinants of Body Elemental Composition 136 Invertebrate Stoichiometry: C:N:P in Zooplankton and Insects 138 Determinants of C:N:P in Invertebrates: The Growth Rate Hypothesis 142 Molecular Biology and the C:N:P Stoichiometry of Growth, or Ecosystem Scientists Go Astray 150 A Simple Molecular-Kinetic Model of the Growth Rate-C:N:P Connection 160 Structural Investment and the Stoichiometry of Vertebrates 168 Elemental Composition and Body Size 171 Catalysts for Ecological Stoichiometry 175 Summary and Synthesis 178 5. Imbalanced Resources and Animal Growth 179 Mass Balance in Growth Processes 180 Maximizing Yield in Chemistry and in Ecology 185 Limiting Factors for Heterotroph Growth: Development of Threshold Element Ratio Theory 189 A New Minimal Model of the Stoichiometry of Secondary Production 197 Some Real World Problems in Stoichiometric Balance 205 Growth Efficiency 222 Catalysts for Ecological Stoichiometry 227 Summary and Synthesis 229 6. The Stoichiometry of Consumer-Driven Nutrient Recycling 231 A Brief History of Studies of Consumer-Driven Nutrient Recycling 232 Stoichiometric Theories of Consumer-Driven Nutrient Recycling 235 Evidence That Consumers Differentially Recycle Nitrogen and Phosphorus 245 Microbial Mineralization 249 The Stoichiometry of Consumer-Driven Nutrient Recycling by Vertebrates 252 Catalysts for Ecological Stoichiometry 259 Summary and Synthesis 260 7. Stoichiometry in Communities: Dynamics and Interactions 262 Species Interactions 264 Positive Feedbacks and Multiple Stable States 277 Trophic Cascades 291 Light: Nutrient Effects at the Community Level 298 Feedbacks Owing to the "Constraints of Stuff": C:N Ratios in Tall-Grass Prairie 307 Catalysts for Ecological Stoichiometry 308 Summary and Synthesis 310 8. Big-Scale Stoichiometry: Ecosystems in Space and Time 313 Empirical Patterns in Ecosystem Stoichiometry 315 Linkages in the Stoichiometry of Biomass Yield: Using One Substance to Obtain Another 336 Nutrient Use Efficiency at the Ecosystem Level 341 The Stoichiometry of Food-Chain Production: A New Term, Carbon Use Efficiency 348 The Fate of Primary Production 350 Global Change 354 Catalysts for Ecological Stoichiometry 364 Summary and Synthesis 366 9. Recapitulation and Integration 370 Recapitulation 370 Integration: Toward a Biological Stoichiometry of Living Systems 376 Appendix 382 Literature Cited 385 Index 431

Robert W. Sterner is Professor and Head of the Department of Ecology, Evolution, and Behavior at the University of Minnesota. James J. Elser is Professor of Biology at Arizona State University. Peter Vitousek, one of the world's leading ecosystem ecologists, is Morrison Professor of Population and Resources at Stanford University.

"Few, if any, details of stoichiometry seem to have been overlooked by Sterner and Elser, and their book will be a useful reference to me for many years to come... The hundreds of references in the bibliography are worth the price of the book alone."--David W. Schindler, Nature "Robert Sterner and James Elser take a giant stride in knitting together perspectives across scales, biomes, and disciplines to craft an integrative and predictive vision of the topic... It knits together such a broad range of relevant topics that anyone interested in the connections between biology and elemental cycles should give it a try."--Anthony F. Michaels, Science "I believe that his is one of the most important books written in ecology in the last 10 years... I predict that in a few years it will be inconceivable to ignore the stoichiometric perspective when tackling an ecological problem... Ecological Stoichiometry is well written in colloquial and friendly prose. The authors strived to explain their arguments clearly and in detail. The many mathematical models are explained with laudable lucidity and the figures that illustrate them are consistently good."--Carlos Martinez Del Rio, Ecology "How often do you read a book that has a large number of 'aha!' moments in every chapter? This is a significant piece of synthesis and scholarship that brings together a very large number of disciplines and disparate chunks of data into a very satisfying whole... Never before have I seen a book which spans the scales from molecular biology to ecosystems so effectively... It is sure to become a classic."--Graham Harris, Journal of Plankton Research "Sterner and Elser adopt a simple, incremental logic in the book and follow it with fidelity... The book is an excellent introduction to ecological stoichiometry for graduate students; and for those already in the field, it is a thorough review of the complexities and nuances of stoichiometric ecology by two of its best practitioners."--Robert E. Hecky, Limnology and Oceanography Bulletin "A fresh and stimulating perspective on this area of ecology. The clear, readable style renders this text highly accessible."--Bulletin of the British Ecological Society "[This] book is seminal and presents a clear, well-reasoned overview of ES, thereby providing the biological community with an essential piece to the puzzle of integrating micro- and macro-level biology."--Van Savage, Complexity

• List of Figures ix List of Tables xiii Foreword xv Preface xvii 1. Stoichiometry and Homeostasis 1 Scope 3 Stoichiometry and Homeostasis 8 Yield 25 The Redfield Ratio 27 Conventions and Concerns about Element Ratios 31 Some Conventions about Growth Rate 34 A Logical Framework 35 The Structure of This Book 40 Summary and Synthesis 41 Key Definitions 42 2. Biological Chemistry: Building Cells from Elements 44 The Basis for Selection of Carbon, Nitrogen, and Phosphorus in Biochemical Evolution 45 The Elemental Composition of Major Biochemicals 51 Cell Components: The Elemental Composition of Cellular Structures 66 Summary and Synthesis 78 3. The Stoichiometry of Autotroph Growth: Variation at the Base of Food Webs 80 Cellular and Physiological Bases 81 C:N:P Stoichiometry of Entire Higher Plants 87 Autotrophs in Captivity 89 Theories of Autotroph Stoichiometry 107 Autotrophs in the Wild: Oceans, Lakes, and Land 120 Causes of Variation in Autotroph C:N:P in Nature 127 Catalysts for Ecological Stoichiometry 132 Summary and Synthesis 133 4. How to Build an Animal: The Stoichiometry of Metazoans 135 Biochemical and Biological Determinants of Body Elemental Composition 136 Invertebrate Stoichiometry: C:N:P in Zooplankton and Insects 138 Determinants of C:N:P in Invertebrates: The Growth Rate Hypothesis 142 Molecular Biology and the C:N:P Stoichiometry of Growth, or Ecosystem Scientists Go Astray 150 A Simple Molecular-Kinetic Model of the Growth Rate-C:N:P Connection 160 Structural Investment and the Stoichiometry of Vertebrates 168 Elemental Composition and Body Size 171 Catalysts for Ecological Stoichiometry 175 Summary and Synthesis 178 5. Imbalanced Resources and Animal Growth 179 Mass Balance in Growth Processes 180 Maximizing Yield in Chemistry and in Ecology 185 Limiting Factors for Heterotroph Growth: Development of Threshold Element Ratio Theory 189 A New Minimal Model of the Stoichiometry of Secondary Production 197 Some Real World Problems in Stoichiometric Balance 205 Growth Efficiency 222 Catalysts for Ecological Stoichiometry 227 Summary and Synthesis 229 6. The Stoichiometry of Consumer-Driven Nutrient Recycling 231 A Brief History of Studies of Consumer-Driven Nutrient Recycling 232 Stoichiometric Theories of Consumer-Driven Nutrient Recycling 235 Evidence That Consumers Differentially Recycle Nitrogen and Phosphorus 245 Microbial Mineralization 249 The Stoichiometry of Consumer-Driven Nutrient Recycling by Vertebrates 252 Catalysts for Ecological Stoichiometry 259 Summary and Synthesis 260 7. Stoichiometry in Communities: Dynamics and Interactions 262 Species Interactions 264 Positive Feedbacks and Multiple Stable States 277 Trophic Cascades 291 Light: Nutrient Effects at the Community Level 298 Feedbacks Owing to the "Constraints of Stuff": C:N Ratios in Tall-Grass Prairie 307 Catalysts for Ecological Stoichiometry 308 Summary and Synthesis 310 8. Big-Scale Stoichiometry: Ecosystems in Space and Time 313 Empirical Patterns in Ecosystem Stoichiometry 315 Linkages in the Stoichiometry of Biomass Yield: Using One Substance to Obtain Another 336 Nutrient Use Efficiency at the Ecosystem Level 341 The Stoichiometry of Food-Chain Production: A New Term, Carbon Use Efficiency 348 The Fate of Primary Production 350 Global Change 354 Catalysts for Ecological Stoichiometry 364 Summary and Synthesis 366 9. Recapitulation and Integration 370 Recapitulation 370 Integration: Toward a Biological Stoichiometry of Living Systems 376 Appendix 382 Literature Cited 385 Index 431