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Behaioral Medicine

Ornish D, Scherwitz L W, Billings J H, Brown S E, Gould K L, initially termed comparative psychology. For many
Merritt T A, Sparler S, Armstrong W T, Ports T A, Kirkeeide years the flagship journal devoted to comparative
R L, Hogeboom C, Brand R J 1998 Intensive lifestyle changes behavior and its biological bases was the Journal of
for reversal of coronary. Journal of the American Medical
Comparatie and Physiological Psychology, published
association 280: 2001–7
Pavlov I P 1927 Conditioned Reflexes (trans. Anrep G V). by the American Psychological Association. But in-
Oxford University Press, London creasingly the fields of comparative studies of behavior
Potter J, Steinmetz K 1996 Vegetables, fruit and phytoestrogens and physiological analysis of behavioral phenomena
as preventive agents. In: Stewart B W, McGregor D, Kleihues went separate ways. Consequently, in 1983 the journal
P (eds.) Principles of Chemopreention. International Agency was divided in two separate journals. Following is a
for Research on Cancer, Lyon, France portion of the editorial justifying this action by the
Rooney B L, Murray D M 1996 A meta-analysis of smoking then editor:
prevention programs after adjustment for errors in the unit of
analysis. Health Education Quarterly 23: 48–64 The separation of the Journal of Comparatie and Physio-
Shumaker S, Grunberg N 1986 Introduction to proceedings of logical Psychology into two companion journals, Behaioral
the National Working Conference on Smoking Relapse. Neuroscience and the Journal of Comparatie Psychology,
Health Psychologist 5: 1–2 acknowledges the current state of the sciences concerned with
Spiegel D, Bloom J R, Kraemer H C, Gottheil E 1989 Effect of biology and behavior. Behavioral neuroscience is the broader
psychosocial treatment on survival of patients with metastatic contemporary development of physiological psychology. In
breast cancer. Lancet. 2(8668): 888–91 all animals above the level of the sponge, the nervous system
Wolff H G 1953 Stress and Disease. Thomas, Spring- is the organ of behavior. All biological and behavioral
field, IL variables that exert any influence on behavior must do so by
acting on and through the nervous system.
B. N. Henderson and A. Baum Traditionally, physiological psychology emphasized certain
approaches and techniques, particularly lesions, electrical
stimulation, and electrophysiological recording. Study of the
biological bases of behavior is now much broader and
includes genetic factors, hormonal influences, neuro-
Behavioral Neuroscience transmitter and chemical factors, neuroanatomical sub-
strates, effects of drugs, developmental processes, and en-
vironmental factors, in addition to more traditional
Behavioral and cognitive neuroscience are two sides of approaches. All these variables act ultimately through the
the same coin. The field as a whole is concerned with nervous system. All these areas of investigation are entirely
the neuronal and biological bases of behavior and appropriate for Behaioral Neuroscience – all studies in which
experience, that is to say, psychology. Indeed, the biological variables are manipulated or measured and relate
distinction between ‘behavioral’ and ‘cognitive’ is to behavior, either directly or by implication.
The contemporary meaning of the term ‘behavioral neuro-
rather arbitrary. Operationally, behavioral neuro- science’ is almost as broad as ‘behavior’ itself. Behavioral
science might be defined by the kinds of papers neuroscience is the field concerned with the biological
published in the journal Behaioral Neuroscience and substrates of behavior. Judging by its current rate of
like journals. The emphasis is on biological studies of development, behavioral neuroscience could well become a
such basic phenomena as sensation and perception, dominant field of science in the future (Thompson 1983, p. 3).
motivation, learning, and memory, phenomena that
occur similarly in humans and other animals. Most By the time this occurred, the term ‘behavioral
studies are done on infrahuman animals, including neuroscience’ was increasingly used to describe that
non-human primates. By the same token, most of the aspect of the interdisciplinary field of neuroscience
papers in the journal Cognitie Neuroscience and like concerned with the biological basis of behavior. By the
journals are on humans, with imaging studies pre- same token, ‘cognitive neuroscience’ was adapted
dominating. But many behavioral neuroscientists more recently to emphasize that aspect of neuroscience
study basic psychological processes in humans, and concerned with the more complex phenomena of the
many who consider themselves cognitive neuro- ‘mind.’
scientists study such processes in animals.
The field of neuroscience is relatively recent as
a ‘unified’ discipline; the first meeting of the Society 1. Historical Deelopments
for Neuroscience was held in 1971. The goal was
to merge the disparate fields concerned with the The development of the two aspects of the study of
study of the nervous system, e.g., neuroanatomy, ‘brain, mind and behavior’ is best viewed historically.
neurophysiology, neurochemistry, neuropharmaco- As noted above ‘behavioral neuroscience,’ the subject
logy, neuroendocrinology, physiological psychology, of this article, is the modern equivalent of ‘physio-
clinical neurology, etc., into one interdisciplinary logical psychology.’ Since Wilhem Wundt, the founder
field. of the modern field of psychology in the 1870s and
There was a very long tradition in psychology of the himself a physiologist, titled his epoch making text
study of animal behavior and its biological bases, GrundzuW ge der Physiologische Psychologie (1874), we

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begin with his era. Wundt insisted on the experimental clinical neurology and neuroanatomy, were vigorous
approach to all questions of psychology: the challenge and exciting fields at the beginning of the twentieth
was to find experimental methods to understand century.
psychological processes. The most appropriate The development of instrumentation was still
methods, e.g., measures of brain activity, did not of another very important factor. The human EEG was
course exist in his day. Indeed, the only methods rediscovered in 1929 (Berger) and the method applied
available were simple measurements of behavior like to animal research in the 1930s. At the turn of the
reaction time, and verbal descriptions of experience by century the major experimental techniques for the
introspection, both of which were used in Wundt’s study of brain function were ablation and electrical
laboratory. stimulation. Neuroanatomy was in its descriptive
Wundt’s career illustrates well the roots of the phase; the monumental work of Ramon y Cajal was
developing field of physiological psychology. After a published over a period of several decades beginning
year of study at Heidelberg in basic medical sciences, near the end of the nineteenth century. Neuro-
he moved to Berlin and worked for two years with chemistry was in its purely descriptive phase.
Johannes Mu$ ller, a founder of modern, experimental Merely because techniques and basic knowledge
physiology. Mu$ ller, incidentally, was a vitalist and were limited does not mean that the field was quiescent.
believed that it would never be possible to measure the On the contrary, there was ferment over a number of
speed of the nerve impulse; his own student Helmoltz basic issues ranging from the mind-brain problem to
proved him wrong. Wundt then took his doctorate in localization of function and the nature of neuronal
medicine and was appointed Dozent in physiology at interactions. Add to this the ferment developing in
Heidelberg. Helmholtz then joined the same institute psychology as John Watson took on the older es-
and they were colleagues, but not collaborators, for a tablishment and behaviorism began to dominate. This
period of 13 years. Wundt later moved to Leipzig and is the background from which modern physiological
established the first formal psychological laboratory, psychology developed.
in 1879.
Wundt’s text and research at Leipzig represented a
brilliant experimental physiologist grappling with 2. Lashley and the Engram
problems of the mind and mental events, applying the
experimental methods of science as best he could. In Karl S. Lashley is the most important figure in the
fact, the bulk of the research done in the Leipzig development of physiological psychology in America
laboratory was on sensation and perception, and on (see Lashley, Karl Spencer (1890–1958)). He obtained
reaction time. his Ph.D. in zoology at Johns Hopkins University. At
Another laboratory of physiological psychology Hopkins he studied with John Watson and was heavily
was founded at about the same time, that of William influenced by Watson’s developing notions of beha-
James. James went to Harvard and Harvard Medical viorism. While there he also worked with Shepherd
School and studied with the great naturalist, Louis Franz at a government hospital in Washington; they
Agassiz. James was then, after a year of study abroad, published a paper together in 1917 on the effects of
appointed an instructor in physiology at Harvard cortical lesions on learning and retention in the rat.
College in 1872. He actually established an informal Lashley then held teaching and research positions at
physiological psychology laboratory there in 1877 and the University of Minnesota (1917–26), the University
taught a graduate course on the relations between of Chicago (1929–35) and then at Harvard from 1935
physiology and psychology. until his death in 1958. During the Harvard years
In sum, the two founders of modern psychology, he spent much of his time at the Yerkes Primate
Wundt in Germany and James in America, were both Laboratory in Orange Park, Florida.
physiologists by training. More than a third of William Lashley devoted many years to an analysis of brain
James’ extraordinary and influential text, Principles of mechanisms of learning, using the lesion-behavior
Psychology (1890), was devoted to the nervous system. method, which he developed and elaborated from the
However, this does not necessarily mean that Wundt work with Franz. During this period, Lashley’s theor-
and James attempted to reduce psychology to physi- etical view of learning was heavily influenced by two
ology; rather they proposed that the subject matter of congruent ideas: localization of function in neurology
psychology should be studied scientifically, as in and behaviorism in psychology.
physiology. This view could equally apply to be- Localization of function was the major intellectual
havioral and cognitive neuroscience as it exists today. issue concerning brain organization at the turn of the
There were powerful intellectual pressures moving century. An extreme form of localization was popular
bright young physiologically inclined psychologists at at the beginning of the nineteenth century with Gall
the turn of the century. The theory of evolution was a and phrenology. The neurologist Flourens moved
major force. The field of physiology, in particular away from that position by arguing for no specific
neurophysiology, in the work of Sir Charles localization within the cerebrum. However, Broca’s
Sherrington and then Lord Adrian, together with discovery of a speech center in 1861 began to move the

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pendulum back. The critical and classic study on ization of Behaior (1949) is a landmark, as is Sperry’s
localization was that of Fritsch and Hitzig (1870). work on consciousness.
They stimulated the cerebral cortex electrically and We now treat the development of the major areas of
defined the motor area: not only was it localized to a research that characterize modern behavioral neuro-
particular region of the cortex but there were specific science, namely neural and biological substrates of
organization and localization within it. learning and memory, motivated behaviors, and sen-
In Watson’s behaviorism, the learning of a par- sory processes.
ticular response was held to be the formation of a
particular set of connections, a series set. Conse-
quently, Lashley argued, it should be possible to
localize the place in the cerebral cortex where that 3. Learning and Memory
learned change in brain organization was stored: the
engram. (It was believed at the time that learning 3.1 Palo
occurred in the cerebral cortex). Thus, behaviorism
and localization of function were beautifully con- Lashley’s pessimistic conclusions in his 1929 mono-
sistent: they supported the notion of an elaborate and graph put a real damper on the field concerned with
complex switchboard where specific and localized brain substrates of memory. But there were other
changes occurred when specific habits were learned. major traditions developing. Perhaps the most im-
Lashley set about systematically to find these portant of these was the influence of Pavlov. His
learning locations—the engrams—in a series of studies writings were not readily available to Western
culminating in his 1929 monograph Brain Mechanisms scientists, particularly Americans, until the pub-
of Intelligence. In this study he used mazes differing in lication of the English translation of his monumental
difficulty, and made lesions of varying sizes in all work Conditioned Reflexes in 1927. It is probably fair
different regions of the cerebral cortex of the rat. The to say this is the most important single book ever
results of this study profoundly altered Lashley’s view published in the field of behavioral neuroscience.
of the brain organization and had an extraordinary Pavlov developed a vast and coherent body of em-
impact on the young field of physiological psychology. pirical results characterizing the phenomena of con-
The locus of the lesion is unimportant; the size is ditioned responses, what he termed ‘psychic reflexes.’
critically important, particularly for the more difficult He argued that the mind could be fully understood by
mazes. These findings led to Lashley’s two theoretical analysis of the higher order learned reflexes and their
notions of equipotentiality and mass action, i.e., all brain substrates.
areas of the cerebral cortex are equally important (at W. Horsley Gantt, an American physician, worked
least in maze learning), what is critical is the amount with Pavlov for several years and then established a
removed. Pavlovian laboratory at Johns Hopkins. He trained
Lashley’s interpretations stirred vigorous debate in several young psychologists, e.g., Roger Loucks and
the field. Walter Hunter, an important figure in Wolf Brogden, who became very influential in the
physiological-experimental psychology at Brown Uni- field. Perhaps the most important modern behavioral
versity who developed the delayed response task, analyses of Pavlovian conditioning are the work of
argued that in fact the rat was using a variety of Robert Rescorla and Allan Wagner.
sensory cues: as more of the sensory regions of cortex Although Pavlov worked with salivary secretion,
were destroyed, fewer and fewer cues became avail- most studies of classical conditioning in the West
able. Lashley and his associates countered by showing tended to utilize skeletal muscle responses, a' la
that removing the eyes has much less effect on maze Bechterev. Particularly productive have been Pav-
learning than removing the visual area of the cortex. lovian conditioning of discrete skeletal reflexes (e.g.,
Others argued that Lashley removed more than the the eyeblink response) characterized behaviorally by
visual cortex. Out of this came the long series of lesion- Isadore Gormezano and Allan Wagner and analyzed
behavior studies analyzing behavioral ‘functions’ of neuronally by Richard Thompson and his many
the cerebral cortex. Initially, studies focused on sen- students; and fear conditioning, first characterized
sory areas of the cerebral cortex. Beginning in the behaviorally by Neal Miller and analyzed neuronally
1940s several laboratories, including Lashley’s and by several investigators (see Sect. 3.6).
those of Harry Harlow at the University of Wisconsin
and Karl Pribram at Yale, took up the search for the
more complex functions of association cortex using
3.2 Consolidation
monkeys. Lashley’s interests were not limited to brain
mechanisms of learning. His influence is felt strongly Carl Duncan’s discovery of the effects of electro-
through the many eminent contemporary behavioral convulsive shock on retention of simple habits in the
neuroscientists who worked or had contact with him; rat, in 1949, began the modern field of memory
Austin Riesen, Donald O. Hebb, Roger W. Sperry and consolidation. Hebb and Gerard were quick to point
Karl Pribram are examples. Hebb’s book The Organ- out the implication of two memory processes, one

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transient and fragile and the other more permanent fish tail flip escape: Krasne, Kennedy, see Krasne and
and impervious. James McGaugh and his associates Bryan 1973) have all arrived at the same underlying
have done the classic work on the psychobiology of synaptic mechanism: a decrease in the probability of
memory consolidation. McGaugh and his colleagues transmitter release from presynaptic terminals of the
(see McGaugh 2000) demonstrated memory facili- habituating pathway. Habituation is thus a very
tation with drugs and showed that these effects were satisfying field: agreement ranges from defining be-
direct and not due to possible reinforcement effects of havioral properties to synaptic mechanisms. In a sense,
the drugs (and similarly for ECS impairment). the problem has been solved. Habituation also pro-
Chemical approaches to learning and memory are vides a most successful example of the use of the model
recent. The possibility that protein molecules and biological systems approach to analysis of neural
RNA might serve to code memory was suggested some mechanisms of behavioral plasticity (see Groves and
years ago by such pioneers as Gerard and Halstead. Thompson 1970).
The RNA hypothesis was taken up by Hyden and
associates in Sweden and by several groups in
America. An unfortunate byproduct of this approach
was the ‘transfer of memory’ by RNA. These exper-
3.4 Hippocampus
iments, done by investigators who shall remain name-
less, in the end could not be replicated. Study of the role of the hippocampus in learning and
At the same time, several very productive lines of memory has become an entire field. It began with
investigation of neurochemical and neuroanatomical Brenda Milner’s extraordinary studies of patient
substrates of learning were developing. In 1953, Krech H. M., who following bilateral temporal lobectomy
and Rosenzweig began a collaborative study on lost the ability to remember his own life experiences,
relationships between brain chemistry and behavior. dating to a year or so before his operation and
Their initial studies concerned brain levels of AChE in continuing to the present (see Milner 1966). However
relation to the hypothesis behavior and included he is able to learn motor and even complex sensory-
analysis of strain differences (see, e.g., Krech et al. motor skills normally. A large animal literature grew
1954). More recently, they and their collaborators, in unsuccessful attempts to develop animal models of
Bennett and Diamond, discovered the striking differ- H. M.’s syndrome. In 1978 Mortimer Mishkin, at
ences in the brains of rats raised in ‘rich’ vs. ‘poor’ NIMH, developed a striking analogue of H. M’s
environments. William Greenough, at the University syndrome in monkeys by removing the hippocampus
of Illinois, replicated and extended this work to and amygdala bilaterally. The animals still had normal
demonstrate dramatic morphological changes in the visual pattern discriminations but were markedly
structures of synapses and neurons as a result of impaired in recognition memory (delayed non match-
experience. ing to sample task). From recent work, e.g., by
Mishkin, Larry Squire, Stuart Zola and others, it is
clear that the most extreme form of the syndrome in
both monkeys and humans involves the hippocampus
and related cortical areas but not the amygdala. Study
3.3 Model Systems
of hippocampal function, particularly in humans in
The use of model biological systems has been an the context of declarative or ‘what’ memory is a major
important tradition in the study of neural mechanisms topic in cognitive neuroscience today.
of learning. This approach has been particularly Another facet of hippocampal study in the context
successful in the analysis of habituation, itself a very of behavioral neuroscience is long-term potentiation
simple form or model of learning (see Habituation). (LTP), discovered by Bliss and Lomo in 1970. Brief
Sherrington did important work on flexion reflex tetanic stimulation of monosynaptic inputs to the
‘fatigue’ in the spinal animal at the turn of the century. hippocampus causes a profound increase in synaptic
Sharpless and Jasper (1956) established habituation as excitability that can persist for hours or days. Many
an important process in EEG activity. Modern Russ- view it as a leading candidate for a mechanism of
ian influences have been important in this field: the key memory storage, although direct evidence is still
studies of Evgeny Sokolov, first on habituation of the lacking.
orienting response in humans and more recently on Yet another major impetus to study of the hip-
mechanisms of habituation of responses in the simpli- pocampus is the remarkable discovery of ‘place cells’
fied nervous system of the snail (see Sokolov 1963). by John O’Keefe (1976). When recording from single
The defining properties of habituation were clearly neurons in the hippocampus of the behaving rat, a
established by Thompson and Spencer in 1966, and given neuron may respond only when the animal is in
the analysis of mechanisms began. Several laboratories a particular place in the environment (i.e., in a box or
using different preparations (Aplysia withdrawal maze) reliably and repeatedly. There is great interest
reflex: Kandel and associates, see Kandel 1976; ver- now in the possibility that LTP may be the mechanism
tebrate spinal reflexes: Thompson and Spencer; cray- forming place cells. A number of laboratories are

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making use of genetically altered mice to test this years, the fields of ‘motivation’ and ‘emotion’ have
possibility. tended to go separate ways. However, motivation and
emotion have a common historical origin. In the
3.5 Cerebellum seventeenth and eighteenth centuries, instinct doctrine
served as the explanation for why organisms were
Yet another brain structure that has become a minor driven to behave (at least infrahuman organisms
industry in brain substrates of learning and memory is without souls). Darwin’s emphasis on the role of
the cerebellum. Masao Ito and his associates in Tokyo adaptive behavior in evolutionary survival resulted in
discovered the phenomenon of long-term depression the extension of instinct doctrine to human behavior.
(LTD) in cerebellar cortex in 1982. Repeated con- Watson rebelled violently against the notion of instinct
junctive stimulation of the two major projection and rejected it out of hand, together with biological
systems to the cerebellum, mossy-parallel fibers and mechanisms of motivation.
climbing fibers, yields long-lasting decreases in ex- In 1928, Bard showed that the hypothalamus was
citability of parallel fiber-Purkinje neuron synapses. responsible for ‘sham rage.’ In the 1930s, Ranson and
Ito developed considerable evidence that this process his associates at Northwestern, particularly Magoun
underlies plasticity of the vestibulo-ocular reflex. and Ingram, published a classic series of papers on the
Thompson and associates and others showed that the hypothalamus and its role in emotional behavior.
memory traces for classical conditioning of discrete Somewhat later, Hess and his collaborators in Switzer-
responses (e.g., eyeblink conditioning) are stored in land were studying effects of stimulating the hypo-
the cerebellum (Thompson 1986). LTD appears to be thalamus in freely moving cats.
a critical process of such memory formation in the
cerebellar cortex. The output of the cerebellum to
other brain structures is from the cerebellar nuclei and 4.1 Motiated Behaiors
Purkinje neurons inhibit nuclear neurons; hence cor- It is against this backdrop that the modern field of
tical LTD would decrease inhibition on nuclear psychobiology of motivation developed (the term
neurons. ‘motivated behaviors’ is now preferred to ‘motiv-
ation’). Karl Lashley was again the prime mover. His
3.6 Amygdala paper on The experimental analysis of instinctie
behaior in 1938 was the key. In it, he argued that
Still another structure to assume a major role in the motivated behavior varies, and is not simply a chain of
behavioral neuroscience of learning and memory is the instinctive or reflex acts, that it is not dependent upon
amygdala. It was known for some time that electrical any one stimulus, and that it involves central state. His
stimulation of the amygdala could elicit autonomic conclusions, that ‘physiologically, all drives are no
and emotional responses. An elegant literature de- more than expressions of the activity of specific
veloped by, e.g., Michael Davis, Joseph Le Doux, mechanisms’ and that hormones ‘activate some central
Michael Fanselow and others demonstrated clearly mechanism which maintains excitability and activity’
that the amygdala was the key structure in conditioned have a very modern ring.
fear (see Davis 1992). Indeed, this literature is at least A most important series of studies was originated in
consistent with the view that the essential memory a paper by Klu$ ver and Bucy on Psychic blindness and
traces for classical fear conditioning are established in other symptoms following bilateral temporal lobectomy
the amygdala. in rhesus monkeys in 1937. This came to be known as
The amygdala is also critical for instrumental the Klu$ ver–Bucy syndrome. The animals exhibited
learning of fear. James McGaugh and his associates marked changes in motivation and aggressive be-
demonstrated that for both passive and active avoid- havior. The Klu$ ver–Bucy syndrome, together with the
ance learning (animals must either not respond, or suggestion of Papez (1937) that the structures of the
respond quickly, to avoid shock) amygdala lesions limbic system formed a basic circuit for motivation
made immediately after training abolished the learned and emotion, led to the extensive field concerned with
fear (see McGaugh 1989). Surprisingly, if these same the behavioral functions of the ‘limbic system’ (see
lesions were made a week after training, learned fear Isaacson 1982).
was not abolished, consistent with a process of Lashley’s general notion of a central mechanism
consolidation. The apparent differences in the role of that maintains activity was developed by Beach in an
the amygdala in aspects of learned fear is an interesting important series of papers in the 1940s into a central
issue. excitatory mechanism and ultimately a central theory
of drive. This view was given a solid physiological
4. Motiation and Learning basis by Donald Lindsley from the work he and
Magoun and associates were doing on the ascending
Physiological and neural mechanisms of motivation reticular activating system. Lindsley sketched his
and emotion have been a particular province of activation theory of emotion in his important chapter
physiological psychology since the 1930s. In recent in the Steens Handbook (1951). D. O. Hebb and Elliot

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Stellar pulled all these threads together into a general 5. Sensory Processes
and influential central theory of motivation.
In his empirical work, Beach focused on brain A major topic area in physiological psychology has
mechanisms of sexual behavior. As the study of sexual been the study of sensory processes: sensation and
behavior developed, hormonal factors came to the perception. Indeed, this is perhaps the original field of
fore and the modern field of hormones and behavior psychology, dating back at least to Newton. Although
developed. Beach played a critical role in the de- specialization has resulted in separation between the
velopment of this field. Even within the field of fields of psychophysics and sensory physiology, in the
hormones and behavior, several fields have developed. sense that few individual scientists do research in both
Sexual behavior has become a field unto itself. Another fields, they remain closely interlocked. From the
important field today is the general area of stress. The beginning, explorations of sensory and perceptual
endocrinologist Hans Selye was an important intel- phenomena have always involved hypothetical physio-
lectual influence. Kurt Richter is the pioneering figure logical mechanisms, e.g., the Young–Helmholtz 3
in this field in physiological psychology. Richter took receptor theory of color vision and the Hering op-
his Ph.D. from Johns Hopkins in 1921 and established ponent process theory (see Hurvich and Jameson
his laboratory there. He worked in several areas, 1974).
including the role of the adrenal gland in stress. The This has been a field of extraordinary progress in the
modern field of stress focuses on hormonal-behavioral twentieth century. Techniques have been critically
interactions, e.g., the work of Seymore Levine. important. Early in the century there were really no
Neal Miller represents a uniquely important tra- tools, other than rather crude anatomical methods, for
dition in behavioral neuroscience. From the beginning analyzing the organization of sensory systems in the
of his career, Miller was interested in physiological brain. The pioneering studies of Adrian (1940) in
mechanisms of both motivation and learning. He took England and Marshall, Woolsey and Bard (1941) at
his Ph.D. from Yale in 1935 and stayed on at Yale for Johns Hopkins were the first to record electrical
many years, with a year out in 1936 at the Vienna evoked potentials from the somatic sensory cortex in
Psychoanalytic Institute. Miller was a pioneer in early response to tactile stimulation. Woolsey and his
studies of punishing and rewarding brain stimulation associates developed the detailed methodology for
and their roles in learning. He was the first to evoked potential mapping of the cerebral cortex. In an
demonstrate conditioned fear. In more recent years, extraordinary series of studies, Woolsey and his
his work focused on mechanisms of instrumental colleagues determined the localization and organ-
conditioning of autonomic responses. The impact of ization of the somatic sensory areas, the visual areas
his work is much wider than physiological psychology, and the auditory areas of the cerebral cortex in a
influencing psychiatry and clinical medicine as well. comparative series of mammals. They initially defined
two areas (I and II) for each sensory field.

4.1.1 Electrical self-stimulation of the brain. James


5.1 Organization of the Sensory Systems
Olds, whose untimely death in 1976 cut short an extra-
ordinary career, made perhaps the most important In the 1940s and 1950s the evoked potential method
discovery yet made in the field of behavioral neuro- was used to analyze the organization of sensory
science: rewarding electrical self-stimulation of the systems at all levels from the first order neurons to the
brain. He got his Ph.D. at Harvard and worked with cerebral cortex. The principle that emerged was
RichardSolomon.Solomon,incidentally,althoughpri- strikingly clear and simple; in every sensory system the
marily a behavioral student of learning, had great nervous system maintained a receptotopic map or
impact on physiological psychology through his projection at all levels from receptors to cerebral
theoretical-experimental analysis of hypothetical cortex: skin surface, retina, basilar membrane. The
central factors in learning. As a graduate student same organization held for the second sensory areas.
Olds read and was much influenced by Hebb’s The The receptor maps in the brain were not one-to-one,
Organization of Behaior and obtained a post- rather they reflected the functional organization of
doctoral with Hebb at McGill in 1953. He began each system: fingers, lips, and tongue areas were much
work with Peter Milner and they discovered electrical enlarged in primate somatic cortex, half the primary
self-stimulation of the brain. The recent history of visual cortex represented the forea, and so on.
this field is well known to everyone. The brain The evoked potential method was very well suited to
reward system now appears closely similar if not analysis of the overall organization of sensory systems
coextensive with the ‘addiction’ system in the brain: in the brain. However, it could reveal nothing about
the medial forebrain bundle dopamine projection what the individual neurons were doing. This had to
system to the nucleus accumbens, prefrontal cortex, wait development of the microelectrode. Indeed, the
and other structures. microelectrode has been the key to analysis of the fine-

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grained organization and feature detector properties developments in genetics and molecular biology,
of sensory neurons. Metal microelectrodes were behavioral genetics has become a component of
developed in the early 1950s: Davies at Hopkins de- neurogenetics.
veloped the platinum-iridium glass coated micro- The field of behavioral neuroscience ( physiological
electrode, Hubel and Wiesel at Harvard developed the psychology) has a long history of asking fundamental
tungsten microelectrode, and the search for feature questions about biology, behavior, and experience
detectors was on. The pioneering studies were those of and developing approaches to deal with these
Mountcastle and associates at Hopkins on the organ- questions. The consequence is often that entire new
ization of the somatic-sensory system, those of Hubel fields of scientific endeavor develop and became
and Wiesel at Harvard on visual system and Rose, distinct subspecialties in their own right. Perhaps in a
Hind, Woolsey and associates at Wisconsin on the more general sense the same can be said of psychology.
auditory system. Thanks to the microelectrode and to
modern pathway tracing techniques, we now know See also: Amygdala (Amygdaloid Complex); Cere-
that each sensory modality is represented multiply in bellum: Cognitive Functions; Cognitive Neuroscience;
the cerebral cortex. Comparative Neuroscience; Emotion, Neural Basis
Meanwhile, pioneering work was being done on of; Hebb, Donald Olding (1904–85); Hippocampus
receptors. Hartline and Ratliff analyzed receptor and Related Structures; Lashley, Karl Spencer
responses in a simple visual system, Limulus (horse- (1890–1958); Learning and Memory: Computational
shoe crab), and discovered lateral inhibition. Von Models; Learning and Memory, Neural Basis of;
Bekesy discovered the standing wave patterns in the Mind—Body Dualism; Motivation, Neural Basis of;
cochlea (working in the psychology department at Pavlov, Ivan Petrovich (1849–1936); Psychophysio-
Harvard). Dark adaptation was explained in bio- logy; Visual Perception, Neural Basis of; Wundt,
chemical terms by Wald at Harvard. The role of eye Wilhelm Maximilian (1832–1920)
movements in visual perception was elucidated by
Riggs and associates at Brown.
In the recent past, progress in analysis of sensory Bibliography
processes has been quite remarkable. The ability to
Adrian E D 1940 Double representation of the feet in the
measure, with the microelectrode, the activity of single sensory cortex of the cat. Journal of Physiology 98: 16
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stimulus control, has been matched by the great rage with special reference to the sympathetic nervous system.
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Enzyme concentration in brain and adjustive behavior significantly molded the character of American pol-
patterns. Science 120: 994–6 itical science for the next 15–20 years. Since then, there
Lashley K S 1929 Brain Mechanisms and Intelligence. University have been hardly any self-declared behavioralists, but
of Chicago Press, Chicago the spirit of empiricism and methodical profession-
Lashley K 1938 The experimental analysis of instinctive alism has shaped the discipline of political science
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Annual Reiew of Neuroscience 12: 255–87 political science, behavioralism mainly represents a
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moving rat. Experimental Neurology 51: 78–109 dominant institutionalist approach, known as ‘institu-
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Neurology and Psychiatry 38: 725–43 approach centered on the analysis of the actual (as
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University Press, London branches of government, political parties, and interest
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fact-finding, and a strict distinction between factual
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Thompson R F, Spencer W A 1966 Habituation: A model the New Deal and especially during World War II
phenomenon for the study of neuronal substrates of behavior. revealed the shortcomings of the realist approach. The
Psychological Reiew 73: 16–43 gulf seemed too wide between what was required for
Watson J B 1913 Psychology as the behaviorist views it. effective political consulting and what political scien-
Psychological Reiew 20: 158–77 tists were able to provide. In particular, the limited
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coming (Dahl 1961). For example, realists fell short in
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R. F. Thompson institutional analysis only. To many political scien-
tists, the insights taken from neighboring disciplines,
Copyright # 2001 Elsevier Science Ltd. especially from psychology, sociology, and the econ-
omics, seemed theoretically and methodologically
All rights reserved. more sophisticated than those taken from political
science.
Primarily younger political scientists became sharp
Behavioralism: Political critics of the realist approach but none formulated it
with more urgency than David Easton. In program-
Behavioralism is a current or approach within the matic writings dating back to the early 1950s, he
discipline of political science. In very general terms, it criticized the theoretical deficit of the research at that
can be described as theory-led empiricism which aims time, the lack of methodological reflection and of

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International Encyclopedia of the Social & Behavioral Sciences ISBN: 0-08-043076-7

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