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Perceiving in depth Volume 2, Stereoscopic vision PDF

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PERCEIVING IN DEPTH OXFORD PSYCHOLOGY SERIES 1. Th e Neuropsychology of Anxiety 18. Perceptual and Associative Learning 34. Looking Down on Human Intelligence J. A. Gray G. Hall J. Deary 2. Elements of Episodic Memory 19. Implicit Learning and Tacit Knowledge 35. From Conditioning to Conscious E. Tulving S. Reber Recollection H. Eichenbaum and N. J. Cohen 3. Conditioning and Associative Learning 20. Neuromotor Mechanisms in Human N. J. Mackintosh Communication 36. Understanding Figurative Language D. Kimura S. Glucksberg 4. Visual Masking B. G. Breitmeyer 21. Th e Frontal Lobes and Voluntary Action 3 7. Active Vision R. Passingham M. Findlay and I. D. Gilchrist 5. Th e Musical Mind J. A. Sloboda 22. Classifi cation and Cognition 38. Th e Science of False Memory W. K. Estes C. J. Brainerd and V. F. Reyna 6. Elements of Psychophysical Th eory J.-C. Falmagne 23. Vowel Perception and Production 39. Th e Case for Mental Imagery B. S. Rosner and J. B. Pickering S. M. Kosslyn, W. L. Th ompson, and G. Ganis 7. Animal Intelligence L. Weiskrantz 24. Visual Stress 40. Seeing Black and White Wilkins Gilchrist 8. Response Times R. D. Luce 25. Electrophysiology of Mind 41. Visual Masking, 2e Edited by M. D. Rugg and M. G. H. Coles B. Breitmeyer and H. Öğmen 9. Mental Representations Paivio 26. Attention and Memory 42. Motor Cognition N. Cowan M. Jeannerod 10. Memory, Imprinting, and the Brain G. Horn 27. Th e Visual Brain in Action 43. Th e Visual Brain in Action D. Milner and M. A. Goodale D. Milner and M. A. Goodale 11. Working Memory Baddeley 28. Perceptual Consequences of Cochlear 44. Th e Continuity of Mind Damage M. Spivey 12. Blindsight B. C. J. Moore L. Weiskrantz 45. Working Memory, Th ought, and Action 29. Perceiving in Depth, Vols. 1, 2, and 3 Baddeley 13. Profi le Analysis I. P. Howard with B. J. Rogers D. M. Green 46. What Is Special about the Human Brain? 30. Th e Measurement of Sensation R. Passingham 14. Spatial Vision D. Laming R. L. DeValois and K. K. DeValois 47. Visual Refl ections 31. Conditioned Taste Aversion M. McCloskey 15. Th e Neural and Behavioural Organization J. Bures, F. Bermúdez-Rattoni, of Goal-Directed Movements 48. Principles of Visual Attention and T. Yamamoto M. Jeannerod C. Bundesen and T. Habekost 32. Th e Developing Visual Brain 16. Visual Pattern Analyzers 49. Major Issues in Cognitive Aging J. Atkinson N. V. S. Graham T. A. Salthouse 33. Th e Neuropsychology of Anxiety, 2e 17. Cognitive Foundations of Musical Pitch J. A. Gray and N. McNaughton C. L. Krumhansl PERCEIVING IN DEPTH VOLUME 2 STEREOSCOPIC VISION Ian P. Howard Brian J. Rogers CENTRE FOR VISION RESEARCH DEPARTMENT OF EXPERIMENTAL PSYCHOLOGY YORK UNIVERSITY OXFORD UNIVERSITY TORONTO 1 1 Oxford University Press, Inc., publishes works that further Oxford University’s objective of excellence in research, scholarship, and education. Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi N ew Delhi Shanghai Taipei Toronto With offi ces in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Th ailand Turkey Ukraine Vietnam Copyright © 2012 by Oxford University Press, Inc. Published by Oxford University Press, Inc. 198 Madison Avenue, New York, New York 10016 www.oup.com Oxford is a registered trademark of Oxford University Press 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, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of Oxford University Press. ____________________________________________ A copy of this book’s Cataloging-in-Publication Data is on fi le with the Library of Congress. ISBN: 978-0-19-976415-0 ____________________________________________ 9 8 7 6 5 4 3 2 1 Printed in the United States of America on acid-free paper CONTENTS OF VOLUME 2 11. Physiology of disparity detection 1 21. Depth contrast 433 12. Binocular fusion and rivalry 51 22. Stereopsis and perceptual organization 470 13. Binocular summation, masking, and transfer 107 23. Th e Pulfrich eff ect 515 14. Binocular correspondence and the horopter 148 24. Stereoscopic techniques and applications 538 15. Linking binocular images 182 16. Cyclopean vision 210 References 564 17. Stimulus tokens for stereopsis 249 Index of cited journals 621 18. Stereoscopic acuity 287 P ortrait index 624 19. Types of binocular disparity 363 Subject index 625 20. Binocular disparity and depth perception 385 v A NOTE ON VIEWING THE STEREOGRAMS Th e stereograms presented in this book can be fused with well worth acquiring, since it is oft en the only way to achieve the aid of the prisms provided. Th e prisms should be held fusion with displays presented at vision conferences or close to the eyes and about 12 cm above the page. Th e viewer when you have lost your stereoscope. should be parallel to the plane of the page and correctly ori- A pair of images has one sign of disparity when fused by ented within that plane. Incorrect orientation shows as an convergence and the opposite sign of disparity when fused elevation of one image with respect to the other. Th e act of by divergence. A change in the sign of disparity reverses the fusing a side-by-side pair of images by diverging the eyes is apparent depth relationships in the fused image. For stereo- known as divergent fusion or uncrossed fusion. Th e act of grams in which the sign of disparity does not matter for the fusing images by converging the eyes is c onvergent fusion illustration of a phenomenon, only one pair of images is or crossed fusion. Th e prisms fuse the images by divergent provided. When the eff ect depends on the sign of disparity, fusion. Stereograms may also be free-fused by diverging or two stereogram pairs are provided — one pair for readers converging the eyes. In learning to free-fuse, it helps if the who prefer to converge the eyes, and the other for readers eyes converge on a pencil point held at the correct distance who prefer to diverge. Note that the provided lenses fuse by between the stereogram and the eyes. For divergence, one divergence only. Some stereograms in the book have triple may place a piece of clear plastic over the stereogram and images in a row. Th ese create two side-by-side fused images fi xate the refl ection of a point of light seen beyond the plane with opposite signs of disparity, plus fl anking monocular of the stereogram. Th e correct distance can be found by images. Th erefore, four images are seen when the images are observing how the images move as the pencil is moved in correctly fused. In some cases, it is instructive to compare depth. the image formed by convergent fusion with that formed Aft er some practice, readers will fi nd that they can con- by divergent fusion. In other cases, only one of the fused verge or diverge the eyes without an aid. When stereograms images is of interest. In this case, the location of the are free-fused, one sees each eye’s image on either side of the fused image of interest is indicated by a cross for those fused image. Th e presence of three pictures confi rms that who fuse by convergence and by two parallel lines for those correct vergence has been achieved. Free fusion is a skill who fuse by divergence. vi 11 PHYSIOLOGY OF DISPARITY DETECTION 11.1 Introduction 1 11.5.1 Disparity detectors in V2 and V3 24 11.1.1 Basic terms 1 11.5.2 Disparity detectors in the dorsal stream 26 11.1.2 Discovery of disparity detectors 2 11.5.3 Disparity detectors in the ventral stream 28 11.2 Subcortical disparity-tuned cells 4 11.5.4 Parvo- and magnocellular disparity detectors 29 11.2.1 Disparity tuning in the pulvinar 4 11.6 Higher-order disparities 30 11.2.2 Disparity tuning in the nucleus of the optic tract 4 11.6.1 Detection of horizontal disparity gradients 31 11.2.3 Disparity tuning in the superior colliculus 4 11.6.2 Detection of vertical disparity gradients 31 11.3 Disparity detectors in cats 5 11.6.3 Spatial modulations of disparity 33 11.3.1 Disparity detectors in areas 17 and 18 5 11.6.4 Joint tuning to disparity and motion 33 11.3.2 Disparity detectors in higher visual areas of cats 6 11.6.5 Joint spatial and temporal disparities 34 11.4 Disparity detectors in primate V1 6 11.7 Evoked potentials and stereopsis 34 11.4.1 Disparity tuning functions 6 11.8 PET, fMRI, and stereopsis 37 11.4.2 Number and homogeneity of detectors 14 11.8.1 Stationary stimuli 37 11.4.3 Position- and phase-disparity detectors 16 11.8.2 Motion in depth 38 11.4.4 Detection of vertical disparity 18 11.9 Detection of midline disparity 39 11.4.5 Orientation and disparity tuning 20 11.9.1 Eff ects of midline section of the chiasm 39 11.4.6 Disparity tuning and eye position 21 11.9.2 Eff ects of callosectomy 40 11.4.7 Disparity in contrast-defi ned stimuli 22 11.10 Models of disparity processing 40 11.4.8 Dynamics of disparity detectors 23 11.10.1 Energy models 40 11.5 Disparity detection in higher visual centers 11.10.2 Neural network models 49 of primates 24 11.1 INTRODUCTION stimulus falls outside the fi eld of view of the other eye, or (c) the stimulus is occluded to the other eye by a nearer stimulus. 11.1.1 BASIC TERMS A binocular stimulus is one seen at the same time by Th e term “ stereoscopic vision ” means literally, “solid sight.” both eyes. Th e term “ dichoptic ” was originally used to Strictly speaking, it refers to the visual perception of the describe the well-separated eyes of insects in contrast to 3-D structure of the world, when seen by one eye or by two. holoptic eyes, which have overlapping visual fi elds. Th e However, the term is generally used to refer to 3-D depth term “dichotic” was coined by Stumpf ( 1916 ) to denote perception arising from binocular disparities. the stimulation of each ear by a distinct sound. By analogy, S everal terms referring to binocular vision are in the term “dichoptic” has come to mean stimulation of the common use, but their meanings vary from author to two eyes by distinct distal stimuli (see Wade and Ono author. Strictly speaking, all animals with two eyes have 2005 ). binocular vision. Even animals with laterally placed eyes A dichoptic stimulus consists of distinct distal stimuli, integrate the information from the two eyes to form a one presented to one eye and one to the other, which an coherent representation of the fi eld of view. Also, the fi eld experimenter can control independently. Th ere are two of view of almost all mammals has some region in which the basic procedures for gaining dichoptic control. Th e fi rst is monocular fi elds overlap. However, the term “ binocular to present distinct stimuli to the two eyes in a stereoscope vision ” is usually reserved for animals possessing a large area or by an equivalent procedure such as free fusion. Th e other of binocular overlap within which diff erences between the procedure is to place diff erent fi lters or lenses in front of the images are used to code depth. two eyes. Dichoptic stimuli usually diff er in some defi ned A m onocular stimulus is a distal display seen by only way specifi ed by an experimenter. Th e diff erence may be one eye because (a) the other eye is closed or absent, (b) the (a) a disparity of position, size, or orientation between parts 1 or the whole of the stimuli, or (b) a diff erence in luminance, contrast, color, shape, or motion. Th e term “ dioptic stimulus ” has been used to mean a pair of identical stimuli in a stereoscope, in contrast to dichoptic stimuli, which diff er in some respect (Gulick and Lawson 1976 ). In the masking literature, a dichoptic stimu- lus is one in which a mask and a test stimulus are shown to diff erent eyes. In a dioptic stimulus they are shown to both eyes, and in a monoptic stimulus they are shown to one eye. Th e term “ monoptic depth ” has been used to denote an impression of depth created by a single eccentric stimulus in one eye (Section 17.6.5). 11.1.2 DISCOVERY OF DISPARITY DETECTORS Before the 1960s many scientists, including Helmholtz, believed that stereopsis does not involve conjunction of inputs from the two eyes at an early stage (Section 2.10.5). Ramón y Cajal ( 1911 ) proposed that inputs from corre- sponding retinal regions converge on what he called “isody- namic cells” and that this forms the basis of unifi ed binocular vision. Th is idea was confi rmed when Hubel and Wiesel ( 1959 , 1 962 ) reported that cells in the cat’s visual Figure 11.2. Colin Blakemore. Born in Stratford-upon-Avon, England in 1944. He obtained a B.A. in medical sciences from Cambridge University in 1965 and a Ph.D. in physiological optics from Berkeley in 1968. He also holds a Sc.D. (Cantab) and D.Sc. (Oxon). He was lecturer in physiology at Cambridge from 1972 to 1979, and then Waynfl ete Professor of Physiology at Oxford University. Between 1990 and 2003 he was director of the McDonnell-Pew Center for Cognitive Neuroscience in Oxford. He is now chief executive of the Medical Research Council. He has been the recipient of the Robert Bing Prize from the Swiss Academy of Medical Sciences, the Netter Prize from the French Académie Nationale de Médecine, the Royal Society Michael Faraday Award, the G.L. Brown Prize from the Physiological Society, and the Charles F. Prentice Award from the American Academy of Optometry. cortex receive inputs from the two eyes and that the recep- tive fi elds of these b inocular cells occupy corresponding positions in the two eyes. But consider what would happen if the monocular receptive fi elds of each binocular cell occupied identical positions in the retinas and were identi- cal in all other respects. Each binocular cell would respond optimally to similar stimuli with zero disparity. Such cells would not be diff erentially tuned to diff erent disparities and would therefore be incapable of coding relative depth. If images falling on corresponding locations diff ered, inputs to binocular cells could sum or rival. For stereopsis, the visual system needs binocular cells that are maximally responsive to inputs from receptive fi elds that are displaced Figure 11.1. Horace B. Barlow. Born in England in 1921. He graduated by diff erent amounts from exact correspondence. Any such from Trinity College, Cambridge. He was a research fellow at Trinity College between 1950 and 1954 and a lecturer at King’s College, cell would respond optimally to a stimulus with disparity Cambridge, between 1954 and 1964. Between 1964 and 1973 he was of a given magnitude and sign (crossed or uncrossed). A set professor of physiological optics and physiology at the University of of such cells with diff erent receptive-fi eld off sets in one California at Berkeley. He then returned to the physiological laboratory sign or the other could code diff erent disparities and hence at Cambridge University as a Royal Society Research Professor. He became a fellow of the Royal Society of London in 1969. relative depth. 2 • STEREOSCOPIC VISION In 1967 Jack Pettigrew discovered binocular cells with these properties in the cat’s visual cortex. Such cells are now known as disparity detectors. Pettigrew was a student of Peter Bishop at Sydney University, Australia (Section 2.10.5). He then joined Barlow and Blakemore in Berkeley, California. Working together, they confi rmed the existence of disparity detectors in the cat (Barlow et al. 1967 ) (Portrait Figures 11.1 , 11.2 , and 11.3 ). Similar fi ndings were reported about the same time from Sydney by Pettigrew, Nikara, and Bishop ( 1968 ) (Portrait Figure 11.4) . In 1977, Gian Poggio and his coworkers at Johns Hopkins University discovered dispar- ity detectors in monkey V1. Th e search for binocular cells tuned to diff erent dispari- ties was beset with the problem of ensuring that the images in the two eyes are in register. If the images are slightly out of register, a cell tuned to zero disparity will appear to be tuned to a disparity equal to the image misregistration. Also, any movement of the eyes during the recording intro- duces artifacts. Several procedures have been used to solve this problem. In the anesthetized animal, eye movements Figure 11.4. Peter O. Bishop. Born in Tamworth, New South Wales, are controlled by paralyzing the eye muscles and attaching Australia in 1917. He obtained the M.B. and B.S. in 1940 and the D.Sc. the eyeball to a clamped ring. A rotating mirror or a prism in 1967 from the University of Sydney. Aft er serving as a surgeon during the war he studied at University College London. He held of variable power controls the eff ective direction of gaze. In academic appointments at the University of Sydney from 1950 to 1967 the reference-cell procedure, diff erent electrodes record when he became professor of physiology at the Australian National University in Canberra. He retired in 1983. He is a fellow of the Australian Academy of Sciences, fellow of the Royal Society of London, offi cer of the Order of Australia, and joint winner of the Australia Prize in 1993. responses of a test cell and a reference binocular cell, each with receptive fi elds in the central retinas. Changes in the response of the reference cell indicate when eye movements have occurred (Hubel and Wiesel 1970 ). In a related proce- dure, eye drift is monitored by the response of a reference cell to monocular stimulation (Maske et al. 1986a ). Image stability can also be indicated by responses of LGN cells of foveal origin, one from each eye (LeVay and Voigt 1988 ). Th ese procedures indicate when eye drift has occurred, but they do not specify when test stimuli have zero dispar- ity, since the reference cell may not be tuned to zero dispar- ity. One solution to this problem is to use the mean response of several reference cells to defi ne zero disparity (Nikara et al. 1968 ). Another procedure is to use an ophthalmo- scope to project images of retinal blood vessels onto the screen on which the stimuli are presented (Bishop et al. 1962 ; Pettigrew et al. 1 968) . Th e problem is simplifi ed when testing is done on alert monkeys trained to converge Figure 11.3. John D. Pettigrew. Born in Wagga Wagga, Australia, in 1943. their eyes on defi ned targets. He obtained his B.Sc. in physiology from the University of Sydney in I n identifying a disparity detector one must ensure that 1966 and an M.B. from Sydney University Medical School in 1969. He has worked at Berkeley with H. Barlow and C. Blakemore, at the changes in responses are not due to incidental changes in California Institute of Technology, Queens University in Canada, and stimulation. For example, the stimulus in one eye may move the Zoologisches Insititüt in Munich. Since 1988 he has been director, outside the receptive fi eld of the binocular cell when the of the Vision, Touch, and Hearing Research Centre at the University of experimenter changes the disparity of the stimuli. Eff ects Queensland, Australia. He is a fellow of the Royal Society of London and of the Australian Academy of Science. of monocular position can be separated from eff ects of PHYSIOLOGY OF DISPARITY DETECTION • 3

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