Extrastriate Cortex in Primates PDF
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CEREBRAL CORTEX Volume 12 Extrastriate Cortex in Primates CEREBRAL CORTEX Series Editors: Edward G. Jones and Alan Peters Advisory Committee Antonio R. Damasio, Iowa City, Iowa Irving T. Diamond, Durham, North Carolina Patricia Goldman-Rakic, New Haven, Connecticut Jon H. Kaas, Nashville, Tennessee John H. Morrison, New York, New York Kathleen S. Rockland, Iowa City, Iowa Adam M. Sillito, London, England Wolf Singer, Frankfurt, Germany R. D. Terry, La]olla, California P. Ulinksi, Chicago, Illinois Volume 1: Cellular Components of the Cerebral Cortex Volume 2: Functional Properties of Cortical Cells Volume 3: Visual Cortex Volume 4: Association and Auditory Cortices Volume 5: Sensory-Motor Areas and Aspects of Cortical Connectivity Volume 6: Further Aspects of Cortical Function, Including Hippocampus Volume 7: Development and Maturation of Cerebral Cortex Volume SA: Comparative Structure and Evolution of Cerebral Cortex, Part I Volume 8B: Comparative Structure and Evolution of Cerebral Cortex, Part II Volume 9: Normal and Altered States of Function Volume 10: Primary Visual Cortex in Primates Volume 11: The Barrel Cortex of Rodents Volume 12: Extrastriate Cortex in Primates Series Editors: Edward G. Jones and Alan Peters CEREBRAL CORTEX Volume 12 Extrastriate Cortex in Primates Edited by KATHLEEN S.ROCKLAND The University of Iowa Iowa City, Iowa JON H. KAAS Vanderbilt University Nashville, Tennessee and ALAN PETERS Boston University School ofM edicine Boston, Massachusetts Springer Science+B usiness Media, LLC The Library of Congress cataloged the first volume of this title as follows: Main entry under title: Cerebral cortex. Includes bibliographical references and index. Contents: v. 1. Cellular components of the cerebral cortex. 1. Cerebral cortex---Collected works. !. Peters, Alan, 1929- Il. Jones, Edward G ., 1939- . [DNLM: 1. Cerebral cortex-Anatomy and histology. 2. Cerebral cortex-Physiology. WL 307 C4136] QP383.C45 1984 612'.825 84-1982 ISBN 978-1-4757-9627-8 ISBN 978-1-4757-9625-4 (eBook) DOI 10.1007/978-1-4757-9625-4 © 1997 Springer Science+Business Media New York Originally published by Plenum Press, New York in 1988 Softcover reprint of the hardcover 1st edition 1988 http:// www.plenum.com 10 9 8 7 6 5 4 3 2 1 Ali 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 Contributors Jean Bullier Cerveau et Vision INSERM 371, 69675 Bran Cedex, France Stephanie Clarke Institut de Physiologie, Universite de Lausanne, CH-1005 Lausanne, Switzerland Catherine G. Cusick Department of Anatomy SL49, Tulane School of Medicine, New Orleans, Louisiana 70112 Charles G. Gross Department of Psychology, Princeton University, Princeton, New Jersey 08544 Balazs Gulyas Division of Human Brain Research, Department of Neuroscience, Karolinska Institute, S-17177 Stock holm, Sweden Jon H. Kaas Department of Psychology, Vanderbilt University, Nashville, Tennessee 37240 Tirin Moore Department of Brain and Cognitive Sciences, Mas sachusetts Institute of Technology, Cambridge, Massachusetts 02139 Lionel G. Nowak Section of Neurobiology, Yale University School of Medicine, New Haven, Connecticut 06510 Guy A. Orban Laboratorium voor Neuro- en Psychofysiologie, KU Leuven, Medical School, Campus Gasthuis berg, B-3000 Leuven, Belgium Esther Peterhans Department of Neurology, University Hospital Zurich, CH-8091 Zurich, Switzerland v vi Heather L. Read Center for Molecular and Behavioral Neuro CONTRIBUTORS science, Rutgers University, Newark, New Jersey 07102 Kathleen S. Rockland Department of Neurology, 2007 RCP, College of Medicine, University of Iowa, Iowa City, Iowa 52242-1053 Hillary R. Rodman Department of Psychology and Yerkes Regional Primate Research Center, Emory University, Atlan ta, Georgia 30322 Anna Wang Roe Section in Neurobiology, Yale University School of Medicine, New Haven, Connecticut 06510 Marcello G. P. Rosa Vision, Touch and Hearing Research Centre, De partment of Physiology and Pharmacology, The University of Queensland, Queensland 4072, Aus tralia Jeffrey D. Schall Vanderbilt Vision Research Center, Department of Psychology, Vanderbilt University, Nashville, Ten nessee 37240 Peter H. Schiller Department of Brain and Cognitive Sciences, Mas sachusetts Institute of Technology, E25-634 MIT, Cambridge, Massachusetts 02139 Ralph M. Siegel Center for Molecular and Behavioral Neuro science, Rutgers University, Newark, New Jersey 07102 Keiji Tanaka Brain Science Institute, RIKEN, 2-1 Hirosawa, Wako-shi, Saitama, 351-01, Japan Daniel Y. Ts'o Department of Neurobiology, Rockefeller Univer sity, New York, New York 10021 S. Zeki Wellcome Department of Cognitive Neurology, University College London, London WClE 6BT, England Karl Zilles C. & 0. Vogt-Institute of Brain Research, Univer sity of Dusseldorf, D-40225 Dusseldorf, Germany Preface Over the last twenty-five years, there has been an extensive effort, still growing for that matter, to explore and understand the organization of extrastriate cor tex in primates. We now recognize that most of caudal neocortex is visual in some sense and that this large visual region includes many distinct areas. Some of these areas have been well defined, and connections, neural properties, and the functional consequences of deactivations have been studied. More recently, non invasive imaging of cortical activity patterns during visual tasks has led to an expanding stream of papers on extrastriate visual cortex of humans, and results have been related to theories of visual cortex organization that have emerged from research on monkeys. Against this backdrop, the time seems ripe for a review of progress and a glance at the future. One caveat important to emphasize at the very onset is that the reader may be puzzled or confused by the use of different terminologies. Individual investi gators commonly tend to favor different terminologies, but in general some prove more advantageous than others. As discussed by Rowe and Stone (1977) as well as by others, there is an unfortunate tendency for role-indicating names to lead to fixed ideas about function, in contrast to those that are more neutral and adaptable to new findings. Terminology differences may often seem trivial in that they reflect no basic disagreements (the names "MT," "V5," and "motion area" designate the same area; "blobs," "patches," and "puffs" designate the same subdivisions in area VI). Other differences reflect unresolved issues and disparate viewpoints. For example, is the region called "ventral V3" part of a larger visual area, V3, or is it a separate visual area, VP? Likewise, "dorsal V3" has been conceptualized as part of another area, DM, rather than as a compo nent of V3. As another example, it makes a great deal of difference for compari sons across cortical areas and across species if the sublayer in V 1 that projects to "MT" or "V5" is considered as layer IIIC or IVB. Terminology also reflects and, even more, guides ways of thinking about the visual system as a whole. Does the "extrastriate" rubric include visually related vii viii cortex in the parietal lobe? In the frontal lobe? Probably the most popular conceptualization of visual cortex has been a fixed hierarchy of processing areas. PREFACE However useful this approach may be, ambiguities exist in the assignment of individual areas to levels, as graphically portrayed in any number of M. C. Escher prints (for example, the 1960 lithograph Ascending and Descending, which is discussed by D. Hofstadter [1979, p. 13] in the context of strange loops and tangled hierarchies). Ultimately what is "ascending" or "descending" may be less than certain. Moreover, the terminology of hierarchy does not easily incorporate other aspects of visual processing, such as remporal factors or the possibility which the field is just beginning to consider-of dynamic interactions and groupmgs. The present volume covers a range of relevant topics. The first chapter, by Charles Gross, reviews the historical development of ideas and concepts about extrastriate visual cortex. As we approach the symbolic milestone of the next millennium, this chapter provides a gracious tribute to the antecedents of cur rent views and a reminder of the future progress we can expect. The following five chapters review and examine general features of cortical organization. Peter Schiller discusses the implications of conceptualizing processing at the level of single neurons and further considers the question of how functional specificity might emerge from neurons that are multifunctional within dynamically adjust able rather than dedicated systems. Chapters by Jon Kaas and Marcello Rosa consider theories of how cortex is divided into areas, criteria for identifying fields, and the nature of cortical representations. Lionel Nowak and jean Bullier cover the important and somewhat neglected topic of the temporal characteris tics of cortic-al processing (see also Guy Orban, Chapter 9). These authors sum marize evidence for fast and slow pathways as a compilation of earlier distinc tions between serial and parallel processing. Kathleen Rockland reviews recent advances in understanding of the microarchitecture of cortical connections, including features of spatial convergence and divergence, as well as laminar patterns of terminations, and discusses how these data might relate to theories of hierarchical organization. Chapters 7-13 focus on features of specific visual areas or groups of related visual areas. The second visual area, V2, is reviewed in separate chapters by Anna Roe and Dan Ts'o and by Esther Peterhans. Roe and Ts'o discuss modular specialization with V2, the remapping of visual space within the different classes of modules, and interaction between modules. Peterhans stresses the complex response properties of neurons in V2 and relates response features to aspects of perception. The structural and functional characteristics of area MT/V5 and associated areas are thoughtfully considered by Guy Orban, who also develops the issue of homologous areas in monkeys and humans. "Higher order" visual areas in the temporal lobe, including the region of the "superior temporal poly sensory area," and subdivisions of classical inferotemporal cortex are covered in separate chapters by Catherine Cusick and Keiji Tanaka. Ralph Siegel and Heather Read consider the functions of visual and visuomotor areas of posterior parietal cortex, while jeffrey Schall extensively reviews visuomotor areas of espe cially the frontal lobe. A single chapter, by Hillary Rodman and Tirin Moore, addresses the nor mal development of connections and function in extrastriate cortex, as well as the reorganization that occurs in extrastriate cortex as the result of injury or ix expenence. PREFACE The last group of chapters, 15-17, concentrates on the organization of extrastriate cortex in humans. The scope and detail contained in these chapters, largely based on research over the last few years, testify to the gathering momen tum of research on visual cortex of humans, both from functional mapping and from anatomical studies. From this perspective, the authors again treat the difficult issues of how to identify cortical areas, functional specializations, and network interactions. Results in human experiments benefit from and even re quire the conceptual context that has emerged from earlier studies in monkeys and other mammals, and the importance of continuing investigations in nonhu man mammals is clear. The closing chapter by Semir Zeki questions whether there is "one overall consciousness or many" while considering the possible con tributions of several visual areas or regions to human experience, such as those concerned with color or motion perception. (An additional invited chapter, "Topography and Function of Human Visual Cortex," had been prepared by Dr. Edgar A. DeYoe, but owing to unforeseeable circumstances could not be included in this volume.) Volume 12 of Cerebral Cortex on extrastriate cortex of primates complements Volume 10, Primary Visual Cortex in Primates. Volume 3 in this series, entitled Visual Cortex, covered aspects of organization and function in several different species. Important features of visual cortex development were discussed in Vol ume 7, entitled Development and Maturation of Cerebral Cortex. Overall, the authors of the present volume have provided an extensive and detailed review of extra striate cortex of primates. Some emphasis has been placed on features that are likely to exist in both monkey and human brains. Several relevant topics could have been considered more fully in this already large volume, and more points of view could have been expressed. However, some comparative and evolution ary aspects of visual cortex organization have been addressed in Volume 8 of this series, and future chapters and volumes on visual cortex can be anticipated. Kathleen S. Rockland Jon H. Kaas Alan Peters Iowa City, Nashville, and Boston References Hofstadter, D. R., 1979, Godel, Escher, Bach: An Eternal Golden Braid, Basic Books, New York. Rowe, M. H., and Stone, J., 1977, Naming of neurons, Brain Behav. Evol. 14:185-216.
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