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Brains as Engines of Association: An Operating Principle for Nervous Systems PDF

217 Pages·2019·101.949 MB·English
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Brains as Engines of Association Brains as Engines of Association AN OPERATING PRINCIPLE FOR NERVOUS SYSTEMS Dale Purves 1 1 Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide. Oxford is a registered trade mark of Oxford University Press in the UK and certain other countries. Published in the United States of America by Oxford University Press 198 Madison Avenue, New York, NY 10016, United States of America. © Oxford University Press 2019 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, by license, or under terms agreed with the appropriate reproduction rights organization. Inquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above. You must not circulate this work in any other form and you must impose this same condition on any acquirer. Library of Congress Cataloging- in- Publication Data Names: Purves, Dale, author. Title: Brains as engines of association : an operating principle for nervous systems / Dale Purves. Description: New York, NY : Oxford University Press, [2019] | Includes bibliographical references and index. Identifiers: LCCN 2018035410 | ISBN 9780190880163 Subjects: LCSH: Brain. | Neurobiology. | Neural circuitry. Classification: LCC QP356.25 .P86 2019 | DDC 612.8/ 2— dc23 LC record available at https://l ccn.loc.gov/ 2018035410 9 8 7 6 5 4 3 2 1 Printed by Sheridan Books, Inc., United States of America CONTENTS Preface ix Acknowledgments xiii What Nervous Systems Do for Animals PART I } 1. Putting the Question in Perspective 3 Introduction 3 Life on Earth 3 Defining Life 5 Energy 6 Evolution 7 Mechanisms 8 Teleology 10 Conclusion 11 Suggested Reading 11 2. Organisms Without Nervous Systems 13 Introduction 13 Bacteria 13 Protists 16 Plants 18 The General Strategy 21 Conclusion 21 Suggested Reading 21 3. Organisms With Nervous Systems 23 Introduction 23 Defining Nervous Systems 23 The Emergence of Nervous Systems 26 The Emergence of Central Nervous Systems 28 What Do Brains Add? 28 Conclusion 33 Suggested Reading 33 vi { Contents Nervous Systems as Engines of Association PART II } 4. The Organization of Nervous Systems 37 Introduction 37 Stimuli 37 Preneural Processing 38 Neural Processing 39 Behavioral Output 43 Neural Systems and Subsystems Are Interactive 45 Conclusion 47 Suggested Reading 47 5. The Problem 48 Introduction 48 Vision as an Example 49 The Basic Challenge 49 An Answer in General Terms 51 Qualia Determined by Empirical Ordering 54 Perceptual Discrepancies 54 Mechanisms 55 Other Modalities 56 Conclusion 57 Suggested Reading 58 6. Neural Associations 59 Introduction 59 Associations Wrought by Evolution 59 Associations Wrought by Lifetime Learning 61 Associations Wrought by Culture 65 Behavioral Categories of Associations 67 Reward 68 Behavioral Responses as Reflexes 69 What Is Associated? 70 Counterarguments 70 Conclusion 71 Suggested Reading 71 Evidence that Neural Systems Operate Empirically PART III } 7. Evidence from Lightness and Color 75 Introduction 75 Luminance and Lightness 75 Analyzing the Occurrence of Luminance Patterns 78 Effects of Other Luminance Patterns 80 Contents } vii Spectral Energy and Color 84 The General Strategy 86 Conclusion 88 Suggested Reading 88 8. Evidence from Geometry 89 Introduction 89 Seeing Intervals 89 Seeing Angles 93 Seeing Object Sizes in Two Dimensions 96 Seeing Object Sizes in Three Dimensions 97 Seeing Stereo Depth 100 Conclusion 103 Suggested Reading 103 9. Evidence from Motion 104 Introduction 104 Apparent Motion 104 The Perception of Speed 106 Implications for the Perception of Time 110 The Perception of Direction 112 Conclusion 115 Suggested Reading 115 10. Evidence from Audition 117 Introduction 117 Sound Signals 117 Sources of Tones 118 Sound Signal Spectra 119 The Problem in Audition 120 An Empirical Approach 122 Evidence from Speech 123 Evidence from Music 127 Implications for Any Sensory System 129 Conclusion 129 Suggested Reading 130 Alternative Concepts of Neural Function PART IV } 11. The Major Options 133 Introduction 133 Neural Function as Feature Detection 133 Neural Function as Statistical Inference 136 Neural Function as Predictive Coding 137 viii { Contents Neural Function as Efficient Coding 138 Neural Function as Computation 140 Conclusion 143 Suggested Reading 143 12. Summing Up 145 Introduction 145 Obstacles and a Way Around Them 145 Empirical Ranking 146 Winning Games 147 Biological and Artificial Intelligence 148 Consequences for Neuroscience 150 The Status of Reasoning 152 Novel Situations 153 Choice 154 Culture 155 Frequency of Stimuli 155 Conclusion 156 Suggested Reading 156 Glossary 159 Bibliography 169 Index 187 PREFACE This book concerns a question that has bothered me— and no doubt many others— for a long time: What does the human brain and the rest of the nervous system actually do? Based on subjective experience, the question seems gratuitous: Isn’t the brain the ultimate organizer of our behavior and the seat of the “I” when we think and act? The term “behavior,” however, includes an enormous range of responses to internal and external stimuli, and what “organizing” this welter of reactions means is not at all clear. Likewise, what the “I” might be in this conception of neural function has been debated for centuries without resolution. If the goal of biology is to understand how living things succeed in the world and the mechanisms they use to do so, then this sort of answer only papers over our ignorance about the function of the nervous system compared to other organs. By middle school, kids have learned that the respiratory system oxygenates the blood, that the digestive system infuses blood with nutrients, that the cardiovas- cular system conveys oxygen and nutrients to body tissues, and so on. An oper- ating principle of one sort or another has been documented for every major organ system save one: the brain and the rest of nervous system.1 Although textbooks and an overwhelming original literature testify to the wealth of neurobiological infor- mation gleaned over the past century or more, a comparable statement about the nervous system will not be found. This deficiency has led some investigators to imagine that the human brain is so specialized that understanding it will require approaches that have not been needed to fathom the operating principles of other organ systems. The implication is that conventional anatomical and physiological thinking is not up to understanding the human brain, sometimes described as “the most complex object in the known Universe.”2 Some of the disciplines envisioned as coming to the rescue are com- puter science, mathematics, statistics, network theory, game theory, graph theory, cybernetics, systems engineering, cognitive psychology, physics, quantum physics, philosophy, and “big data” mining. The big data approach (called “connectomics”) rests on the idea that, if all else fails, documenting the connectivity of the estimated 1 The term “brain” is often used as if the rest of the nervous system doesn’t count for much. In fact, there are about as many nerve cells in the spinal cord and peripheral nervous system as there are in the cerebrum, cerebellum, and brainstem (the usual definition of the brain). 2 This truism was recently asserted by Christof Koch, chief scientific officer of the Allen Institute for Brain Science (June 14, 2013, on NPR’s “Science Friday” with Ira Flatow). ix

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