The Limbic System SECOND EDITION The Limbic System SECOND EDITION Robert L. Isaacson State University of New York at Binghamton Binghamton, New York Springer Science+Business Media, LLC Library of Congress Cataloging in Publication Data Isacson, Robert Lee, 1928- The limbic system. Bibliography: p. Includes index. 1. Limbic system. I. Title. [DNLM: 1. Limbic system. WL 307 I73L] QP383.2.1821982 599.01'88 82-9077 ISBN 978-1-4757-6757-5 ISBN 978-1-4757-6755-1 (eBook) AACR2 DOI 10.1007/978-1-4757-6755-1 © 1982 Springer Science+Business Media New York Originally published by Plenum Press, New York in 1982 Sofleover reprint of the hardcover 2nd edition 1982 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 Preface to Second Edition Like the first edition of The Limbic System, this book was written as an introduction to the neurobehavioral study of the limbic system; it is intended for students of brain-behavior relationships in psycho biology, neurosciences, physiology, and anatomy, as well as those with clinical interests in the brain and its actions. Also, like the first edition, the book is flavored with my own orientation to the field based on some 22 years of active research. My own perspective on the limbic system is described in the final chapter. It is an overview of how the different portions of the brain may work in concert to produce mental and behavioral actions. I do not hold to the ideas presented there with strong commitment. This chapter is only an overview and I am sure that other and better comprehensive approaches will be generated. Nevertheless, it helps to have some perspective to integrate diverse facts and information. In regard to other biases, I still believe that the greatest gains in understanding brain organization will come from studies using the rat, the cat, and the rabbit. Research using nonhuman primates takes even more time and expense now than it did in 1974, and the gen eration of useful information with such species must therefore pro ceed slowly. Research using humans which is directed toward un derstanding brain function must depend on examination of the effects of disease or accidents. Because of this limitation, the study of indi viduals with similar damage can occur only rarely-if at all. Further more, as emphasized throughout this edition, the observable con sequences of brain damage depend on genetic endowment, pre- and postdamage experiences, and other factors, as well as on the nature v vi Preface to Second Edition of the damage. Therefore, I doubt that much fundamental knowledge of brain damage will come from studies with the human. As I men tioned in the preface to the first edition, I am convinced that the use of the lesion technique will provide the mainstay of future research, although the goals and methods of lesion studies will be different in the next 10 to 20 years. Despite its difficulties, which are really similar to those using brain stimulation or other approaches,l there is no other method that offers a better way to determine just how large portions of the central nervous system interact with each other. In the future such studies will use new neurochemical techniques that offer the promise of distinguishing cellular from axonal damage and more biochemical and pharmacological methods to determine the le sion-induced effects. In 1974 I admitted that the book could not be considered com prehensive. Only some 500 references were cited at that time, and it was clear that many more could have been included. I tried to con centrate on the ones that seemed most important for current research and theory. In this second edition there are almost 1,400 references and, once again, I am sure that they are not comprehensive. The large increase in the number of references that needed to be noted is a sign of the scientific information explosion in general and an increased interest in limbic system structure and function. In the original preface I expressed my appreciation to the graduate students who were then in the laboratory. Their thoughts, ideas, and energies about our research were always exciting. Now, another group of students is in the laboratory, helping to unravel the secrets of the brain and behavior. These are John Hannigan, Jr., Joe E. Springer, and Jeanne Ryan. Daniel K. Reinstein, now at the Massachusetts Institute of Technology, was with me for most of the period I was rewriting the book and has helped shape my ideas about the limbic system. To them, my thanks. They keep me young, at least in spirit. In the last year or so, I have also had the benefits of collaboration of two more senior associates, David Hertzler (SUNY-Oswego) and Alex Poplawsky (Bloomsburg State College), who visited the laboratory for extended periods. 1 Isaacson, R. L. Brain stimulation effects related to those of lesions. In M. M. Patterson & R. P. Kesner (Eds.), Electrical stimulation research techniques. New York: Academic Press, 1981, pp. 205-220. Preface to Second Edition vii When starting this second edition, I wrote to several friends who are recognized authorities in different areas of limbic system research to solicit their ideas about what should be emphasized. They all re sponded with their ideas and I have tried to follow their suggestions. However, it has taken so long to finish this second edition that I am sure they would make additional or different suggestions now if asked. I appreciate their help in orienting me to their perspectives. Those to whom I wrote originally were Graham Goddard, Sebastian P. Gross man, and Gary W. Van Hoesen. While writing the book I benefited from discussions of the content and ideas with a number of people, especially Peter Donovick, Linda Spear, Paul MacLean, Per Andersen, Larry Swanson, Daniel P. Kimble, and Gyorgy Buzsaki. It may be hard to recognize the specific contributions made by any of these people to the book itself but they produced strong influences on my thinking and my writing. No book can be prepared without excellent secretarial assistance. In the first edition it was Mrs. Virginia Walker, at the University of Florida, who was indispensable. This time it was Mrs. JoAnn Kovalich at SUNY-Binghamton. Without her help, the book would not have been finished. Thanks. ROBERT L. ISAACSON Binghamton, New York Preface to First Edition While this book is intended to be an introduction to the neuroanatomy of the limbic system and to studies of the behavior of animals in which the limbic system is stimulated or damaged, it is primarily intended for advanced students of brain-behavior relationships. I have as sumed the reader to have some understanding of the structure of the brain, of basic neurophysiology, and of modern behavioral tech niques. It has been written for students in graduate programs in psychobiology, physiological psychology, and the neurosciences, but it also should be of interest to some medical students and to others with catholic interests in the biology of behavior. In the first chapter, I review the structure of the limbic system and in subsequent chapters consider the behavioral effects of lesions and stimulation of components of the limbic system. Supplemental information derived from recording the electrical signals of the brain is included where it seems appropriate. The final chapter presents a perspective of the limbic system related to brain stem mechanisms and the neocortex. Understanding the behavioral contributions of the limbic system presupposes understanding how the limbic system in teracts with other systems of the brain. Even though there is only one chapter overtly devoted to theo retical issues, various biases of mine influence all chapters. Anyone reading the book with a critical attitude will soon be aware of them. I would like to alert the reader to some of them ahead of time. Simply put: the book is flavored by my own orientation to re search and reading. For the past 15 years, I have been involved in research programs which were directed at elucidating the effects of limbic system destruction on behavior in the rat, cat, and rabbit. ix x Preface to First Edition Therefore, studies using these species and using lesion techniques probably are overly represented. Other biases of mine will be appar ent. In many later chapters, consideration is given to the behavioral correlates of electrical rhythms recorded from various limbic struc tures. In these chapters, the effects of electrical stimulation of various limbic structures on behavior are also discussed. Yet, while these topics are discussed, much less emphasis is placed on them and on studies using other species with which I am less well acquainted. My biases, the topics and areas given emphasis, come in part from my own research. This research is based on beliefs pertaining to research strategies. I believe that the most substantial knowledge available about the limbic system comes from the use of stimulation and lesion techniques in rats and cats. There is far less useful infor mation available today arising from work using other techniques and other species. Since a great deal more information is needed before an adequate conceptualization can be achieved, this information must come from careful studies using species like the cat and the rat. These animals are amenable to laboratory experimentation and can be used in sufficient numbers so as to provide this information relatively quickly. If we had to rely on studies using nonhuman primates, our under standing of the behavioral contributions made by the limbic system could be greatly delayed. The rat is a necessary laboratory tool, both for neuroanatomists and for those interested in the behavioral con tributions of the limbic system. The cat makes its greatest contribution to neurophysiological research. The other obvious orientation of the book is the emphasis placed on understanding the functions of the limbic system as revealed by destruction of its parts. For 30 years and more, investigations have been underway of the behavior of animals which have had some portion of the limbic system destroyed by aspirative or by electrolytic lesions. Seeking to understand a structure by destroying it is a useful technique for the behaviorally oriented neuroscientist, although it has many faults, dangers, and traps for the unwary experimenter. Never theless, equally dangerous are the faults and traps awaiting those using other techniques in the neurobehavioral sciences. In reality, the lesion technique is no better or worse than any other, such as re cording electrical rhythms or recording from single units. Satisfactory understanding of the limbic system will not come from the studies of animals with lesions of this or that part of the limbic system or Preface to First Edition xi from the analysis of single cell activity but from theoretical contri butions derived from the use of all available techniques. Theories must be evaluated on the basis of whether they make sense of the facts to be explained. A theory must provide a meaningful and useful synthesis of information derived from any and all techniques. Even though the book has a large number of references to pub lished work on the limbic system, it is not a truly comprehensive review of the literature. I doubt that a totally comprehensive review of the experimental literature can be done by anyone. Still, I have tried to represent fairly most of the important studies which have influenced present research directions. Nevertheless, some of my col leagues will be offended by not finding their favorite studies in the book. To them I apologize for the oversight. On the other hand, some studies were omitted because I felt that they had not added a great deal to our understanding of the limbic system. I have tried to select articles from the literature which highlight information of the greatest importance for the understanding of the limbic system. Theories of limbic function have not been emphasized in this book, since I do not think there are any adequate theories of limbic system function. My theory, presented in the last chapter, is offered in an apologetic fashion. At best, it offers only the broadest outlines for a schema of limbic system function. In almost a playful spirit, it is offered as a potential stimulant to others. In a lecture given at the University of Florida in the spring of 1973, Paul MacLean suggested that scientists did not belong in the laboratory if their work and the generation of ideas were not fun. All of my days in the laboratory have been rewarding. I cannot imagine a more fascinating or more interesting life than struggling to find out how the brain works. There fore, I hope the reader will evaluate the last chapter with understand ing. But my research and study of the limbic system is motivated by far more than the joy it provides. It is motivated by the belief that learning at least some of the secrets of the nervous system will be of value to mankind. This is not an intangible or abstract motive. Re tarded children are very real, as are people with other brain disorders. While much of my present work is directed toward problems of gen eral interest rather than toward retardation specifically, the ultimate goal is to better understand the human condition and to help provide information on which effective therapies and treatments for the brain damaged can be based.
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