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Neurosciences Research Program Neurosciences Research Program Volume 6, Number 1 May 1, 1968 BIOLOGY OF DRIVES A Report of an NRP Work Session chaired by Elliot S. Valenstein NEUROSCIENCES RESEARCH PROGRAM 280 Newton Street, Brookline, Massachusetts 02146 Telephone: Area Code 6 17,522-6700;Cable: NEUROCENT NRP CENTER STAFF Chairman Francis 0. Schmitt Progra m Director Communications Director Gardner C. Quarton Theodore Melnechuk Business Manager Managing Editor & Librarian L. Everett Johnson George Adelman A dministra tive 0fficer Katheryn Cusick Assistant to the Chairman Harriet E. Schwenk Resident Scientists Frank Morrell Joe M. Parks Writer-Editors Frederick E. Samson, Jr. Yvonne M. Homsy Catherine M. LeBlanc Secretaries Julie Sofield Frances M. Delagi Veronica Twomey Patricia Anne Dimond Patricia E. West Cynthia J. Mello Jane I. Wilson Library Assistant Audio- Visual Technician Linda T. Knowles Wardwell F. Holman SPONSORSHIP AND SUPPORT The Neurosciences Research Program, sponsored by the Massachusetts Institute of Technology, is an interdisciplinary, interuniversity organization the primary goal of which is to facilitate the investigation of the physical basis of mental processes includ- ing memory and learning. To this end the NRP, as one of its activities, conducts scientific meetings to explore crucial problems in the neurosciences and publishes the results of these Work Sessions in this Bulletin. Nbis supported in part by National Institutes of Health Grant No. GM 1021 1-06, National Aeronautics and Space Admin- istration Grant No. Nsg 462 Amendment 4, Office of Naval Research Grant Nonr(G)- 0002647, The Rogosin Foundation, and Neurosciences Research Foundation, Inc. Neurosciences Res. Prog. Bull., Vol. 6, No. 1 1 BIOLOGY OF DRIVES A report of an NRP Work Session held April 3-5, 1966 Elliot S. Valenstein The Fels Research Institute Yellow Springs, Ohio Yvonne M. Homsy and Anne H. Rosenfeld NRP Writer-Editors CONTENTS List of Participants ..................3 I. Introduction .................... 5 11. The Diversity of Motivational Mechanisms and the Definition of Drive .................8 A. The Diversity of “Motivational” Mechanisms .....8 B. Comparative and Developmental Considerations in the Study of Motivation ............15 C. Potentiating Operations .............19 111. Drive as a General Activating Factor .......... .22 A. Drive and Arousal .............. .23 B. Fractionation and Differentiation of “Arousal” Processes ..................27 C. Neuroendocrine Setting of Behavioral Propensities ...3 1 2 Biology of Drives IV . Drive and Brain Mechanisms ..............37 A . Central Nervous System : Species-specific Behavior and Drive ..............37 B . Can Studies of Feeding Centers Provide a Model forDrive? .................45 C . Drive Centers and a Drive State ..........50 V . Drive. Motivation. Reinforcement. and Learning .......58 A . The Relationship between Drive and Learning .....58 B. Reinforcement and Motivation ..........62 C . Drive. 1ncentive.Motivation. and Reinforcement ....67 VI . Biological Mechanisms of Reinforcement .........73 A . Neurochemical Substrates of Reinforcement .....73 VI1 . Epilogue ...................... 85 Bibliography .................... 91 Index ....................... 108 Neurosciences Res. Prog. Bull., Vol. 6, No. 1 3 LIST OF PARTICIPANTS Dr. Daniel E. Berlyne Dr. Daniel S. Lehrman Department of Psychology Institute of Animal Behavior University of Toronto Rutgers University Toronto, Canada Newark, New Jersey Dr. Dalbir Bindra Dr. Neal E. Miller Department of Psychology The Rockefeller University McGill University New York, New York Montreal, Canada Dr. Carl Pfaffmann Dr. Joseph V. Brady The Rockefeller University Division of Neuropsychiatry New York, New York Walter Reed Army Institute of Research Dr. Warren W. Roberts Washington, D. C. Department of Psychology University of Minnesota Dr. Robert W. Goy Minneapolis, Minnesota Department of Reproductive Physiology and Behavior Dr. Francis 0. Schmitt Oregon Regional Primate Center Department of Biology Beaver ton, Oregon Massachusetts Institute of Technology Dr. Sebastian P. Grossman Cambridge, Massachusetts Department of Psychology The University of Chicago Dr. Larry Stein Chicago, Illinois Department of Psycho- pharmacology Dr. Robert A. Hinde Wyeth Laboratories Sub-Department of Animal Radnor, Pennsylvania Behavior University of Cambridge Dr. Philip Teitelbaum Madingley, Cambridge, England Department of Psychology University of Pennsylvania Dr. John I. Lacey Philadelphia, Pennsylvania Department of Psychophysiology- Neurophysiology Dr. Elliot S. Valenstein The Fels Research Institute The Fels Research Institute Yellow Springs, Ohio Yellow Springs, Ohio NOTE: NRP Work Session summaries are reviewed and revised by participants prior to publication. Neurosciences Res. Prog. Bull., Vol. 6, No. 1 5 I. INTRODUCTION A Work Session on the “Biology of Drives” was held on April 3 to 5, 1966. The purpose of this Work Session was to attempt to find some agreement, or at least to explore different views, on the concepts of drive and motivation. The term drive is closely related to biological needs and may apply to behavior ranging from eating to that characterized by some as exploration. The need for novel sensory input may be as physiologically based as the more obvious need for nutrients. As drives are viewed as representing physiological states, they may also serve as interoceptive stimuli to activate and direct behavior, and changes in “drive stimuli” may be critically involved in the learning process. It is useful in delineating the construct of drive as a central determinant of response variability to exclude the short-duration reversible changes in responsivity, such as those attributed to fatigue or habituation, and the long-duration changes, such as those considered to be learning. It has been suggested that the term drive is similar to the word force as used by physicists, and it is worthwhile to note their illuminating similarities. In both cases, the terms represent conditions that impart movement, or have the potential to impart movement, to an object or an organism, and both are inferable from observable events. Neither the physicist nor the psychologist expects to see a force or a drive, but here the similarities seem to end. Force can be quantified and, whether it is measured in terms of pounds, newtons, or dynes, the reference is to the movement of some specified weight at a specified speed. Since force possesses a unitary dimension, one can make meaningful quantitative comparisons. The situation is quite different with respect to the term drive. It is anything but a unitary concept as is explicitly elaborated in Hinde’s presentation, “The Diversity of ‘Motivational’ Mechanisms.” We use drive to refer to the state of the organism which produces activity although we are compelled to admit that there may be times when at least some would suggest the existence of a high drive level in the absence of any overt activity. Hinde sees the existence of “nonactive” behaviors, such as some fear responses and incubation in birds, as one of the objections to thinking of drive as an energy concept. Brady maintains that this lack of a specific referent for the terms drive and motivation indicates that there may be little, if any, scientific value in employing the concepts. Teitelbaum responds to the issue by suggesting an operational definition to distinguish 6 Biology of Drives between drive and motivation. According to Teitelbaum’s view, one is justified in using the term motivation only when an animal can learn an arbitrary response to obtain an appropriate goal object. In this way it may be possible to separate the consummatory behavior related to drive states from motivation. A blowfly’s responsiveness to sweet solutions may vary with periods of deprivation, but, since attempts to train this animal to perform an instrumental response in order to obtain a sweetened solution have failed, one can only speak of changes in drive level but not motivation. Lehrman and Hinde argue that, since the comparative liter- ature is replete with examples of such diverse mechanisms underlying plasticity, it would be most difficult (if not impossible) to apply any operational definition of motivation to all animals. Gnimals can exhibit plasticity in one situation and not in others. In the absence of an exhaustive study of an animal’s adjustive mechanisms in response to different environmental demands, it would be dangerous to conclude that a slight change in the testing situation would not result in the learning of an instrumental response. In part precipitated by studies of the “reticular activating system,” other terms have crept into our vocabulary which have much in common with drive as a nonspecific activator. We speak of “activation levels” and “arousal levels” and consider their relationship to learning, reinforcement, and ultimately performance. Berlyne’s presentation elaborates on the usefulness of these concepts and the role of the environment-organism interaction in modulating “arousal,” while Lacey, in stressing the diffi- culties in measuring any generalized “arousal,” is forced to question the value of the concept. It is a common observation that both nonspecific and some goal-directed behaviors go through periods of high and low activity, which it is only natural to presume are related to changes in the state of the organism. These changes in the state of the organism are not only determinants of fluctuations in activity, but also account for the fact that the
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