Color Vision C O L OR V I S I ON Perspectives from Different Disciplines Editors Werner G. K. Backhaus Reinhold Kliegl John S. Werner W Walter de Gruyter DE G Berlin · New York 1998 Editors Werner G. K. Backhaus John S. Werner Theoretical and Experimental Biology Department of Psychology Department of Biology University of Colorado Freie Universität Berlin Boulder, CO 80309 Thielallee 63 U.S.A. D-14195 Berlin e-mail: [email protected] Germany e-mail: [email protected] Reinhold Kliegl Department of Psychology University of Potsdam P.O. 60 15 33 D-14415 Potsdam Germany e-mail: [email protected] With 237 mostly four-colored figures © Printed on acid-free paper which falls within the guidelines of the ANSI to ensure permanence and durability Library of Congress Cataloging-in-Publication Data Color vision: perspectives from different disciplines/editors Werner G. K. Backhaus, Reinhold Kliegl, John S. Werner. Includes bibliographical references and index. ISBN 3-11-016100-1 (cloth: alk. paper). ISBN 3-11-015431-5 (pbk.: alk. paper) 1. Color vision. I. Backhaus, Werner. II. Kliegl, Reinhold, 1953- III. Werner, John Simon. QP483.C646 1998 612.8'4~dc21 97-46863 CIP Die Deutsche Bibliothek - Cataloging-in-Publication Data Color vision: perspectives from different disciplines/ ed. Werner G. K. Backhaus ... - Berlin; New York: de Gruyter, 1998 ISBN 3-11-015431-5 brosch. ISBN 3-11-016100-1 gb. © Copyright 1998 by Walter de Gruyter & Co., D-10785 Berlin All rights reserved, including those of translation into Reproductions and typesetting: City-Repro, Berlin foreign languages. No part of this book may be repro- Printing: Druckhaus Berlin-Mitte GmbH, Berlin duced or transmitted in any form or by any means, elec- Binding: Heinz Stein, Berlin tronic or mechanical, including photocopy, recording, or Cover design: Hansbernd Lindemann, Berlin any information storage and retrieval system, without Printed in Germany permission in writing from the publisher. Preface This book, Color Vision — Perspectives from Spillmann, and Michael Wertheimer. We also Different Disciplines, originated in an interdisci- thank Gary Smith, director of the Einstein Forum plinary "Symposium on Color Vision" that took in Potsdam, whose initial encouragement and con- place at the Einstein Forum in Potsdam in tinued support launched the book project. Sybille February 1996, with workshops in Berlin at the De Vito-Egerland (Division for External Affairs, Freie Universität Berlin (FUB), the Federal FUB), Mary Gaebel, Johannes Haack (Inter- Institute of Material Research and Testing (Bun- disciplinary Center for Cognitive Studies, Uni- desanstalt für Materialforschung und -prüfung) versity of Potsdam), Matthias Kross (Einstein and the School of the Arts (Hochschule der Forum), Katharina Misch (FUB), Konstanze Künste). The main goal of the symposium was to Pistor (Ministry of Science, Research, and review the current state of color vision research, Culture, Brandenburg), and Christiane von Alten bringing together scientists from such different (The Permanent Commission for Research and disciplines as neurobiology, psychology, color Academic Recruitment, FUB) contributed much metrics, medicine, philosophy and art. time and energy both to the organization of the symposium and the realization of the book. The The resulting book is conceptualized in the first symposium as well as the color illustrations in the place as a text book for introductory courses at the book were made possible by the generous finan- graduate level. It provides an introduction to the cial support of the Freie Universität Berlin, the exciting field of color vision for students and read- Einstein Forum Potsdam, the Interdisciplinary ers who are new to the field. The second intention Center for Cognitive Studies at the University of of this book is to give established scientists an Potsdam, the Brandenburg Ministry for Science, overview and an update on research activities in Research and Culture, and the Council on related fields. We hope that the material gathered Research and Creative Work of the University of from the diversity of fields will motivate students Colorado. Christiane Bowinkelmann and Mario and experts alike to take a broader perspective Noyer-Weidner, of Walter de Gruyter Publishers with regard to color vision from the different dis- Berlin, brought the book to completion. We are ciplines. grateful to each of them. J. S. W. is especially grateful to the Alexander von Humboldt Foun- dation and the University of Potsdam for a guest Acknowledgements professorship related to a senior scientist award during the time of the symposium. The successful completion of this project depend- ed on the commitment of a number of valued col- leagues. We want to thank the contributing authors who prepared their chapters according to the aims Berlin, 31 July 1997 Werner G. K. Backhaus of this book and who responded constructively to Reinhold Kliegl the suggestions for revisions that were made to John S.Werner them. Valuable comments were provided by Hans Irtel, Rainer Mausfeld, Brooke Schefrin, Lothar Contents Introduction xiii 2.1.1 Phenomenology of Color Vision .... 45 Werner G. K. Backhaus, Reinhold Kliegl 2.1.2 Disciplines 45 and John S. Werner 2.1.3 Psychophysical Simulations 46 2.1.4 Physiological Simulations 46 2.2 Color Stimuli 47 I. Color Vision in Art and Science 2.3 Psychophysics of Color Vision 48 2.3.1 Psychophysical Judgments 50 1. Aging through the Eyes of Monet 2.4 Psychophysical Color Spaces 51 John S. Werner 2.4.1 The Color Similarity (MDS) Space. . 51 1.1 Introduction 1 2.4.2 The Elementary Color Space (Color 1.2 A Link between Sunlight and Aging . 5 Sensations Space) 52 1.3 The Trivariance of Color Mixture: 2.4.3 The jnd Scale 54 Maxwell and Helmholtz 9 2.5 Neurophysiology of Color Vision ... 54 1.4 Monet's Early Impressionistic Style . 12 2.5.1 Humans and Other Vertebrates 54 1.4.1 Possible Influences of Turner and 2.5.2 Honeybees and Other Invertebrates. . 56 Goethe 13 2.6 Physiological Color Spaces 57 1.4.2 Possible Influences of Chevreul 2.6.1 Physical Description of the Color and Delacroix 17 Stimulus 57 1.5 Monet's Years in Argenteuil and 2.6.2 The Color Stimulus Space 57 Vetheuil 23 2.6.3 The Photoreceptor Sensitivity 1.6 The Opponent Code for Color (Light Absorption) Space Appearance: Hering 23 (1st Physiological Color Space) 60 1.7 Monet's Response to Pointillism and 2.6.4 The Photoreceptor Excitation Space Divisionism 27 (2nd Physiological Color Space).... 61 1.8 Hay Stack and Cathedral Series 30 2.6.5 The Color-Opponent Coding Space 1.9 Monet Returns to London 33 (3rd Physiological Color Space) .... 61 1.10 Water Lilies and Cataracts 33 2.6.6 Color Spaces and jnd Scales 62 1.11 Summary 38 2.7 Psychophysical and Physiological References 39 Simulations of Color Vision 62 2.7.1 The Psychophysical (MDS) Color Space in Honeybees 62 II. Physiology and Neuroethology 2.7.2 Neuronal Color-Coding and Color-Choice Behavior in Honeybees 64 2. Physiological and Psychophysical 2.7.3 Identification of the Physiological Simulations of Color Vision in COC Space and the Psychophysical Humans and Animals MDS Space 65 Werner G. K. Backhaus 2.8 Conscious vs. Unconscious 2.1 Introduction 45 Judgments 71 viii Contents 2.8.1 Color Sensations in Animals 71 5.5 Variation in Normal Color Vision ... 115 2.9 Conclusions 74 5.6 What Can Visual Pigment Gene References 75 Expression Tell Us about the Architecture of the Retina? 116 References 117 3. Receptors, Channels and Color in Primate Retina Barry B. Lee 6. Source Analysis of Color-Evoked Potentials in a Realistic Head Model 3.1 Introduction 79 Confirmed by Functional MRI 3.2 Physiology and Anatomy in the Walter Paulus, Renate Kolle, Retina 80 Jürgen Baudewig, Nora Freudenthaler, 3.3 Conclusions 86 Mathias Kunkel, Michael Finkenstaedt References 87 and Hans-Heino Rustenbeck 6.1 Introduction 121 4. Chromatic Processing in the Lateral 6.2 Methods 127 Geniculate Nucleus of the Common 6.3 Results 128 Marmoset (Callithrix jacchus) 6.4 Discussion 129 Jan Kremers, Eberhart Zrenner, 6.5 Summary 129 Stefan Weiss and Sabine Meierkord References 129 4.1 Introduction 89 4.2 Spectral Responsivities 91 7. Wavelength Information 4.3 Responses of LGN Cells to Various Processing versus Color Perception: Photoreceptor Contrasts 93 Evidence from Blindsight and 4.4 Selective Photoreceptor Stimulation Color-BIind Sight in Human Observers 95 Petra Stoerig 4.5 Summary 98 References 98 7.1 Introduction 131 7.2 Wavelength Information Processing . 134 7.2.1 Wavelength Information Processing 5. Molecular Genetics and the in Cortical Blindness 134 Biological Basis of Color Vision 7.2.2 Wavelength Processing in Maureen Neitz and Jay Neitz Cortical Color Blindness 138 5.1 Introduction 101 7.3 Segregation of Wavelength and 5.2 Background 101 Intensity Information and Constancy. 141 5.2.1 Types of Congenital Color Vision 7.4 Color Perception 143 Defects 101 References 145 5.2.2 Genome Organization and Inheritance Patterns of Color Vision Defects .... 102 8. Color Vision in Lower Vertebrates 5.2.3 Genes and Gene Expression 103 Christa Neumeyer 5.3 Spectral Tuning of M- and L-Cone Pigments 104 8.1 Introduction 149 5.4 Color Vision Defects 108 8.2 Wavelength Discrimination in Lower 5.4.1 What Distinguishes Normal from Vertebrates 150 Anomalous Pigments? Ill 8.2.1 Goldfish 150 5.4.2 What Distinguishes Photopigments 8.2.2 Turtles 151 Underlying Dichromacy from Normal 8.2.3 Amphibia 154 Pigments? 113 8.3 Color Constancy and Color Contrast. 155 Contents ix 8.4 Color Vision and Other Visual 11. Basic Color Terms and Basic Color Functions: Evidence for Parallel Categories 207 Processing of Visual Information ... 157 Clyde L. Hardin 8.5 Color Perception 159 Discussion and Summary 215 8.6 Summary 160 References 216 References 161 9. Color Vision: Ecology and 12. Color Perception: From Grassman Evolution in Making the Best of Codes to a Dual Code for the Photic Environment Object and Illumination Colors Peter G. Kevan and Rainer Mausfeld Werner G. K. Backhaus 12.1 Introduction 219 9.1 Introduction 163 12.2 Elementaristic vs. Ecological 9.2 Palaeontological Record 164 Perspectives in Color Research 220 9.3 Daylight and Color Vision 165 12.3 Attributes of Color 222 9.4 Colorimetry 167 12.4 Early Color Coding and the 9.5 Color Spaces 168 Elementaristic Approach 224 9.6 Evolution of Floral Colors and Color 12.4.1 Newton and Helmholtz's Approach Vision 171 to Color Perception 224 9.7 Color Patterns in Flowers 175 12.4.2 The Young-Helmholtz Theory and 9.8 Trichromacy and Tetrachromacy .... 177 Grassmann's Laws 225 9.9 Conclusions 178 12.4.3 Opponent-Color Theory 226 References 178 12.4.4 Relating Psychophysical and Neurophysiological Color Codes. . . . 228 12.4.5 Elementary Color Codes Accounting III. Psychology and Philosophy for Variations in Spatial and Temporal Context 231 10. The Perception of Blackness: 12.5 Ecological and Computational An Historical and Contemporary Perspectives 236 Review 12.5.1 The Problem of Approximate Color Vicki J. Volbrecht and Reinhold Kliegl Constancy from a Computational 10.1 Introduction 187 Point of View 238 10.2 The Phenomenology of Blackness. . . 187 12.5.2 Qualitative Observations on the Dia- 10.2.1 Helmholtz: Trichromatic Theory of lectic Relationship of Illumination Color Vision 189 and Object Color 240 10.2.2 Hering: Opponent-Process Theory of 12.6 Center-Surround Configurations as Color Vision 189 Minimal Stimuli for Triggering a 10.2.3 Criticism and Other Theories 191 Dual Code for Object Colors' and 10.3 Historical Review 194 'Illumination Colors' 242 10.3.1 Induction Experiments 194 12.6.1 Laminar Segmentation and a Dual 10.3.2 Blackness-Induction Experiments .. . 195 Code for Object Color' and 10.4 Physiological Mechanisms 201 'Illumination Color' 243 10.5 Conclusion 202 12.6.2 Segregation of Object Color' and References 202 'Illumination Color' in Minimal Seurat-type Configurations 245 References 248 χ Contents 13. Color Contrast Gain Control IV. Color Metrics and Application Michael D'Zmura 16. Dichromacy - 13.1 Introduction 251 The Simplest Type of Color Vision 13.1.1 What is Contrast Gain Control? 251 Horst Scheibner 13.1.2 Selectivity for Spatial Frequency, Orientation and Color 252 16.1 Introduction: An Initial Overview . . . 285 13.1.3 Feed-Forward, Matrix-Multiplicative 16.2 The Trichromatic Instrumental Color Circuitry 253 Space 3V 286 bgr 13.1.4 Spatial Pooling of Contrast 254 16.3 Measuring the Deuteranopic Missing 13.2 Model Components 254 Color and Reducing Trichromacy 13.3 Color Image Processing 256 to Deuteranopia 287 13.3.1 Channel Responses 256 16.4 The Transition from the Instrumental 13.3.2 Channel Contrasts 257 Trichromatic Space to the 13.3.3 Channel Interaction 261 Instrumental Deuteranopic Space . . . 290 13.3.4 Channel Gains 261 16.5 The Transformation from the Tri- 13.3.5 Multichannel Contrast Gain Control . 263 chromatic Instrumental Color Space 13.4 Discussion 264 to the Deuteranopic Opponent-Color 13.5 Summary 265 Space 291 References 265 16.6 The Role of the Fundamental Color Space 294 16.7 Construction of the Fundamental 14. Binocular Brightness Combination: Color Spaces 3V and 2V and PTD PX A Mechanism for Combining the Deuteranopic Opponent-Color Two Sources of Rather Similar Channels 295 Information 16.8 A Synopsis of Deuteranopia 298 Hans Irtel 16.9 A Synopsis of Dichromacy 299 14.1 Intensity Invariance of Binocular 16.10 A Lattice-Theoretical Classification of Dichromacy and Other Color Brightness 267 Deficiencies 301 14.2 Methods 269 14.3 Results 270 16.11 Concluding Remarks 301 14.4 Discussion 271 16.12 Summary 302 14.5 Summary 273 References 302 References 273 17. Current CIE Work 15. Inferences about Infant to Achieve Physiologically-Correct Color Vision Color Metrics Kenneth Knoblauch, Jänos Schanda Michelle L. Bieber and 17.1 Introduction 307 John S. Werner 17.2 Cone Excitation Spectra 308 15.1 Introduction 275 17.2.1 Choice of the Color-Matching 15.2 Inferences from Luminosity 275 Functions 308 15.3 Inferences from Silent Substitution . . 277 17.2.2 Deriving L-, M-, S-Cone Excitation 15.4 Inferences about Rod Intrusion 278 Spectra from Color-Matching Data . . 309 15.5 Inferences about M- and L-Cones . . . 279 17.2.3 Intra-Ocular Screening 310 15.6 Summary 281 17.2.4 Derivation of the Fundamental References 281 Response Curves 311