LAIROTIDE YROSIVDA EETTIMMOC Giovanni Berlucchi Mary .B Bunge Robert .E Burke Larry E Cahill Stanley Finger Bernice Grafstein Russell .A Johnson Ronald .W Oppenheim Thomas .A Woolsey (Chairperson) The History of Neuroscience pargo~botuA i" n " by VOLUME 5 Edited by Larry R. Squire AMSTERDAM (cid:12)9 BOSTON (cid:12)9 HEIDELBERG (cid:12)9 LONDON NEW YORK (cid:12)9 OXFORD ~ PARIS (cid:12)9 SAN DIEGO SAN FRANCISCO (cid:12)9 SINGAPORE (cid:12)9 SYDNEY (cid:12)9 TOKYO REIVESLE Academic Press is an imprint of Elsevier Elsevier Academic Press 30 Corporate Drive, Suite 400, Burlington, Massachusetts 01803, USA 525 B Street, Suite 1900, San Diego, California 92101-4495, USA 84 Theobald's Road, London WC1X 8RR, UK This book is printed on acid-free paper. O Copyright (cid:14)9 2006 by the Society for Neuroscience. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permissions may be sought directly from Elsevier's Science & Technology Rights Department in Oxford, UK: phone: (+44) 1865 843830, fax: (+44) 1865 853333, E-mail: [email protected]. uoY may also complete your request on-line via the Elsevier homepage (http://elsevier.com), by selecting "Support & Contact" then "Copyright and Permission" and then "Obtaining Permissions." Library of Congress Catalog Card Number: 2003 111249 British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library ISBN 13:978-0-12-370514-3 ISBN 10:0-12-370514-2 For all information on all Elsevier Academic Press publications visit our Web site at www.books.elsevier.com Printed in the United States of America 06 07 08 09 01 11 9 8 7 6 5 4 3 2 1 Working together to grow libraries in developing countries moc.reivesle.www ] ww.bookaid.org ] gro.erbas.www =LSEVI ER BOOK AlIntD ...... ,~Sttib FC" ...... I t;oun" datlOl " Contents Samuel H. Barondes 1 Joseph E. Bogen 47 Alan Cowey 125 David R. Curtis 171 Ennio De Renzi 227 John S. Edwards 271 Mitchell Glickstein 301 Carlton C. Hunt 353 Lynn T. Landmesser 383 Rodolfo R. Llinãs 413 Alan Peters 453 Martin Raft 505 Wilfrid Rall 551 Mark R. Rosenzweig 613 Arnold Bernard Scheibel 657 Gerald Westheimer 697 Arnold Bernard Scheibel BORN: New York, NewYork January ,81 1923 :NOITACUDE Columbia College, B.A. (1944) Columbia University College of Physicians and Surgeons, M.D. (1946) University of Illinois, M.S. (neuroanatomy) (1953) APPOINTMENTS: University of Tennessee College of Medicine (1952) University of California, Los Angeles, (UCLA) School of Medicine (1955) Director, Brain Research Institute, UCLA (1987-1995) HONORS DNA SDRAWA (SELECTED): John Simon Guggenheim Fellow (1953-1954) George H. Bishop Lectureship, Washington University (1967) Distinguished Graduate Student Teaching Award, UCLA (1977) Fellow, American Academy of Arts and Sciences (1981) Fellow, Royal Norwegian Academy of Sciences (1983) Scientist of the Year, Alzheimer's Disease Research Association (1988) The Harriet and Charles Luckman Distinguished Teaching Award, UCLA (1997) The California Biomedical Research Association Distinguished Service Award (1998) Distinguished Life Fellow, American Psychiatric Association (2003) Research Pioneer Award, Alzheimer's Disease Research Association (2004) Arnold Scheibel is known for his studies of the detailed architecture of the spinal cord, brain stem, and cerebral cortex and introduced the module concept into central nervous system research. His description of the recurrent axonal projection of the nucleus reticularis thalami led to the gatelet theory of selective attention. His Golgi studies of human brain tissue extended our knowledge about the nature of neuronal changes in senile brain disease and in schizophrenia. He demonstrated correlations between human cognitive activity and structural change, and emphasized the role of plasticity in the living reactive brain. Arnold Bernard Scheibel p atterns in Neuropil, or "passion in neuropil" as some friends inter- preted it, was the title of my first and longest lasting research grant from the National Institutes of Health. The title spelled out the thrust and excitement that I have always felt for the fine structure in the nervous system. The fact that definable organizations of neurons, dendrites, and axons~the formal minuet of cerebellar Purkinje cells, the stately files of neocortical pyramids with their cathedraI-like dendritic arches, the overlapping swirls of inferior olive cells, or the town and coun- try spotting of cell villages throughout the brainstem core~might serve as vital substrates for cognition and behavior has, over the years, held me spellbound. With this in mind, I am grateful to the editor of The History of Neuroscience in Autobiography and the Society for Neuroscience for their invitation to look back over a period of more than half a century and share memories. In the Beginning Both sides of my family arrived in the United States in the decades after the Civil War. I should know when, but when one is young, one's forebear- ers seem uninteresting, and when one is old enough to care, the sources are gone. My maternal grandmother's family came from what was then part of southern Germany. My father's family emigrated from the old Austro- Hungarian Empire. Dad's family may have been vintners and there is apparently still a Scheibel aperatif or liqueur that is locally available in cen- tral Europe. Several of Dad's uncles were architects who practiced in the Cleveland-Youngstown area and this flair with a pencil did not escape him. (I often wonder whether my own preference for neurohistology and neural circuitry, the fine architecture of the nervous system, was not another and somewhat more derivative expression of that same gene line.) Although Dad trained to be an architect, he quickly found that he was better able to support his family by working in the "front office". And so he became an advertising and sales manager at a time when that was still a fairly exotic occupation. Among the experiences he shared with me was his being at the airfield on a foggy morning in 1927 when Lindbergh took off for Paris. Lindbergh was wearing a Bulova watch that Dad had just strapped Arnold Bernard Scheibel 659 on his wrist, a model that was then sold widely and successfully as the "Lone Eagle." Although Mother's education was cut short by financial reverses in her family (my maternal grandfather was saintly but not a business man), she remained until almost the end of her life an indefatigable reader and much of my interest in history and biography comes from her. During Mother and Dad's lives together, they had many differences but were in complete accord as to the importance of education for their young people. I was born in the northwest part of New York City and spent the first 24 years of my life in Manhattan and the Bronx. Although one of the largest cities in the world, it was still thought of (in its pre-Big Apple days) as "little old New York" by its inhabitants and that is still the way I remember it. I was my parents' only child but when I was 5, my aunt died in childbirth and her newborn son was raised by my parents. The grief surrounding the death of my aunt pervaded the family for a number of years and, coming as it did in the latter part of 1928, it is somehow mixed in my mind with the gathering anxiety and suffering that followed the stock market crash and Great Depression, which followed shortly there- after. As a "child" of the Depression, I still fancy I see its carryover in my unwillingness to spend money on myself. My cousin Milton, who became in every sense my brother was always gifted, always passionate, seldom pre- dictable. Unlike me, he was excellent in chess, good in mathematics, and a bit of a rebel. He became an economist and spent part of his adult life in Washington as Executive Assistant Secretary for Defense under several administrations. The most vivid memories of my childhood are those of the books that my dad arranged to come my way once I learned to read. The Mysterious Universe by Sir James Jeans leaves a special impress, with its discussions of recently discovered red giants like Betelguese and Antares, and the great matrix of dust that seemed to fill the galaxy. I also recall short biographies of the presidents, several lives of Lincoln, and Compton's Pictured Encyclo- pedia, which became my bible (and then Milton's) for several years. Inter- estingly, I remember no introductions to biology or much curiosity on my part in this direction (with the possible exception of Paul deKruif's Microbe Hunters). Outdoor sports were far down the list of encouraged activities. For my parents, life was a serious business and play was at best a slightly disreputable way of taking a necessary break from meaningful work. I am afraid that I was raised with the idea that professional athletes were lit- tle better than ne'er-do-wells or "bums." Under these conditions, it is not surprising that my graduation from Columbia College hung in the balance until I learned to swim (still a prerequisite for graduation at that time). I remain perpetually grateful to my parents for their commitment to my education. The middle 1930s were marked by two educative experiences that still live vividly for me. One was the trip around the country that we 660 Arnold Bernard Scheibel took by car in the spring of 1934. Dad was "between position" in those deep Depression days and they accordingly decided that it might be the time to kick over the traces for a few months. They assumed that I would learn more this way than I could possibly gain in my 6th grade school class. Considering that the vehicle of choice was a 1928 Nash without springs and an effective top speed of 38 mph, it was a valiant decision. In the nation's capital, it was a fortunate ll-year-old who could sit and listen while the United States Senate debated the proposed independence of the Philippine Islands or watch great jurists such as Brandeis, Cardoza, and Hughes (still meeting in the old Senate Chamber) consider the con- stitutionality of New Deal legislation. Further in our course toward the southwest lay the battlefields of the Civil War, Mississippi paddle wheel- ers, the gathering threat of the Dust Bowl, the memories of Tombstone, Arizona, and the citrus-loaded valleys of California. Luckiest of all was the fact that we made the trip just before the "homogenization of America" began. Aside from a few of the "new fangled" tourist cabins, the hotels or boarding houses we stayed in were often those that dated from frontier days. Another 10 years and they would all be gone, swallowed up by the omnipresent Hiltons, Sheratons, Best Westerns, and Holiday Inns. The other lucky break was winning a scholarship at Horace Mann, a small, private middle school-high school at the northern edge of New York City. Rigorous discipline, sometimes great teaching (with the unfortunate exception of biology), small classes (five of us struggled through 4 years of Latin together), and a very active sports and extracurricular program helped me begin to spread my wings. I liked drama, writing, and illustration and actually took up track and tennis. The pattern continued at Columbia College, the (then) surprisingly small college arm of Columbia University, to which I also won a scholarship. Immature as I was through my college years, I still recognized the privilege of being exposed to instructors such as Lionel Trilling, Mark Van Doren, and Charles Frankel. It was a heady time, but I still was not sure where I was going. As a liberal arts major, I took a short science sequence put together for those who would presumably have nothing further to do with these fields. The opening salvo here was a semester of physics taught by Professor John Dunning, then already deeply involved in the mysterious "Manhattan Project" going on behind plywood partitions just down the corridor in Pupin Hall. Things began to look up with geology and improved still more as fossils and life forms became the focus. I think I began to realize at that point that the study of living things was more interesting than I could have guessed. The strongly negative impression that had followed me from my unfortunate high school biology classes began to dissolve and I realized~ almost reluctantly~that perhaps what I had been reading in my Great Books classes, about the appropriate study of mankind being man, might just be true.