Motivation and Emotion 57agreed with James and Lange that the initial event had to be
perception of an emotion-arousing situation but argued that
the development of autonomic (sympathetic) responses—
release of epinephrine and other bodily changes—occurred
concomitantly with the subjective feelings (see Cannon,
1927). However, his primary interest was in the physiology,
particularly the peripheral physiology. Cannon’s view was
championed by the distinguished Johns Hopkins physiologist
Philip Bard, who stressed the key role of the brain, particu-
larly the thalamus and hypothalamus, in both emotional
behavior and experience (see Bard, 1934). Cannon, inciden-
tally, also contributed the notion of homeostasis, which he
developed from Bernard’s concept of the milieu interieur.
A key issue in these theories was the role of sympathetic
arousal or activation in the experience of emotion. This issue
was tested in a classic study by Stanley Schachter and Jerome
Singer at Columbia University in 1962. They injected human
subjects with either effective doses of epinephrine or a
placebo. The epinephrine activated the sympathetic signs of
emotions (pounding heart, dry mouth, etc.). Both groups of
subjects were told they were receiving a shot of a new vita-
min. Stooges acted out euphoria or anger in front of the sub-
jects. The subjects were either informed of what the injection
might do, for example, the autonomic side effects, or not in-
formed. Results were dramatic. Uninformed epinephrine
subjects reported emotional experiences like those of stooges
but informed epinephrine subjects did not report any emotion
at all. Emotion is more than sympathetic arousal—cognitive
factors are also important.
Experimental work on brain substrates of emotion may be
said to have begun with the studies of Karplus and Kreidl in
1910 on the effects of stimulating the hypothalamus. In 1928,
Bard showed that the hypothalamus was responsible for
“sham rage.” In the 1930s, S. W. Ranson and his associates at
Northwestern, particularly H. W. Magoun, published a clas-
sic series of papers in the hypothalamus and its role in emo-
tional behavior (Ranson & Magoun, 1939). In the same
period, W. R. Hess (1957) and his collaborators in Switzer-
land were studying the effects of stimulating the hypothala-
mus in freely moving cats. A most important paper by H.
Klüver and P. Bucy reported on “psychic blindness and other
symptoms following bilateral temporal lobectomy in rhesus
monkeys” in 1937. This came to be known as the Klüver-
Bucy syndrome. The animals exhibited marked changes in
motivation and aggressive behavior.
Pribram (Bucy’s first resident in neurosurgery) developed
the surgical methods necessary to analyze the Klüver-Bucy
syndrome. This analysis led to his discovery of the functions of
the inferotemporal cortex in vision and to the exploration of the
suggestions of J. W. Papez (1937) and P. D. MacLean (1949)
that the structures of the limbic system (the “Papez” circuit) are
concerned with motivation and emotion. However, modern
neuroanatomy deconstructed the Papez circuit. The emphasis
is now on the hypothalamus-pituitary axis, on descending
neural systems, and on the amygdala.MotivationToday most workers in the field prefer the term motivated
behaviorsto emphasize the specific features of behaviors re-
lating to hunger, thirst, sex, temperature, and so forth. Karl
Lashley was again a prime mover. His 1938 paper, “Experi-
mental Analysis of Instinctive Behavior,” was the key. He ar-
gued that motivated behavior varies and is not simply a chain
of instinctive or reflex acts, is not dependent on any one stim-
ulus, and involves central state. His conclusions, that “physi-
ologically, all drives are no more than expression of the
activity of specific mechanisms” and that hormones “activate
some central mechanism which maintains excitability and ac-
tivity,” have a very modern ring.
Several key figures in the modern development of the
psychobiology of motivation are Clifford Morgan, Eliot
Stellar, Kurt Richter, Frank Beach, Neal Miller, Philip
Teitelbaum, and James Olds. Morgan went to graduate
school at Rochester, where his professors included E. A. K.
Culler and K. U. Smith and his fellow graduate students in-
cluded D. Neff, J. C. R. Licklider, and P. Fitts. He then be-
came an instructor at Harvard, where he first worked in
Lashley’s laboratory in 1939. He later moved to Johns Hop-
kins, where he remained until 1959. As a graduate student
and later at Harvard, Morgan came to doubt Cannon’s then
current notion that hunger was the result of stomach con-
tractions. Morgan did a series of studies showing this could
not be a complete or even satisfactory account of hunger
and feeding behavior. Eliot Stellar and Robert McCleary,
then undergraduates at Harvard, worked with Morgan. They
focused on hoarding behavior and completed a classic
analysis of the internal and environmental factors control-
ling the behavior.
Lashley’s general notion of a central mechanism that
maintains activity was developed by Beach in an important
series of papers in the 1940s and by Morgan in the first edi-
tion of his important text, Physiological Psychology(1943),
into a central excitatory mechanism and ultimately a central
theory of drive. This view was given a solid physiological
basis by Donald B. Lindsley from the work he and H. W.
Magoun, G. Moruzzi, and associates were doing on the as-
cending reticular activating system. Lindsley sketched his ac-
tivation theory of emotion in his important chapter in the
StevensHandbook(1951). Hebb (1955) and Stellar (1954)