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The Concept of Levels of Organization in the Biological Sciences

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The Concept of Levels of Organization in the Biological Sciences The Concept of Levels of Organization in the Biological Sciences PhD Thesis Submitted August 2014 Revised June 2015 Daniel Stephen Brooks Department of Philosophy Bielefeld University Reviewers: Martin Carrier, Maria Kronfeldner Dedicated in friendship to Jan and Magga “He who has suffer'd you to impose on him knows you.” Table of Contents Table of Contents........................................................................................................................1 Chapter One: The Intuitive Appeal and Ubiquity of 'Levels of Organization'............................5 1.1 Introduction ................................................................................................................5 1.2 Analyzing 'Levels of Organization' in Biological Science..........................................7 1.2.1 Level Claims.....................................................................................................10 1.2.2 Wimsatt’s Characterization of Levels ..............................................................14 1.3 Initial Distinctions.....................................................................................................16 1.3.1 Erroneous Concepts of Levels...........................................................................18 1.4 Depictions of Levels in Biological Textbooks...........................................................20 1.4.1 The Character of 'Levels' in Biological Science................................................24 1.4.2 The Significance of 'Levels' in Biological Science...........................................30 1.5 The Concept of Hierarchy.........................................................................................34 1.5.1 'Hierarchy' Does Not Exhaust 'Levels of Organization'....................................37 1.6 The Structure of This Dissertation.............................................................................40 Chapter Two: Philosophical Conceptions of Levels.................................................................43 2.1 Introduction...............................................................................................................43 2.2 The Layer-Cake Account of Levels...........................................................................43 2.2.1 Global Scope and Comprehensive Character....................................................46 2.2.2 Stepwise Compositional Continuity .................................................................47 2.2.3 Linearity of Levels Strata..................................................................................50 2.2.4 Correspondence Between Nature and Science..................................................52 2.2.5 The Continued Influence of the Layer-Cake Account.......................................55 2.3 The Mechanistic Account of Levels..........................................................................59 2.3.1 Local Scope and Radically Contextualized Character......................................62 2.3.2 Principled Rejection of Correspondence Between Science and Nature...66 2.3.3 Branching Structure ..........................................................................................68 2.3.4 Constitutive Relevance......................................................................................69 2.4 Conclusion.................................................................................................................71 Chapter Three: Levels Skepticism............................................................................................76 3.1 Introduction...............................................................................................................76 3.2 Levels – A Flawed, Misleading, and Irrelevant Concept? ........................................76 3.2.1 The Levels Concept is Flawed..........................................................................77 3.2.2 The Levels Concept is Misleading....................................................................81 3.2.3 Are Levels of Organization Irrelevant to Science?...........................................84 3.3 Considerations of Levels Skepticism.........................................................................86 3.3.1 A Straw-Man Conception of Levels..................................................................86 3.3.2 Guilt by Association: The Layer-Cake Account as the Default Conception of Levels..............................................................................................................................89 3.3.3 Association of 'Levels' with 'Layer-cake Levels'...............................................90 3.3.4 Association of 'Levels' with Layer-Cake Reductionism....................................92 3.4 Conclusion ................................................................................................................98 Chapter Four: A Fragmentary Concept...................................................................................101 4.1 Introduction.............................................................................................................101 4.2 Semantic Variation and the Levels Concept............................................................103 4.2.1 Fragments of the Levels Concept ...................................................................107 4.2.2 A Common Standard for Comparing Level Claims in Biology......................109 4.3 A Framework for Analyzing Semantic Variation in Biological Concepts...............113 4.3.1 Components of Semantic Content and the Molecular Gene Concept.............116 4.3.2 Epistemic Goals of Concept Usage.................................................................119 4.3.3 Unifying Fragmentary Concepts under their Epistemic Goal.........................121 4.4 The Epistemic Goal of 'Levels of Organization'......................................................126 4.4.1 How Levels of Organization Structure Problems ..........................................128 4.4.2 Descriptive Level Claims................................................................................128 4.4.3 Hypothetical Level Claims .............................................................................130 4.5 Case Study for the Use of Levels: The Explanation of Oxidative Phosphorylation133 4.5.1 Ox-phos as a Multi-level Phenomenon...........................................................135 4.5.2 The Shift from Chemical to Biological Problem............................................138 4.6 Conclusion...............................................................................................................147 Chapter Five: The Organicist Roots of the Levels Concept....................................................149 5.1 Introduction.............................................................................................................149 5.2 The Historical Context of the Levels Concept........................................................154 5.3 The Organicist Program and the Levels Concept....................................................161 5.3.1 The Tenets of the Organicist Program.............................................................161 5.3.2 The Organicists' Use of the Levels Concept ..................................................171 5.3.3 The Epistemic Goal behind the Organicists' Use of Levels ...........................177 5.4 The Integrative Account of Levels of Organization................................................184 5.4.1 Global Scope and Contextualized Character...................................................187 5.4.2 Incomplete Epistemic Continuity in Levels of Organization..........................191 5.4.3 Weak Association Between Science and Nature.............................................195 5.5 Integrative Levels & Layer-cake Levels..................................................................199 5.6 Organicist Influence on Mitchell’s Chemiosmotic Hypothesis...............................202 5.7 Conclusion ..............................................................................................................205 General Conclusion.................................................................................................................208 References...............................................................................................................................211 Acknowledgments...................................................................................................................224 Table of Figures Figure 1.1 Different Overlapping K-Decompositions...............................................................11 Figure 1.2 Levels of Organization in Campbell Biology..........................................................22 Figure 1.3 Common Definitional Criteria for Levels................................................................26 Figure 1.4 Levels of Organization in Solomon et al.'s Biology................................................27 Figure 1.5 Formal Hierarchies..................................................................................................36 Figure 1.6 Types of Branching Structures in Hierarchies.........................................................37 Figure 2.1 Layer-Cake Account of Levels................................................................................45 Figure 2.2 Microreduction Relations in the Layer-cake Account.............................................48 Figure 2.3 Multi-level Mechanisms..........................................................................................62
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