Evolution from a Thermodynamic Perspective

During the 1940s and early 50s, Carl F. Jordan spent boyhood summers at his uncle’s hunting and fishing lodge in northern Maine. He enjoyed the wilderness there, especially canoe trips on the Allagash and Penobscot rivers, and deplored the  cutting of the spruce-fir forests by the pulp and paper companies. In 1953, he enrolled at the University of Michigan and majored in forestry, because he believed that it could help him conserve the forests, but in those days, forestry was all about “getting out the cut”. After he acquired his Ph.D. in plant ecology from Rutgers Univ.in 1966, he joined H.T. Odum in an Atomic Energy Commission project in Puerto Rico, looking at the dynamics of radioactive isotopes in the rain forest following the world-wide atmospheric testing of nuclear weapons.  In 1969, Carl moved to Argonne National Laboratory where he continued studies of radioactive pollution from nuclear power plants.  In 1974,he had the opportunity to lead an ecology project for the University of Georgia to determine energy flow and nutrient cycling in the Amazon Region of Venezuela. In 1980, Carl returned to the School of Ecology in Athens Georgia while continuing tropical research in Brazil, Costa Rica, Mexico and Thailand. In 1993, Carl acquired a farm near Athens Georgia that had once been part of a pre-Civil War cotton plantation and began research on more sustainable ways of farming. He originated the first University course in Georgia on organic farming, and opened the farm to tours and classes interested in sustainable agriculture. Carl retired as Professor Emeritus in 2009, and took his new freedom to begin research for Evolution from a Thermodynamic Perspective, and recently to develop a forum where the controversies raised in that book could be discussed. The forum is available at the website Thermodynamic-Evolution.org

• C​ontents.- Part 1. Theory.- To Understand Economics, Follow the Money: To Understand Ecosystems, Follow the Energy.- Two Views of Ecology, Evolution, and Conservation.- Why I Wrote this Book.- Dualities Still Impede Conservation Efforts.- The Intergovernmental Science-Policy Platform of Biodiversity.- Targets for Conservation.- Evolving Objectives.- Literature Review.- Updating Ecosystem Ecology.- References.- What Can We Learn by Studying Ecosystems that We Can’t Learn from Studying Populations?.- The Predator-Prey Conundrum.- The Serengeti Ecosystem.- Evolution in the “Ecological Theater”.- Predator-Prey Interactions Tell Only Part of the Story.- Evolution in the “Thermodynamic Theater”.- References.- A Thermodynamic Definition of Ecosystems.- Ecosystems in the 20th Century.- Cycling of Strontium-90.- Cesium-137 in Food Chains.- Recycling of Isotopes in Norwegian Sheep.- Ecological Energetics.- Is it Time to Bury the Ecosystem Concept?.- A Thermodynamic Definition of Life.- A Thermodynamic Definition of Ecosystems.- The Phase Transition between Order and Chaos.- References.- Thermodynamic Characteristics of Ecosystems.- Equilibrium.- The Equilibrium Law.- Thermodynamic Equilibrium.- Open Thermodynamic Systems.- Ecosystems are Thermodynamically Open Non-Equilibrium Systems.- Work is Performed by Non-equilibrium Systems.- Advantage of a Thermodynamically Open System.- 4.3 Ecosystems are Entropic.- 4.4 Ecosystems are Cybernetic.- Cybernetic Systems.- Economic Systems are Cybernetic Ecosystems are Cybernetic.- The Ecosystem Feedback Function.- Indirect vs. Direct Feedback.- Deviation Dampening and Amplifying Feedback.- Set Points.- Ecosystems are Autocatalytic.- Ecosystems have Boundaries.- Ecosystems are Hierarchical.- Hierarchy in Physical Systems.- Hierarchy in Ecological Systems.- Common Currencies.- Macro-and Micro-System Models.- Why an Ecosystem Model that Includes Everything is not Possible.- A Nested Marine Community.- Ecosystems are Deterministic.- Ecosystems are Information Rich.- An Engineering Definition of Information.- Information to Facilitate Exchange.- High Energy Information.- Low Energy Information.- Information Theory.- Genetic Information.- Ecosystems are Non-Teleological.- Criticisms of Ecosystem Models.- References.- Ecosystem Control: A Top-Down View.- Two Ways to Look at Systems.- Composing and Decomposing Trophic Webs.- Decomposers in Soil Organic Matter.- Decomposers in Marshes and Mangroves.- Control of Systems.- Top-Down vs. Bottom-Up.- Top-Down Exogenous Control.- Exogenous Impacts and Stability.- Top-Down Endogenous Control.- Endogenous Control through Nutrient Recycling.- Autocatalysis.- Control of Microbial Activity.- Inhibition of Microbial Activity by Leaf Sclerophylly.- Inhibition of Microbial Activity Chemical Defenses.- Inhibition of Microbial Activity by Ecological Stoichiometry.- The Synchrony Principle.- The Decay Law.- Direct Nutrient Cycling.- The Role of Animals.- Indirect Interactions.- Marine Systems.- Nutrient and Energy Recycling.- Exogenous Control.- Control in Lakes.- Control in Managed Ecosystems.- References.- Ecosystem Control: A Bottom-Up View.- Species as Arbitrageurs of Energy.- Relation Between Rate of Flow and Mass in Hydraulic Systems.- Relation Between Population Biomass and Rate of Energy Flow.- Equilibrium.- Mechanisms of Adjustment.- Adjustments and Climate Change.- Bird Populations.- Dis-equilibrium.- Population Instability vs. Ecosystem Instability.- Control by Interactions: Direct vs. Indirect.- Indirect Interactions.- Direct Interactions.- Predator – Prey.- Mutualisms.- Competition.- Decomposition.- Parasitism and Disease.- Commensalism and Amensalism.- Persistence of Negative Interactions.- References.- Ecosystem Stability.- Background.- A Thermodynamic Definition.- Regime Shift.- Metastability.- Pulsed Stability.- Resistance and Resilience.- Species Richness and Functional Stability.- Species Richness and Cultural Values.- Keystone Species, and Population and Ecosystem Stability.- 7.5.1 Keystone Species in the Yellowstone region of Wyoming.- References.- 8. Case Studies of Ecosystem Control and Stability.- Walden.- “Harmony in Nature”.- Feedback Produces Nature’s “Harmony”.- Feedback Mechanisms.- Perturbations in Amazon Rain Forests.- Top-Down Control.- The San Carlos Project: A Small-scale, Low Intensity, Short Duration Disturbance.- 8.3.2 The Jarí Project: A Large-scale, High Intensity, Long Duration Disturbance.- Bottom-Up Control.- The El Verde Project.- The Long-Term Ecological Research Project in Puerto Rico.- The Lago Guri Island Project.- The Biological Dynamics of Tropical Rainforest Fragments Project.- What have Case Studies Taught us about Stability of Tropical Ecosystems?.- Tropical Ecosystems are Stable.- Tropical Ecosystems are Unstable.- Energy Flow in Tropical Savannas and Rain Forests.- Insects in Tropical Ecosystems.- Application of Lessons to Other Regions.- Relevance to Temperate Zones.- Relevance to Aquatic Ecosystems.- The Experimental Lakes Project (Ecosystem Control of Species).- Lake Mendota Studies (Species Control of Ecosystems).- 8.7 Case Studies as Tests of Thermodynamic Theory.- References.- Entropy and Maximum Power.- Entropy.- 9.2 Entropy in a Steel Bar.- Thermodynamic Equilibrium.- Entropic Gradients.- Capturing and Storing Entropy.- Evapotranspiration and Entropy Reduction.- Life is a Balance between Storing and Releasing Entropy.- The Law of Maximum Entropy Production.- Energy for Metabolism as well as Growth.- Unassisted Entropy Capture is a Unique Characteristic of Life.-9.6Entropy Storage by Ecosystems.- 9.6.1 What Causes Entropy to be Stored?.-  9.7 Capturing Pressure.-  9.8 Entropy and Time.- 9.8.1 Time’s Speed Regulator.- Efficiency of Energy Transformations.- Passage of Time for Cats.- 9.9The Maximum Power Principle.-9.10 Optimum Efficiencies for a Truck and its Driver.-9.11 Sustainability.- References.- A Thermodynamic View of Succession.- 10.1 The Population View.- 10.2 The Thermodynamic View.- 10.2.1 Leaf Area Index and Succession.- 10.2.2 Power Output as a Function of Leaf Area Index.- 10.2.3 What Causes Changes in Leaf Area Index?.- 10.2.4 Maximum Entropy Production Principle.- 10.2.5 Successional Ecosystems Move Further from Thermodynamic Equilibrium.- 10.2.6 Entropy Storage by Animals.- 10.3 The Strategy of Ecosystem Development.- A Problem with Odum’s Strategy.- Why Power Output Continues to Increase.- Revised Definition of Maximum Power.- Costs of Ecosystem Stabilization.- Transactional Costs.- Succession, Power Output, and Efficiency.- 10.5.1 Kleiber’s Law.- Are Ecosystems Spendthrifts?.- Interactions Between Species Facilitate Increase in Power Output.- Facilitation.- Tolerance.- Inhibition.- Intermediate Disturbance Hypothesis.- Nutrient Use Efficiency during Succession.- Succession Following Logging vs Following Agriculture.-  10.10 Thermodynamic View of Succession: Implications for Resource Management.- References.- Panarchy.- The Universal Cycle of Systems.- Panarchy.- Thermodynamic Interpretation of the Sacred Rules.- 11.2.1 Growth and Consolidation.- 11.2.2 Collapse.- Renewal.- Sub-systems.- Panarchy over 2 Billion Years of Evolution.- Consolidation, Bureaucracy and System Collapse.- Bureaucracy in Action (Case Studies).- Case Study: Panarchy in the Georgia Piedmont.- Thermodynamic Interpretation.- References.- 12. A Thermodynamic View of Evolution.- 12.1 Life – A Physicist's View.- 12.1.1 Life is Produced by Capturing Entropy.- 12.1.2 The Origin of Life.- 12.2 Two Approaches to Evolution.- 12.2.1 The Eco-Evo-Devo View.- 12.2.2 The Thermodynamic View.- 12.2.3 Fitness.- 12.2.4 The “Goal” of Evolution.- 12.3 The Relationship between Species and Environment.- 12.3.1 Evolution’s “Theater”.- 12.3.2 Is Evolution Stochastic or Deterministic?.- 12.4 Ecosystem Evolution.- 12.4.1 Succession was the Clue.- 12.4.2 Ecosystems Moved away from Equilibrium.- 12.4.3 Thermodynamic Mechanisms.-  12.4.4 Biological Mechanisms.- 12.4.5 Ecosystem Fitness.- 12.4.6 Ecosystems Evolve One Step at a Time.- 12.5. The Origin of Ecosystems   .- 12.5.1 Origin of Feedback Loops.- 12.5.2 Origin of Trophic Levels.- 12.5.3 Why are there Trophic Levels?.- 12.6 The “Goal” of Ecosystem Evolution.- 12.6.1 Conflicting Goals?.- 12.6.2 “Motivations” of Species.- 12.6.3 The Earth Ecosystem.- 12.6.4 Why is there Resistance to the Idea of Ecosystem Evolution?.- 12.6.5 Evolution of Economic Systems.- 12.7 A Thermodynamic Model of Ecosystem Evolution.- 12.7.1 Network Models.- 12.7.2 Increase in Complexity of Trophic Webs.- 12.7.3 Evolution of Trophic Webs.- 12.7.4 Life Moves Ashore.- 12.8 Biodiversity and the Five Great Extinctions.- 12.8.1 The Cretaceous-Tertiary (K-T) Boundary Extinction.- 12.8.2The Amazing Sustainability of Trophic Chains.- 12.8.3 A Test of Thermodynamic Theory.- 12.9 Panarchy and Evolution.-  12.10 Thermodynamic Requirements for Living Systems on Other Planets.- References.- .-Why is Species Diversity Higher in the Tropics?.- 13.1 Tropical Explorations.- 13.2 A Few Theories.- 13.3 A Thermodynamic Explanation.- 13.3.1 The Latitudinal Energy Gradient.- 13.3.2 The Latitudinal Productivity Gradient.- 13.3.3 The Data.- 13.3.4 Other Factors Affecting Productivity.- 13.4 Empirical Evidence for a High Productivity High Diversity Correlation.- 13.5 Humboldt’s Enigma.- 13.5.1 Are Productivity and Species Richness Correlated on Tropical.- Mountains?.- 13.6 The Mechanism Linking Productivity and Diversity.- 13.7 Answer to “Why is Species Diversity Higher in the Tropics?”.- 13.7.1 Differences within the Tropics.- 13.8 Why is Species Diversity Low at High Latitudes?.- 13.9 An Economic Perspective on Diversity.- 13.9.1 Energy Flow, Economic Growth and Professional Diversity.- References.- What Have We Learned by Viewing Evolution from a Thermodynamic Perspective?.- 14.1 What we have Learned.- 14.1.1 Fitness Means Maximization of Power Output.- 14.1.2 Feedback is Essentialfor Sustainability.- 14.1.3 Control of Energy Flow Occurs both Top-down and Bottom-up.- 14.1.4 Storage of Entropy is a Powerful Characteristic of Living Systems.- 14.1.5 Evolution is the Storage of Entropy.- Objections to the Ecosystem Concept.- 15.1 Criticisms of the Ecosystem Concept.- 15.1.1 Ecosystems are Abstractions.- 15.1.2 Ecosystems are Ephemeral.- 15.1.3 Ecosystems are Oversimplifications.- 15.1.4 The Ecosystem Concept is Merely a Paradigm.- 15.1.5 The Ecosystem Concept is Not Based on Facts.- 15.2 Ecosystems are not Cybernetic.- 15.3 Inappropriate Machine Analogies.- 15.4 Objections to Ecosystem Evolution.- 15.4.1 No Measure of Fitness.- 15.4.2 Evolution has no Goals.- 15.4.3 The Theory Can’t be Tested.- 15.4.4 No Mechanisms.- 15.4.5 Contradicts Neo-Darwinism.- 15.4.6 Restricted Definition.- 15.5 Setting up a Straw Man.- 15.6 Harmony in Nature?.- 15.7 Conservatism.- References.- What has Thermodynamics Taught us about Conservation?.- 16.1 “Habitat” is not Synonymous with “Ecosystem”.- 16.2 Conservation and Feedback.- 16.3 A Few Case Studies.- 16.3.1 The Serengeti.- 16.3.2 Black Footed Ferret.- 16.3.3 Golden Lion Tamarin.- 16.3.4 Whooping Cranes.- 16.3.5 Puerto Rican Parrot.- 16.4 Conserving Feedback Loops.- 16.5 The Importance of Reservoirs for Recovery of Feedback Loops.- 16.6 Biodiversity Hotspots.- 16.7 A Conservationist’s Dilemma.- 16.8 Conservation and Feedback: A Final Word.- References.- Part 2. Application.- Thermodynamic Laws and Agriculture.- 17. A Farmer’s Dilemma.- 17.1 How Ecosystems and Economic Systems are the Same.- 17.2 How Ecosystems and Economic Systems are Different.- 17.3 Planet Earth is a Feedback System.- References.- 18. Agricultural Problems are Systems Problems.- 18.1 The Morrill Land-Grant Acts.- 18.2 The Evolution of Agricultural Research.- 18.2.1 Reductionism.- 18.2.2 The Empirical Approach.- 18.2.3 The Analytical Approach.- 18.3 Agricultural Development Models.- 18.3.3 The Ratchet Effect.- 18.4 The Systems Approach.- 18.4.1 Adaptive and Deterministic Cycles.- 18.4.2 Business Cycles and New Paradigms.- References.- Instability in Economic Food Systems.- 19.1 Pressures for Economic Expansion.- 19.1.1 Political Pressures.- 19.1.2 Humanitarian Challenges.- 19.1.3 Invested Academic Interests.- 19.2 Instability of Economic Food Systems: External Factors.- 19.2.1 Booms and Busts.- 19.2.2 Vulnerability of Farmers.- 19.3 Instability of Economic Food Systems: Internal Factors.- 19.3.1 Source of Energy for Yield.- 19.4 Lack of Feedback: Case Study.- 19.4.1 Data.- 19.4.1.1Energy Inputs.- 19.4.1.2 Energy Returned on Energy Invested.- 19.5 Control in Ecological vs. Economic Systems.- 19.6 The Emergence of Feedback and Control.- 19.7 Stability of Economic Food Systems.- References.- 20 Energy Efficiency in Agricultural Systems.- 20.1 Two Kinds of Energy.- 20.2 Early Comparisons of Energy Use Efficiency.- 20.3 Energy Returned on Energy Invested.- 20.4 EROI for Industrial Corn.- 20.4.1 Production Functions.- 20.4.2 An Energy Production Function.-  20.5 Economic Considerations.- 20.5.1 Income.- 20.5.2 Costs.- 20.5.3 Profit.-  20.6 A Farmer’s Dilemma.- 20.7 The Maximum Power Principle and Economic Theory.- References.- The First Law of Thermodynamics and Genetic Engineering (There is no Free Lunch).- 21.1 Source of Energy for Increased Crop Yield.- 21.1.1 Endosomatic Energy.- 21.1.2 Hybridization.- 21.1.3 High Yield Rice.- 21.2 Domestication of Balsas teosinte.- 21.2.1 Calculations.- 21.2.2 Mechanisms.- 21.3 Other Tradeoffs.- 21.4 The Free Lunch has already been Eaten.- References.- 22 Top-down vs. Bottom-up Control in Resource Management Systems.- 22.1 Background.- 22.2 Experimental Site and Hypotheses.- 22.2.1 Methods.- 22.2.2 Traditional Agroforestry (Hierarchical level – the Ecosystem).- 22.2.3 Organic Production (Hierarchical level - local economic community).- 22.2.4 Sun Coffee Plantation (Hierarchical level – the corporate economy) 22.3 System Comparisons.- 22.3.1 Energy Input.- 23.3.2 Output.- 22.3.3 Results.- 22.4 Discussion.- 22.4.1 Effect of energy sources on system outputs.-  22.5 Conclusions.- 22.5.1 Feedback and Environmental Sustainability.- 22.5.2 Economic sustainability.-  References.- 23 Services of Nature in Agricultural Systems.- 23.1 Services of Nature.- 23.2 The Nutrient Recycling Service of Nature.- 23.2.1 Erosion Prevention.- 23.3 Energy in Agricultural Systems.- 23.3.1Embedded Energy.- 23.3.2 Embodied Energy.- 23.4 The Systems Analyzed.-  23.5 Summary of Results.- 23.6 Discussion.- 23.6.1Shifting Cultivation.- 23.6.2. High Rates of Return on Exosomatic Inputs.- 23.6.3 Low Rates of Return on Exosomatic Inputs.- 23.6.4 Rates of Return on Endosomatic Inputs.- 23.6.5 Sustainability.- 23.6.6 Benefits and Costs of Herbicides.-  References.- 24. Optimizing Sustainability.- 24.1 Two Views of Sustainability.- 24.2 A Compromise for Agriculture.- 24.2.1 Value of Services of Nature (Endosomatic Inputs).- 24.2.2 Value of Exosomatic Inputs.- 24.2.3 Energy vs. Dollars as a Measure of Sustainability.- 24.3 An Economic Model for Compromise.- 24.4 Case Studies.- 24.4.1 Pest Control by Services of Nature.- 24.5 Trends.-  References.- 25 Agriculture that Incorporates Services of Nature.- 25.1 Environmentally Benign Agriculture.- 25.2 Intercropping.- 25.3 Regenerative Agriculture.- 25.4 Agroforestry.- 25.4.1 Agroforestry in Tropical Regions.- 25.4.2 Agroforestry in the Temperate Zone.-  25.5 Disadvantages of Agroforestry.- 25.6 The Governmental Perspective.- References.- 26 Rebuilding Natural Capital: A Case Study.- 26.1 The Nature of Capital.- 26.1.1 Depletion of Natural Capital.- 26.2 Rebuilding Natural Capital.- 26.3 An Obstacle to Rebuilding Natural Capital.- References.- Can Organic Agriculture Feed the World?.- 27.1 Organic Agriculture vs. Low- Energy-Input Agriculture.- 27.2 Organic Agriculture vs. Conventional Agriculture.- 27.3 Can Agriculture Dependent on Low Energy Input Feed the World?.- 27.4 Yield is Notthe Problem.- References.- What has Thermodynamics Taught us about Sustainability?.- What We Have Learned by Viewing Resource Management from a Thermodynamic Perspective?.- 28.1.1Services of Nature are Not Free.- Valuing Nature’s Services and Natural Capital.- Natural Capital is not Recognized.- National Capital.- Energetic Value of Nature’s Services.- 28.4 Taxes, Fees, and Reimbursements.- 28.5 Case Studies.- 28.5.1 Agriculture (The Plowman’s Folly).- 28.5.2 Fisheries.- 28.5.3 Forestry.- 28.5.4 Species Conservation.- 28.5.5 Wildlife Management.- 28.5.6 Flood Control.- 28.5.7 Landscape Management.- 28.5.8 Climate Change.- 28.6 Natural Resources and the Free Market System.- References.- Part 3 Conservation.- Conservation of Resource Systems Means Preserving the Services of Nature.- In Wilderness is the Preservation of the World.- 29.1 Sacred Groves.- 29.2 Information is Stored in Sacred Groves.- 29.3Wilderness as a Resource Bank for Nature’s Services.