Animal Psychophysics: the design and conduct of sensory experiments

Animal Psychophysics: the design and conduct of sensory experiments

Animal thedesignandconduct Springer Science+Business Media, LLC

P sychl())]jj>h ysics~ of sensory experd.m.enis EDITOR: WILLIAM C. STEBBINS Kresge Hearing Research Institute, and Departments of Otorhinolaryngology and Psychology, University of Michigan, Ann Arbor, Michigan

Main entry under title: library of Congress Cataloging in Publication Data Animal psychophysics: the design and conduct of sensory experiments. Reprint of the ed. published by Appleton-Century-Crofts, New York. Includes bibliographies. 1. Senses and sensation-testing. 2. Psychology, Physiological-Methodology. I. Stebbins, William C., 1929- ed. [DNLM: 1. Ethology-Congresses. 2 Perception-Congresses. 3. Psychophysics-Congresses. 4. Sensation-Congresses. QP441 A598 1969] [QP435.A571974] 156'.2'8 74-16468 ISBN 978-1-4757-4516-0 ISBN 978-1-4757-4514-6 (ebook) DOI 10.1007/978-1-4757-4514-6 1970 Springer Science+Business Media New York Originally published by Plenum Press, New York in 1970. Softcover reprint of the hardcover 1st edition 1970 All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher

SUSAN BARROW Department of Psychology, Brown University, Providence, Rhode Island. MARK A. BERKLEY Psychology Department, Florida State University, Tallahassee, Florida. HARRY J. CARLISLE Department of Psychology, University of California at Santa Barbara, Santa Barbara, California. BEN CLOPTON Department of Physiology and Biophysics, and Regional Primate Research Center, University of Washington Medical School, Seattle, Washington. JOHN I. DALLAND Division of Social Sciences, Richmond College, The City University of New York, Staten Island, New York. EBERHARD FETZ Department of PhYSiology and Biophysics, and Regional Primate Research Center, University of Washington Medical School, Seattle, Washington. MITCHELL GLICKSTEIN Department of Psychology, Brown University, Providence, Rhode Island. GEORGE GOUREVITCH Hunter College, The City University of New York, New York, New York. JOSEPH KIMM Departments of Otolaryngology, Physiology and Biophysics, and Regional Primate Research Center, University of Washington Medical School, Seattle, Washington. v

vi Contributors VICTOR G. LATIES Department of Radiation Biology and Biophysics, The University of Rochester School of Medicine and Dentistry, Rochester, New York. ERICH LUSCHEI Department of Physiology and Biophysics, and Regional Primate Research Center, University of Washington School of Medicine, Seattle, Washington. RICHARD W. MALOTT Psychology Department, Western Michigan University, Kalamazoo, Michigan. MARILYN KAY MALOTT Psychology Department, Western Michigan University, Kalamazoo, Michigan. JOSEF M. MILLER Departments of Otolaryngology, Physiology and Biophysics, and Regional Primate Research Center, University of Washington Medical School, Seattle, Washington. W. LLOYD MILLIGAN Psychological Research Laboratory, Veteran's Administration Hospital, Columbia, South Carolina. DA VID B. MOODY Kresge Hearing Research Institute, and Department of Psychology, University of Michigan, Ann Arbor, Michigan. JOHN A. NEVIN Psychology Department, Columbia University, New York, New York. BARBARA A. RAY Neurology Research, Massachusetts General Hospital, Boston, Massachusetts. ROBERTW. REYNOLDS Department of Psychology, University of California at Santa Barbara, Santa Barbara, California. PETER B. ROSENBERGER Pediatric Neurology Unit, Massachusetts General Hospital, Boston, Massachusetts.

Contributors THOMAS R. SCOTT Psychological Research Laboratory, Veteran's Administration Hospital, Columbia, South Carolina. JAMES SMITH Psychology Department, Florida State University, Tallahassee, Florida. WILLIAM C. STEBBINS Kresge Hearing Research Institute, and Departments of Otorhinolaryngology and Psychology, University of Michigan, Ann Arbor, Michigan. SYLVIA THORPE Psychology Department, Brown University, Providence, Rhode Island. BERNARD WEISS Department of Radiation Biology and Biophysics, The University of Rochester School of Medicine and Dentistry, Rochester, New York. DEAN YAGER Psychology Department, Brown University, Providence, Rhode Island. vii

Preface In May of 1969, the contributors to this book gathered at the University of Michigan in Ann Arbor for three days to talk about their work in the behavioral analysis of animal sensory function and to share their research experiences in the laboratory with particular emphasis on methodology in behavioral training, testing, and instrumentation. It was their feeling and mine as a consequence of this meeting that we had sufficient substance to justify a book which we hoped would be of interest and even of pragmatic value to any biologic or biomedical scientist whose work deals with sensory function. Clearly, there is no aspect of an organism's behavior that is not to some extent controlled by environmental stimuli. In recent years, due in large part to technical advances in microscopy and histology and in electrophysiology, there have been several extremely informative published proceedings from conferences and symposia concerned with some of the early and very basic stages in the reception of environmental energy by the sense organs and its processing by the nervous system. Transduction at the receptor and stimulus coding by the nervous system, cell membrane changes, and the basic structure of the receptor and related tissue as seen through the electron and phase contrast microscope have received major attention, and exciting new discoveries in sensory function and structure have been reported. Ultimately, such discoveries must be related to an intact behaving organism. Questions ranging from the resolving power of a sensory system (the concept of threshold) to a wide variety of complex perceptual phenomena can only be directed to such a behavioral preparation. For all of their obvious value in understanding nervous function, potentials recorded in end organ and brain take on new meaning when their relationship to behavior has been made clear and unequivocal. Although data on sensory acuity are available for man, it is only in recent years that we have improved our techniques to be able to obtain similar data from lower animals. Since much of the research on physiologic function and anatomic structure has been done with these animals, it becomes particularly important to know in some detail their sensory and perceptual acuity as revealed by their behavioral responses to sensory stimulation. Training and testing procedures based on the principles of operant conditioning have shown that animals can report on their sensory capabilities in as precise and reliable a fashion as humans. In the past decade in the laboratory, we have acquired considerable normative data on sensory acuity in animals. Second, developments in other areas of inquiry have reached a stage of readiness for such behavioral technology. For example, the information to be gained from electrical recording and stimulating in neural tissue and direct infusion of substances in the awake animal in neurophysiology and neuropharmacology is greatly increased if the animal has been trained as a precise observer and reporter of sensory events. The conference and the book, then, had at least two primary objectives. The first was in fact to demonstrate the efficacy of these operant ix

x Preface conditioning methods for the analysis of sensory function in animals, and the second was to show, where possible, the use of these methods in conjunction with methods from other biologic research areas in an attempt to better understand some aspect of sensory function (normal or pathological). I think it is safe to say, in conclusion, that the success of Ollr venture-conference and book-rests to a large extent on the data that these behavioral procedures have produced. There did not appear to us to be any very obvious order of arrangement for the chapters in the book, although the reader will see that some attempt at grouping has been made. The introductory chapter was written primarily for our colleagues in other research areas of biology and biomedicine in the hope that our methods and our findings might be of some interest to them and even applicable to their work. In that chapter, I have tried to show some of the background for the development of animal psychophysics from both the literature of human psychophysics and animal conditioning. I have also tried to introduce some of the terminology that we use, some typical experimental designs and arrangements, and some idea of the problems encountered. All this is done far too briefly, but reference is made to other sources for more extended background and to the chapters themselves where procedures and findings are described with the same clarity, rigor, and thoroughness with which the original experiments were carried out in the laboratory. Although the chapters are based on papers presented within a somewhat formal structure at the conference, a general attitude of informality and enthusiasm prevailed at the time. Hopefully some of this attitude remains in the book, which has not been severely edited. It is an attitude which characterizes the field. We who work there have a good time doing what we do and talking with our colleagues about it. The support for the research, the conference, and the book came from many sources. Granting agencies are appropriately listed with the individual chapters in the book. Partial support for the conference and subsequent preparation of the book came from Appleton-Century-Crofts. A contract from the Office of Naval Research (Nonr(G)- 00024-67) to E. R. Galanter supported a two-day planning session in the spring of 1968 at Columbia University; unfortunately, Dr. Galanter was unable to attend our conference in Ann Arbor. There are many people who played an important part, directly and indirectly, in putting together both the conference and the book. I would dedicate the book to the contributors who made it an exciting and rewarding conference and sent their chapters in quickly, to the students from our laboratory at the University of Michigan who were so helpful in making the conference a success: Bill Clark, Julie Costin, Doris Foster, Swayzer Green, Elbert Magoun, Dick Pearson, and Bud Wilkinson, to Karen Pritula who played a major role in taking care of all the many problems that arose between original manuscript and page proofs, to my colleague Dave Moody for keeping the lab going, to Dick Van Frank for his help as Appleton-Century-Croft's Editor-in-Chief, Scientific Books Department, to my wife and daughters who provided inspiration and rarely complained about late hours and lost weekends, and to those of my teachers from whom I learned, among many other things, that a liberal attitude may be suitable for politics but a conservative one with caution and healthy skepticism is sometimes more appropriate for science: W. W. Cumming, F. S. Keller, H. D. Patton, W. N. Schoenfeld, and A. L. Towe.

Contents Contributors Preface v ix 1. Principles of Animal Psychophysics William C. Stebbins 1 2. The Measurement of Ultrasonic Hearing John 1. Dalland 21 3. Studies of Hearing and Hearing Loss in the Monkey William C. Stebbins 41 4. Detectability of Tones in Quiet and in Noise by Rats and Monkeys George Gourevitch 67 5. Psychophysical Testing of Neurologic Mutant Mice Barbara A. Ray 99 6. Conditioned Suppression as an Animal Psychophysical Technique J ames Smith 125 7. Response-Adjusting Stimulus Intensity Peter B. Rosenberger 161 8. The Psychophysics of Pain and Analgesia in Animals Bernard Weiss and Victor G. Laties 185 9. Thermal Reinforcement and Temperature Regulation Harry J. Carlisle 211 10. Visual Discriminations in the Cat Mark A. Berkley 231 xi

xii Contents 11. Vision in Monkeys with Lesion of the Striate Cortex Mitchell Glickstein, Susan Barrow, and Erich Luschei 249 12. Investigations of Goldfish Color Vision Dean Yager and Sylvia Thorpe 259 13. Reaction Time as an Index of Sensory Function David B. Moody 277 14. Sensory Neurophysiology and Reaction Time Performance in Nonhuman Primates Josef M. Miller, Joseph Kimm, Ben Clopton, and Eberhard Fetz 303 15. The Use of Reaction Time in Monkeys for the Study of Infonnation Processing Robert W. Reynolds 329 16. The Psychophysical Study of Visual Motion Aftereffect Rate in Monkeys Thomas R. Scott and W. Lloyd Milligan 341 17. Perception and Stimulus Generalization Richard W. Malott and Marilyn Kay Malott 363 18. On DiHerential Stimulation and DiHerential Reinforcement John A. Nevin 401 Author Index 425 Subject Index 431

Animal Psychophysics~ the design and conduct of sensory experiments