Ecological Stoichiometry
The Biology of Elements from
Molecules to the Biosphere
Robert W. Sterner and James J. Elser
lechnische UniversitSt Oarmstadt
FACHBEREtCH 10 — BlOLOGiE
B i b! Iot hek —
.Schnittspahnstrafie 10
D-8 4 2 8 7 D a r m s t a d t
PRINCETON
UNIVERSITY
PRESS
• PRINCETON
AND
OXFORD
Contents
ix
List of Figures
List of Tables
->
Foreword
Preface
1. Stoichiometry and Homeostasis
Scope
Stoichiometry and Homeostasis
Yield
The Redfield Ratio
Conventions and Concerns about Element Ratios
Some Conventions about Growth Rate
A Logical Framework,
The Structure of This Book
Summary and Synthesis
Key Definitions
2. Biological Chemistry: Building Cells from Elements
The Basis for Selection of Carbon, Nitrogen, and Phosphorus
in Biochemical Evolution
The Elemental Composition of Major Biochemicals
Cell Components: The Elemental Composition of
Cellular Structures
Summary and Synthesis
3. The Stoichiometry of Autotroph Growth: Variation at
the Base of Food Webs
Cellular and Physiological Bases
C:N:P Stoichiometry of Entire Higher Plants
Autotrophs in Captivity
xiii
XV
xvii
1
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89
VI
CONTENTS
Theories of Autotroph Stoichiometry
Autotrophs in the Wild: Oceans, Lakes, and-'Land
Causes of Variation in Autotroph C:N:P in Nature
Catalysts for Ecological Stoichiometry
Summary and Synthesis
107
120
127
132
133
4. How to Build an Animal: The Stoichiometry
of Metazoans
;
Biochemical and Biological Determinants of Body
Elemental Composition
Invertebrate Stoichiometry: C:N:P in Zooplankton
and Insects
Determinants of C:N:P in Invertebrates: The Growth
Rate Hypothesis
Molecular Biology and the C:N:P Stoichiometry of Growth, or
Ecosystem Scientists Go Astray
A Simple Molecular-Kinetic Model of the Growth RateC:N:P Connection
Structural Investment and the Stoichiometry of Vertebrates
Elemental Composition and Body Size
Catalysts for Ecological Stoichiometry
Summary and Synthesis
5. Imbalanced Resources and Animal Growth
Mass Balance in Growth Processes
Maximizing Yield in Chemistry and in Ecology
Limiting Factors for Heterotroph Growth: Development of
Threshold Element Ratio Theory
A New Minimal Model of the Stoichiometry of
Secondary Production
Some Real World Problems in Stoichiometric Balance
Growth Efficiency
Catalysts for Ecological Stoichiometry
Summary and Synthesis
6. The Stoichiometry of Consumer-Driven
Nutrient Recycling
A Brief History of Studies of Consumer-Driven
Nutrient Recycling
Stoichiometric Theories of Consumer-Driven
Nutrient Recycling
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171
d75
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197
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235
CONTENTS
Evidence That Consumers Differentially Recycle Nitrogen
and Phosphorus
Microbial Mineralization
The Stoichiometry of Consumer-Driven Nutrient Recycling
by Vertebrates
Catalysts for Ecological Stoichiometry
Summary and Synthesis
7. Stoichiometry in Communities: Dynamics and
Interactions
Species Interactions
Positive Feedbacks and Multiple Stable States
Trophic Cascades
>
Light:Nutrient Effects at the Community Level
Feedbacks Owing to the "Constraints of Stuff": C:N Ratios in
Tall-Grass Prairie
Catalysts for Ecological Stoichiometry
Summary and Synthesis
8. Big-Scale Stoichiometry: Ecosystems in Space
and Time
Empirical Patterns in Ecosystem Stoichiometry
Linkages in the Stoichiometry of Biomass Yield: Using One
Substance to Obtain Another
Nutrient Use Efficiency at the Ecosystem Level
The Stoichiometry of Food-Chain Production: A New Term,
Carbon Use Efficiency
~>
The Fate of Primary Production
Global Change
Catalysts for Ecological Stoichiometry
Summary and Synthesis
9. Recapitulation and Integration
Recapitulation
Integration: Toward a Biological Stoichiometry of
Living Systems
vii
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376
Appendix
382
Literature Cited
385
Index
431