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AN ORAL HISTORY of NEUROPSYCHOPHARMACOLOGY the FIRST FIFTY YEARS Peer Interviews AN ORAL HISTORY OF NEUROPSYCHOPHARMACOLOGY THE FIRST FIFTY YEARS Peer Interviews Volume Two: Neurophysiology Copyright © 2011 ACNP Thomas A. Ban (series editor) AN ORAL HISTORY OF NEUROPSYCHOPHARMACOLOGY Max Fink (volume editor) VOLUME 2: Neurophysiology All rights reserved. No part of this book may be used or reproduced in any manner without written permission from the American College of Neuropsychopharmacology (ACNP). Library of Congress Cataloging-in-Publication Data Thomas A. Ban, Max Fink (eds): An Oral History of Neuropsychopharmacology: The First Fifty Years, Peer Interviews Includes bibliographical references and index ISBN: 1461161452 ISBN-13: 9781461161455 1. Electroencephalography and neurophysiology 2. Pharmaco-EEG 3. Sleep EEG 4. Cerebral blood flow 5. Brain metabolism 6. Brain imaging Publisher: ACNP ACNP Executive Office 5034A Thoroughbred Lane Brentwood, Tennessee 37027 U.S.A. Email: [email protected] Website: www.acnp.org Cover design by Jessie Blackwell; JBlackwell Design www.jblackwelldesign.com AMERICAN COLLEGE OF NEUROPSYCHOPHARMACOLOGY AN ORAL HISTORY OF NEUROPSYCHOPHARMACOLOGY THE FIRST FIFTY YEARS Peer Interviews Edited by Thomas A. Ban Co-editors Volume 1: Starting Up - Edward Shorter Volume 2: Neurophysiology - Max Fink Volume 3: Neuropharmacology - Fridolin Sulser Volume 4: Psychopharmacology - Jerome Levine Volume 5: Neuropsychopharmacology - Samuel Gershon Volume 6: Addiction - Herbert D. Kleber Volume 7: Special Areas - Barry Blackwell Volume 8: Diverse Topics - Carl Salzman Volume 9: Update - Barry Blackwell Volume 10: History of the ACNP - Martin M. Katz VOLUME 2 NEUROPHYSIOLOGY ACNP 2011 VOLUME 2 Max Fink NEUROPHYSIOLOGY Preface Thomas A. Ban Dedicated to the Memory of Jonathan O. Cole, President ACNP, 1966 PREFACE Thomas A. Ban In Volume 1 of this series, 22 clinicians and basic scientists reflected on their contributions to the “starting up” of neuropsychopharmacology (NPP). The cen- tral theme was the adoption of a behavioral methodology for the detection of psychotropic properties of drugs (see Brady, Cook, Dews and Stein, Volume 1). In Volume 2, the emphasis shifts to the detection of the action of drugs on the functional activity of the brain. There are two complementary approaches, with corresponding methodologies for neurophysiological research in NPP. One is focused on the electrical activity of the brain, using electroencephalography (EEG), and the other on cerebral blood flow (CBF) and metabolism, using brain imaging. The bridge between neurophysiological and behavioral (psychiatric) research is the conditioned reflex (CR). The origin of the idea that the condi- tioned reflex is at the core of mental activity is in Wilhelm Griesinger’s adoption of “reflex activity,” as the basis of all brain activity, and his introduction of the concept of “psychic reflex,” in the mid-1840s.1, 2 His notion that mental activity is reflex activity became a feasible proposition by the end of the 19th century after the delineation of the structural underpinning of reflex activity in the brain by Camillo Golgi’s description of “multi-polar cells” in the cerebral cortex,3 Ramón y Cajal’s recognition of the “neuron,” as a structural and functional unit of the nervous system,4, 5 and Charles Sherrington’s postulation that the “synapse” is the functional site of transmission from one neuron to another.6, 7, 8 The concept of “psychic reflex” became accessible to scientific study in the early 20th century through Ivan Petrovich Pavlov’s discovery of the CR.9 His findings that verbal signals could replace sensory signals in CR formation opened a path for research in which mental activity is perceived as CR activity in the second (verbal) signal system and studied with conditioning methods (see Ban, Volumes 4 and 9). Pavlov’s postulation that CR formation is based on the opening of formerly non-operating paths in the brain10, 11, 12 became demonstrable by the 1990s13 (see Kandel, Volume 3). Between the mid-1930s and late 1950s, a series of centrally acting substances, including alcohol,14 amphetamine,15 adrenaline and acetylcholine,16 mescaline,17 chlorpromazine and reserpine,18 was studied on CR activity in animals19, 20, 21 by Horsley Gantt and his associates in the United States. In the 1960s interest shifted from the study of the effect of psychotro- pic drugs on the CR in animals to the effect of these drugs on the human electroencephalogram. Recognition that electrical activity is a natural property x AN ORAL HISTORY OF NEUROPSYCHOPHARMACOLOGY – NEUROPHYSIOLOGY of the living brain dates back to Emil Du Bois-Reymond’s detection of electric currents (action potentials), from the peripheral nerves and brain of frogs with a galvanometer in the late 1840s..22, 23 His discovery that living brain generates electricity was substantiated independently in the mid-1870s by Richard Caton and Vasilij Jakovlevich Danilevskii, who recorded electrical currents and the fluctuations of these currents from the cerebral hemispheres of rabbits, mon- keys,24 and dogs.25 It was Hans Berger, in the 1920s, who first recorded electrical activity from the human brain through the intact skull. He introduced EEG, a technique for recording the electrical activity of the brain,26 and showed that the spontaneous waking EEG was “sensitive to” hypoxia, hypocapnia, barbiturates, bromides, caffeine, cocaine, chloroform, morphine, scopolamine, and insulin coma.27, 28 By the time the new psychotropic drugs were introduced in the 1950s, research on the electrical activity of the brain also included “sleep EEG,”29 with the rec- ognition of rapid eye movement (REM) sleep,30, 31 “topography,”32 “functional EEG,”33, 34 and the study of “evoked (event related) potentials” (ERP).35, 36, 37, 38 In the 1980s research was extended to the study of brain metabolism, as measured by cerebral blood flow. The roots of this research lie in the assump- tion frequently held in the 1920s, that deficient oxidative processes play an important role in the pathogenesis of mental syndromes and especially of schizo- phrenia.39 To stimulate oxygen consumption, carbon dioxide inhalation was tried in “catatonic” patients,40, 41 and nitrous oxide inhalation in “psychoses,”42 with equivocal results.43 Juda Quastel’s report in 1939 that oxygen uptake in the brain is much greater in vivo than in vitro,44 and the introduction of the first quantitative in vivo measurement of CBF by Dumke and Schmidt in 1943,45 led in the 1970s to the introduction of brain imaging techniques, suitable for the study of structure- activity relationships in the brain. Prior to their introduction, information on the living human brain was limited to structural changes revealed by pneumoencephalogra- phy46 and carotid angiography,47 and the only relatively consistent finding relevant to psychiatry was enlarged cerebral ventricles in schizophrenia first reported by Jacobi and Winkler in 1921.48, 49 Early studies on structure-function relationships, as for example the linking of “motor aphasia” to a lesion of the posterior part of the frontal lobe by Paul Broca in 1861,50 and of “sensory aphasia” to the posterior part of the temporal lobe by Carl Wernicke in 1874,51 were based on pathological anatomical find- ings, mainly in stroke patients.52 Research from the morgue moved to the study of the living brain of animals with the introduction of electrical stimulation by Fritch and Hitzig in 1870.53 Electrical stimulation was adopted in the study of the structure-function relationship in patients undergoing brain surgery by Wilder Penfield in the early 1950s.54 His findings, that electrical stimulation of certain parts of the temporal Preface xi lobe (TL) evoked déjà vu experiences and old memories, contributed to the recognition of the role of the TL in memory storage.55 Introduction of the chronic implanted electrode technique in animals by Walter Rudolf Hess in 1932, and his studies of the diencephalon, thalamus and hypothalamus,56 extended the scope of the use of electrical stimulation in the study of structure-function relationships in the brain. In 1954, James Olds and Peter Milner found that rats with metal electrodes implanted in their nucleus accumbens, repeatedly pressed a lever controlling the delivery of an electric current, as if they were seeking pleasure by electrical activation of the region.57 The discovery of “a pleasure center,” involved in the “reinforcement” of behav- ior by “reward,” provided the key for the identification of the cerebral structures involved in “operant (instrumental) conditioning,” a form of learning, in which a motivational (emotional) factor, i.e., pleasure or punishment, is involved in the acquisition of CR.58 In 1939 James Papez suggested that emotions are processed through a reverberating circuit in the brain – which consists of the cingulate gyrus, hippo- campus, amygdala, mammillary bodies, hypothalamus and anterior thalamus – before becoming subjectively-experienced feelings in the cerebral cortex.59, 60 The “Papez circuit” is also referred to as “limbic lobe,” because its site corre- sponds with Paul Broca’s “great limbic lobe,”61 and “visceral brain,” because of its numerous connections with the autonomic nervous system.62 The subjective experience of feelings depends on the state of arousal of the cerebral cortex regulated by brain stem structures. In 1949 Moruzzi and Magoun reported that electrical stimulation of the ascending reticular forma- tion in the brain stem of animals produced a generalized activation and desyn- chronization of the EEG with
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