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Refining the Boundary Conditions of the Darwinian Concept of Adaption: the Affirmation of Darwinism Through Evo- Devo Charles Joseph Alt

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Refining the Boundary Conditions of the Darwinian Concept of Adaption: the Affirmation of Darwinism Through Evo- Devo Charles Joseph Alt Florida State University Libraries Electronic Theses, Treatises and Dissertations The Graduate School 2010 Refining the Boundary Conditions of the Darwinian Concept of Adaption: The Affirmation of Darwinism Through Evo- Devo Charles Joseph Alt Follow this and additional works at the FSU Digital Library. For more information, please contact [email protected] THE FLORIDA STATE UNIVERSITY COLLEGE OF ARTS AND SCIENCES REFINING THE BOUNDARY CONDITIONS OF THE DARWINIAN CONCEPT OF ADAPTATION: THE AFFIRMATION OF DARWINISM THROUGH EVO‐DEVO By CHARLES JOSEPH ALT A Dissertation submitted to the Department of Philosophy in partial fulfillment of the requirements for the degree of Doctor of Philosophy Degree Awarded: Fall Semester, 2010 The members of the Committee approve the Dissertation of Charles Joseph Alt defended on July 1, 2010. ________________________________________________ Michael Ruse, Ph.D Professor Directing Dissertation ________________________________________________ Matthew Day, Ph.D Outside Committee Member ________________________________________________ Russ M. Dancy, Ph.D Committee Member Approved: ____________________________________________________________ J. Piers Rawling, Chair, Department of Philosophy ____________________________________________________________ Joseph Travis, Dean, College of Arts and Sciences The Graduate School has verified and approved the above named committee members. ii For Mom, Dad, and Matthew; Dad, I could live 10 million years, each day trying to become a better man. But never in 10 million years could I even approximate the man that you are. My only hope is that one‐day I can say that I was like my father. That truly will have been a life well lived. Mom, in my lifetime I have been lucky enough to cross paths with Nobel Laureates, named professors, eminent scholars, etc. I have learned a great deal from these individuals, but it has been you who has been my greatest teacher and mentor. You have always been my best friend, even if I have never expressed this sentiment verbally. I cannot even begin to articulate the ways in which you have affected me, and I am eternally grateful that I have been able to share my life with you. Matt, I could not possibly imagine a better friend. You have been tolerant of my many flaws, and have forgiven my many mistakes. I have always admired the man that you are (even though you are my younger brother) and I am so proud of all that you have accomplished. iii ACKNOWLEDGEMENTS This dissertation would not have been written without the encouragement and help of many friends, family members, and colleagues. First, I would like to thank Michael Ruse, whose support and advice have been invaluable to me throughout my graduate education. I could not have asked for a better mentor, teacher, or friend. I would also like to thank the Ruse family (Lizzie, Emily, Oliver, and Ed) for taking me into their home and treating me as member of the family. I cannot possibly thank you enough for the warmth, hospitality, and support that you have given me. I would like to thank Russ Dancy and Matthew Day, for providing me with useful feedback throughout the dissertation process. I would like to thank the Philosophy Department at the Florida State University, for providing me with a top quality graduate education, and allowing me to complete this dissertation from my home in CT. I would like to thank the Greenwich Academy for its continued support throughout this process. You have helped to provide me with the motivation to continue my education, and your support has enabled me to complete this work. I would like to thank Fr. Tom Regan, S.J., who first encouraged me to pursue philosophy, and who has remained an outstanding mentor and friend. You took notice of my abilities when I didn’t know they were there, and this small act has profoundly changed the course of my life. You truly are the embodiment of Fr. Arrupe’s message, and the life of St. Ignatius of Loyola. You have lived your life in service of others, and I am truly lucky that I have been a part of the wonderful life that you have lived. I would like to thank Dale Houle, for challenging me to challenge myself, and for teaching me to do, rather than think about doing. Lastly, and most importantly, I would like to thank my family (Matt, Lisa, Mom, Dad, and Murphy) for their continued support throughout this process. Not only have they motivated me to never give up on what seemed an endless search for comprehension, they have provided me with compassion, love, and understanding along the way. In all of the possible worlds, there is not one in which I could have a better family! iv TABLE OF CONTENTS List of Figures.............................................................................................................................. viii Abstract.......................................................................................................................................... ix 1. MOTIVATIONS, DIRECTIONS, AND INTENTIONS ........................................................1 1.1 Motivations ....................................................................................................................4 1.2 Chapter Summaries........................................................................................................9 2. WHAT IS DARWINIAN EVOLUTION..............................................................................25 2.1 What is Darwinian Evolution.......................................................................................27 2.1.1 Darwin’s Predecessors.....................................................................................27 2.1.2 Why Darwin?...................................................................................................32 2.2 The Role of Adaptation in Darwin’s Theory of Evolution by Natural Selection ........38 2.2.1 The Design Question........................................................................................39 2.2.2 The Problem of Final Causes (Aristotle to Darwin) ........................................41 2.3 Darwin’s Adaptation....................................................................................................43 2.3.1 The Evolution of Darwin’s Adaptation Concept .............................................46 2.4 Adaptation of the term “Adaptation” Since Darwin....................................................47 3. COMMON CRICITICSMS OF THE CONCEPT OF ADAPTATION................................52 3.1 Gould & Lewontin: Spandrels of San Marco and the Panglossian Paradigm .............53 3.1.1 Spandrels..........................................................................................................55 3.1.2 The Adaptationist Response ............................................................................59 3.2 Additional Challenges..................................................................................................62 3.2.1 Genetic Drift ....................................................................................................63 3.2.2 Kimura’s Neutral Theory.................................................................................63 3.2.3 Structural and Developmental Constraints ......................................................65 3.3 The New Adaptation Concept......................................................................................67 3.3.1 The Non-historical Concept of Adaptation......................................................68 3.3.2 The Optimization Approach ............................................................................71 3.3.3 Empirical / Experimental Approaches to Studying Adaptation.......................75 3.3.4 Laboratory Evolution.......................................................................................79 3.4 Summary......................................................................................................................82 4. INTRO TO THE EVO-DEVO DEBATE .............................................................................84 4.1 Embryology (1920-1950).............................................................................................87 4.1.1 The Structure / Function Divide ......................................................................88 4.1.2 The Modern Synthesis .....................................................................................89 4.2 Why are Accounts of Inheritance Incomplete without Development?........................92 v 4.3 What Evo-Devo Brings to the Table............................................................................94 4.3.1 Evo-Devo Brings Development Back to the Table..........................................95 4.4 Some Goals of the New Evo-Devo..............................................................................97 4.4.1 Reviving the Old Debates ................................................................................97 4.4.2 Developmental Constriants..............................................................................99 4.4.3 Micro and Macro-evolution ...........................................................................101 4.4.4 A More Informed Evolution ..........................................................................102 4.5 Conclusion .................................................................................................................103 5. SELF-ORGANIZATION IN BIOLOGICAL SYSTEMS ..................................................105 5.1 Self-Organization.......................................................................................................108
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