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Holism and Reductionism in Biology and Ecology the Mutual Dependence of Higher and Lower Level Research Programmes

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Holism and Reductionism in Biology and Ecology the Mutual Dependence of Higher and Lower Level Research Programmes RIJKSUNIVERSITEIT GRONINGEN Holism and Reductionism in Biology and Ecology The mutual dependence of higher and lower level research programmes Proefschrift ter verkrijging van het doctoraat in de Wijsbegeerte aan de Rijksuniversiteit Groningen op gezag van de Rector Magnificus, dr. F. van der Woude, in het openbaar te verdedigen op donderdag 9 april 1998 des namiddags te 4.15 uur door Richard Christiaan Looijen geboren op 27 augustus 1956 te Ifar (West Irian) Eerste promotor: Prof. dr. T.A.F. Kuipers Tweede promotor: Prof. dr. J. van Andel HOLISM AND REDUCTIONISM IN BIOLOGY AND ECOLOGY The mutual dependence of higher and lower level research programmes © 1998, Rick Looijen Voorkant: het verhaal van de vier blinden en de olifant; ontwerp: Joan Looijen, Rick Looijen, Ubel Smid CONTENTS ACKNOWLEDGEMENTS .......................................... 11 CHAPTER 1 GENERAL INTRODUCTION ....................................... 13 1.1 The holism-reductionism dispute ....................... 13 1.2 Division of the book ............................... 15 CHAPTER 2 HOLISM AND REDUCTIONISM IN BIOLOGY .......................... 23 2.1 Introduction ..................................... 23 2.2 The reduction problem in biology ...................... 24 2.2.1 The hierarchical structure of nature ................ 24 2.2.2 Complex hierarchy ........................... 26 2.3 Epistemological aspects of the problem .................. 27 2.3.1 Reduction and the unity of science ............... 29 2.3.2 Autonomism versus provincialism ................. 29 2.4 Ontological aspects ................................ 30 2.4.1 Materialism and causal determinism ............... 30 2.4.2 Vitalism versus constitutive reductionism ............ 31 2.4.3 ’Ontological’ reductionism: atomism ............... 32 2.4.4 Ontological holism: emergentism and organicism ...... 33 2.5 Methodological aspects ............................. 35 2.6 Ethical aspects ................................... 36 2.7 Concluding remarks: radical, moderate and anti-reductionist research strategies ................................. 37 CHAPTER 3 REDUCTION OF LAWS AND THEORIES ............................. 41 3.1 Introduction ..................................... 41 3.2 Nagel’s general reduction model ....................... 42 3.2.1 Homogeneous reductions ....................... 42 3.2.2 Heterogeneous reductions ...................... 44 3.3 Kuipers’s reduction model ........................... 45 3.3.1 The reduction of the ideal gas law ................ 46 3.3.2 Explanation and reduction steps .................. 47 3.3.3 Types of reduction ........................... 49 3.4 Concluding remarks ............................... 55 3.4.1 Epistemological versus ontological reduction ......... 55 3.4.2 Eliminative versus non-eliminative reduction ......... 55 CHAPTER 4 REDUCTION OF CONCEPTS ....................................... 57 4.1 Introduction ..................................... 57 4.2 Ontological identity relations ......................... 57 4.2.1 Identities versus correlations .................... 57 4.2.2 Criteria for ontological identity relations ............ 58 4.2.3 Thing-identities and attribute-identities ............. 59 4.2.4 Types and tokens ............................ 59 4.2.5 Type-identities and token-identities ................ 60 4.3 Ontological identities versus micro-reductions ............. 61 4.3.1 Introduction ................................ 61 4.3.2 An example: ’water = H2O’..................... 61 4.3.3 Constitutive reductionism and relations between wholes and parts .................................. 63 4.3.4 Causal explanation and substitution ................ 64 4.3.5 Other examples ............................. 65 4.4 Ontological identities versus ontological reduction .......... 66 4.4.1 Introduction ................................ 66 4.4.2 Epistemological verificationism .................. 67 4.4.3 Things, concepts, and terms ..................... 67 4.4.4 Epistemological reduction ...................... 68 4.5 Conclusions ..................................... 69 CHAPTER 5 EMERGENCE, REDUCTION AND CO-OPERATING RESEARCH PROGRAMMES 71 5.1 Introduction ..................................... 71 5.2 The doctrine of emergence ........................... 71 5.2.1 The whole is more than the sum of its parts .......... 71 5.2.2 Emergent evolution ........................... 72 5.2.3 Empirical and epistemological claims .............. 73 5.3 Emergence and reduction: a contradiction? ................ 74 5.3.1 Emergent properties .......................... 74 5.3.2 Emergence: a trivial thesis? ..................... 76 5.3.3 Reduction of emergent ’properties’ ................ 77 5.4 Ontological and epistemological aspects of emergence and reduction ....................................... 78 5.4.1 Emergence versus ontological identity .............. 78 5.4.2 Reduction without levelling ..................... 80 5.5 Methodological aspects of reduction .................... 81 5.5.1 Analysis versus synthesis? ...................... 81 5.5.2 Co-operating research programmes ................ 82 5.5.3 Partial reduction ............................. 83 5.6 Summary and conclusion ............................ 84 CHAPTER 6 CO-OPERATING RESEARCH PROGRAMMES: REDUCTION OF THE BOHR EFFECT85 6.1 Introduction ..................................... 85 6.2 Oxygen uptake by the blood .......................... 85 6.2.1 The Bohr effect ............................. 85 6.2.2 The role of hemoglobin ........................ 86 6.2.3 Allosteric effects ............................. 87 6.3 Reconstruction ................................... 88 6.3.1 Co-operating research programmes ................ 88 6.3.2 Reduction steps ............................. 90 6.3.3 Application or correlation? ...................... 92 6.3.4 Emergence ................................. 93 6.4 Concluding remarks ............................... 95 6.4.1 Reduction: construction or reconstruction? ........... 95 6.4.2 ’Holism’ and ’reductionism’ ..................... 96 6.4.3 Partial reduction ............................. 97 6.4.4 The function of hemoglobin ..................... 98 CHAPTER 7 FUNCTIONAL EXPLANATIONS IN BIOLOGY .......................... 99 7.1 Introduction ..................................... 99 7.2 Analyses of functional explanations .................... 100 7.2.1 Functions, goals and intentions .................. 100 7.2.2 Examples of functional explanation ............... 101 7.2.3 Functions, causes and effects ................... 102 7.2.4 The standard analyses of Nagel and Hempel ........ 103 7.2.5 Kuipers’s non-standard analysis ................. 105 7.2.6 Are functional explanations deductive arguments? .... 106 7.3 The problem of functional equivalents .................. 108 7.3.1 The specificity of biological laws ................ 108 7.3.2 Functional equivalents are irrelevant .............. 109 7.3.3 Functional properties as INUS-conditions .......... 110 7.4 The unity of causal and functional explanations ........... 111 7.4.1 The emergence of functional properties ............ 111 7.4.2 Functional properties as adaptations .............. 112 7.4.3 Natural selection and the principle of functionality .... 113 7.4.4 Exceptions to the functionality principle? ........... 114 7.4.5 Co-operating research programmes ............... 115 7.5 Conclusions .................................... 116 CHAPTER 8 THE REDUCTION PROBLEM IN ECOLOGY .......................... 121 8.1 Introduction .................................... 121 8.2 Holism and reductionism ........................... 122 8.2.1 Radical holism ............................. 123 8.2.2 Radical reductionism ......................... 126 8.2.3 The moderate position ........................ 127 8.2.4 Summary of approaches ....................... 129 8.3 The intellectual immaturity of ecology .................. 130 8.3.1 Reduction and co-operation .................... 130 8.3.2 The anomalous status of ecology ................ 131 8.3.3 The inhibitory effect of the holism-reductionism dispute .................................. 132 8.4 Remainder of the book ............................ 133 CHAPTER 9 ECOLOGICAL COMMUNITIES: CONCEPTUAL PROBLEMS AND DEFINITION 135 9.1 Introduction .................................... 135 9.2 The problem of ambiguity .......................... 136 9.2.1 Different concepts of community ................ 136 9.2.2 Holistic and reductionistic views ................. 137 9.2.3 Communities and biocoenoses .................. 138 9.2.4 Interactions between species .................... 139 9.3 The boundary problem and the problem of heterogeneity ..... 141 9.3.1 The boundary problem ....................... 141 9.3.2 The problem: communities as groups of populations . 143 9.3.3 Boundary communities ....................... 144 9.3.4 The problem of heterogeneity ................... 145 9.3.5 The solution: communities as sets of individuals ..... 147 9.4 Possible objections to the solution ..................... 148 9.4.1 Structure and classification of communities ......... 149 9.4.2 Community dynamics ........................ 150 9.5 An empirical example ............................. 152 9.5.1 Salt marsh plant communities ................... 153 9.5.2 Conceptual schemes ......................... 154 9.6 Animal communities .............................. 157 9.7 Conclusion ..................................... 157 CHAPTER 10 THE DISTINCTION BETWEEN HABITAT AND NICHE .................. 159 10.1 Introduction ...................................
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