Clonal Amplification of Behavior: A Simple Interpretation of the Effect of Reinforcement Open Access

Abstract

The theory of neuronal group selection (Edelman, 1987) is an account of neural development and dynamics that has been used as the theoretical basis for autonomous agents that are capable of an impressively wide range of adaptive behaviors (e.g., Edelman, 2007; Krichmar & Edelman, 2002; 2005; Krichmar, Nitz, Gally, & Edelman, 2005; Krichmar, Seth, Nitz, Fleischer, & Edelman, 2005; Seth & Edelman, 2007). Edelman’s theory draws parallels between natural selection and the adaptive dynamics of neuronal groups in response to environmental consequences. Critics have focused on the theory’s use of clonal amplification as the reproduction method, which they see as insufficiently adaptive (Crick, 1989; Fernando, Karishma, & Syathmary, 2008; Fernando, Goldstein, & Syathmary, 2010; Fernando, Szathmary, & Husbands, 2012). When comparing Edelman’s theory to the evolutionary theory of behavior dynamics (McDowell, 2004), McDowell argued that the theories differ in their reproduction methods and that a simulation that more purely models the clonal amplification dynamic may assess its viability (2010). This dissertation reports the results of the proposed simulations, which indicate that an implementation of the theory of neuronal group selection using clonal amplification can produce patterns of behavior that are quantitatively and qualitatively like humans and animals in operantly reinforcing environments. However, the range of viable parameters is smaller than for the evolutionary theory of behavior dynamics. There are also differences in the patterns of behavior predicted by the two theories that would need to be assessed with human or animal experiments to determine which is the better account.

Table of Contents

TABLE OF CONTENTS

Chapter 1: General Introduction ................................................................................................ 1

1.1. The Matching Law ............................................................................................................... 2

1.2. The Evolutionary Theory of Behavior Dynamics (ETBD) .................................................. 4

1.3. Theory of Neuronal Group Selection (TNGS) ..................................................................... 6

1.4. Practical Importance of Adaptive Models of Behavior ........................................................ 9

1.5. Objective of this Dissertation ............................................................................................. 11

Chapter 2: General Methods ..................................................................................................... 13

2.1. ETBD Creatures ................................................................................................................. 13

2.1.1. Representation of potential behaviors within the ETBD ............................................. 14

2.1.2. Step 3A: Beneficial selection ...................................................................................... 17

2.1.3. Step 4: Reproduction ................................................................................................... 19

2.1.4. Step 5: Variation .......................................................................................................... 20

2.2. Translating the TNGS to the ETBD: Three Algorithmic Variations ................................. 20

2.2.1. Algorithmic variant of step 4: Cloning reproduction .................................................. 23

2.2.2. Algorithmic variation of step 5: Phenotypic variation ................................................ 24

2.2.3. Algorithmic variant of step 3A: Roulette-continuous selection .................................. 26

2.3. Virtual Environments ......................................................................................................... 29

2.4. Apparatus ........................................................................................................................... 32

Chapter 3: ETBD and TNGS Behavior on Concurrent RI RI Schedules ............................. 33

3.1. Matching to Rates of Reinforcement on Single Schedules ................................................ 33

3.2. Methods .............................................................................................................................. 39

3.2.1. Participants .................................................................................................................. 39

3.2.2. Procedures ................................................................................................................... 40

3.2.3. Analyses ...................................................................................................................... 41

3.2.3.1. Data pooling and averaging .................................................................................. 42

3.2.3.2. Weighted ensemble fitting .................................................................................... 42

3.2.3.3. Analytic approach to ensemble fits ....................................................................... 46

3.2.3.4. Changeover profiles .............................................................................................. 48

3.3. Results ................................................................................................................................ 49

3.3.1. Best quantitative law of effect model .......................................................................... 49

3.3.2. Best fitting model parameters ...................................................................................... 51

3.3.2.1. Exponent (a) values .............................................................................................. 51

3.3.2.2. Asymptote (k) values ............................................................................................ 52

3.3.2.3. Rate of the quantitative law of effect’s ascent ...................................................... 53

3.3.3. Quadratic description of changeover profiles.............................................................. 55

3.3.4. Post-hoc analysis of changeover profiles .................................................................... 56

3.4. Discussion .......................................................................................................................... 62

3.4.1. Conformance to the matching law and the quantitative law of effect ......................... 62

3.4.2. Parameter values .......................................................................................................... 64

3.4.3. Changeovers ................................................................................................................ 65

3.4.4. Conclusion ................................................................................................................... 68

Chapter 4: Matching to Rates and Magnitudes of Reinforcement ........................................ 69

4.1. Methods .............................................................................................................................. 71

4.1.1. Participants .................................................................................................................. 71

4.1.2. Procedures ................................................................................................................... 72

4.1.3. Analyses ...................................................................................................................... 72

4.1.3.1. Data pooling and averaging .................................................................................. 73

4.1.3.2. Bivariate matching law equation .......................................................................... 73

4.2. Results ................................................................................................................................ 73

4.3. Discussion .......................................................................................................................... 75

4.3.1. Conclusion ................................................................................................................... 78

Chapter 5: General Discussion .................................................................................................. 79

REFERENCES ............................................................................................................................ 84

EQUATION SUMMARY .......................................................................................................... 96

FIGURES AND TABLES .......................................................................................................... 99

APPENDICES ........................................................................................................................... 135

Appendix A: Experiment 1 Fitting Measures of the Exponential-Bitwise-Bitflip Creature Type ................................................................................................................................................. 136

Appendix B: Experiment 1 Fitting Measures of the Exponential-Clone-Bitflip Creature Type ................................................................................................................................................. 152

Appendix C: Experiment 1 Fitting Measures of the Exponential-Clone-Pheno-Uniform Creature Type .......................................................................................................................... 168

Appendix D: Experiment 1 Fitting Measures of the Exponential-Clone-Pheno-Linear Creature Type ......................................................................................................................................... 184

Appendix E: Experiment 1 Fitting Measures of the Exponential-Clone-Pheno-Exponential Creature Type .......................................................................................................................... 200

Appendix F: Experiment 1 Fitting Measures of the Exponential-Clone-Pheno-Gaussian Creature Type .......................................................................................................................... 216

Appendix G: Experiment 1 Fitting Measures of the Linear-Bitwise-Bitflip Creature Type .. 232

Appendix H: Experiment 1 Fitting Measures of the Linear-Clone-Bitflip Creature Type ..... 248

Appendix I: Experiment 1 Fitting Measures of the Linear-Clone-Pheno-Uniform Creature Type ......................................................................................................................................... 264

Appendix J: Experiment 1 Fitting Measures of the Linear-Clone-Pheno-Linear Creature Type ................................................................................................................................................. 280

Appendix K: Experiment 1 Fitting Measures of the Linear-Clone-Pheno-Exponential Creature Type ......................................................................................................................................... 296

Appendix L: Experiment 1 Fitting Measures of the Linear-Clone-Pheno-Gaussian Creature Type ......................................................................................................................................... 312

Appendix M: Experiment 2 Bivariate Matching Fitting Measures ......................................... 328

About this Dissertation

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School	Laney Graduate School
Department	Psychology
Degree	Ph.D.
Submission	Dissertation
Language	English
Research Field	Psychology, Behavioral Sciences Psychology, Behavioral Psychology, Experimental
Keyword	Evolutionary computation Agent-based modeling Matching law Steady-state behavior Quantitative law of effect Operant behavior
Committee Chair / Thesis Advisor	Jack J McDowell, Emory University
Committee Members	Nathan A. Call , Marcus Autism Center Eugene K. Emory, Emory University Daryll B. Neill, Emory University Elaine F. Walker, Emory University

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