THE BEHAVIORAL AND BRAIN SCIENCES (1982) 5, 407-467

Printed in the United States of America

Toward a general psychobiological

theory of emotions
Jaak Panksepp
Department of Psychology, Bowling Green
State University, Bowling Green, Ohio 43403

Abstract: Emotions seem to arise ultimately from hard-wired neural circuits in the visceral-limbic brain that facilitate diverse and
adaptive behavioral and physiological responses to major classes of environmental challenges. Presumably these circuits developed
early in mammalian brain evolution, and the underlying control mechanisms remain similar in humans and "lower" mammals. This
would suggest that theoretically guided studies of the animal brain can reveal how primitive emotions are organized in the human
brain. Conversely, granted this cross-species heritage, it is arguable that human introspective access to emotional states may provide
direct information concerning operations of emotive circuits and thus be a primary source of hypotheses for animal brain research. In
this article the possibility that emotions are elaborated by transhypothalamic executive (command) circuits that concurrently activate
related behavior patterns is assessed. Current neurobehavioral evidence indicates that there are at least four executive circuits of this
type - those which elaborate central states of expectancy, rage, fear, and panic. The manner in which learning and psychiatric
disorders may arise from activities of such circuits is also discussed.

Keywords: affect; brain theory; command circuits; emotion; expectancy; fear; hypothalamus; panic; psychiatric disorders; rage;
reinforcement

The scientific study of emotions is beset by more than the
usual number of methodological and conceptual prob-
lems. It is difficult to agree how, within the constraints of
scientific objectivity, we can derive substantive under-
Standing of phenomena that appear intimately linked to
the internal experiences of organisms. In animal re-
search, the traditional solution (and compromise) has
been to study the objective behavioral and physiological
manifestations of presumed emotional states, with an
exclusive focus on behavior and a calculated disregard of
the underlying states. Conversely, at the human level
there is abundant discussion of the various emotional
states and little knowledge concerning their sources in
the brain. Thus, while major concepts concerning the
nature of emotions arise from human introspection, most
knowledge concerning the mechanisms underlying emo-
tionality arises from animal brain research. This paper is
based on the conviction that the study of emotions, in
both humans and animals, continues to be impoverished
by our failure to blend these two sources of knowledge.
The major obstacles to achieving the above synthesis
are our inability to read the animal mind as directly as we
can read our own and the subtleties of emotional nuance
that can evolve during the course of social learning in
humans. Accordingly, introspection and the consequent
tendency to anthropomorphize remain anathema in ani-
mal research. However, the fundamental organization of
emotions in the brain may be quite similar in humans and
other mammals. If this is so, recent advances in brain
research may permit anthropomorphism to become a
more useful strategy for understanding certain primitive
psychological processes in animals than it has been in the
past. Through the conjoint application of an-
thropomorphic reasoning and animal brain research,
more knowledge concerning the nature of emotions may
be derived than from either approach taken alone. This is
not to say that we can measure the subjective awareness
of other animals, but to assert that basic emotions may
have obligatory internal dynamics, which humans share
with other mammals. By using our subjective sources of
insight concerning these dynamics, we may be able to
resolve some of the subtleties of brain organization more
readily than if we follow the dictum that all we can know
are the behavioral and physiological symptoms of emotive
states.
In any event, it should be self-evident that the use of
anthropomorphism in the study of mammalian emotions
cannot be arbitrarily ruled out. Although its application
may be risky under the best of circumstances, its validity
depends on the degree of evolutionary continuity among
brain mechanisms that elaborate emotions in humans and
animals. Hence, the degree of anthropomorphism that
can have scientific utility in mammalian brain research
should be directly related to the extent that emotions
reflect class-typical mechanisms as opposed to species-
typical ones.
Although a definitive statement concerning the degree
of similarity between neural systems that mediate emo-
tions in animals and humans cannot be made, available
evidence suggests that fundamental emotional circuits
are inherited components of the limbic brain, which are
to a substantial degree a shared mammalian heritage
(Crosby & Showers 1969; MacLean 1973; Parent 1979).
Species-typical instinctual behaviors remain intact even
after radical decortication soon after birth (e.g., Murphy,
MacLean, & Hamilton 1981), and, with few exceptions,
trauma to subcortical tissue in humans yields behavioral
changes similar to those which result from experimental

© 1982 Cambridge University Press 0140-525X1821030407-61 IS06.00 407

Panksepp: Psychobiology of emotions
manipulation of these circuits in other mammals. For
instance, animals with ventromedial hypothalamic le-
sions exhibit voraciousness and rage (Heatherington &
Ranson 1942; Wheatley 1944), as do humans (Reeves &
Plum 1969). Posterior hypothalamic damage brings on
somnolence, and anterior hypothalamic damage insom-
nia, in both animals and humans (McGinty & Sterman
1968; Nauta 1946; Ranson 1939; von Economo 1930).
Electrical stimulation of the amygdala can yield rage in
both (de Molina & Hunsperger 1963; King 1961). Al-
though exceptions to such apparent homologies could
also be cited, the general conclusion appears to be that
the functional terrain of the subcortical limbic brain
across mammalian species is remarkably similar, in kind if
not in precise organization. Accordingly, the study of
these parts of the emotional brain in animals should
provide a credible outline for understanding the primi-
tive emotional functions of the human brain. Conversely,
it should follow that some properties of these circuits in
the animal brain could be deciphered by judicious intro-
spection. Although it is not evident how a clear path
through the possibilities of human imagination is to be
achieved, anthropomorphic excesses can be checked by
the constraint that verifiable consequences must be gen-
erated with the accepted empirical tools of behavioral
brain research. By insisting that introspectively derived
conceptions of animal emotion be linked simultaneously
to specific behaviors and brain processes, the testability
of the resulting ideas could help stave off the terminally
debilitating scholastic debates, which have immobilized
this important area of inquiry in the past.
Although behavioral and physiological effects of emo-
tional states may be all that can ever be empirically
measured in animals, some of these could be used as
indicator variables for more hidden central processes that
may be operative in certain situations. Indeed, such an
indirect theoretical analysis of certain neural systems may
be essential, because the field of inquiry is confronted by
experimental difficulties similar to those encountered in
particle physics, where the clarification of intranuclear
forces has to proceed through indirect observations that
yield only indirect affirmative or negative evidence about
the nature of the underlying processes. In a similar way,
the functions of certain brain circuits may be sufficiently
internalized in the fabric of the brain so that their normal
operations and intersystemic cleavage lines need to be
derived initially by introspectively based theoretical in-
ference. Of course, our initial conceptualizations will
invariably be simplifications, which fail to do full justice to
the actual circuit configurations in the brain. With such
qualifications in mind, the overall aim of this paper is to
outline a testable theory of how emotional processes may
be organized in the central nervous system of mammals.
Because they will receive little further emphasis here,
I shall reiterate the theoretical assumptions underlying
this proposal. They are: (1) Distinct emotional processes
reflect activity in specific hard-wired brain circuits. (2)
Humans share essentially similar primitive emotional
processes with all other mammals and perhaps some
related vertebrates. (3) The number of fundamental emo-
tional circuits is limited, but through their intermixtures
and social learning, emotions in humans have a much
richer phenomenological texture than is contained in the
408 BEHAVIORAL AND BRAIN SCIENCES (1982)3
individual emotive circuits. (4) The most efficient and
reasonable way to obtain an initial neurotaxonomy of the
hard-wired emotional processes in the mammalian brain
is through introspection. (5) The scientific understanding
of emotional processes must arise from a clarification of
how they are organized in the brain (and hence, out of
"ethical" necessity, the animal brain).
On the basis of these assumptions, the substantive aims
of this paper are to propose a taxonomy of emotions that is
supported by existing neurobehavioral data; to provide an
approximate neural scheme of how emotions may be
organized in the brain; and to discuss the implications of
such circuits for the understanding of learning and emo-
tional disorders. Admittedly, each successive aim is
based on increasingly large measures of speculation. In
general, I shall argue that "the mammalian brain" con-
tains at least four primitive executive (command) circuits
that initiate the distinct subjective emotional states of
expectancy, fear, rage, and panic in humans and that
trigger the corresponding emotive processes in animals.
First, however, I shall trace some of the historical roots of
this endeavor.
Historical overview
A great deal has been written on emotions through the
ages, and one who attempts the journey through that
material may soon concur with James (1892, p. 241) that
"the varieties of emotion are innumerable" and that the
descriptive literature on the subject "is one of the most
tedious parts of psychology. And not only is it tedious, but
you feel that its subdivisions are to a great extent either
fictitious or unimportant, and that its pretences to ac-
curacy are a sham," so that "reading of classic works on
the subject" is as interesting as "verbal descriptions of the
shapes of the rocks on a New Hampshire farm."
Still, at the conceptual Jevel, an inquiry into emotions
must start with a taxonomy of emotional experience in
humans. Were it not for the consensus that such subjec-
tive states exist in humans, emotion as a construct would
never have arisen in the study of animal behavior. Many
plausible taxonomies of emotions have come from the
consideration of such subjective states, and it is reason-
able to expect that a cogent physiological theory of emo-
tion should subsume the best organizational principles
generated from that perspective. Historically, the con-
struction of taxonomies of emotional experience, though
not strictly empirical, led directly to the study of emo-
tional expression in humans and kindred animals and,
recently, to attempts to specify how various emotional
processes are related to physiological changes in both
brain and body.
Taxonomies of emotion. In one of the earliest major
treatises to attempt to relate brain and mind, Willis (1683,
p. 49) began his ninth chapter on "Two Discourses
Concerning the Soul of Brutes" as follows:
Concerning the Number of the Passions, as it hath
been variously disputed among Philosophers, so in
famous Schools, this Division into Eleven Passions,
long since grew of use; to wit, the Sensitive Appetite is
distinguished into Concupiscible and Irascible, to the

first, are counted commonly six Passions, viz. Pleasure
and Grief, Desire and Aversions, Love and Hatred; but
to the latter five, viz. Anger, Boldness, Fear, Hope,
and Desperation, are wont to be attributed.
Foreshadowing future controversies, Willis disagreed:
But this distribution of the Affections is not only in-
congruous, for that Hope is but ill referred to the
Irascible Appetite, and Hatred and Aversion, seem
rather to belong to this, than to the Concupiscible: But
it is also very insufficient, because some more noted
Affections, as Shame, Pity, Emulation, Envy, and
many others, are wholly omitted: Wherefore the An-
cient Philosophers did determinate the Primary to a
certain Number, then they placed under their several
Kinds, very many indefinite Species.
Perhaps as a reaction to the categorical excesses of
medieval scholastics, Descartes, in his discourse on "the
passions of the soul" (1650), reduced emotional life to six
primary passions: wonder, love, hate, desire, joy, and
sadness, with all the rest being "made up of some of these
six or at least are species of them" (cf. Ruckmick 1936, p.
39). The idea that primary emotional processes are lim-
ited to a relatively small group of fundamental systems is
still with us, with the number of primary emotions usually
ranging from three to eight; the father of behaviorism
suggested that three, namely, fear, rage, and sexual love,
characterized the fundamental nature of man (Watson
1924). However, except for general suggestions of where
emotions may be elaborated in the brain (e.g., Papez
1937), as well as highly concrete neurophysiological hy-
potheses concerning the genesis of specific emotions
(e.g., anxiety, in Gray, 1979), there has been little at-
tempt to relate any of the well articulated taxonomies of
emotion to specific brain circuits. The aim of this paper is
to attempt such a synthesis. [See also BBS multiple book
review of Gray: The Neuropsychology of Anxiety, BBS 5
(3) 1982, this issue.]
After the ideas expressed in this paper had crystallized,
I discovered a forgotten system closely resembling it in
form, if not specific content. This system, proposed by
Cogan (1802) and summarized in Table 1, viewed emo-
tions from the subjective perspective, and thus contained
the existential virtues (or mentalistic excesses, depending
on one's point of view) that have traditionally proved
troublesome in the scientific analysis of emotion. Howev-
er, it will be argued that the scheme contains the seed of
truth that any credible psychobiological theory of emo-
tion must ultimately nurture. Cogan suggested that hu-
mans inherit five primitive passions - those of desire,
anger, fear, sorrow, and joy. Under somewhat different
labels, expectancy, rage, fear, and panic, I shall attempt
to relate the first four of these passions to the circuitry of
the visceral brain.
Since Cogan's time, many other taxonomies have been
proposed, but in the absence of concurrent analyses of
brain mechanisms, they remain on the fringes of science.
Recent thought on the subject has been well summarized
by Ruckmick (1936) and Plutchik (1980). Without phys-
iological anchor points, theories of emotion remain gra-
tuitous because they cannot escape from fatal cir-
cularities, although they can generate useful experimen-
tation (Strongman 1973). With clearly specified
physiological anchor points, however, circularity can be
Panksepp: Psychobiology of emotions
Table 1. Taxonomy of the passions (Cogan, 1802)
Class I Passions arising from principle of self-love
Order I From the idea of good
Genus 1 Good in possession (JOY: gladness, cheerful-
ness, mirth, satisfaction, contentment, com-
placency)
Genus 2 Good in expectancy—DESIRE
Order II From the idea of evil
Genus 1 Losses and disappointment—SORROW
Genus 2 Evils which are apprehensive—FEAR
Genus 3 Conduct which deserves reprehension—
ANGER
Class II Passions and affections derived from the so-
cial principle
Order I Excited by ideas of benevolence
Genus I Benevolent desires (generosity, condolence,
pity, charity, liberality)
Genus 2 Good opinion (gratitude, thankfulness, admi-
ration)
Order II Excited by idea of evil
Genus 1 Malevolence (malice, envy, rancor, cruelty,
prejudice, jealousy, resentment, revenge)
Genus 2 Unfavorable opinion (horror, contempt, dis-
dain)
overcome, straightforward, testable hypotheses can be
generated, and refutations can be decisive.
The expression of emotion. Perhaps as a response to the
endlessly equivocal nature of introspectively derived
taxonomies of emotions, as well as the development of
new intellectual disciplines, such as physiognomy (Lava-
ter 1789-98) and phrenology (Gall 1835; Spurzheim
1832), the nineteenth century saw the rise of descriptive
analysis of emotional expression, ranging from detailed
studies of the facial musculature generating the emotional
countenance (Bell 1806) to the recognition that the ste-
reotyped character of emotional expression in animals
and humans indicates the common inheritance of mecha-
nisms that govern the elaboration of emotions (Darwin
1872). At the end of the century, the emphasis on emo-
tional expression, as opposed to emotional experiences or
processes, led to the renowned James-Lange theory
(James 1884), which entertained placing anger inside the
fist rather than behind it. By proposing that the bodily
changes occurring during emotive acts might be causes
rather than consequences of emotions, the James-Lange
theory challenged common reason and reaped a harvest
of negative evidence and critical counterargument (Can-
non 1927, 1931; Sherrington 1900), though perhaps none
sufficiently compelling as to be decisive, especially if one
wished to entertain an infinite regress into the potential
subjective awareness of a brain disconnected of its inputs
and outputs. Although it is likely that peripheral auto-
nomic feedback influences the labeling of emotional
states in humans and provides reafferent feedback that
intensifies emotional experience (Schachter & Singer
1962) and behavior (Baccelli, Guazzi, Libretti, & Zan-
chetti 1965), it now seems likely that the James-Lange
BEHAVIORAL AND BRAIN SCIENCES (1982)3 409
Panksepp: Psychobiology of emotions
theory reversed primary cause and consequence. Yet,
unlike earlier inquiries such as Darwin's, in which little
heed was given to the nature of proximal causes, the
James-Lange perspective emphasized the need for such
a level of analysis if emotions were ever to be probed
experimentally.
The study of emotional expression, as reflected in
somatic and visceral changes, has remained a hallmark of
the scientific analysis of emotions (Ax 1953; Eibl-
Eibesfeldt 1979; Ekman 1973; Grings & Dawson 1978).
Unfortunately, brain research generally continues to
cling to the tenet that the only credible scientific ap-
proach to this area is the study of emotional expression.
Although the approach has yielded many elegant facts
(Flynn 1976; Hess 1957), it is my contention that such a
single-minded focus on emotional expression without
concurrent consideration of underlying emotional pro-
cess has outlived its usefulness in psychobiology and may
be currently hindering progress in understanding the
neural organization of emotion.
Brain organization of emotive processes. Since range
could be precipitated in animals by decerebration (Goltz
1892), the suspicion arose that certain subcortical parts of
the brain were important for organizing emotions. How-
ever, it was not until after the turn of the century that
experimental evidence was found that indicated that
certain areas of the brain, especially those near the
ventral surface, are critical for organizing the autonomic
changes that accompany emotional states and behaviors.
Further study of decorticate rage by Bard (1928; 1934),
Cannon (1929), and others established that the expression
of anger could be initiated subcortically, suggesting that
the neocortex tonically inhibits circuits that mediate this
emotional behavior. This discovery led to the hypothesis
that endogenous brain systems elaborate emotions and,
specifically, that thalamic circuits mediate emotional ex-
periences (Cannon 1929). Concurrently, Karplus and
Kreidl (1909; 1927), followed by Kabat (1936), Ranson and
Magoun (1939), and others, were demonstrating dramatic
changes in the autonomic responsivity of anesthetized
cats during localized electrical stimulation of the hypo-
thalamus. This response suggested that the ventral part of
the upper brainstem was a head ganglion of the autonom-
ic nervous system, which participated in the genesis of
emotional behavior and experience. The fact that
thalamic damage did not eliminate decorticate rage in
cats (Bard & Macht 1958) affirmed the probability that the
hypothalamus was essential for elaborating emotional
responsivity. Hess's studies (1925-49, and English sum-
mary in 1957) established that rage and other well coordi-
nated emotive behaviors could be instigated by localized
electrical stimulation of the hypothalamus in a conscious
animal. This work solidly established the concept that the
emotional nervous system is embedded in the visceral
nervous system. Later, with the recognition of the di-
verse anterior connectivities of hypothalamic systems
(Papez 1937) and the emotional changes resulting from
damage to those circuits (Bard & Mountcastle 1948;
Kluver & Bucy 1939), the concept of a visceral-emotional
brain was generalized to the entire limbic system (Mac-
Lean 1949).
Further study of restricted decortications in cats indi-
cated an insensitivity to emotion-provoking stimuli when
410 BEHAVIORAL AND BRAIN SCIENCES (1982)3
only medial and temporal cortex were left intact, amend-
ing the earlier idea that the cortex generally inhibited
emotional responsivity (Bard & Mountcastle 1948). Since
heightened emotionality emerged gradually following
damage to those remaining cortical areas, it was sug-
gested that inhibition of emotional responsivity might be
due specifically to limbic cortical influences, but this
conclusion remained tenuous, because it was outwardly
inconsistent with the emotional placidity that appeared
after temporal lobe damage in monkeys (Kluver & Bucy
1937) and cats (Schreiner & Kling 1953). More recently,
wildness has been induced by cingulate cortical damage
in neodecorticate animals (Murphy et al. 1981), suggest-
ing that disinhibition of rage may have resulted from
damage to medial limbic cortex.
Although many questions concerning decorticate rage
remain, it is now clear that both excitatory and inhibitory
influences on emotional behavior are elaborated through
limbic cortex and ganglia (Ursin 1965; Ursin & Kaada
1960). The early studies definitively localized the elabora-
tion of primitive emotional tendencies in the vis-
ceral-limbic brain, and the old controversies concerning
whether the emotional changes observed in some of those
early studies were devoid of emotional content, whether
they were "sham" or "real," have become less pertinent
with the recognition that hierarchical levels of organiza-
tion exist within functionally continuous psycho-
behavioral organ systems of the brain (Bernston & Micco
1976). Indeed, questions of animal awareness were
shelved as basically unanswerable, and the mainstream of
investigation continued to focus on the actual brain cir-
cuits that participate in the expression of "emotive"
behaviors. It became clear that the functions of higher
brain areas >vere mediated through lower systems (de
Molina & Hunsperger 1962), and it came to be widely
assumed by psychobiological investigators that the elu-
cidation of hard-wired brainstem circuits would provide
the basic information through which higher functions
were ultimately to be understood. At the same time, the
likely relevance of these circuits to the emotional life of
humans was implicitly accepted but generally treated
with benign neglect. The translation between the two was
understood to be empirically impossible, and hence, in
some ultimate logical sense, gratuitous.
Thus, even though there has been a great deal of
excellent work on brainstem mediation of emotive behav-
iors, especially aggression, there has been little attempt
to integrate concepts from the taxonomy of human emo-
tional experience with data concerning the limbic circuit-
ry of the animal brain. Although the work on brain
organization of aggression has yielded a harvest of knowl-
edge (Adams 1979; Flynn 1976; Siegel & Edinger 1981),
as has the analysis of autonomic changes that typically
accompany such agonistic emotive states (Mancia &
Zanchetti 1981), other affective processes that do not
have such dramatic somatic and autonomic manifesta-
tions remain largely unstudied.
A definition of emotions
On the assumption that emotions arise from the activities
of brain circuits, a general definition of emotion would
consist of a list of attributes such brain circuits possess. In

the absence of extensive knowledge concerning these
circuits, the list must first be constructed rationally. I
propose that such circuits have at least the following six
attributes: (1) They are genetically hard-wired and de-
signed to respond unconditionally to stimuli arising from
major life-challenging circumstances (Figure 1). (2) They
organize behavior by activating or inhibiting classes of
related actions (and concurrent autonomic/hormonal
changes) that have proved adaptive in the face of those
types of life-challenging circumstances during the evolu-
tionary history of the species (Figure 2). (3) Through
recurrent feedback, emotive circuits change the sen-
sitivities and responsivities of sensory systems relevant
for those behavior sequences. (4) The activity in the
underlying neural systems can outlast the precipitating
circumstances (i.e., there is either substantial positive
feedback or long-term neuromodulation in the circuits).
(5) Activity in these circuits can come under the condi-
tional control of emotionally neutral environmental stim-
uli via a reinforcement process that is a design feature of
each emotive circuit. (6) Activity in such circuits has
access to and reciprocal interactions with brain mecha-
nisms that elaborate consciousness (to provide higher-
order response selection among the specific behavioral
acts that are sensitized by arousal of an emotive system).
It is only this last aspect of emotions, so evident in human
mental life, which remains so untestable in animal re-
search. However, since all mammals seem to share funda-
mentally similar emotive brain circuits, this last attribute
may prove to be the "Rosetta stone" via which the general
organization of the primitive hard-wired emotional sys-
tems of the mammalian brain can be deciphered.
Thefirstfour criteria should help delimit the number of
emotive circuits in the mammalian brain and help gener-
ate distinctions among the many vague terms that have
been used in discussions of the affective life. For instance,
PAIN .ndTHREAT
Of M>RM
DESTRUCTION
Figure 1. Major classes of environmental stimuli that require
rapid behavioral adjustments presumably converge upon com-
mand systems of the brain to trigger species-typical sensory-
motor patterns.
Panksepp: Psychobiology of emotions
a distinction between affective "feelings" and "emotions"
may be made on the basis of the degree of arousal within
an emotive system. Thus, environmental stimuli that
provoke weak interactions with primitive emotive cir-
cuits without generating sustained reverberatory feed-
back may be considered to have simply provoked an
affective "feeling." Furthermore, the fact that there are a
variety of homeostatic interoreceptors in the hypo-
thalamus, designed to monitor various bodily states,
which not only interact with emotive systems but which
surely also have distinct sensory attributes, further ampli-
fies the variety of discriminable "feelings" that can be
elaborated by the brain. By assuming that most of the
internal and external stimuli encountered in everyday life
interact with a limited number of primitive emotive
circuits, the list of distinct emotions can be kept much
shorter than the list of distinct feelings. Accordingly,
questions of how the affective coloring of each human
subjective experience is achieved can be shelved until the
primitive emotional circuitries of the brain are deline-
ated, and the analysis of basic emotive systems need not
be paralyzed by the vast variety of human affective
experience in the real world.
Moreover, several primitive affective states, which
have been incorporated in past taxonomies of emotional
life, will be ignored because they do not appear to fulfill
all of the criteria mentioned above. For instance, neither
"surprise" and "disgust," both of which have clear behav-
ioral referents in animals (e.g., Davis 1979; Grill &
Norgren 1978) and may be controlled by specific brain-
stem circuits (the "stimulus-bound" startle and retching
described by Hess, 1957), are included as major emo-
tional states in the present context (powerful affects as
they may be in humans), because they do not activate
diverse classes of species-typical behaviors (permitting
adaptive response selection among alternatives) and be-
cause their underlying states appear to be largely time-
locked to the precipitating stimuli, with no sustained
regenerative feedback in the underlying brain systems.
In other words, "startle" and "disgust" may be more
reflexive in character than the brain processes that are
herein considered to be truly emotional.
For similar reasons, the concepts of pleasure/relief and
displeasure/distress are not included in the present analy-
sis. Such states do not appear to generate unique behav-
ioral manifestations, and such generalized affects may cut
across many distinct emotive states and probably arise
secondarily from changes in activity of the discrete emo-
tive systems to be discussed. For instance, pleasure and
relief may be states which accompany stimuli that tend to
return organisms toward physiological homeostasis and
emotional equilibrium, while unpleasantness and dis-
tress may arise from stimuli that remove an organism
from those respective conditions (Aristotle in "Nic-
homachean Ethics"; Cabanac 1971). Although such con-
cepts can yield general behavioral predictions in animal
research (as generalized response guiding principles) and
may even operate through discrete neurochemical sys-
tems of the brain, such as dopamine (Wise 1982) or
endorphins (Panksepp 1981b; Stein & Belluzzi 1978),
they will not be discussed further here because currently
their behavioral indicators in animals are rather subtle
and diffuse; incisive approaches to the neurobiological
analysis of these more generalized phenomena may first
BEHAVIORAL AND BRAIN SCIENCES (1982)3 411
 Panksepp: Psychobiology of emotions

FOREWARD LOCOMOTION, SNIFFING, INVESTIGATION

STIMULUS-BOUND-
APPETITIVE
BEHAVIOR
AND
SELF-STIMULATION

a.
U

o STIMULUS-BOUND
FLIGHT
"STIMULUS-BOUND
DISTRESS VOCALIZATION
AND AND
ESCAPE BEHAVIORS EXPLOSIVE BEHAVIORS

i
N
Ui
Ul

STIMULUS-BOUND-
BITING
AND
AFFECTIVE ATTACK

ATTACK, B I T I N G , FIGHTING

Figure 2. Emotive-command systems are defined primarily by neural circuits from which well-organized behavioral sequences can
be elicited by localized stimulation of brain tissue. The approximate locations of these systems are depicted on hemifrontal
hypothalamic sections of the rat brain; cp - cerebral peduncle; dm - dorsomedial nucleus; mf - medial forebrain bundle; mt
-mammillothalamic tract; ot - optic tract; re - nucleus reuniens; vm - ventromedial nucleus; x - fornix; zi - zona incerta. The major
behavioral outputs of each command system are indicated. The various command systems are anatomically close to each other and
presumably interact so as to maintain behavioral focus on a single class of behaviors. It is assumed that the various possible
interactions among systems lead to second-order emotive states consisting of blended activities across the primary systems. Some
possible interactions are outlined. Some systems may also have biphasic interactions depending on the level of activity within an
emotive system and the past experiences of the animal.

require clarifying the mechanisms that organize the more
discrete emotive states of the brain.
Within these constraints, the analysis of the brain
mechanisms mediating emotions becomes a more man-
ageable undertaking. Only certain neural circuits begin
to fulfill the definitional requirements of emotive sys-
tems, and many of the fuzzier dimensions of affective life
can be temporarily excluded from consideration.
Emotive circuits of the brain: The theory
Emotional concepts can be tied to specific brain mecha-
nisms through a variety of maneuvers. The idea that
emotions simply arouse (Duffy 1941) served as a theoreti-
cal basis for proposing that the ascending reticular activat-
ing system is a physiological substrate for emotion
(Lindsley 1950). The fact that localized stimulation of the

412 BEHAVIORAL AND BRAIN SCIENCES (1982)3


limbic system could induce both positive and negative
affective states, as indicated by species-typical approach
and escape behaviors, suggested that emotions and rein-
forcements are intrinsically organized by dichotomous
affective circuits of the brain (Glickman & Schiff 1967).
Although the most illuminating results, with regard to
how emotions are organized in the nervous system, have
been obtained using localized electrical stimulation of
brain tissue with indwelling electrodes in awake animals
(Hess 1957), I believe the evidence supports the exis-
tence of a more highly articulated emotive "phrenology"
in the limbic system than has been postulated before.
It is astounding that the simple application of electrical
"noise" to certain brain areas can yield a variety of well-
coordinated species-typical behavior sequences, most
notably ones which appear to reflect emotional states. By
stimulating specific areas in the hypothalamus, we can
induce animals to exhibit ragelike behaviors, explorato-
ry-investigative behaviors, flight, and several other be-
havior patterns indicative of complex neuropsychological
integration. The waveform of the electricity applied to
the brain tissue does not seem to be orchestrating the
actual sequences of the artificially elicited behavior. The
simplest explanation for these remarkable effects is that
we are stimulating neural networks that can facilitate
(command) activity in distant sensory filtering systems
and motor patterning circuits governing the sequencing
of emotive acts.
The activation of these systems not only induces emo-
tional behaviors and concurrent visceral and hormonal
changes (Mancia & Zanchetti 1981), but it also sensitizes
sensoryfieldson the body surfaces. In cats, hypothalamic
stimulation yielding attack, increases perioral biting re-
flexes, the probability that touch to the arm will induce
lashing out, and aggressive responsivity of the con-
tralateral eye (Flynn 1976). In rats, hypothalamic stimula-
tion that induces stimulus-bound appetitive behaviors
can increase perioral (Smith 1972) and visual (Beagley &
Holley 1977) sensitivity, and damage to these systems is
accompanied by sensory-motor neglect (Balagura,
Wilcox, & Coscina 1969; Frigyesi, Ige, Iulo, & Schwartz
1971; Marshall & Teitelbaum 1974). Also, even though
the most spectacular kinds of stimulus-bound behavior
are strictly limited to the duration of brain stimulation,
more subtle behavioral changes persist for a while. For
instance, Hess (1957) and Nakao (1958) detected affective
changes in their cats for minutes following hypothalamic
stimulation. Similarly, the increased motor activity in-
duced by stimulation of lateral hypothalamic reward sites
decays gradually during the minute following stimulation
(Rolls & Kelly 1972). The general functions of these
circuits appear to be genetically coded (Roberts & Ber-
quist 1968), although the exact behavioral expressions of
each system may be quite malleable (Valenstein, Cox, &
Kakolewski 1970).
This paper is based on the assumption that these
"stimulus-bound" behaviors reflect species-typical ex-
pressions of class-typical brain circuits which mediate
emotions, and, hence, that a realistic and scientifically
useful taxonomy of emotions, in humans as well as other
mammals, could be based upon the number of distinct
behavioral control systems that can be activated in this
manner. However, difficulties arise in enumerating the
number of emotive command systems, for the number of
Panksepp: Psychobiology of emotions
discrete behaviors that can be observed during such
stimulation appears to be greater than the number of
discrete command systems actually present under the
stimulating electrodes. Of course, this is reasonable if one
considers that the function of such executive systems may
be to facilitate simultaneously a number of related behav-
ioral options so that reinforcement processes can mold
adaptive behavior sequences for a variety of related life
circumstances (Figure 1), which differ in detail.
In any event, this discrepancy poses great problems for
accurate categorization, as is apparent in recent contro-
versies over the nature of the circuits which mediate
"stimulus-bound" appetitive behaviors in the hypo-
thalamus. For instance, from a relatively diffuse zone
centered around the dorsolateral hypothalamus of the rat,
we can evoke feeding, drinking, gnawing, hoarding, pup
retrieving, predatory aggression, and copulation. Are
there discrete circuits under our electrodes for each of
these behavioral acts (the specificity hypothesis), or are
they all induced by a unitary neural influence (the non-
specificity hypothesis)? Without going into details, which
can be found elsewhere (Panksepp 1981a), current evi-
dence overwhelmingly supports the nonspecificity hy-
pothesis for such stimulus-bound appetitive behaviors.
Aside from the basic demonstration of behavioral mal-
leability after lateral hypothalamic stimulation (i.e., one
appetitive behavior can be changed to another through
experience: Valenstein et al. 1970), the most important
evidence supporting such a conclusion is: (1) "Stim-
ulus-bound" appetitive behaviors do not change when an
electrode is passed through the relevant lateral hypothal-
amic field of a single animal (Wise 1971). (2) The type of
behavior elicited from an electrode does not change when
the tissue under the electrode is damaged; instead, the
threshold for elicitation of the behavior is increased
(Bachus & Valenstein 1979). (3) The temperament of the
test animal may be more important in determining the
stimulus-bound appetitive behavior observed than the
exact location of the stimulating electrode in the active
zone within the dorsolateral hypothalamus (e.g., stim-
ulus-bound predatory attack in rats is most easily ob-
tained in animals that exhibit some tendency to spon-
taneously emit this behavior: Panksepp 1971a).
Still, this is not to suggest that several distinct emotive
circuits do not course through the lateral hypothalamic
field where stimulus-bound behaviors are most fre-
quently encountered, for there appear to be several
distinct classes of behavior that can be activated (Figure
2). Also, the present conception of nonspecificity does not
imply that the functions of the underlying circuits are not
genetically determined. Although each emotive circuit
may be nonspecific in the sense that it can provoke
several distinct emotive actions, they may all be specific
(hard-wired), in the sense that each generates a single,
homogeneous psychobehavioral tendency (Figures 1 and
2). Presumably, an analysis of these partially interdigitat-
ing, transhypothalamic emotive command circuits pres-
ently provides the best foundation for neurophysiological
understanding of emotions in both animals and humans.
To some investigators in the field this may be self-
evident, but the scarcity of neuro-psycho-behavioral
investigations based on this assumption suggests that it
has not yet been widely accepted.
Herein I propose four fundamental transdiencephalic
BEHAVIORAL AND BRAIN SCIENCES (1982)3 413
Panksepp: Psychobiology of emotions

emotion-mediating circuits passing between midbrain,
limbic system, and basal ganglia of the mammalian brain.
Based on the extreme emotional experiences that these
systems are presumed to mediate in humans, they are
labeled the "expectancy," "rage," "fear," and "panic"
circuits. Their specific neuronal projections and interac-
tions can only be surmised, but from localized
brain-stimulation studies we do know the approximate
locations of these circuits in the several species. Although
sites have been identified in diverse areas of the brain, at
present the hypothalamus is the best site in which to
pursue systematic study of the properties of these cir-
cuits. Such executive circuits (Figure 3) are likely to be
activated by sensory information concerning various
classes of environmental events (Figure 1), as well as
stimuli arising from internal states (autonomic reafferents
and homeostatic states of the body). Once activated,
these circuits arouse and organize related arrays of soma-
tic (Figure 2), hormonal, and visceral sensory-motor
processes. This paper will focus on the psychobehavioral
attributes of these circuits.
Expectancy. It is proposed that through the medial fore-
brain bundle of the lateral hypothalamus passes a gener-
alized "foraging-expectancy" circuit, which is sensitized
by major homeostatic imbalances in the body and their
respective environmental incentives. The major adaptive
function of this circuit is to produce a specific type of
motor arousal - characterized by exploratory and investi-
gative (i.e., foraging) activities - which induces an animal
to move from where it is to where it should be in order to
consume the substances needed for survival. In artificial
experimental situations, this circuit can also mediate a
highly energized form of electrical self-stimulation of the
brain. The term expectancy has been selected as the
major label for this system, because it is assumed that the
circuit readily comes under the conditional control of
environmental cues and is thereby essential for elaborat-
ing anticipatory appetitive behaviors. Accordingly,
throughout this paper expectancy is intended only in the
positive sense, as hope, desire - joyful anticipation,
rather than in the negative sense in which it can also be
used.
Fear. It is proposed that sites in the hypothalamus from
which flight and unconditional escape behaviors can be
elicited represent the trajectory of fear circuitry, the
major adaptive function of which is to respond to all
external stimuli that have the potential of harming or
hurting the body.
Rage. It is proposed that sites from which angry emo-
tional displays and affective attack can be elicited repre-
sent the location of a rage circuit, the major adaptive
function of which is to invigorate behavior when activity
in the expectancy circuit diminishes rapidly as well as
when the body is irritated or uncomfortably restrained.
Panic. It is proposed that sites from which distress vocal-
izations and explosive agitated behavior can be elicited
represent the approximate trajectories of panic circuitry,
the major adaptive function of which is to sustain social
cohesion among organisms whose survival depends on
reciprocity of care-soliciting and care-giving behaviors.
The approximate hypothalamic locations of these cir-
cuits are depicted in the hemifrontal sections through a
rat hypothalamus in Figure 2. It is assumed that these
four life-challenges (Figure 1) are archetypal, and that
the evolution of such extensive systems in the brain was
dictated by the adaptive advantage of rapidly acting

Figure 3. A simplified schematic of the trajectory of a transdiencephalic emotive command system on a parasaggital view of a rat
brain. The system collects various type of processed sensory information along its trajectory; for instance: (1) perceptual influences
from temporal and frontal lobes and various basal ganglia; (2) somatosensory inputs from the thalamus; (3) homeostatic inputs from
body-state detectors in the hypothalamus; (4) decoded olfactory, vomeronasal, and other perceptual inputs from midline thalamic
nuclei; and (5) simple sensory controls such as those arising from taste, touch, sound, and pain from lower brainstem levels. The
system is conceived as elaborating motor control by ascending influences (e.g., behavior sequencing mechanisms of basal ganglia) as
well as by descending control of emotion-appropriate autonomic and somatic reflex control circuits (6).

414 BEHAVIORAL AND BRAIN SCIENCES (1982)3


command circuits, which could simultaneously recruit
multiple body systems to meet major environmental
emergencies.
The command-system construct has been much used
in the analysis of stereotyped motor patterns in inverte-
brates, although it remains surrounded by many contro-
versies (Kupfermann & Weiss 1978). The analogous na-
ture of "stimulus-bound" behaviors evoked by
hypothalamic stimulation suggests that this principle can
guide investigation of certain brain systems in mammals.
Although the neural trajectories and the exact functions of
these circuits have yet to be mapped accurately, I shall
attempt to summarize existing knowledge of these puta-
tive emotive command systems. This review is highly
selective and emphasizes only materials most influential
in my own thinking. A more comprehensive coverage of
past work related to many of these issues, can be found in
Panksepp (1981a).
The "expectancy" command system. A unified executive
sensory-motor system for the mediation of explora-
tion-approach-investigation (i.e., foraging) is proposed
to be isomorphic with neural circuits mediating electrical
self-stimulation of the lateral hypothalamus (LH). In rats,
LH self-stimulation is characterized by increased forward
locomotion with persistent sniffing, exploration, and ma-
nipulation of prominent objects in the environment
(Christopher & Butter 1968). Activity in this circuit may
mediate the incentive attributes of rewards (Trowill,
Panksepp, & Gandelman 1969), and it is now known that
its cell are highly responsive to both external incentives
and internal homeostatic imbalances (Oomura 1980;
Panksepp 1981a; Rolls 1980). It is apparent that activity in
this circuit promotes an animal's capacity to seek out
biologically important stimuli in the environment, in part
by invigorating motor behavior (Wayner, Barone, &
Loullis 1981) and also by modifying sensory acuity
(Gellhorn, Koella, & Ballin 1955; Marshall 1979; Smith
1972). A fundamental requirement for survival is that an
animal move from where it is to where life-sustaining
objects are located. Lateral hypothalamic stimulation
potentiates the requisite behavioral tendencies in all
vertebrates studied, suggesting that a primitive executive
system designed to trigger foraging activities courses
through the forebrain bundles of the ventral dien-
cephalon. Although the behavioral expressions of arousal
in this circuit, such as genus-typical consummatory ac-
tivities (including feeding, drinking, gnawing, object car-
rying, hoarding, predatory aggression, and copulatory
behavior), may vary substantially across genera, the com-
mand structure appears to have been comparatively re-
sistant to evolutionary modification; this persistance sug-
gests that an effective solution for coordinating bodily
activities in response to archetypal survival needs was
sustained, functionally intact, during the diversification
of vertebrate species into different ecological niches.
However, the variety of genus-typical expressions of
activity in this system could explain the relative ease or
difficulty with which self-stimulation has been observed
in various species - it being readily obtained in rats
(because they exhibit energetic foraging) but not so read-
ily in cats (because they exhibit a more restrained form of
stalking).
Although the phenomenon of self-stimulation has been
Panksepp: Psychobiology of emotions
used to derive biological significance for such traditional
behavioral concepts as reward and reinforcement, as well
as for psychological states such as pleasure (see Panksepp,
1981a, for review of the literature), the highly energized
behavior of an animal electrically stimulating its own
lateral hypothalamus has remained enigmatic from such
perspectives. Although there may be considerable diver-
sity in the response topographies that individual animals
exhibit during self-stimulation (perhaps partially because
of overlap with other emotional command systems), most
animals move forward, sniff intensely, and interact with
many objects in the environment, especially those ob-
jects with strong intrinsic connections with the activated
circuits (e.g, odors). The invigorated explorato-
ry-investigative activity of such animals is just what
would be expected if a nonspecific circuit specialized for
the generation of incentive-directed (foraging) behavior
were activated. On a priori grounds, it would be advan-
tageous for activity in such a circuit to be amplified by
internal need states as well as in response to positive
incentives, until contact with an appropriate consumma-
tory object was achieved and until internal homeostatic
detectors provided satiety and inhibition of further
foraging.
Many of the unusual behavioral characteristics that can
be obtained by stimulating this emotive system (e.g., the
unusual motivational characteristics discussed by Trowill
et al., 1969, and the behavioral "plasticity" described by
Valenstein et al., 1970) can be explained by the way in
which normal control gates in the circuit are superseded.
Artificial activation of the circuit could override normal
input signals, such as the inhibition that may arise di-
rectly from consummatory acts as well as those arising
from replenishment of body needs. This would not only
make stimulus-bound appetitive behavior more agitated
than that normally seen in animals (i.e, foraging arousal
being sustained during reward consummation), but due
to the absence of accruing negative feedback signals,
there would be a relative absence of satiation during self-
stimulation of the circuit. Also, since the underlying
system is nonspecific - subserving a general behavioral
function, which can be accessed by various positive en-
vironmental incentives as well as by homeostatic need
detectors - enforced activation of this system could lead
to a variety of specific stimulus—bound consummatory
behaviors. These would be interchangeable, depending
not only on the availability of goal-objects but also on
each animal's past foraging history. Furthermore, since
the stimulation evokes a generalized emotive state, which
is artificially uncoupled from homeostatic inputs, the
specific stimulus-bound behaviors observed during ar-
tificial activation of the circuit could develop characteris-
tics that are independent of momentary physiological
needs.
Expectancy was selected as a label for this neuronal
network to emphasize the fact that this system mediates
anticipatory appetitive behaviors and, presumably, the
corresponding subjective state in humans. In animals,
the activity of this system comes spontaneously under the
conditional control of correlated environmental cues
(Clarke & Trowill 1971; Olds 1975; Rolls 1980). Lateral
hypothalamic cells of rats fire with increasing frequency
when the animal is approaching food; they slow down
dramatically when the animal starts to eat, but promptly
BEHAVIORAL AND BRAIN SCIENCES (1982)3 415
Panksepp: Psychobiology of emotions
resume rapid activity when its food is taken away (Ham-
burg 1971). Moreover, anticipatory enhancement of
motor activity induced by signals predicting food is
markedly attenuated by lateral hypothalamic lesions
(Campbell & Baez 1974).
The postulated existence of a discrete circuit, designed
to energize appetitive foraging behaviors, readily ex-
plains most of the behavioral changes that have been
observed following manipulations of the lateral hypo-
thalamus and suggests a variety of behavioral and neu-
rophysiological hypotheses. At a behavioral level, manip-
ulation of the system should generally modify the capacity
of an organism to seek out biologically important stimuli
in its environment; and changes in the cellular activities
of the system during appetitive tasks (e.g., Hamburg
1971) should generalize across different positive rewards.
Perhaps the most objective physiological-behavioral in-
dicator of the state of this system in rats and other species
in which olfaction predominates would be the level of
sniffing, not only during spontaneous exploration, but as
correlated with environmental cues that predict environ-
mental rewards (Clarke, Panksepp, & Trowill 1971;
Clarke & Trowill 1972). Indeed, for the past decade we
have routinely identified lateral hypothalamic self-stim-
ulation sites in anesthetized animals during surgery by
monitoring sniffing evoked by electrical stimulation of
hypothalamic tissues. This measure could be used as a
straightforward behavioral assay for tapping the momen-
tary state of the "foraging-expectancy" system.
This conception does not preclude the possibility that
specific neuronal circuits for the various motivational
states (e.g., thirst, hunger, salt appetite, sexual behavior)
exist in the hypothalamus. Homeostatic receptor systems
appear to exist in medial strata of the hypothalamus; these
systems presumably modulate activity in the generalized
foraging system of the lateral hypothalamus and probably
have drive-specific interconnections to brain circuits
which control consummatory acts (for a review, see Pank-
sepp, 1981a).
The "rage" command system. The first hypothalamic
emotive system to be discovered was the one mediating
rage (Hess 1932; Ranson 1934), although the idea that this
system elaborated a true emotional state was not accepted
by all (Masserman 1941). It is now clear that, even though
occasional electrode sites are encountered, which may
generate affective displays apparently void of true emo-
tional content, most electrodes placed in this circuit
generate an "affective" attack, which is well integrated
with environmental events (Nakao 1958). Indeed, in
monkeys, hypothalamically elicited aggression is di-
rected along dominance lines established prior to stim-
ulation. Robinson, Alexander, and Browne (1969) report
that submissive animals direct their attacks primarily
toward dominant ones, even to the point where stable
dominance reversals are established. Delgado (1969) and
Alexander and Perachio (1973) have also found stim-
ulus-bound aggression to be guided by preexisting social
rank, but they observed attack to be directed most fre-
quently toward submissive animals. In general, most
recent studies indicate that artificial activation of the
hypothalamic rage circuit precipitates aggression, which
is well integrated with an animal's normal life circum-
416 BEHAVIORAL AND BRAIN SCIENCES (1982)3
stances, rendering untenable the conclusion that this
circuit simply triggers vacuous emotional displays.
Hypothalamic stimulation that generates affective at-
tack has been found to be aversive in cats (Nakao 1958),
monkeys (Plotnick, Mir, & Delgado 1971), and rats
(Panksepp 1971a). However, this circuit overlaps sub-
stantially with expectancy systems. Thus, both stim-
ulus-bound "affective" attack and agitated self-stimula-
tion (characterized by vigorous biting of the
manipulandum) can be obtained from hypothalamic
zones in which these systems presumably intermingle.
Circuit overlap appears to be especially dense in tissue
just ventrolateral to the fornix columns, with the active
field for rage extending ventrally and medially and the
most sensitivefieldfor self-stimulation extending dorsally
and laterally (Panksepp 1971a).
Such mingling may complicate the dissection of under-
lying circuitries, but it also points to the probable interac-
tion of these systems in the control of behavior. Adaptive
considerations suggest that rage and expectancy circuits
should strongly inhibit each other. A high level of forag-
ing and anticipation should dampen aggressive impulses,
whereas anger, and the resulting increased vigor of be-
havior, should be intensified when expected rewards are
not forthcoming. Thus, the increased responding ob-
served at the beginning of extinction (i.e., the "frustra-
tion effect") may be due to a release of the rage circuit,
and selective damage to that circuit should reduce the
effect. Electrical stimulation applied to zones of overlap
in these circuits may yield other unusual behavioral
symptoms, for, typically, the two systems should not be
active concurrently. For instance, self-stimulation at
these sites may be sensitive to stimulus duration, with
animals exhibiting vigorous responding for short pulse
durations and poor self-stimulation for long durations.
Also, such animals may exhibit vigorous biting of the
manipulandum during self-stimulation.
The most objective and easily quantified behavioral
indicators of activity in the rage system are vocal displays,
such as growling (Jiirgens & Ploog 1970), hissing, and the
potentiation of striking and biting behaviors (Flynn, Ed-
wards, & Bandler 1971). This last measure has yielded
data that are highly consistent with the proposed reci-
procity of the rage and expectancy systems. Both the
activation of hypothalamic attack sites and the deactiva-
tion of self-stimulation sites induce biting (Hutchinson &
Renfrew 1978). Accordingly, with the use of such mea-
sures (summarized in Figure 2), clear predictions can be
made concerning the interactions that should be ob-
served during concurrent activation of "pure sites" within
these systems.
Few investigations have sought to evaluate the effect
on naturally occurring aggression of precise surgical
damage to the rage command system at the hypothalamic
level. This omission may be due, in part, to the fact that
lateral hypothalamic lesions precipitate a severe gener-
alized amotivational syndrome (Ranson 1939; Strieker &
Zigmond 1976), which makes it difficult to interpret
experiments with large LH lesions (e.g., Karli & Vergnes
1964; Panksepp 1971c). Still, no thorough investigation
has been conducted that has behaviorally verified the
placement of electrodes before making of lesions; from a
command-system perspective, such a verification seems

essential for deriving meaningful conclusions from lesion
work.
Although the rage command system is viewed as the
substrate for affective attack, diverse forms of aggression
can be distinguished in terms of their eliciting conditions
(Moyer 1976). Presumably many of these stimuli con-
verge upon a common executive system, although certain
forms of aggression, such as predation, need not arise
from the activity of this system. The hypothalamic circuit-
ry mediating these behaviors has been distinguished
anatomically (Flynn 1976) and affectively (Panksepp
1971a). Indeed, from the present perspective, quiet-
biting attack forms of predatory aggression are considered
to be expressions of the foraging-expectancy system, or
perhaps a mixture of activity in several systems. Similar-
ly, defensive aggression may be a mixture of concurrent
activity in the fear and rage systems (Panksepp 1979a).
The "fear" command system. In comparison to lateral
hypothalamic expectancy and rage systems, the proper-
ties of circuits that mediate escape and flight behaviors
(i.e., the fear emotive system) are poorly understood.
Although the feeling of anxiety has been precipitated in
humans by stimulation of several brain areas, especially
in the temporal lobe (Chapman 1958; Heath, Monroe, &
Mickle 1955; Jasper & Rasmussen 1958), the most ex-
pressive displays of fear in rats are observed during
stimulation of the anteroventral hypothalamus (Panksepp
1971a). Anxiety is also precipitated by electrical stimula-
tion of this zone in humans (Grinker & Serota 1938;
Monroe & Heath 1954).
Hypothalamic sites that provoke flight surround the
core system for which rage (threat behavior) is elicited (de
Molina & Hunsperger 1962), and there appears to be
substantial anatomical overlap and functional interaction
between these systems. Accordingly, many of the behav-
ioral patterns elicited from this area are characterized by a
mixture of flight and threat (i.e., defensive behavior).
Although stimulation of the intermediate zones typically
elicits flight, defensive attack often ensues, if no avenue of
escape is available. Concurrent activation of the two
systems with independently placed electrodes can yield
synergistic effects, with either threat or flight being
intensified (Hunsperger, Brown, & Rosvold 1964).
The affective state elicited by stimulation of sites pro-
ducing flight is aversive, even though self-stimulation can
also be obtained from some sites. However, self-stimula-
tion behavior obtained from such electrodes is accom-
panied by flight responses (Panksepp 1971a). In rats, such
self-stimulation is characterized by crescendos of lever
pressing interspersed with frequent running away from
the lever and energetic leaps upward.
Because the simple behavioral measure of aversion
does not readily discriminate between several trans-
hypothalamic emotive systems, the best indicator vari-
ables for the state of the fear system may be derived from
the response topographies of species-typical flight re-
sponses. In the rat, one highly characteristic response
elaborated by this system is an upward leap. At anterior
lateral hypothalamic sites, this response is so systematic
that it can be used quantitatively with repeated brief
stimulations (Panksepp 1971b).
Although the existence of flight patterns elicited from
Panksepp: Psychobiology of emotions
the hypothalamus has been recognized since Hess's pi-
oneering work, the fact that this behavior has not been
extensively used to analyze the brain organization of fear
(and, hence, anxiety) suggests that it has not generally
been regarded as reflecting a true emotional state, but
rather only as a motor patterning mechanism. Such an
interpretation has probably been reinforced by the many
failures to condition avoidance behaviors with hypothala-
mically elicited flight as the unconditioned stimulus (e.g.,
Bower & Miller 1958; Roberts 1958; Wada & Matsuda
1970). But such conditioned avoidance has been obtained
by others (Delgado, Roberts, & Miller 1954; Nakao 1958),
and it may be essential to have "pure" fear sites, or
perhaps some concurrent stimulation of pain inputs, to
obtain efficient avoidance behavior. This may explain
why flight behavior obtained from the periventricular
gray has proved more readily conditionable (Wada, Mat-
suda, Jung, & Hamm 1970).
Even though "anxiety has been considered the alpha
and omega of psychiatry" (Delgado 1969, p. 131), hypo-
thalamically elicited flight remains to be used as a general
model for analyzing neural substrates of anxiety. Nor has
any substantial attempt been made yet to analyze the role
of this neural system in the elaboration of natural avoid-
ance and conditioned emotional responses. If such a
command system plays a cardinal role in the elaboration
of fear, it is to be expected that selective damage to the
circuit (as might be obtained with the use of behaviorally
verified electrode placements) would markedly attenuate
the development of conditioned fear, as well as those
unconditional fear responses that are initiated by percep-
tual inputs rostral to the level of brain damage. There is
substantial evidence that deficits in the elaboration of fear
result from damage to ascending projections of this sys-
tem in the amygdala (Kliiver & Bucy 1937; Ursin 1965).
The "panic" command system. This may be the most
controversial of the emotive systems to be proposed in
this paper. It is not typically found among the traditional
lists of emotional responses attributed to the visceral
brain via brain stimulation techniques. However, Cogan
(1802) and many others have entertained the idea that
sorrow may be a fundamental emotion (Table 1), and, in
the present context, sorrow (defined as the diminutive
form of panic, in the same manner that anxiety may be a
diminutive form of fear, and anger a diminutive form of
rage) appears to reflect activity in a basic emotive system
of the brain. Our past work on separation-induced dis-
tress in various species has been premised on the exis-
tence of such an emotion. The characteristic and univer-
sal emotional consequence of socially isolating young
animals is protest, especially as reflected in the objective
behavior of crying. This response appears to be mediated
by an innate brain system that is attuned to the presence
and absence of social stimuli.
Our work in this area (summarized in Panksepp, Her-
man, Vilberg, Bishop, & DeEskinazi 1980; Panksepp
1981b) affirms that similarities exist between the underly-
ing neurochemical dynamics of opiate addiction and of
social dependence. This suggests that some of the neural
processes of this emotive system may involve brain
opioids; this has been confirmed by brain stimulation
studies (Herman 1979;.Herman & Panksepp 1981). Dis-
BEHAVIORAL AND BRAIN SCIENCES (1982)3 417
Panksepp: Psychobiology of emotions
tress vocalizations in guinea pigs can be elicited by
stimulating many opioid-rich sites, including the dor-
somedial thalamus, ventral septal, and preoptic areas,
and less dense zones in the amygdala and medial hypo-
thalamus. However, these vocalizations follow the termi-
nation of brain stimulation, and therefore may not clearly
indicate the localization of the command circuit for this
emotive system. Stimulus-bound distress calls have
been elicited in the vicinity of the central gray, suggesting
that the executive system for this emotional state inter-
mingles with the other emotive systems at the level of the
medial mesencephalon.
Although rats do not exhibit marked separation-in-
duced crying, stimulation of the anterior basal hypoth-
alamus yields a stimulus—bound aversive response,
which seems distinct from fear- and rage-related behav-
iors. This response consists of a freezing reaction, accom-
panied by apparently strong negative affect, which culmi-
nates suddenly in explosive behavior, during which the
animal exhibits violent leaping about the test chamber
(Panksepp 1971a). Unlike stimulus-bound flight, this
behavioral response typically habituates rapidly, and,
with closely spaced trials, ever-increasing current levels
are needed to reelicit the explosive component of the
behavior sequence. These behaviors may be homologous
to the emotional disturbance known as "panic" attacks
(Klein 1981), and this response may thereby indicate the
hypothalamic location of the emotive command system
that is attuned to social loss. It is noteworthy that similar
explosive behavior patterns occur in addicted animals
during opiate withdrawal.
Although activity of the panic system may be reflected
in a variety of agitated behavioral patterns, separation-
induced distress vocalization is currently the best indica-
tor of arousal in this system. The activity of this emotive
system should also be expressed in the tendency of
animals to exhibit social cohesion, and decreased gregari-
ousness has been observed in rats following ventromedial
and anterolateral hypothalamic lesions (Enloe 1975). Fur-
ther analysis of social cohesion and separation distress
after damage to the various brain areas where stim-
ulus-bound crying and explosive behaviors can be elic-
ited will be needed to clarify the behavioral characteris-
tics of this emotive system.
Properties of emotive systems
Anatomy of the emotive systems of the visceral
brain. The actual neuronal trajectories of the proposed
emotive systems can only be sketched in broad outline.
Although it is clear that lateral hypothalamic neuropil
contains widely ramifying, long-axoned systems, both
ascending and descending, with multiple interneuronal
feedback loops (Millhouse 1979; Palkovits & Zaborszky
1979), our knowledge of functional circuits in such a
reticular substance remains too imperfect to attempt
filling in the details of system connectivities. The gross
anatomy of some systems can be surmised from anatomi-
cal studies of self-stimulation and stimulus-bound appe-
titive (e.g., Gallistel, Karreman, &Reivich 1977; Roberts
1980; Routtenberg & Malsbury 1969; Wise 1978) and
attack behaviors (Siegel & Edinger 1981) as well as from
studies analyzing the effect of brain lesions on behaviors
418 BEHAVIORAL AND BRAIN SCIENCES (1982)3
evoked from the hypothalamus (e.g., Berntson 1972).
Although the details of the underlying systems remain
to be determined, these putative command systems
probably run from the mesencepahlon, through the re-
ticular fields of the hypothalamus and, perhaps,
thalamus, to basal ganglia and higher limbic areas (Figure
3). Each command system probably emits ascending and
descending controls throughout its trajectory, not only to
interconnect with other behavior control circuits and to
bias sensory and motor processes, but also to coordinate
the activity of each system with homeostatic and autono-
mic states of the body. Thus the trajectory of emotive
command circuits through the hypothalamus may repre-
sent a design feature that permits diverse bodily states to
be brought in line rapidly with behavioral demands.
Some of the above issues are highlighted in the sche-
matized command network in Figure 3. Were this to
represent the command system for panic, it would be
activated normally by social separation. It might bias
ascending sensory projection areas in the cortex, tuning
certain systems to be especially sensitive to social stimuli;
it might activate behavior sequencing mechanisms in the
basal ganglia specialized to generate discrete social sig-
nals (e.g., MacLean 1981); it might establish hypoth-
alamo-pituitary balances to increase ability to cope bet-
ter with stress; and perhaps, via descending influences, it
might bias specific somatic motor tendencies, such as
increased agitation, distress vocalization, and the sensi-
tization of other care-soliciting behavior patterns. At the
same time, overall autonomic balance may be shifted
toward parasympathetic dominance so as to conserve
bodily energy resources. The command system itself
would be modulated by various sensory inputs along its
trajectory as illustrated in Figure 3. Since the system can
be accessed at various levels of the neuroaxis, learned as
well as unconditional influences could modulate the ac-
tivity of the command circuit at diverse points along the
trajectory of the system. Conceptualizing emotive cir-
cuits in this way would permit many rudiments of goal-
directed emotive behaviors to remain operational even
after substantial damage to both limbic forebrain and
hypothalamic tissues (e.g., Ellison & Flynn 1968; Huston
& Borbely 1974).
Neurochemistry of the emotive command sys-
tems. Whether the various emotive control systems mod-
ulate the activities of output circuits via single- or multi-
ple-command transmitters remains unknown. From
available evidence, dopamine and acetylcholine are rea-
sonable candidates as key transmitters in the expectancy
and rage systems respectively.
The catecholamine hypothesis of self-stimulation re-
ward has received substantial support (for reviews, see
German & Bowden 1974; Stein 1978; Wise 1978), and
both norepinephrine and dopamine facilitate self-stim-
ulation at various sites in the lateral hypothalamus as
determined by pharmacologic studies (e.g., Herberg,
Stephens, & Franklin 1976; Phillips & Fibiger 1973;
Zolovick, Rossi, Davies, & Panksepp 1982), even though
recent anatomical studies do not support the importance
of norepinephrine in the self-stimulation phenomenon
(e.g., Clavier, Fibiger, & Phillips 1976; Corbett & Wise
1979). Stimulus-bound object-carrying (foraging)
evoked from the lateral hypothalamus is reduced follow-
 Panksepp: Psychobiology of emotions
ing destruction of dopamine systems (Phillips 1975), and
pharmacological facilitation of brain dopamine activity
with d-amphetamine invigorates exploratory activities,
including forward locomotion and sniffing. A command
function for dopamine systems seems apt, because selec-
tive receptor agonists applied to the striatum can restore
behavioral competence in animals whose ascending DA
systems have been damaged (Marshall, Berrios, &
Sawyer 1980). This suggests that the system works as a
permissive response-biasing mechanism rather than as a
precise information-transfer system.
Acetylcholine may subserve a command function in the
rage system, since applying cholinomimetics into the
hypothalamus and amygdala can provoke rage responses
(Baxter 1968; MacPhail & Miller 1968; Myers 1964). The
observation that cholinergic agonists decrease self-stim-
ulation (Stark & Boyd 1963), whereas anticholinergics
increase such behavior (Pradhan & Kamat 1973) is con-
sistent with the proposed reciprocal inhibitory relations
between the expectancy and rage systems (Figure 2).
Additional evidence for such an interaction is provided by
the tendency of catecholamine depletion in the brain to
make animals irritable (Coscina, Seggie, Godse, &
Stancer 1973; Eichelman, Thoa, & Ng 1972; Nakamura &
Thoenen 1972). Furthermore, a reciprocal action of these
neurochemical systems is apparent in the nigrostria-
tal-striatonigral neuronal loop where dopamine appears
to inhibit acetylcholine cells, and acetylcholine, through
a GABA interneuron, inhibits dopamine cells (see Carls-
son 1965; Groves, Wilson, Young, & Rebec 1975; Iversen
1978; Pradhan & Bose 1978).
Command transmitter candidates for panic and fear
systems remain more elusive, but major inhibitory influ-
ences on these emotive systems may arise via the ben-
zodiazepine receptor (Braestrup & Squires 1977) and Figure 4. Biogenic amines that are widely dispersed through-
endorphin systems (see Panksepp et al., 1980, for re- out the brain may provide nonspecific excitatory and inhibitory
view). Further investigation of the various peptide net- control over emotive command circuits. The frontal sections
works of the limbic-enteric nervous system may reveal depict approximate locations of brain serotonin (5-HT) and
command transmitters for these circuits. norepinephrine (NE) systems at the mid-hypothalamic level as
If emotive systems have strong reciprocal interactions determined by autoradiographic localization of (14C), 5-HT,
with each other, a broad range of specificity in neu- and NE uptakes in the mouse brain. (Photomicrographs accord-
rochemical controls can be expected, ranging from non- ing to Masuoka and Alcaraz, 1975, reprinted by permission).
specific modulatory controls to quite precise coding with-
in a single emotive pathway. It appears that the major
brain amines, serotonin and norepinephrine, exert non- behavioral functions (for review, see Panksepp, 1981b).
specific modulatory control over these circuits (Figure 4). Still, it may eventually be possible to extract functional
Facilitation of brain serotonin activity typically reduces subsystems from such transmitter thickets by the use of
all waking activities, whereas decreases in serotonin ac- new approaches such as "subtractive autoradiography"
tivity can increase many motivated behaviors (see Pan- (Panksepp & Bishop 1981).
ksepp, 1981a, for review). Modulation of brain nor-
epinephrine generally has opposite effects, even though Interactions among emotive systems. The limitless vari-
the evidence is not so clear-cut as for serotonin, especially ety of human emotional experience no doubt beggars
with respect to the fear and panic systems. No doubt detailed description. Yet the rich tapestry of human
there are exceptions to this generalization, but the classic emotion may arise largely from interactions among primi-
idea that these amines provide nonspecific inhibitory and tive emotive systems. For instance, emotions that Cogan
excitatory control over behavior (Brodie & Shore 1957) (1802) included in his Class II are likely to be socially
remains generally consistent with existing evidence. Sim- learned interblendings of the basic emotions of Class I
ilarly, it is to be anticipated that most of the other putative (Table 1), yielding impressive subtleties of human feel-
transmitters that have been identified have multiple ing. The emotion of jealousy may arise from some admix-
functions, and this overlapping may preclude identifica- ture of panic, rage, and expectancy. Resentment may
tion of functional systems by the simple analysis of overall result from a blend of activities in the rage, fear, and
neurochemical geographies in the brain. For instance,
expectancy systems. Joy may be the fulfillment of expec-
brain opiate receptor systems are widespread, and pres-
ently the endorphins are being implicated in a host of tancies, and so on with other affective alchemy. How
complex emotions may arise from the intermixtures of

BEHAVIORAL AND BRAIN SCIENCES (1982)3 419

Panksepp: Psychobiology of emotions
more fundamental elements has received considerable
attention in literature on humans (e.g., Izard 1972; Plu-
tchik 1980).
Although experimental work testing such speculations
remains to be initiated, the phenomenological richness
that may arise from blended activities of primitive emo-
tive systems could provide useful clues and hypotheses
concerning neural interactions that transpire among the
emotive circuits of the brain. For instance, it seems to be
universal human experience that positive expectations
and anger are psychologically incompatible states. From
this, one might deduce that two hypothalamic cells,
electrophysiologically identified to belong to these two
emotive systems, would exhibit reciprocal inhibition.
Similar predictions might be generated for other system
interactions (some personal opinions are summarized in
Figure 2), but such work should be preceded by system-
atic empirical analysis of interactions that occur among
emotional states in thoughtful human observers.
Learning, reinforcement, and transhypothalamic emotive
systems. Although stimulation of hypothalamic emotive
systems can be reinforcing or punishing, as defined by the
"law of effect," this fact provides no information concern-
ing either the nature of reinforcement or the underlying
learning that can be elaborated by these systems. Pre-
sumably, much of learning is based on the capacity of
affectively neutral stimuli (i.e., those which have weak
unconditional interactions with emotive command cir-
cuits) to obtain increasing influence over emotive cir-
cuits. The likelihood that learning ability is an intrinsic
circuit property of emotive systems has been strikingly
confirmed for the expectancy system. LH-stimulation-
elicited sniffing comes to be spontaneously linked to
predictive cues of the environment [for a theoretical
summary of the work of Clarke and colleagues (Clarke
1971; Clarke, Panksepp & Trowill 1970; Clarke & Trowill
1971) in this area, see Panksepp 1981a].
Although there may be many types of learning (ranging
from habituation to "cognitive" mapping), for didactic
purposes, only one type will be considered here: cue
learning (whereby an emotive circuit comes under the
conditional control of previously neutral stimuli). Cue
learning may arise from the capacity of biologically neu-
tral stimuli to have weak interactions with emotive sys-
tems coupled with the capacity of emotive systems to
modify the sensitivities of sensory systems. As "biolog-
ically relevant stimuli" (i.e., incentives) have strong in-
teractions with emotive command systems (Figure 1),
sensory fields which encode neutral environmental stim-
uli are sensitized, and this neural conjunction may forge
ever stronger relations between the most salient environ-
mental stimuli and the aroused emotive system. Specifi-
cally, rapid changes in the activity of an emotive system
may lead to disinhibition of reinforcement-programming
networks situated at the terminal arborizations of each
command system. Thereby, neutral stimuli, which are
temporally linked to the fluctuating activities of emotive
systems, may develop stronger feedback routes to trigger
zones of a command circuit. Similarly, behavior patterns
that just precede substantial reductions in the activity of
an emotive system may become part of the intrinsic habit
hierarchy of an organism. In other words, habit formation
may operate through a rapid reduction in the activity of
420 BEHAVIORAL AND BRAIN SCIENCES (1982)3
emotive systems, leading to temporary disinhibition of
reinforcement-programming networks, which establish
hierarchies among the behavior patterns just evoked by
activity in the command network; this process could
concurrently tag salient neutral stimuli present in a situa-
tion with biological values. From this perspective, the
analysis of many types of learning may have to be based on
an understanding of emotive circuits of the brain, and
there should be considerable emotive state-dependence
for retrieval of memories, a phenomenon recently docu-
mented in humans (Bower 1981).
In light of recent trends toward rejection of reinforce-
ment mechanisms in the genesis of learned behavior
(Bindra 1978), the existence of emotive circuits might
restore vitality to traditional stimulus-response (S-R) and
stimulus-stimulus learning theories. The command sys-
tem conception readily solves the "response-equiv-
alence problem" that has plagued traditional S-R rein-
forcement learning theory. Also, it can easily handle
many of the striking characteristics of animal learning that
have been revealed in recent years, for instance, auto-
shaping, which may be conditioned foraging, and pre-
paredness, which may reflect the intrinsic inputs and
outputs of emotive systems.
Still, there will surely be many complexities to be faced
when one views the activity of these systems in the
life-span of the organism. Through learning, the degree
to which emotions emerge directly from activities of
emotive command systems could change. Although
transhypothalamic emotive command circuits may be
essential for emotions to become manifest in the develop-
ing organism, higher brain areas may assimilate some
A MANIA
SCHIZOPHRENIA
(PARANOID)
Figure 5. If emotional disorders ultimately arise from un-
balanced activities among primitive emotive systems of the
subcortical brain, then a natural taxonomy of major psychiatric
disorders should be derivable from an analysis of these systems.
At the simplest level, emotional disorders may arise from
under-(—) and over-(+) activity of emotive command systems. A
preliminary suggestion of the types of psychiatric disorders that
may arise from such changes is outlined.

functions of the lower circuits with experience. By such
translocations, as well as direct reafferents to the com-
mand circuits, cognitive appraisal may come to be an
especially important influence in the genesis of adult
emotions. Indeed, a growing body of evidence suggests
that such translocation of instinctive behavior patterns
can occur in the animal brain. For instance, behavioral
deficits resulting from damage to the lateral hypoth-
alamus can be overcome by preoperative experiences
(Ahem, Landin & Wolf 1978; Ruger & Schulkin 1980;
Schwartz & Teitelbaum 1974). Thus, the ontogenetic
approach may be as important for understanding the
functional elaboration of emotive processes in the brain as
it is in behavior (Izard & Buechler 1979).
Diseases of emotive systems. Psychiatric disorders
probably reflect imbalanced activities among basic emo-
tive circuitries of the visceral brain. For heuristic pur-
poses, possible relations between the emotive systems
herein discussed and major emotional disorders are sum-
marized in Figure 5. There is substantial evidence for
some of these possibilities and next to none for others.
Because of the complex interactions that can occur among
systems (Figure 2), this scheme is offered as a stimulus to
further thinking, and will not be elaborated on here. I
would note only that an especially large number of ail-
ments may arise from disorders of the system that medi-
ates social affect.
Concluding remarks
Although the complexity of brain neuropil will most
assuredly humble such a simplistic attempt to understand
emotions as was undertaken here, nevertheless, I think
such analyses may help us progress to more sophisticated
perspectives more rapidly than those which seek to tackle
the full complexity of the problem. Moreover, there is a
distinct advantage to simple schemes in that they are
usually testable, permitting clear refutations, which can
facilitate construction of more viable models and
theories.
The present approach has been based on two funda-
mental assumptions. The first, commonly accepted in this
area of research, is that the evolutionary pressure for the
genesis of emotive circuits emerged from general life-
challenging circumstances, which are to some degree the
shared evolutionary heritage of all mammalian species.
The other, perhaps not so commonly accepted, is that in
order to be scientifically substantive, psychological emo-
tional constructs need to be tied, to some extent, to
specific circuits within the brain. Certainly such designa-
tions must be based initially on circular reasoning and
anthropomorphism. However, further specification and
testing of the biobehavioral ramifications of these circuits
and their interactions should permit an escape from
circularity.
The anthropomorphic terms that I selected for discus-
sion of emotive systems may be objectionable to those
committed to the behaviorist tradition, but they were
chosen because it is assumed that, even though the
species-typical expressions and species-characteristic
vigor of these executive systems may have undergone
great diversification during evolution, their basic natures
Panksepp: Psychobiology of emotions
remain relatively stable in the limbic systems of all
mammals. These labels, therefore, recognize commu-
nalities in executive organization of these systems across
mammalian species. Granted the foregoing, the perspec-
tive is advocated that human introspection can be a useful
source of information concerning the underlying nature
of the systems. In other words, with some luck, our
conscious mind may be able to see the dynamics of our
subcortical heritage clearly. The risks attendant upon
admitting such sources of information into the scientific
endeavor are great, but failure to approach the brain with
realistic experiential concepts is equally problematic.
The stance taken herein leads inexorably toward a the-
oretically based analysis of brain function, and as long as it
remains linked to identifiable neural systems and objec-
tive behaviors, the enterprise should prove as scien-
tifically feasible as particle physics - comfortless as that
may be.
Of course, there are alternative ways of viewing the
organization of emotional processes in the brain, and the
usefulness of any approach will ultimately rest upon the
degree to which it promotes an empirical understanding
of the emotions and their role in behavior. At the most
general level - a level at which too much biological
research on the topic has become stranded - emotion
may be seen to be a generalized arousal state from which
individual emotions evolve via a process of social learn-
ing. As Redmond and Hunag (1979, p. 2159) have put it:
"In normal individuals, the emotions associated with
increasing activation, therefore, might be called atten-
tion, carefulness, interest, surprise, alerting, astonish-
ment, alarm, anxiety, fear, panic, or terror. Which was
identified would depend on a number of situational,
semantic, intensity, or personality factors." Although the
role of general arousal is of clear importance in the genesis
of emotions (Figure 4), the fact that an array of emotional
behaviors, such as those discussed herein, can be elicited
by hypothalamic stimulation, at various brain sites but in
similar environmental circumstances, suggests that there
is substantial intrinsic articulation in the emotional cir-
cuits of the brain. As an alternative, one might wish to
seek separate and distinct hard-wired neural representa-
tions for each and every emotional nuance that exists, and
surely single-unit studies could bear out those concep-
tualizations. However, those approaches may also be
misguided if additional evidence indicates an overriding
coherent operation of brain systems in which these indi-
vidual cells are located. At this stage, it seems most
practical to steer a middle course between these ex-
tremes. This middle way means recognizing that certain
classes of distinct behaviors always go together in the
normal organism, as though basic control of each rested in
a common circuit with its genetically determined activity
modulated by prior experiences, momentary perceptual
inputs, and prevailing homeostatic conditions, thereby
affording the possibility of innumerable specific behav-
ioral expressions of activity in the circuit.
In summary, then, the main thesis of this paper is that
the natural order of brain emotive systems is to be found
among a limited number of translimbic command circuits
that act to bring a variety of specific behavioral acts and
concurrent physiological changes under common execu-
tive influence. Of course, the existence of emotional
states in animals must remain an assumption based on the
BEHAVIORAL AND BRAIN SCIENCES (1982)3 421
Commentary/Panksepp: Psychobiology of emotions
recognition of such processes in our human experience
and on the probability that similar mechanisms exist in
the limbic systems of related species. Without such an
assumption, one is left with few scientific options but to
tally the movements of animals and their constitutent
parts in the hope that predictive algorithms will emerge
from accumulated facts. However, if one makes that
assumption (and, to my knowledge, no data exist to
compromise its credibility), then subjective human expe-
rience should be a useful guide for categorizing and
analyzing the emotive circuitries of the brain. Such a
source of knowledge can not only provide a list of primi-
tive emotive states that the brain elaborates (e.g., Table
1), but it can also help establish some dynamic properties
of such circuits. A substantial amount of past work in this
area has been implicitly guided by conceptions such as
these. Yet, because of methodological and conceptual
limitations imposed on the study of behavior by an era
predating the flowering of neuroscience, the implications
of such approaches for understanding how the brain
organizes behavior (and mind) remain largely unex-
plored.
ACKNOWLEDGMENTS
I appreciate comments by Robert Conner, Michael Doherty,
John Jalowiec, and Kenneth Pargament on an earlier draft of this
paper, and I thank Ryan Tweney for introducing me to the
original versions of Cogan (1802) and Willis (1683).
Open Peer Commentary
Commentaries submitted by the qualified professional readership of reason to believe that stimulation of specific neuronal circuits
this journal will be considered for publication in a later issue as
Continuing Commentary on this article. Integrative overviews and
syntheses are especially encouraged.
Assessing internal affairs
Hymie Anisman and Robert M. Zacharko
Department of Psychology, Carleton University, Ottawa, Ontario, Canada stimulation. When rats are given long pulse stimulation (30 sec.)
K1S 5B6
In his provocative paper, Panksepp has provided an exciting
formulation of a difficult issue. Yet it seems that there may be too doses of amphetamine will not show a decline of feeding rates
few equations to solve for too many unknown variables. Even if
one were to assume that emotions, being derivedfromprimitive
systems, are fundamentally similar in human and infrahuman
species, it is questionable whether we can identify these emo-
tions in infrahumans with the precision that would be necessary
to assess unequivocally the propositions advanced by Panksepp.
Particular behavioral events are taken by Panksepp as indices of digms, the consummatory acts came to be directed toward the
certain emotions; freezing and flight, for example, are assumed
to represent fear; attack and fighting reflect rage; distress vocal- not necessary to hypothesize that these behaviors arose because
ization is indicative of panic; forward locomotion and sniffing
signify expectation. Such operational definitions are fine if one
wishes either to distinguish between or to categorize a series of stimulus-bound feeding is decreased.
behaviors, but it would be inappropriate to assume that these
behaviors actually represent the emotions as humans might
interpret them.
422 BEHAVIORAL AND BRAIN SCIENCES (1982)3
Following application of an aversive stimulus, such as
footshock, some strains or lines of mice exhibit intense immo-
bility, whereas other strains display high levels of locomotor
activity (Anisman 1975; Anisman, Grimmer, Irwin, Remington,
& Sklar 1979). Likewise, some strains of mice tend to vocalize,
while others do not (Wahlsten 1972). Does this imply that a
given stressor provokes different levels of emotions or different
emotions entirely across strains, or that the stimuli are not
equally aversive across strains, or that the same emotional
response is generated in each strain, but the behavioral corre-
lates (the defensive repertoire) of these strains differ? As a
second example, rats exposed to footshock display a transient
period of response excitation followed by freezing. Pinel and his
associates (Pinel &Treit 1978; Treit, Pinel, & Fibiger 1981) have
reported that if shock is delivered through a probe, rats will
approach and sniff the probe (expectancy?), back away from it
(fear?), and then bury the probe (rage?) with material that might
be at hand (at paw). The behavior of the organism varies
temporally, and as a function of the stimuli present in the
environment. One could hardly assume, however, that the
behavioral changes were indicative of rapid alterations of emo-
tional state.
The behavioral topography of an animal might be dictated by
antecedent stimulus events and by the opportunities to respond
to environmental stimuli. For instance, cues that signal aversive
events may come to produce an "expectancy" of the primary
aversive stimulus (Bolles 1970) or the conditioning of a fear
response (Rescorla & Solomon 1967). The cognitive or emo-
tional state may lead to immobility or flight, depending on
several situational variables (Bolles 1971). If other defensive
behaviors are unavailable or ineffective in terminating the
aversive stimulation, aggressive behavior may result, particu-
larly if the behavior can be directed at a significant stimulus. The
aggressive act, in this instance, may simply represent a compo-
nent of the animal's defensive repertoire (like freezing or flight),
and it need not be assumed that it was precipitated by rage.
In considering the appetitively motivated behaviors, Pank-
sepp suggests that responding for self-stimulation is representa-
tive of the expectancy/foraging emotion. However, there is
such as those subserving "stimulus-bound" appetitive behav-
iors provide the arousal (activation) necessary to engage in
appetitive acts given the presence of relevant environmental
cues (Valenstein, Cox, & Kakolewski 1970). Further, to the
same point, Panksepp suggests that agitated self-stimulation
(vigorous biting of the manipulandum) reflects the intermin-
gling of the expectancy and rage systems. An alternative and
possibly more parsimonious account of the agitated self-stimula-
tion is that licking and biting the manipulandum is representa-
tive of part of the consummatory sequence induced by brain
with food available, they typically engage in consummatory acts
during the period of stimulation. Animals treated with high
induced by the stimulation, nor will they exhibit a decline of
food-related behaviors (e.g., gnawing at the grid floor, lever, or
foodcup). However, such animals do not consume the available
food (Wishart & Walls 1974). Unpublished data collected by
Wishart and Zacharko have revealed that, as pulse-duration was
shortened in either programmed or self-administered para-
lever (chewing or licking), rather than the available food. It is
of frustration; rather, they are indicative of the consummatory
sequence and are maximally evident as the time available for
Attributing particular emotional changes to animals may be as
misleading as attempts to attribute cognitive changes to animals
exposed to environmental events. When animals are exposed to

unpredictable aversive events over which control is not possi-
ble, the animals, it has been frequently observed, adopt a
passive response style characterized by repeated failures to
avoid or escape the aversive stimulus. Some theorists have
assumed that the passivity is due to "learned helplessness"
(Maier & Seligman 1976), whereas others have attributed the
behavioral changes to response styles engendered by alterations
in central neurochemical lability (Anisman, Kokkinidis, & Sklar
1981; Weiss, Glazer, Pohorecky, Bailey, & Schneider 1979).
Inasmuch as cognitions in animals cannot be directly measured
and neurochemical events can, the former approach appears to
us to be a sterile one that does not lend itself to empirical
validation in infrahuman animals. In the case of the analysis
presented by Panksepp, it may have been preferable simply to
document behavioral changes occurring in various experimental
settings and to delineate the physiological and neurochemical
mechanisms subserving these behaviors. After all, the brain
amine changes that may be related to coping processes or
necessary to prepare the organism for further insults need not
be interpreted in terms of emotional states. The use of labels
such as fear, panic, or rage may, in the long run, be coun-
terproductive. To be sure, Panksepp has repeatedly indicated
the difficulties involved in attempting to infer emotional
changes in animals on the basis of some behavioral events.
Unfortunately, simply recognizing the difficulty of the problem
is not sufficient to eliminate it. Despite our misgivings, we
hasten to add that the polythetic approach adopted by Panksepp
is an exciting one and removes the problem from the narrow
confines within which it is often considered.
Emotions - inferences from hypothetical
hypothalamic circuits?
Magda B. Arnold
5863 Montford Rd., Mobile, Ala. 36608
According to Panksepp, "while major concepts concerning the
nature of emotions arise from human introspection, most knowl-
edge concerning the mechanisms underlying emotionality
arises from animal brain research." Hence he suggests that more
knowledge about emotion might be derived from introspection
combined with the results of brain research than from either
approach alone. I am entirely in accord with this premise, which
underlies the author's theory.
However, there are safeguards that must be observed if these
two approaches are to be valid. First, introspective evidence
must be derived from general human experience. That is, the
emotions proposed in a theory should be generally accepted.
Also, a taxonomy of emotions should include the most important
simple emotions. Finally, the emotion circuits inferred from
brain research should have demonstrable connections with
relevant sensory and motor functions. We know that we must
perceive a situation before we can react to it emotionally and
express these emotions in behavior. Ideally, a theory based on
brain research would indicate how the neural structures and
connections involved in emotion are activated by sensory expe-
rience and in turn activate motor functions.
Panksepp's proposal to "outline a testable theory of how
emotional processes may be organized in the central nervous
system of mammals" seems to promise such an integrated
system. He proposes "a taxonomy of emotions that is supported
by existing neurobehavioral data" and promises "to provide an
approximate neural scheme of how emotions may be organized
in the brain."
However, the execution of this laudable proposal leaves much
to be desired. First of all, Panksepp's taxonomy of emotions is
derived from rather than supported by the results of brain
research. He stipulates that the circuits inferred from the results
Commentary IP&nVsepp: Psychobiology of emotions
of brain stimulation must satisfy several criteria before they can
be identified as emotion circuits. They must be evolutionary
acquisitions; they must be involved in adaptive behavior; they
must affect the sensitivity of relevant sensory systems and be
active beyond the stimulus duration. Since some affects, like
surprise, disgust, pleasure and displeasure, do not fulfill these
criteria, Panksepp simply ignores them.
Panksepp proposes four emotion-mediating circuits, located
mainly in the hypothalamus: expectancy, rage, fear, and panic
command circuits, which are activated by sensory information
from the environment and internal states. He insists that the
emotion of expectancy is involved in anticipatory appetitive
behavior; its circuit is "designed to energize appetitive foraging
behaviors." And he suggests that the sudden violent leaping
seen sometimes in rats after brain stimulation is a form of panic.
Its "diminutive form" is sorrow, inferred from the distress call
evoked by brain stimulation. Such distress calls can be evoked in
various species but not in rats; and only in rats does brain
stimulation evoke the explosive behavior Panksepp calls panic.
There is no evidence for a connection between separation
distress (sorrow) and panic. Panksepp infers rage from attack
evoked by brain stimulation; and fear, from blind running.
These two emotions are the ones most easily recognized in
subjective experience. According to Panksepp's Figure 1, these
four emotions represent extremes. While rage is an extreme
form of anger, and fear may be an extreme form of alarm or
foreboding, as he says, expectancy is hardly an extreme form of
hope or desire, nor is panic an extreme form of sorrow or
separation distress.
Panksepp not only fails to use subjective experience in work-
ing out his taxonomy of emotions, he flatly disregards such
experience if it does not fit the behavior patterns evoked by
brain stimulation. Since he includes sorrow, he should logically
also include joy in his system, yet he never mentions it. He also
has no room for love or affection, although this emotion is
obviously involved in social cohesion. For Panksepp, the panic
circuit functions "to sustain social cohesion," yet distress vocal-
ization (and panic?) surely is a sign of the breakdown of social
cohesion, while love is the bond that maintains it.
Instead of providing a neural scheme of how emotions are
organized in the brain, Panksepp merely mentions the generally
accepted connections from midbrain through the hypothalamus
and thalamus to the basal ganglia and higher limbic area and
suggests that there are also connections with other behavior
control circuits; and that various levels of the neuraxis have
access to emotion circuits. After his explanation we are no closer
to understanding how social separation, for instance, activates
the panic circuit. The infant animal must realize its mother's
absence before making distress calls - but just how does that
realization activate the hypothalamic panic circuit? What is
needed is a viable circuit from sensory areas through the hy-
pothalamus to motor areas to distinguish emotion from motor
circuits. The observed behavior patterns are not themselves
emotions, and the circuits that mediate them are not necessarily
emotion circuits, even if they satisfy the criteria Panksepp has
set up.
Panksepp deplores the failure of earlier theorists to use the
results of brain research in presenting their taxonomies of
emotions. Apparently, he has never examined Arnold's
(1960a;b) theory, which uses subjective experience to identify
emotions, but then outlines specific brain circuits and connec-
tions involved in the sequence from perception to emotion and
action. The theory may need to be corrected or supplemented,
but it does use the results of modern brain research and is the
most extensive attempt to reconcile evidence from brain re-
search with subjective experience. If the theory is inadequate, it
is open to criticism and correction. But to disregard it, and then
deplore the absence of such an attempt, is hardly good scientific
practice.
BEHAVIORAL AND BRAIN SCIENCES (1982)3 423
Commentart//Panksepp: Psychobiology of emotions
Emotions: Hard- or soft-wired?
James R. Averill
Department of Psychology, University of Massachusetts, Amherst, Mass.
01003
Panksepp's argument proceeds on two levels. Thefirstis a thesis
about general strategy for behavioral and brain research, using
emotion as a paradigm; the second is a series of propositions
concerning possible neural circuits that are presumably "hard-
wired" and that mediate four broad classes of emotion (expec-
tancy, rage, fear, and panic). Much could be said about the
argument at both levels, but I will limit my comments to a few
observations on Panksepp's more general thesis and its implica-
tions for the study of emotion. The central proposition of this
thesis is that we should make greater use of our own introspec-
tive awareness as a guide to research and theory. I agree with
this proposition. Computer scientists have profitably used intro-
spective reports in the development of programs (minitheories,
in a sense) that simulate "higher" thought processes. Why cant
neuroscientists do the same when studying the brain? The
analogy between computer and brain sciences is far from exact,
of course. Nevertheless, Panksepp seems to be on firm ground
when he asserts than an unwarranted fear of anthropomorphism
can have - and has had - a stifling effect.
But having granted Panksepp his general thesis, I must
question his application of it. Panksepp cites favorably the great
seventeenth century anatomist, Willis, who used introspective
analyses to guide his neurophysiological investigations of the
emotions. Panksepp does not, however, describe the conclusion
reached by Willis, namely, that emotional processes are lo-
calized within the cerebellum. Willis based this conclusion on
"Analogy and Frequent Ratiocination" followed by "Anatomical
investigation" (1664/1965, p. 111). More specifically, Willis
reasoned that since emotions are often involuntary and irra-
tional, they should be located in a part of the brain that is only
narrowly connected to the cerebrum and that is relatively
similar in animals and humans. His anatomical investigations
suggested that the cerebellum meets these criteria.
The line of reasoning pursued by Willis is both ancient and
modern. A similar view (but with a different localization) can be
found in Plato. (For a historical review of psychophysiological
theories of emotion, see Averill 1974.) Panksepp's article illus-
trates a modern variation on the same theme. That is, introspec-
tion apparently suggests to Panksepp that the emotions are
largely noncognitive and involuntary; he therefore concludes
that the emotions must be mediated by "command circuits" that
are hard-wired into the more primitive parts of the brain, such
as the hypothalamus and the limbic system.
But is this the direction a careful introspective analysis of
emotional phenomena would lead us? Are the emotions as
immutable, as divorced from higher thought processes, and as
primitive as they are often portrayed? Or is this merely a part of
their legitimation, that is, the manner in which they are ra-
tionalized in society (because of the types of functions they tend
to serve)? Obviously, there is not the space here to pursue the
kind of analysis that would be required to answer these ques-
tions. At the risk of sounding dogmatic, therefore, I will simply
state what I believe such an analysis would reveal, namely, that
the emotions are exquisitely variable, both across individuals
and across cultures; that they are fundamentally dependent
upon cognitive processes; and that they are typically quite
purposeful. (For detailed arguments in support of these claims,
see Averill 1980a; 1980b; Solomon 1976.)
Panksepp would, of course, disagree with these last conten-
tions. Where does the source of the disagreement lie? Let us
consider for a moment the case of anger, one of the emotional
"command systems" (rage) postulated by Panksepp. A typical
episode of anger can be manifested in any of a great variety of
ways, depending upon the person and situation; it seldom leads
to physical aggression; it is more often than not constructively
424 BEHAVIORAL AND BRAIN SCIENCES (1982)3
motivated (i.e., intended to help rather than harm the target);
and it is based on the appraisal of events as right or wrong,
justified or unjustified (Averill 1979). Clearly, not all aggressive
acts can be categorized as angry (cf. envy, jealousy, and instru-
mental aggression, to mention a few possibilities). Panksepp,
however, tends to gloss over such differences and indis-
criminately lumps into a single category such diverse states as
"rage," "anger," "wildness," and "affective attack." Moreover,
he assumes that there is a homologous class of behaviors in most,
if not all, mammalian species. Panksepp maybe right, although
I doubt it. The only point I wish to make is that his conclusions
are not based on a careful conceptual or introspective analysis of
human emotions, that is, on the kind of analysis that he is
advocating.
Panksepp might reply that I am quibbling about words. And
indeed I am. Our experience and behavior is intimately related
to the way we categorize the world, as reflected in our language;
and I assume that this fact has important implications for behav-
ioral and brain research.
Panksepp accounts for the "impressive subtleties of human
feeling" in terms of the interblending of the four basic emotions.
Thus, jealousy "may arise from some admixture of panic, rage,
and expectancy . . . and so on with other affective alchemy."
The alchemic metaphor is more appropriate than Panksepp
perhaps realizes. He might have pursued it further. For one
thing, alchemists recognized that, if a baser metal were to be
transformed into a more noble one, the former would cease to
exist in any meaningful sense. In Panksepp's affective alchemy,
the basic emotions are evidently preserved in the admixture.
But preserved or not, the process by which the transformation
occurs remains mysterious.
Panksepp's failure adequately to follow his own program is
also illustrated by his assertion that "it seems to be universal
human experience that positive expectations and anger are
psychologically incompatible states." From this he infers that
the corresponding neural circuits should exhibit reciprocal inhi-
bition. But it is simply not the case that positive expectations are
universally considered incompatible with anger. Aristotle, re-
flecting an even more common view, maintained that anger
"must always be attended by a certain pleasure - that which
arises from the expectation of revenge" ("Rhetoric," 13781').
What kind of neurophysiological theory of emotion might
emerge if we took Panksepp's general thesis seriously and
systematically analyzed our emotional concepts and experi-
ences? A careful analysis would lead, I believe, to a theory that
emphasized the plasticity of neural circuits (as opposed to the
hard-wiring proposed by Panksepp), even in the case of "basic
emotions"; that stressed the hierarchical and heterarchical in-
teractions among circuits (which Panksepp discusses only super-
ficially); and that laid greater importance on socialization than on
natural selection as a systematizing influence (at least as far as
human emotions are concerned). In other words, it would
suggest an approach to the localization of function more along
the lines adumbrated by Luria (1973) than along the lines
proposed by Panksepp.
Specific human emotions are psychobiologic
entities: Psychobiologic coherence between
emotion and its dynamic expression
Manfred Clynes
Music Research Center, N.S.W. State Conservatorium of Music, Sydney
2000, Australia
A plan to study specific emotions as psychobiologic entities is
timely and appropriate, particularly if one attempts, where
possible, to supplement exploration of brain function with
observation of human subjective experience. Specific subjective

entities have long been implicated in the scientific study of
animal sensory physiology: No one doubts the existence of
stereovision, of the perceived sensations of tone, of smell, and so
on, in various animals. In the spectrum of specific emotions,
however, specific sensing or "command" structures are not
obviously at the periphery of the nervous system, but appear to
be discoverable elsewhere in the brain. Categories of subjective
experience here too can help, and can even be essential to
systematic scientific categorization and investigation: As we
progress in the study of brain function, the subjective in-
creasingly becomes objective.
Though the most salient emotions are named in practically all
human languages, and function in dreams, the neuroscientific
study of their nature has not received the attention warranted by
their importance in everyday life, in music, drama, and art
(Langer 1973; Piechowski 1981). Emphasis has been on general
arousal, mediated by catecholamines, on "drives," on "domi-
nance versus submission." [See also Bernstein: "Dominance:
The Baby and the Bathwater" BBS 4 (3) 1981 and Vandervolf &
Robinson: "Reticulo-Cortical Activity and Behavior" BBS 4 (3)
1981.]
One regrets that in the theoretical formulations of Panksepp's
target article "negative" emotions predominate, reflecting the
prevalent orientation of the literature. Three out of four of his
major "distinct" specific categories are rage, fear, and panic
("freezing" is described as occurring both in fear and in panic)!
The fourth, expectancy, is left to shoulder whatever worthwhile
emotional experiences life may have to offer, to animals or to
man. No mention, let alone study, is made of love, of wonder,
courage, for example; and joy is encountered by Panksepp's
system only as a result of fulfilled expectation. This probably
reflects the paucity of readily available animal (and human)
experimental evidence about distinct "positive" emotions
rather than a lack of discernment. But if we indeed share
genetically "hard-wired" emotion programs with mammals, as
Panksepp, probably correctly, suggests, we would hardly be
likely to share only those of "negative" emotions with them.
An important omission in the theoretical formulation is the
inherent communicative function of emotion: the contagion
across individuals. The study of a specific emotion-entity can-
not ignore the auto- and cross-communicative and contagious
aspects of its expression; there is evidence that these are biolog-
ically integral parts of the entity of each specific emotion (e.g.,
sexual mating rituals and dances as genetic programs in many
animals, as well as later-evolved emotion-entities also, as yawn-
ing or laughter). [See also Eibl-Eibesfeldt: "Human Ethology"
BBS 2(1) 1979.]
It is here that there is psychobiologic evidence concerning
human emotions that substantially overlaps and strengthens
Panksepp's position. The characteristic expression and the state
of a specific emotion inherently interact psychobiologically in
ways that can be experimentally clarified.
In our work of over a decade with the expressive generation of
specific emotions through touch and sound, we found, broadly,
that the "purity" of the expressive form strongly affects its ability
to both auto- and cross-generate emotion; it appears that there
are specific dynamic forms of expression to each "basic" emotion
in humans. We have experimentally determined the durations
and dynamic shapes of these forms - called essentic forms
(Clynes 1969; 1973; 1975). We find that there is a biologically
given coherence between the specific dynamic form and the
emotion that it can express. These properties are made evident
also in music and account for the "living quality" of good musical
performances. (Moreover, "mixed" emotions appear to be ex-
pressed not by an algebraic sum but by a temporal telescoping of
separate portions of the component forms into single expressive
forms.)
One aspect of this evidence concerning distinct emotions is
the dynamics of selective satiation that have been observed
when specific emotions are generated through iterative ex-
Commentary/Panksepp: Psychobiology of emotions
pression (Clynes 1973; 1980). It is found that subjects may be
sated with respect to one specific emotion but quite fresh with
respect to another specific one, until sated with respect to the
next one, but ready for a third, and so on through a series of what
appear to be by this criterion basic emotions. (This does not
occur with "mixed' emotions.) This selective satiety was consid-
ered likely to be related to specific chemical receptor sites.
Since then, endorphins and encephalins have been discovered;
however, their possible role in mediating emotions is not ade-
quately treated by Panksepp - he devotes less than a paragraph
to this.
Like Panksepp, we too have found it desirable to discard the
single-dimensional continuum of pleasure/displeasure as a de-
scriptor of specific emotion-entities; in Panksepp's article it
reappears, however, in the different guise of "positive" and
"negative" emotions, a nomenclature that is not further ex-
plained. (A "negative " emotion such as anger, for example, can
give pleasure under some circumstances.) We have named
"negative' those emotions that have a horizontal component of
expressive pressure away from the body.
In humans, specific emotions can be experienced in two
distinct ways or "modes': (1) the "animalistic" Dionysian way
and (2) "'considered' as qualities," the Apollonian way, as in
great music and art. The latter (probably using "command"
routes other than those involving only the limbic system) allows
us, paradoxically, to enjoy a piece of music expressing grief- the
sadder, the better! It is not known if even the highest mammals
are capable of any such degree of "imaginative" use of the
qualities of emotion and their communication. A third mode in
which the expression of specific emotion can be produced in
humans is mimicry. Panksepp's "elaboration" seems an insuffi-
cient concept for the study of the implications of such modes, in
animals or man.
Recent work has indicated that in humans the dynamic forms
of expression of specific emotions appear to be independent of
sensory mode. Touch and sound are found to share the same
dynamic form for specific emotions. Transformations were
found which turn a tactile expression of specific emotion into an
acoustic expression of the same emotion: The transform con-
serves the dynamic form, changing pressure form into frequen-
cy contour with appropriate specific scaling (Clynes & Nettheim
1982; Clynes & Walker 1980; Clynes, Walker, & Nettheim
1981). The sounds so transformed from touch experiences of
seven specific emotions were tested on over three hundred
subjects in the United States and Australia, including a group of
40 central Australian aborigines. Recognition was similar in all
three groups (p < .0001).
It would appear that important parts of the "hard-wired"
programs for the dynamic expression of specific emotions are
independent of sensory mode for both production and recogni-
tion of emotion expression and thus also for communicative
contagion of emotion generation. This also readily lends itself to
study through animal experiments.
In view of these and otherfindings,Panksepp's suggestion of a
cross-fertilization between animal and human studies of taxon-
omy and properties of specific emotion programs should be
productive.
Animal and human emotionality
Jos6 M. R. Delgado
Departamento de Investigacidn, Centro "Ramdn y Cajal," Madrid 34,
Spain
One valuable aspect of Panksepp's paper is the attempt to
identify specific emotions with brain circuitry. This possibility is
testable; and even if his proposed "general psychobiological
theory of emotions" may be debatable, experimental facts are
established, and his approach "can facilitate construction of
BEHAVIORAL AND BRAIN SCIENCES (1982)3 425
Commentan//Panksepp: Psychobiology of emotions
more viable models and theories." In reality, this is the orienta-
tion of many investigators, and when emotional manifestations
are elicited by electrical stimulation of certain brain structures,
it is accepted that some anatomical-functional correlation
exists.
Another interesting aspect of Panksepp's article is the blend-
ing of the two sources of knowledge about emotions, namely
objective manifestations in animals and subjective experience in
humans. It is asserted that "basic emotions may have obligatory
internal dynamics, which humans share with other mammals, "
and also that "control mechanisms remain similar in humans and
'lower' animals." The problem with this thesis, however, is that
the most important human emotions are based on ethics, esthet-
ics, and cultural learning that have no parallels in animal
experience, and also that the most important human expression
is related to language, which, again, cannot be equated with
primitive animal communication.
It is true that the limbic system is a shared mammalian
heritage, but we also share the motor system, reticular forma-
tion, cerebellum, and many other structures. There are sen-
sorimotor neuronal similarities among rats, monkeys, and hu-
mans, but the sensitivity, knowledge, and skills necessary for
outer space travel are not shared with animals. The finding that
ventromedial hypothalamic lesions increase appetite and rage in
both animals and humans means that the hypothalamus does
play some role in some of these behavioral manifestations which
are not necessarily equivalent: the ingestion of food pellets may
be an instinctive act which cannot be compared to the enjoy-
ment of a banquet. Mouse-killing behavior in the rat has very
little in common with organized crime. One of the fallacies to
come out of conferences on animal aggression is the frequent
implication that causality and cerebral mechanisms are similar
in animal fighting and in the strategies of modern warfare.
Area 4 cortical lesions produce contralateral motor paralysis
in different species, but the anatomical, functional, and motor
organization are probably different. Even within the same
species, man, it is doubtful that a farmer and a guitar player have
equivalent motor representation of their hands.
We should not generalize animal experimentation to human
behavior unduly, and we should analyze similarities and dif-
ferences of the phenomenon under consideration. At the core of
this problem is the understanding of the ontogeny of emotions:
Panksepp states, "Emotions seem to arise ultimately from hard-
wired neural circuits in the visceral-limbic brain," and that
"these circuits developed early in mammalian brain evolution."
Certainly there is genetic determination of the main hard-wired
neural circuits, but human beings are born with very immature
brains, and software information received through sensory in-
puts is decisive for neuronal sprouting and for the establishment
of synaptic connections, neurochemical structuring, and other
aspects essential to the final hard-wiring. Also, the frame of
reference necessary for emotional reactivity and many aspects of
emotional expression are learned and not inborn. Motor path-
ways develop early in mammalian brain evolution, but their
anatomical and functional development requires individual
learning and training. Harlow's motherless monkeys were born
with normal brains, but in the absence of maternal care and
suitable sensory inputs, emotional development was disrupted.
Genetic hard-wiring was not enough.
In some of the available literature there is a tendency to
generalize experimental findings among rats and cats to
monkeys and man. In many aspects of nerve depolarization or
muscle contraction this is acceptable, but in many other man-
ifestations, including emotional reactivity and expression, this is
not valid. Rats do not have a sense of humor, and man does not
usually chase mice.
The conclusion I am reaching is that, in agreement with
Panksepp, it is very useful to investigate mechanisms shared by
animals and humans, ranging from unitary neuronal activity to
social behavior, but that at the same time we must investigate
426 BEHAVIORAL AND BRAIN SCIENCES (1982)3
and clarify the many aspects not shared, which are the most
important emotional characteristics of man. Primitive in-
stinctive emotions should be differentiated from cultural, intel-
ligent, cultivated emotional responsiveness. This separation is
essential for the understanding and the education of human
beings and for finding solutions to present emotional conflicts. It
is also essential for any psychobiological theory of emotions.
Another concept on which I would like to comment concerns
the existence of "executive (command) circuits that concur-
rently activate related behavior patterns." This idea is in agree-
ment with our thesis that "brain stimulation triggers physiologi-
cal mechanisms" and that we may consider an "electrical
activation of the will" (Delgado 1969, chaps. 17 and 18). This
idea is also in agreement with our theory of "fragmental repre-
sentation of behavior" (Delgado 1964), and with our proposed
organization of behavioral responses including three functional
sets, namely, the starter, organizer, and performer (Delgado
1969). I do not think, however, that the initial command post for
emotions must necessarily be located in the hypothalamus (and
perhaps not even in the limbic system). As an alternative
hypothesis we have proposed that specific sets of structures are
located in different areas of the brain, forming a functional
constellation with reciprocal facilitatory, and inhibitory rela-
tions (Delgado 1979). The concept of algebraic summation of
dynamic influences triggering a final response is preferable to
the idea of a single specific center. Panksepp indicates in his
article that the command systems may run "from mesen-
cephalon, through the reticular fields of the hypothalamus and,
perhaps, thalamus to basal ganglia and higher limbic areas,"
with other ascending and descending controls. This is the
concept of a functional constellation, and perhaps the difference
of opinion lies only in giving more or less emphasis to the
command value of specific parts of the circuit.
Experimental evidence indicates that, in cats and monkeys,
aggressive behavior can be elicited by electrical stimulation of
the reticular formation, tectum, thalamus, hypothalamus,
amygdala, and other structures; and it can be inhibited by
stimulation of the caudate nucleus, pallidum, and other areas of
the brain. All these structures are functionally interconnected,
and in our studies, we have been able to give to each structure a
mathematical coefficient expressing the value of its activation
with respect to a specific behavioral response (Delgado 1980).
This experimental approach is more neutral and factual than
the attempt to relate brain physiology to ill-defined psychologi-
cal classifications. This is one of the most controversial aspects of
Panksepp's article: Why classify emotions in terms of only four
categories, leaving out pain, pleasure, sex, hunger, and many
others? No convincing explanation is given, nor is this classifica-
tion in agreement with experimental facts. Distress vocaliza-
tions may be associated with fear and rage; they are not ex-
clusively expressions of panic. Our present investigations
(Martin-Ramirez, Blanco, Alonso & Delgado, 1982) show that,
in the cat, hissing can be evoked by stimulation of many points
located from the reticular formation up to the hypothalamus,
with reciprocal potentiating and inhibiting influences; the effect
is in many cases an isolated, out-of-context behavioral module,
and does not involve attacks against cats or rats.
In monkey colonies, electrical stimulation of the central gray
in the same animal with the same parameters evoked different
effects depending on the social status of the animal: aggression
when it was dominant, and submissive behavior, including
grimacing, when its rank was low. [See also Bernstein: "Domi-
nance" BBS 4(3) 1981.] Processing of sensory information
proved essential for the response, and the idea of hard-wiring
cannot be accepted in this case. Aggressive behavior will be very
different depending on whether the subject is a black-belt judo
champion or a sedentary clerk. Learned motor skills and ide-
okinetic formulas are very different, being stored as motor
fragments to be used when needed, for different purposes. It is
very doubtful that motor displays are specific to fear, rage,

panic, and other emotions; it is more probable that, as postu-
lated in our theory of fragmental representation of behavior,
there is an emotionally bound starter, which activates a differ-
ent set: the organizer responsible for the choice of the complex
motor response, which in turn fires another set: the performer,
which puts in action previously learned formulas of motor
response.
One of the merits of Panksepp's paper is that it provides an
excellent review of the subject and a wealth of ideas, which
should trigger emotionality in the readers, and, one hopes
experimental actions to support, refute, and clarify the impor-
tant material that he lucidly presents.
Relating experience to the brain
Joseph de Rivera
Department of Psychology, Clark University, Worcester, Mass. 01610
I enjoyed reading this article. One might take issue with a
number of points: Studies on the recovery of brain function
suggest that even subcortical structures must be less "hard-
wired" than is usually imagined (Fass, Jordan, Rubman, Seibel,
& Stein 1975; Mclntyre and Stein 1973), and the suggestion of
four primary emotional processes rather neglects both
Tomkins's (1962) and Plutchik's (1980) attempts to specify basic
emotional behavior patterns. However, I like Panksepp's treat-
ment of emotions as separate systems with specific structures,
functions, and properties, and I appreciate his audacity in
attempting to combine neurobehavioral and introspective data
to come up with a "psychobiological" theory. As a phe-
nomenological psychologist, I want to respond to this openness
toward the data of human experience. What if we really take
human experience seriously? What is our conscious experience
of emotions? Does this experience suggest any particular organi-
zation to the emotions? Could such an organization be related to
the model proposed by Panksepp? I want to comment briefly on
each of these four questions.
If we take consciousness seriously, we must conclude that it is
inaccurate to say that "eyes see" or "the brain thinks"; rather,
we use our eyes to see with and our brain to think with (cf. Straus
1963). Similarly, from the viewpoint of conscious experience,
brains don't have emotions, persons do. Emotions are not in the
brain, they are how we are in the world. They are not states
produced by neuronal circuits to which we somehow have
introspective access. Rather, they are transformations of how we
are situated in the world. Thus, strictly speaking, we should not
ask how emotions are organized in the brain, but how emotional
organization is related to the organization of neural activity in
the brain.
Our experience of the emotions concurs with Panksepp's
theory insofar as he regards emotions as distinct behavioral
Table 2 (de Rivera). Possible integration of neural and emotional organizations
Panksepp's de Rivera's eight basic states in
subcortical systems dyadic organization of emotions
Emotion transforms other -
"Panic" love (give to other)
"Fear" —fear (pull back from other)-
"Expectancy" desire (go out and get)
"Rage" '—anger (remove other)
Commentary/Panksepp: Psychobiology of emotions
Table 1 (de Rivera). Basic states in a dyadic system of
emotions
The emotion
Transforms
Transforms the other the self
for the self (desire) (elation)
Into something good
in its own right (love) (security)
for the self (fear) (anxiety)
Into something bad
in its own right (anger) (depression)
control or "command" systems that are related to specific
situational events such as the presence of an incentive or a social
loss. However, we do not experience emotions as responses to
environmental stimuli. Rather, emotions transform our rela-
tionship with the world. Thus, anger is not a set of aggressive
responses to frustration, but a transformation of our situation so
that we perceive a challenge to the way things ought to be (cf. de
Rivera 1981). In this regard, the brain model proposed by
Pribram and Melges (1969) is truer to experience. Further,
there is no experiental evidence for a small set of basic emotions.
On the contrary, we have names in English for several hundred
emotions, and close examination of these different emotions
reveals that they are not blends of more primitive basic emo-
tions. Each has unique structures (cf. Lindsay-Hartz's, 1981,
description of the unique properties of joy, elation, and glad-
ness). Thus, rather than a few basic elemental systems, an
investigation of our experience of emotions suggests hundreds
of discrete structures, each involving a unique way of perceiving
the situation, a specific transformation of the way our body
relates to the world, unique "command" instructions about how
to behave, and a particular way of functioning in our lives.
However, our experience does suggest that these hundreds of
discrete emotions are organized in a particular way. Rather than
there being a few relatively unrelated basic emotional systems
that tell the individual organism how to respond to its environ-
ment, our emotions appear to come in mirror-image structures
that reflect the different possible states of one dyadic system
that relates the organism to other organisms, or the self to the
other. At the most rudimentary level, this system may be
described by specifying eight basic transformations of the self
other dyad generated when emotions: (1) transform the other or
the self, (2) into something good or something bad, (3) for the self
or in its own right (cf. de Rivera 1977).
These basic transformations may be illustrated by the emo-
tions listed in Table 1. The hundreds of other emotions reflect
still other permutations of these self/other relations.
• Emotion transforms self
• anxiety (hold on to other)
• security (move out)
• elation (take in)
- depression (remove self)
BEHAVIORAL AND BRAIN SCIENCES (1982)3 427
Commentary/Panksepp: Psychobiology of emotions
Is it possible to relate such an emotional organization, which
is based primarily on conscious experience, to Panksepp's
model of brain organization, which is based primarily on stim-
ulation of the limbic system? I believe so, and some specific
predictions follow from the attempt. We have only to modify his
conceptualization of the four systems so that all of the systems
and not only the "panic" system involve social relationships. We
may then specify two states for each system and view the
separate systems as subsystems of a general system that controls
dyadic relationships. The adaptive significance of such a system
is almost self-evident, at least in mammalian species where the
infant-mother relationship is crucial for species survival.
The proposed system is outlined in Table 2, where the
emotion's "command" is given in parentheses.
In short, if my phenomenological analysis (presented in detail
in de Rivera, 1977) is accurate, and if Panksepp is right in
assuming that "subjective human experience should be a useful
guide for categorizing and analyzing the emotive circuitries of
the brain," then the brain should be organized in a way that
reflects dyadic, mirror-image relationships. Specifically, it
should be possible to delineate at least four other subsystems in
the brain: a "love" system that encourages affiliative behavior, a
"security" system that encourages the exploration of strange
surroundings, a "depression" system that inhibits behavior and
promotes submission by inhibiting aggressiveness, and an "ela-
tion" system that promotes consummatory behavior. I hope
(using my own "expectancy" system) that this commentary will
help guide further exploration of the ways in which brain
organization may be related to our emotions.
Introspection as the Rosetta stone: Millstone
or fifth wheel?
Ronald de Sousa
Department of Philosophy, University of Toronto, Toronto, Ontario, Canada
M4K 2X3
Unlike some other biologically oriented treatments, Panksepp's
definition of emotions involves features crucial to any compre-
hensive theory. These include: (1) coherence and complexity of
associated behavior; (2) an emphasis on the role of emotions in
the determination of differential patterns of salience among
stimuli and facilitated responses (cf. de Sousa 1980); (3) emo-
tional inertia, as in Descartes, (1650, article 74). "The utility of
all the passions consists alone in their fortifying and perpetuat-
ing in the soul thoughts which . . . without that might easily be
effaced from it." I also find it promising, although this point of
contact is not mentioned, that the "panic" system, which Pank-
sepp describes as perhaps "the most controversial" of his pro-
posals, is well supported by the clinical and theoretical work of
John Bowlby (1980) on attachment emotions. [See also Rajecki
et al.: "Toward a General Theory of Infantile Attachment" BBS
1(3) 1978.]
Panksepp advertises a methodological innovation, which is at
first sight attractive: He suggests looking to "judicious intro-
spection" for the '"Rosetta stone' via which the general organi-
zation of the primitive hard-wired emotional systems of the
mammalian brain can be deciphered." But optimism soon
breeds doubt. To me, the phenomenology of emotions evokes
the works of Proust or Freud, whose skepticism about emotional
self-knowledge has been amply confirmed by experimental
evidence (Nisbett & Wilson 1977). In that light, a phrase such as
"to read the animal mind as directly as we can read our own"
rings ironically.
So perhaps one should be glad that Panksepp's practice does
not match his announced method. It turns out, once he gets
down to business, that his taxonomy is based not on conscious
428 BEHAVIORAL AND BRAIN SCIENCES (1982)3
experience but "upon the number of distinct behavioral control
systems that can be activated . . . " (cf. caption to Figure 2).
While the names of these circuits are suggestive of our common-
sense emotional vocabulary, the principles of distinction be-
tween them owe more to general biological and ethological
commonplaces about the nature and kinds of "life-challenging
circumstances" faced by mammals. I see nothing especially
anthropomorphic about those ethological assumptions. No
doubt "emotion as a construct . . . in the study of animal behav-
ior" owes its existence to the analogies of consciousness, but is
this of any more psychological relevance than the analogous
observation about the Newtonian construct of force?
In the detail of its elaboration Panksepp's taxonomy begins to
look rather arbitrary. Why should the desire for social contact
belong to a basic command system different from the one for the
fear (which is not Panksepp's "fear") of its loss? Why should
anxiety be part of "fear" and not of "panic?" How are the
"explosive behaviors" of panic different from manifestations of
rage? (cf. Figure 2). As far as I can tell, answers to these
questions are decided entirely in terms of operational pragmat-
ics. Yet surely here, if anywhere, phenomenology is relevant.
We need criteria of identity and difference among the different
emotions: But all that neurological investigation can identify are
reasonably coherent patterns of behavioral responses. To find
their phenomenological correlates there is no substitute for
direct introspection.
But introspection is no Rosetta stone. The trouble is not only
that it is unreliable, but that what modest access it provides is to
phenomena belonging to a level of analysis different from those
with which they would need "correlating." In general, this
problem of levels is an old philosophical obligate), familiar to
readers of BBS (e.g., Dennet 1978; Fowler & Turvey 1978). In
the present connection it takes two forms.
First, in what sense does Panksepp think the emotions "arise"
out of the command circuits (CCs) he postulates? Consider the
following claims: (a) In certain structural respects (features 1-3
above), CCs parallel the sophisticated emotions, (b) The activa-
tion of these CCs appears to be involved in cases where emo-
tions are present, (c) b explains a, so that we now understand the
mechanism of emotions. Panksepp persuades me of a and b.
Some of his phraseology (Abstract; the early passages on the-
oretical assumptions and aims of the target article) suggests that
he wishes to claim c. But c does not follow. Similarities of
structure at different levels might be due to completely different
mechanisms. I presume, for example, that habituation in an
organism both involves cell habituation in appropriate circuits,
and is functionally analogous to cell habituation. But it certainly
doesn't follow that the mechanism of cell habituation must give
us insight into the mechanism of an organism's habituation.
Second, if we start with a taxonomy of "basic" emotions, we'll
need an explanation of how complex ones arise. Here all that
Panksepp has to offer is "blending." Thus, "jealousy may arise
from some admixture of panic, rage, and expectancy." But it is
not clear how blending could ever add logical structure. And
that - not merely complications of behavioral dispositions - is
what is needed to account for complex emotions. Here, again,
one expects phenomenology to contribute, but in Panksepp's
scheme it has already signed off. What I mean by logical
structure is this: Roughly, an ascription of jealousy involves a
quintary relation of the form: F(t,, t2, p, a,j), whose terms are: a
primary (t2) and a secondary (r2) target (of whom and because of
whom I'm jealous); a prepositional object (p) (what fact about
them makes me jealous); an aim (a) (what I'd like to do about it);
and a formal object (/) (what it is about the putative fact that is
jealous-making?). Other emotions have different polyadicities.
Anger lacks a secondary target; love lacks a propositional object;
perhaps depression lacks an aim. It is not clear what logical
structure the phenomenal correlates of Panksepp's command
circuits have (as far as I can see, the only sort of object to which
they are relevant is their aim, the "well organized behavioral

sequences" to which they give rise); but, whatever the answer,
mere blending among them won't generate different poly-
adicities. So, although Ifindit very plausible to suppose that in
some sense jealousy "may arise from some admixture of panic,
rage, and expectancy," the glue is missing that will make such a
mixture into a single compound.
Emotions are objective events
Elzbieta Fonberg
Department of Neurophysiology, Nencki Institute of Experimental Biology,
02-093 Warsaw, Poland
I would first like to consider Panksepp's theses in terms of my
own general views. I fully agree that "emotions seem to arise
from hard-wired neural circuits of the visceral-limbic brain,'
and that these circuits "developed early in mammalian brain
evolution." I am also convinced that animal investigations can
shed light on the mechanisms of human emotion and that
introspective experience should be taken into account in these
studies. I also agree that transhypothalamic circuits are very
important, although they may not be the only circuits of impor-
tance; the role of other limbic structures, and in particular the
amygdala, may be decisive in the feeling of emotion.
What I do not agree with, however, is Panksepp's classifica-
tion of the emotions in terms of his four arbitrary categories,
which I do not find to be of equal importance.
I shall now attempt to develop my arguments in more detail. I
agree with Panksepp that emotions have their concrete neural
substrates and that they are objective events occurring in
humans as well as in animals. They may be ascribed to definite
brain circuits that are similar (though not identical) in humans
and in higher animals (not only mammals). I have always argued
with those who deny the possibility of investigating emotion and
who ascribe only subjective meaning to this term (Fonberg
1972; 1979a; 1981). I think that such attitudes derive from
idealistic and religious traditions according to which emotions
belong to the soul and not to the body, and are hence inaccessi-
ble to experiment. On the other side, the attitude is reinforced
by the Skinnerian approach: that the only objective way to study
stimulus-response mechanisms is to ignore the underlying
brain substrates. Along with the behaviorists, most physiolo-
gists have objected strongly to using the term "emotion" in
science. (I have always wondered why they have never denied
the objectivity of investigating the pain system while denying
objectivity to all other emotional investigations. Is it not the
same kind of extrapolation to assume that animals see colors and
feel pain like us, as that they feel joy, rage, and fear?
Some years ago, at a conference on emotions and psychoso-
matic illness, I presented my own point of view concerning
emotions (Fonberg 1972): It cannot be denied that emotional
states exist in reality, and that they must therefore depend on
physiological mechanisms; it follows that we should be able to
investigate them by physiological methods. However, the psy-
chological ballast weighs particularly heavily on the definition of
the word "emotional," and various authors use it in different
senses (Fonberg 1972). Panksepp, too, fails to define this term
strictly. He mixes the terms "feeling" and emotion. Feelings are
not a weaker form of emotions, but have a more specific sensory
aspect, whereas the word emotion includes a hedonic valence.
According to my own view (Fonberg 1972), the term "emotion"
comprises (1) negative or positive subjective feelings; (2) exter-
nal emotional symptoms such as motor responses (including
facial expressions), vocalization, and various autonomic changes
specific to different emotional states; and (3) complex forms of
behavior that enable motivated actions to be performed (their
feedback being also of positive or negative emotional character).
I also assume that the appearance of various emotional re-
Commentary/Panksepp: Psychobiology of emotions
sponses depends on the excitation of corresponding brain struc-
tures. These are known as "emotional response centers." The
terms "emotional centers" and "motivational" or "drive cen-
ters" overlap widely and are even used synonymously by some
researchers. Motivation can be defined as the excitation of
certain nervous system structures that evoke definite emotional
states and lead to directed behavior - that is, to the performance
of instrumental acts. It follows from this definition that at the
basis of motivated activity lie the emotional states, which are the
"fuel for the drive. ' On the other hand, the emotional state itself
may not lead to motivated instrumental responses. Certain
conditions are necessary to motivate behavior: (1) a certain level
of excitation of the above mentioned emotional centers; (2) the
presence of the objects that are adequate goals for the particular
emotional state (the nature of these responses in any situation is
determined by accompanying perceptions and by the efficacy of
the ongoing action, which can be called conditioned feedback);
and (3) excitation in the centers that pattern both the innate and
the acquired forms of activity corresponding to specific emo-
tional states (Fonberg 1972).
I am convinced that both the external signs of emotions and
the behavior motivated by emotions are accessible to experi-
ment. Subjective feelings in animals can only be judged by
extrapolation, but it seems plausible that identical motor and
autonomic responses and facial expressions as well as complex
adaptive goal-directed forms of behavior - in both animals and
man, and in both experimental and everyday life situations -
correspond to identical feelings. Studies on the electrical stim-
ulation of the human brain also indicate that stimulation of
structures similar to those from which emotional reactions can
be evoked in animals produces feelings in people (Heath &
Mickle 1960, King 1961; Mark & Erwin 1970, Sem-Jacobsen &
Torkildsen 1960.)
Consider that, after all, our knowledge about human feelings
is likewise based on extrapolation. We cannot be sure whether
the emotions of others, even those of our friends, are similar to
our own. When a friend smiles, shakes your hand, or kisses you,
you assume that he is happy to see you. But your dog is also
"happy" to see you - "smiles," wags his tail, jumps on you, licks
your hand. I would even say that you can be surer about the
emotional state of your dog than that of your friend. The friend
can give you more details about his emotions verbally, but, in
reality, you count on his friendship, not on the basis of his
words, but on the basis of his behavior.
The brain of a friend is more similar to your own, but the
emotional parts of the brain are also well developed in other
mammals. Similar emotional structures with similar functions
can be compared in man and mammals; it is therefore reason-
able to assume that similar behavior and similar brain structures
are the basis for similar emotions. It is true that the amygdala,
for example, is more complicated in humans than in dogs or cats,
but this is because inhibitory and coordinative mechanisms are
more developed in man.
My high estimation of Panksepp's attempt to objectivize the
emotions does not preclude my pointing out that his schema is
not superior to numerous (likewise imperfect) models of prior
investigators, who were already, long ago, convinced that cer-
tain neural circuits were at the basis of emotion. The Panksepp
model consists only of one more hypothetical neural circuit
joined with arbitrary classes of emotions; this does not build a
convincing bridge between the introspective feelings of emo-
tion in his classification and their real brain substrates. Another
weak point is that Panksepp's model ignores the very attractive
hypothesis of the negative and positive hedonic systems as well
as the problem of motivation. Such an approach overlooks
emotional mechanisms very important for both introspective
and behavioral investigations.
I do not accept Panksepp's classification of emotions into four
classes, three negative ones and only one (very controversial)
positive one. Let us comment on the validity of these classes.
BEHAVIORAL AND BRAIN SCIENCES (1982)3 429
Commentary'/Panksepp: Psychobiology of emotions
Expectancy. This term is arbitrarily introduced by Panksepp
and consists of a large drawer into which many noncomparable
items are thrown. First of all, if one is to use the term expectancy
at all, why restrict it to the expectancy of positive events? There
can just as well be expectancy of danger, pain, distress, en-
emies, even of fear; expectancy is therefore the preparatory
period for all the rest of the categories in Panksepp's taxonomy
rather than a separate category. Positive emotional states are not
limited to expectancy: They are chiefly connected with achieve-
ments. One does not feel joy when one possesses a lottery ticket
(even if one expects good fortune); one feels joy at the moment of
receiving the information that the number has won, when one
obtains the money, and, later, spends it. According to
drive-reduction theories, expectancy (which in other terms
would be called drive) is unpleasant (this, however, not always
correct), and only drive-reduction (i.e., satisfaction) is pleasant.
The physiological meaning of expectancy as used by Panksepp is
also very vague. On the one hand it is treated as a kind of general
arousal or nonspecific motivation (with nonspecific external
signs such as searching, sniffing, etc.). On the other hand, it
includes specific behaviors - those involved in feeding, drink-
ing, sexual behavior, etc. The fact that during stimulation
animals can change their behavior according to environmental
stimuli does not indicate that the state evoked by brain stimula-
tion is a nonspecific arousal to do anything or to expect anything;
it indicates that behavior evoked by this stimulation is not
limited to automatic motor patterns but, like normal behavior, is
adaptable, and can be modified.
Panic. Another quite arbitrarily introduced term is panic,
which, according to the dictionary as well as in psychiatry and
social psychological usage means "sudden terror, excessive,
uncontrollable, and illogical fear, infectious fright." This corre-
sponds to a generalized fear state. On the basis of my own
experiments on the generalization of fear stimuli, I have put
forward the hypothesis that there exist separate brain mecha-
nisms for specific fears (fear of a definite imminent noxious
event) and mechanisms of generalized fear (Fonberg 1961),
which are also activated during neurotic states (Fonberg 1958;
1979a). But if we accept the division of emotions into only a few
classes, panic would surely belong to the emotion of fear. Crying
produced by the isolation of young from their mother is also
evoked by fear. Other symptoms of "panic" described by Pank-
sepp are rather similar to signs of general depression.
Fear and rage. These two classes of emotions in Panksepp's
classification are less controversial; they are widely accepted
and based on numerous data. Most authors agree that the fear
and rage systems are separate, although some do think they
form one common system. My experiments on dogs indicate
that there are quite separate points from which rage and fear
responses can be evoked (Fonberg 1966; 1967). These behaviors
were stably evoked for weeks by the stimulation of definite
points and never substituted for one another. I have shown that
avoidance responses are easily formed to a conditional stimulus
by the stimulation of fear points, but not rage points. Other
authors have demonstrated that stimulating rage attack points
can even have a rewarding effect (Fonberg 1979b; Perochio &
Alexander 1974).
The fear system is separate from the pain system (Bolles &
Fanselow 1980; Fonberg 1980), a fact that Panksepp seems to
overlook. But even the pain system (as shown by anatomical and
physiological studies) is not uniform and can be divided into
three different systems (sensory, cognitive, and emotional),
with different anatomical substrates. Mechanisms of aggressive
behavior and rage are also very complicated and depend on
different emotional states (Adams 1979; Fonberg 1979b; Per-
ochio & Alexander 1974), but at least the term rage is much
more understandable than expectancy and panic, and the re-
sults of corresponding investigations are more precise.
Another question I would like to raise is whether the hypo-
thalamus is the most important brain structure for emotions. I
430 BEHAVIORAL AND BRAIN SCIENCES (1982)3
agree with Panksepp in stressing the role of the hypothalamus
and have myself put forward similar ideas (Fonberg 1966; 1967).
However, our experiments on the effect of amygdala lesions
have shown that the dorsomedial part of the amygdala is proba-
bly even more important for emotional states. After receiving
lesions in this area, dogs, who are very good subjects for
studying emotion, are completely indifferent and emotionless.
They are able to perform many difficult tasks, but they rarely do.
They lose their affectionate behavior toward the experimenter,
which is reflected in deterioration on their socially reinforced
tasks (Fonberg 1972; 1981; Fonberg & Kostarczyk 1980).
I would rather think that the amygdala is crucial for emotion
(perhaps for the subjective evaluation of its hedonistic valence)
and that the hypothalamus is involved in integrative and execu-
tive functions connected with emotional behavior. In this re-
spect the hypothalamus could be regarded as a "command"
system.
Inhibition of emotions is a problem that should also be
considered, but it is scarcely discussed by Panksepp. The
existence of two functionally antagonistic systems (i.e., excitato-
ry and inhibitory) within the amygdala (Fonberg 1963; 1968)
suggests that emotions, although also controlled by the neo-
cortex, may be inhibited in their own source within the emotion
system, and this inhibition may have a more intrinsic character.
In conclusion, I did read Panksepp's article with great in-
terest, for he discusses many very important points and fur-
nishes much extremely interesting data. I have not discussed all
the important topics the article called to mind (e.g., the neu-
rochemistry of emotions and the role of emotions in psychiatric
disorders, which is very close to my own interest; see Fonberg
1958; 1979a; 1981). I agree fully with his main thesis that
emotions are objective events and that they should be investi-
gated by combined psychological, behavioral, anatomical, and
neurochemical methods, and ascribed to definite brain sub-
strates. Panksepp's essay did not entirely fulfill my expectations
(in the usual sense of this word), because his arbitrary taxonomy
and model do not clarify these very important issues. The article
was, nevertheless, very interesting and stimulating to
discussion.
Can phenomenology contribute to brain
science?
Gordon G. Globus
University of California Irvine Psychiatry Service, Capistrano by the Sea
Hospital, Dana Point, Calif. 92629
My aim is to comment sympathetically on Panksepp's thesis that
"human introspective access to emotional states may provide
direct information concerning operations of emotive circuits and
thus be a primary source of hypotheses for animal brain re-
search." I shall, however, consider the more general thesis that
human introspective access to all conscious states may provide
such direct information, namely, that the disciplined study of
consciousness (i.e., phenomenology) can contribute to brain
science. If it is so for emotional states, then it is likely so for all
conscious states as well.
I shall expand Panksepp's framework in another way, by
considering some of the conceptual morasses that he (perhaps
wisely) avoids, especially the "world knot" of the consciousness/
brain problems, which entangles us as soon as we try to unpack
what Panksepp means by "direct" information. Now, contem-
porary brain science has provided richly detailed knowledge
regarding the covariation of consciousness and brain. But until
we have a systematic account of the rules governing this covaria-
tion, Panksepp's thesis and its present generalization are se-
riously compromised. In effect, depending on one's particular

view of the consciousness/brain problems, the thesis may or may
not be correct.
For example, if consciousness is an emergent property of
brain functioning, as Sperry (1969) would have it, then Pank-
sepp's taxonomy of emotion does not tell brain science anything,
for Sperry's emergent is different from, more than, and not
reducible to the underlying neural operations; indeed, it would
be a species of category error to make inferences/rom emergent
properties to properties of the interacting parts from which they
emerge. If consciousness is fundamentally distinct from brain,
but interacting with it, as the Cartesian Eccles (1976) would
have it, then a classification in one does not necessarily imply
anything about a classification in the other. Of course, if con-
sciousness is "strictly identical" with brain (Feigl 1967), then
Panksepp's thesis is literally correct.
The claim that phenomenology can contribute to brain sci-
ence is thus relative to a pre-existing philosophical position that
Panksepp does not indicate. When he talks of a conscious mind
that is able to "see clearly" subcortical dynamics, brain circuits
which "mediate" emotions, and brain mechanisms that "elabo-
rate" consciousness, he owes us a viable account of the con-
sciousness/brain relation which will specify the meanings of the
terms in quotation.
There are various attitudes brain scientists may adopt when
faced with this philosophical puzzle. Probably the great majority
would recoil from the philosophical issues entailed by Pank-
sepp's thesis and continue to do the bench science they have
been (quite successfully) doing all along. Some intrepid pragma-
tists may choose to follow Panksepp's lead and actually do some
systematic study of their own consciousness, and then see if it
proves of heuristic value for their research. Others may even be
motivated to work directly on the consciousness/brain problems
so as to give phenomenological contributions to brain science a
firm foundation. Although these problems are traditionally
considered the province of philosophy, and empirical science is
thought to be irrelevant to their resolution, it is worth consider-
ing the possibility (since we seem no closer than ever to resolv-
ing these problems) that a clear understanding of some basic
principles of brain functioning might move philosophical discus-
sion forward at certain sticking points (such as the notorious
causal theory of perception).
A comparison of the merits of these respective attitudes
would require considerable discussion, and would probably
prove indecisive, since these attitudes run deep. On issues like
this, each of us tends to take the path which interests him most,
which is probably salutary, since it maintains scientific plural-
ism. I want to adopt the pragmatic attitude here toward the
general thesis that the study of consciousness can contribute to
brain science, and to supplement Panksepp's illustration of this
by another example.
I shall consider briefly a partial taxonomy of consciousness,
and what value it might have for brain science. I claim (where
"claim" is understood phenomenologically as an invitation to
coreflect; Zaner 1970) that consciousness over time divides into
two incompatible phases. In the ordinary phase of intentional
action, consciousness is always a consciousness-of, whether
conscious of emotions, wants, external objects, thoughts, etc. In
the extraordinary phase in which all intentionality ceases (which
may require some special training in meditation to achieve),
there is no consciousness-of. There is no object of awareness.
There are no distinctions whatsoever. There is only the distinc-
tionless case of an unbroken and unbounded whole. I claim also
that consciousness may vary continuously in "richness'; under
different conditions, there is more or less of it, as when we are
wide awake, drowsy, or in nondreaming sleep. Furthermore,
the state of sleep periodically supports episodes of conscious-
ness (dreaming) that can be so vivid that they are indistinguisha-
ble from waking consciousness.
If these phenomenological observations in fact do provide
"direct information," as Panksepp says, about the brain, then
Commentan//Panksepp: Psychobiology of emotions
we are in a position to theorize that the highest brain system or
"super system" has two primary modes of organization, an
ordinary mode associated with what is presumably a very high-
level neural processing action and an extraordinary mode in
which that action ceases. The complexity of this system's organi-
zation can vary continuously. There is variation in the complex-
ity of this system's organization during waking and periodically
during sleep. The study of the brain during dreaming thus
provides an opportunity to understand how this system is
autonomously constituted, without the complications of on-
going sensory input and behavior. I think it would be quite
difficult to come to such a theory via laboratory investigation - at
least in the present state of the art.
Whatever the value of Panksepp's observations and the pre-
sent phenomenological ones, it can be seen that the study of
consciousness could aid in developing the theory of brain at its
most complex levels of functioning, a level about which brain
science is notoriously silent and - given the complexity of the
system under consideration - is likely to remain silent for some
time. Practically speaking, if we are to theorize at all about the
highest levels of brain functioning, we ought at least to consider
what phenomenologists have to say about consciousness in
devising our theories.
In summary, Panksepp's thesis generalized to the claim that
phenomenology can contribute to brain science can only be
judged pragmatically, in the absence of a solution to the con-
sciousness/brain problems. With the impoverished knowledge
that we presently have of brain functioning at the highest levels,
it would be premature to rule out any phenomenological contri-
bution on dogmatic grounds. In the context of a multifaceted
brain science, it would seem instead salutary to admit a lively
interface between phenomenology and brain science, as Pank-
sepp has demonstrated for the case of emotions.
On the classification of the emotions
Jeffrey A. Gray
Department of Experimental Psychology, University of Oxford, Oxford 0X1
3UD, England
Panksepp has described a general approach to the construction
of a psychobiological theory of emotion with which I am in
substantial agreement. Details of the scheme for classifying the
emotions that Panksepp proposes are also consonant in part with
a scheme put forward previously on different grounds (Gray
1972). However, Panksepp's scheme rests on too narrow an
evidential basis (largely that of experiments on self-stimulation
and experimenter-applied stimulation of the hypothalamus). In
consequence, his suggested set of four basic emotional states
runs into major difficulties when it is confronted with data
obtained by other methods.
These difficulties centre on Panksepp's distinction between
"fear" and "rage," a distinction based upon the experimental
difference between sites in the hypothalamus at which stimula-
tion elicits flight (= fear) and attack (= rage) respectively. But,
even within Panksepp's own terms of reference, the distinction
is weakly supported. For, as he states, "Hypothalamic sites that
provoke flight surround the core systems from which rage
(threat behavior) is elicited (de Molina & Hunsperger 1962), and
there appears to be substantial anatomical overlap and func-
tional interaction between these systems." And, as his discus-
sion elsewhere in the article makes clear, given such overlap and
interaction, it is an arbitrary decision (in the absence of other
evidence) whether flight and attack are alternative expressions
of the same emotional state (as suggested by Gray, 1972) or each
an expression of a different emotional state (as believed by
Panksepp). Fortunately, there is other evidence. But this evi-
dence does not favour two of Panksepp's key assumptions: that
BEHAVIORAL AND BRAIN SCIENCES (1982)3 431
Com?nentan//Panksepp: Psychobiology of emotions
"fear" should be defined in terms of flight, and that flight and
attack reflect different emotional states.
How should one define fear? Defining fear in terms of flight
behaviour leads to conclusions that are at variance with those
that can be derived from most other relevant lines of argument.
Two other behavioural measures have commonly been used in
studies of fear: defecation (especially in the open field test), as in
studies of the genetics of fearfulness (Broadhurst 1960) or sex
differences in fearfulness (Gray 1979b); and the suppression of
responding under threat of footshock, as in the use of Geller and
Seifter's (1960) schedule or passive avoidance paradigms to
quantify anxiety. Neither of these measures relates to flight
behaviour in the way that Panksepp's scheme would lead us to
expect. Thus, antianxiety drugs, which clearly alleviate be-
havioural suppression induced by the threat of shock, do not in
general affect escape (Gray 1977); septal lesions, which reduce
both passive avoidance behaviour and open field defecation, do
not affect skilled escape behaviour and they even improve
escape during agonistic encounters with conspecifics (Gray
1982a); and female rats, who defecate less and show poorer
passive avoidance than males (Gray 1979b), nonetheless run
more than males when they are shocked (Beatty & Beatty 1970;
Wilcock 1968). It follows from such patterns of data (and others
could be cited) that, ifflightis the correct criterion of fear, these
other measures are inappropriate. Yet the case for both defeca-
tion and passive avoidance as measures of fear is strong, and
furthermore (although there are exceptions) the conclusions to
which these two indices give rise are usually concordant with
one another (Gray 1979b). It would seem more parsimonious,
therefore, to abandon the postulate thatflightis a good index of
fear, especially since Panksepp does not offer either evidence or
argument in favour of this postulate.
Do flight and attack reflect different emotional states? As
noted above, the hypothalamic sites at which stimulation elicits
flight and attack closely intermingle. Indeed, they are some-
times the same sites. As Panksepp states, stimulation at some
points "typically elicits flight, [but] defensive attack often en-
sues, if no avenue of escape is available." Under these circum-
stances, one may reasonably assume that both these patterns of
behaviour are expressions of the same emotional state, the
nature of the environment then determining which pattern
prevails on a particular occasion. Such an assumption would lie
comfortably within Panksepp's general line of reasoning (though
he chooses not to make it). It is supported by the general
similarity between the experimental conditions which, in non-
physiological experiments, provoke flight and attack respec-
tively. These two patterns of behaviour may each be elicited not
only (as is well known) by punishing stimuli (e.g., footshock),
but also by frustrative nonreward (e.g., Adelman & Maatsch
1956; Gallup 1965). Furthermore, there appears to be a psycho-
logically important distinction between unconditioned punish-
ment and nonreward, on the one hand, and conditioned stimuli,
which signal the imminent occurrence of these two events, on
the other. Both flight and attack are elicited by unconditioned
aversive events of either (whereas behavioural inhibition or
freezing are not); while conditioned aversive stimuli do not
typically elicit either flight or attack, but freezing and be-
havioural inhibition instead (Gray 1975; Myer 1971).
This distinction between conditioned and unconditioned
aversive stimuli makes it possible to predict many of the effects
of drugs (Feldon, Guillamon, Gray, De Wit, & McNaughton
1979; Gray 1977); and it forms part of the evidence on which I
have based the concept of a "behavioural inhibition system"
specialized to respond to signals of aversive events (whether
punishing or frustrating) but not the aversive events themselves
(Gray 1975; 1977). The resulting model differs from Panksepp's
in the following two principal ways. First, anxiety (which sub-
sumes the concepts of fear and anticipatory frustration as used
by Mowrer, 1960, and Amsel, 1962, respectively) is treated as
activity in the behavioural inhibition system, elicited by condi-
432 BEHAVIORAL AND BRAIN SCIENCES (1982)3
tioned aversive stimuli and manifested (among other ways) as
passive avoidance behaviour. Second, a separate "fight-flight
system" mediates responses to unconditioned aversive stimuli,
and activity in this system may manifest itself as eitherflightor
attack behaviour. I have set out the evidence for this theory
elsewhere (Gray 1972; 1982a; 1982b). Clearly, further experi-
mental work is needed to test the two models. The fact that they
make such clearly different assumptons at these key points
should facilitate the task of discriminating between them.
Panksepp's psychobiological theory of
emotions: Some substantiation
Robert G. Heath
Department of Psychiatry and Neurology, Tulane University School of
Medicine, New Orleans, La. 70112
Panksepp has tackled a subject of tremendous scope. The
manner in which he undertakes the task is to be commended.
He modestly labels his hypothesis as a start, which he hopes will
lead to sound experimental work for broadening the base under-
lying his theory. I am in agreement with many of the ideas he
advances.
A major premise of Panksepp's is the importance of introspec-
tion by human beings, the only species capable of reporting its
thoughts and feelings - essential information in providing leads
concerning the function of neural-emotive circuits. Although
Panksepp is somewhat apologetic in discussing this point, I
consider it a primary strength of his article. It is apparent from
human introspection that sensory perception is closely interre-
lated with emotion. Panksepp cites a number of references, but
the data base he uses to substantiate his hypothesis of brain
circuitry and emotion is, unfortunately, narrow. Whereas he
focuses principally on the effects of stimulation to and ablation of
the lateral hypothalamus in the rat, the existing data base for this
neural-emotional behavioral relationship is much broader. It is
derived from the use of many additional divergent methods and
thereby provides hard data to support some aspects of Pank-
sepp's hypothesis. Moreover, the available data base provides
additional platforms for modified and expanded hypotheses for
the collection of still more data.
The most significant data are from extensive human studies in
which a wide variety of inspective findings have been correlated
with the subjective introspective data reported by patients
(Heath 1974; 1975; Heath, Cox, & Lustick 1974; Heath &
Gallant 1964). These investigations involved implantation of
deep brain electrodes for long-term studies. Electrodes have
been implanted not only into the hypothalamus and numerous
other sites in the conventional limbic system, but also into
several sensory relay nuclei and deep sites involving other
functional systems, as well as over numerous surface brain
areas. With these techniques, the effects of brain stimulation on
feelings have been recorded. Further, it has been possible to
correlate changes in deep brain activity (electroen-
cephalographic recordings) with varied emotional states: spon-
taneous emotion, emotional states induced in association with
mood-altering drugs, and, in a few cases, emotional states
induced with administration of transmitters directly into the
brain parenchyma at sites within the "hard-wired" emotive
circuits. Multiple brain recordings concomitant with patients'
introspective reporting have suggested that correlates with both
pleasurable and aversive emotional states occur elsewhere than
in the lateral hypothalamus. Both recording and stimulation
data indicate that the lateral hypothalamus may well be func-
tioning as a type of final common pathway in the emotional
circuitry. This would suggest that one should use caution in
extrapolating too extensively from rat data (self-stimulation,
ablation). Let us hope that Panksepp has put to rest the long-

prevailing concept that brain stimulation, as observed in ani-
mals, represents sham rage. The emotion generated in patients
by means of brain stimulation has always been an integral part of
consciousness - never a sham response.
Deep electrode studies in intractably ill patients have led to
studies in animals in which it was possible to outline the extent
of the emotional circuitry, as well as the functional relationships
within the system more completely and precisely than cited by
Pankscpp. "Although it is clear that lateral hypothalamic neu-
ropil contains widely ramifying, long-axoned systems, both
ascending and descending, with multiple interneuronal feed-
back loops, . . . our knowledge of functional circuits in such a
reticular substance remains too imperfect to attempt filling in
the details of system connectivities." Numerous anatom-
ic-physiologic studies in animals (derived from the correlated
introspective-inspective studies in human subjects) provide a
base of hard data for Panksepp's speculation about interrelations
of the neural systems for sensation and emotion (Heath 1976;
1977; Heath, Dempesy, Fontana & Fitzjarrel 1980; Heath,
Franklin, & Shraberg 1979; Heath, Llewellyn, & Rouchell
1980). They have led to the demonstration of the important role
of the vestibular proprioceptive cerebellum (midline vermis) in
modulating emotion, as well as the recent demonstration of
cerebellar pathology associated with disordered emotion. In
sensory deprivation experiments, the absence of proprioceptive
input rapidly induces the severe disruptive emotion characteris-
tic of psychosis. Auditory and visual stimuli can profoundly alter
one's emotional state, and, conversely, changes in emotional
state alter sensory perception. Panksepp correctly relates appe-
titive behaviors to emotional circuitry. Basic metabolic states
related to survival selectively alter both emotional and sensory
perception.
In conclusion, I agree with most of the principles in Pank-
sepp's well-outlined hypothesis. To avoid repetition, I have not
focused on many of his major points with which I agree. I have
chosen instead to discuss only a few of the author's points that I
believe could have been better illustrated had he drawn on the
existing broader data base. Like Panksepp, I feel the need to
emphasize that the subject under consideration is much broader
than can be reviewed in this relatively short commentary. Only
a few aspects of this very broad subject have therefore been
considered.
From stimulus-bound emotive command
systems to drive-free emotions
C. E. Izard
Department of Psychology, University of Delaware, Newark, Del. 19711
As a differential emotions theorist, concerned mainly with the
role of discrete emotions and patterns of emotions in human
development, I found myself avidly foraging through Pank-
sepp's article, swept along, at times seemingly uncontrollably,
by my expectancy command system, or rather by its human
homologue. In my theory (Izard 1977), the homologue would be
the discrete emotion of interest-excitement and various other
emotions in patterns or combinations with interest, patterns
that would modify the direction and intensity of behavior as
required by ecological conditions and events.
There is a serious though perhaps not fatal flaw in this
homology. The emotion of interest, although capable of amplify-
ing or attenuating the appetitive behaviors under the control of
Panksepp's expectancy command system, is also capable of
operating quite independently of them. My foray through this
article was accomplished while my appetitive drives were well
satisfied. Thus, the emotion of interest is neither an appetitive
affect nor one that is slave to consummatory behavior. Interest
has the utmost flexibility and generality, capable of motivating
Comm«ntart//Panksepp: Psychobiology of emotions
behavior in any direction and relative to any object or event, but
all the appetitive functions under the expectancy command
system are highly specific with respect to object and consumma-
tion, with the possible exception of exploration. Interest, in
human beings, surely motivates all kinds of exploration - from
environmental to epistemic - and it can in the course of such
exploration interact with other emotions as well as with the
appetitive functions. So here we may have a common ground.
But is it enough? Perhaps it is, for it certainly suggests a common
thread across a wide span of species. And it seems reasonable
that in human phylogeny something like Panksepp's expectancy
command system could have differentiated into various rela-
tively independent motivational systems, including such drives
or affects as hunger and sex, as well as the emotion of interest.
I am suggesting that the human emotion of interest - which,
among other things, promotes attention to biologically as well as
socially important stimuli (Langsdorf, Izard, & Rayias 1981) -
may have had its origins in something like an expectancy
command system. Its evolution as a discrete system indepen-
dent of appetitive functions may have stemmed from mutant
behaviors that had no immediate consummatory goal, were not
stimulus-bound, yet had longer range pay-off in the processes
of adaptation. For example, the distinct advantage of territorial
exploration without the restrictions of appetitive drives is the
greater flexibility of cognitive and motor activities.
In similar fashion, something like the "panic" command
system could have given rise to the drive-free human emotions
of sadness and anger. However, Panksepp's panic command
system would be much more appropriately labeled the distress
command system. While panic connotes overwhelming terror
(the extreme of fear), distress connects both semantically and
historically with sorrow, sadness, or grief. Darwin (1872/1965)
argued that the human expression of sadness is a neuromuscular
pattern that regulates the emergency facial and vocal expression
of physical distress. The young infant's physical-distress facial
pattern in the brow-eye region (brow drawn sharply down and
together and eyes tightly closed) can be converted to a sadness
signal (obliquely raised inner corners of the eyebrows and
slightly squinted eyes) by contraction of the medial frontalis
muscle, which is antagonistic to the orbicularis oculi, corruga-
tor, glabella, and eyebrow depressors that tightly close the eyes
and lower the brows. Interestingly, the pattern in the mouth
region that completes the full-face sadness expression (mouth
corners pulled downward by action of the depressor anguli oris)
is occasionally seen in young infants immediately following
painful medical procedures (Izard, Dougherty, Coss, &
Hembree 1981). This latter pattern tends to reduce the size of
the mouth and is associated with (and may "cause") a dampening
of the distress cry (vocalization). These changes in the facial and
vocal signals probably tend to elicit cuddling and other nur-
turant behaviors from the care-giver.
In terms of observable display, the anger expression is even
closer to the infant's physical distress expression, and in young
infants this expression is often followed by an anger expression.
This suggests possible interconnections between pain or physi-
cal distress, sadness, and anger, with the latter being the
emotional counterpart of the protest seen in mother-infant
separation. Relevant to this point and to Panksepp's discussion
of separation as a stimulus that activates the panic command
system, Shiller and Izard (1981) have found that the most
frequent emotion expression during brief separation of 13-
month-old human infants from their mothers was anger; the
second most frequent expression was sadness, with a small
minority showing this expression predominantly.
Much of what Panksepp describes under the rubric of the
panic command system parallels in some way what we see in
human distress and sadness. As just noted, however, we see
anger and sadness in mother-infant separations, but we have
observed no parallel for the freezing-explosive behavior se-
quence Panksepp describes.
BEHAVIORAL AND BRAIN SCIENCES (1982)3 433
Commentary/Panksepp: Psychobiology of emotions
In early infancy, certain emotion expressions sometimes ap-
pear to function as stimulus-bound behaviors. Pain immediate-
ly elicits the well-defined physical distress pattern, and a nod-
ding human face regularly evokes a smile. While these affective
expressions do not serve appetitive drives, they function as
powerful social signals that typically elicit nurturant and positive
affective behavior on the part of the care-giver. With increasing
age, increasing cognitive and motor capacities, and the impact of
life experiences, the emotion expressions appear less and less
stimulus-bound; they become responsive to a virtually limitless
array of stimuli. It is this great capacity of the human being to
respond with emotion and hence to be appropriately motivated
in relation to highly diverse incentive events that make human
behavior so complex and so adaptive in so many different
situations.
The rage and fear command systems have quite comparable
homologues in human emotions. It is conceivable that at an
early stage in our evolution, the similarity was even greater.
At the present time, of course, comparisons yield several
important differences between these command systems and the
emotions of anger-rage and fear—terror in human beings. In
human beings there are weaker connections between activation
of the anger mechanisms and aggression than is the case in lower
animals. Also in the case of human beings, anger and fear (as is
true of other emotions) rarely occur in "pure" forms for very
long. They typically occur in patterns with other emotions that
are caused by events in the situation and by the original emo-
tion, anger or fear. These patterns of emotion elicit (or occur
with) certain affective-cognitive structures that stem from
learning and life experiences. The other emotions in the pattern
and the affective-cognitive structures serve to inhibit, amplify,
or attenuate the "original" emotion and give human behavior its
great complexity and variable relationships with external
events.
I have a few miscellaneous comments on the paper. (1) M. B.
Arnold (1960a; 1960b) should be credited for the observation
that "phenomenological analysis of emotional experience can
guide us in identifying the brain structures and pathways that
mediate feelings and emotions" (1960a, p. iii). (2) The robust
evidence for the innateness and universality of certain human
emotion expressions (Ekman, Friesen, & Ellsworth 1972; Izard
1971) provides general support for Panksepp's view of emotions
as fundamental evolutionary-biological phenomena. (3) A
rapidly growing body of research on emotions in human devel-
opment is generally supportive of Panksepp's view on the
fruitfulness of an ontogenetic approach in understanding
brain-emotion relationships. (4) the emotional-behavioral pro-
cesses described under the panic command system (at least the
sadness/grief components) are more probably associated with
depressive disorders rather than those specified in Figure 5. (5)
The concept of "emotional disorders" as commonly used is a
misnomer, from Panksepp's vantage point, as well as mine. In
Panksepp's system and in differential emotions theory, the
emotions are basically adaptive. In cognitive and behavioral
dysfunction, the problem is not with the emotions but with
faulty or deficient connections between emotion, cognition, and
behavior. Panksepp's use of the term emotional disorder as
unbalanced activities among emotions (or "emotive systems") is
a reasonable one, but it does not reflect the problem of dishar-
mony between the emotion, cognition, and motor-behavior
systems.
Despite my differences with Panksepp - some of which must
come from differences in perspectives developed from immer-
sion in the study of different phyla - I think his article provides
biopsychological support for the currently increasing accep-
tance of differential emotions theory, which postulates a limited
number of innate fundamental emotions, each with a unique set
of neurophysiological substrates, expressions, and quality of
consciousness (Izard 1977). The last is basically a feeling state
that has inherently motivational-adaptive functions, including
434 BEHAVIORAL AND BRAIN SCIENCES (1982)3
the vastly important one of recruiting and sustaining appropri-
ate cognitions and instrumental behaviors.
A two-tiered theory of emotions: Affect and
feeling
Julian Jaynes
Department of Psychology, Princeton University, Princeton, N.J. 08544
No theory of emotion can make sense out of the psychological
data unless it is absolutely clear about what is conscious in
emotional experience and what is not.
Let me state briefly my own point of view.
All mammals, including ourselves, have a basic set of genet-
ically organized affects, specific aptitudes to respond to charac-
terizable classes of stimuli or events in certain characterizable
ways. Back in evolutionary history, I believe, the core of most of
these aptitudes, their "aptic paradigms," were actual overt
behaviors of reptiles. But in mammals these overt behaviors
have become inhibited by more recently evolved parts of the
brain, leaving them more diffusely related to actual overt behav-
ior in a more general energizing way (MacLean 1973). Most
emotional behaviors of mammals are thus paleoreptilian behav-
iors transformed on both the stimulus and the response side by
limbic and cortical selective inhibition. And just as the behav-
iors in the repertoire of any animal cannot be set out as similar
things, with the same characteristics - being different in their
nature, timing, pervasiveness, situational dependence, interre-
lations with other behaviors, and dependence or independence
of them - so mammalian emotions also cannot be set out in a
linear array. A list of mammalian emotions or affects, although
nonlinear and often interlocking, could start off clearly enough
with the usual items like fear, anger, disgust, and the various
affectional systems (Harlow 1971) and then end up somewhat
vaguely, but no less appropriately, with those behaviors that are
less commonly called emotional because of their lesser inten-
sity, such as drowsiness and curiosity. Since expectancy is
common to all molar mammalian behavior, I would not include
it in such a list.
Are all behaviors emotions? It is not the behaviors themselves
that we denote by emotions, but their inferred organizations,
which usually include four characteristics: (1) emotions have a
clear variation in intensity, so that every emotion has a more or
less to it; (2) even if not immediately expressed, emotions are
associated with a distinct class of behaviors; (3) emotions have
bodily and often facial expressions, which are distinguishable
from more usual states; (4) there exist distinct classes of stimuli
or situations that occasion emotions.
This list is to be compared with Panksepp's list under his
heading "A definition of emotions." What Panksepp has done is
to suggest a theory of the brain circuitry involved that is
interesting in itself, but is so restricted in an ad hoc way as to
eliminate many behavioral responses that most observers would
include under the term emotion.
But the real point of my commentary concerns consciousness
and how consciousness, once established in history, adds some-
thing new to genetically programmed mammalian affects that
we can call/eeftngs. I believe this to be an historical change that
occurred in the ancient Mediterranean and Near Eastern world
sometime after 1000 B.C. which, as I have tried to show else-
where, is the beginning of the period when human beings
learned consciousness on the basis of metaphor (Jaynes 1976).
By consciousness I mean precisely what Panksepp does in his
few references to it, namely, all our introspective experience,
including retrospection and imagination. Once human beings
have these capacities, they can think out consciously and pri-
vately what they are to do, in contrast to a more ancient
mentality called the "bicameral mind," in which what we call

volition was a matter of hearing and obeying auditory hallucina-
tions called gods.
With this metaphor-generated consciousness, there comes
the metaphor of time as a space, or spatialized time, in which
human beings 'see' themselves as embedded in their own
histories. This human sense of a lifetime begins over the same
period as the development of the idea of historical time (for
evidence for this, see Starr 1968), as well as the appearance of
ideas of justice, retribution, doing wrong, remission of wrongs,
and forgiveness, all very curious behaviors when we think of
them against the background of the evolution of mammalian
behavior, and all occurring in world history for the first time.
The result is a new set of social and political problems. Behind
these social changes are the changes in personal emotion that
arisefromthe new human capacity to stretch out the affects over
the spatialized time, or, in other words, to dwell on past
behaviors or on possible future behaviors and respond to them
as if they were presently occurring, with copies of the affects
themselves. These are our feelings. My language here is difficult
and metaphoric, because these processes are not fully under-
stood at present. But let us consider some examples.
Shame is one of the most powerful affects. (For the best
discussion, see Tomkins 1963). It occurs in many mammals,
such as canines, primates, and early man. Observation of pre-
sent-day children's development or reminiscences about our
own will demonstrate the tyrannous power of humiliation, of
rejection from a social group, in shaping behavior to some norm
or other. In fact, we have all been so shaped by shame into our
customary behavior that, as adults, we rarely have this affect in
its pure state. But once human beings can reminisce about
shameful actions or imagine possible shameful actions, until
they imagine real or possible humiliation and ostracization in a
general way, we have guilt: Guilt is thus a conscious feeling that
is generated from the biological affect of shame.
This is obvious, I think, in the development of consciousness
in the child (see Kohlberg 1976). It is also evident in the
development of consciousness in ancient history. Consider, for
example, the story of Oedipus. In its earliest form (in two lines
from the Iliad and two lines from the Odyssey - pretty meager
evidence, I agree), it seems to be the story of a king who had
killed his father and married his mother and had two daughters-
sisters by her, and who (since the incest taboo is the reason the
story is mentioned at all) certainly felt some shame, but then
apparently lived a long remorseless life with his somewhat
strange family until he was buried with military honors at
Thebes. But just a few centuries later, in the fifth century B.C.,
consciousness has transformed the story into the tragic sense of
guilt that savages its way through the great tragic trilogy of
Sophocles, (on this point, see Dodds 1951).
Similarly with other genetically organized affects. Fear, when
consciously reminisced about and projected into the future,
becomes the conscious feeling of anxiety. The affect of anger
stretched out over spatialized time in consciousness becomes
hatred. Excitement becomes the conscious feeling of joy; dis-
gust, contempt; affiliation, love. And so on. All of these changes
need to be carefully documented in historical data and in child
development.
Perhaps most interesting are the sexual emotions, which were
usually omitted from Victorian lists of the emotions (including
Darwin's) because their bodily expression is primarily genital
rather than facial. We do know that mating in the anthropoid
apes, in contrast to ourselves, is casual and almost minimal, with
observations of mating in gibbons, chimpanzees, orangutans,
and gorillas in the wild being extremely rare. Paralleling these
facts, tomb and wall paintings, sculpture and the writings of
bicameral civilizations rarely if ever have any sexual references.
All classicists will agree with this, that all Mycenean and Minoan
art, in particular before 1000 B.C. is what seems to us as severely
chaste. But after the advent of consciousness, say, about 700
B.C., Greek and Etruscan art is rampant with sexual references,
Commentary/Vanksepp: Psychobinlogy of emotions
very definitely demonstrating that sexual feelings were a new
and profound concern in human development in these regions.
We can perhaps appreciate this change in ourselves if we try to
imagine what our sexual lives would be like if we could not
fantasize about sexual behavior.
By this two-tiered theory, I mean to imply that because of our
ability to fantasize, both by reminiscing about the past and
imagining the future, our present affectional experience consists
of far more than the basic affects of mammals. I should like to
point out here in passing how well this development fits in with
the James-Lange theory (with some slight shifts in terminology).
We see a bear, which occasions the genetically organizaed affect
of fear, and we run away, which we are then conscious of,
turning the fear into the feelings of extreme anxiety. So the
James-Cannon controversy may be resolved.
Emotional terminology is a considerable problem. We have
an excessive vocabulary of emotional terms that arise from our
own experience and the culture in which we live. In observing
animal behavior, when we see a behavior similar to our own in
emotional circumstances, we usually label it with that emotion.
This is, of course, all we can do; otherwise the behavior of
animals is merely topological. But if we have not made a clear
distinction between the affects and conscious feelings, we are
likely to make mistakes and project our own conscious feelings
into the simple affects of lower animals. Panksepp is making the
methodological point that we have to use our own consciousness
of our own behavior to label the emotions of lower animals.
True. But it certainly needs the added caution I have just
outlined. The two-tiered theory also suggests that studies of
hypothalamic stimulation or other presently known neural sub-
strates of emotion in animals, although largely important for
understanding basic mammalian affects, are limited in their
application to human emotional experience.
Parting's sweet sorrow: A pain pathway for
the social sentiments?
Leonard D. Katz
Department of Philosophy, Princeton University, Princeton, N.J. 08544
The general theoretical and methodological framework em-
ployed by Panksepp should be relatively uncontroversial within
the community of physiological psychologists, ethologists, and
comparative neuroanatomists interested in the functional
neuroscience of emotion. There, at least, this way of proceeding
comes naturally, even when candor about the use of our own
wired-in emotional categories in interpreting animal behavior
does not. On the other hand, those accustomed to the study of
emotion at a more molar level - and especially social psychol-
ogists, computational cognitivists, and holistic functionalists -
will be understandably disposed to doubt that physiology and
neurochemistry can speak to their subject at all. I shall first
attempt to argue and cajole these skeptics out of their specialist
prejudice. I then turn to the substantive question of Panksepp's
limbic command system for a basic emotion of panic or sorrow,
in terms of which, for the sake of both convenience and con-
creteness, my entire discussion proceeds.
Suppose something like Panksepp's proposed central circuit
is involved in isolation distress as well as in the more elaborate
social emotions. What follows, then? Not necessarily that the
attachment of a child for his mother is subcortically (or even
subcutaneously) localized. We could still say that this is a
relation between persons (and that love for an imaginary person
is only sham love). Or, alternatively, that the child's attachment
is a state of the whole person (or mind or brain), that its
intentional object (underdetermined by the subcortical circuit-
ry) is essential to its being the state it is, and that it must have
some such object to be that sort of social attachment at all. But
however we may answer such subtle questions about attach-
BEHAVIORAL AND BRAIN SCIENCES (1982)3 435
Commentary/Panksepp: Psychobiology of emotions
ment and love, Panksepp's point should remain unchanged. He
would have found a neural center to match his hunch that these
social emotions have a common core. The mammalian basic
emotion of sorrow would then also figure at the center of our
thinking about what makes longing for an absent person what it
is, as distinct from feeling toward that same person gruding
respect, rage, or sympathetic pain. Moreover, we should have
that much more reason to believe that the affective states are
among those psychological states that have a clearly biological
basis - rather than being mere figments of self- or social
attribution, or fallout from the holistic programming or global
activity of the corticalized human brain.
But whereas the general program is plausible, the particulars
of localization and interpretation seem premature. And es-
pecially in the case of panic-sorrow. The possibilities open are
legion; I shall discuss but a single simple hypothesis that must be
refuted before we have reason to believe in a discrete system for
social distress and cohesion. It is that there is no such system
distinct from the central pathways for pain. For all we have been
told, most of Panksepp's forebrain sites are best construed as
sites for central analgesia, and the distress vocalizations (along
with agitated behavior) upon the offset of stimulation, as normal
responses to pain. (Obviously, an ethologist's eye for the to-
pography and significance of the species-typical behavioral ex-
pressions of motivational states is called for here.) The same goes
all the more for the central gray, so far as we know from data
cited here. We may have only central pain, and the rebound on
stimulation offset of the opioid-inhibited affective dimension of
pain (Melzack & Casey, 1970); and no special system for social
pain.1
As for the hypothalamic response that Panksepp suggests
"may . . . indicate the hypothalamic location of the emotive
command system that is attuned to social loss," (in "The 'Panic'
Command System") this seems to have been interpreted as
stimulus-bound attack in the original publication (Panksepp
1971a).2 While the reinterpretation seems to fit better the
original description of behavior, an ethologist's eye, once more,
is needed to rule out alternatives.3 I should like to know, for
example, the significance of the original denial of jumping and
the current mention of leaping. Is this like running or bounding,
and is it compatible with reinterpretation as an escape response,
a sexual approach, or perhaps along the lines of the normal way
an immature rat solicits play? The gaping, chattering, and
squealing described in the 1971 paper require similar careful
examination. Finally, Panksepp's doubts (in "A Definition of
Emotions") about the usefulness of the pleasure-distress dimen-
sion notwithstanding, approach and avoidance are among the
clearest of behavioral signs. Here, as happens quite frequently,
interpretation at the pleasure-pain level is almost all that re-
mains constant through change of hermeneutic view. And, after
all, Panksepp's typology sits well with a hedonic partition of
emotion into positive emotions (lumped here under "expectan-
cy") and the negative emotions, which he calls by their specific
names.
Pleasure and pain are phylogenetically and ontogenetically
old. It seems only reasonable to suppose that the later and
specifically mammalian developments are outgrowths of the
earlier, and that the introspectively easy and theoretically natu-
ral partition of affect into positive and negative marks a biolog-
ically important functional distinction as well. These newer
components, and the older systems on which they are built,
work together. The hierarchically organized mammalian brain is
not an army or bureaucracy in which a command system is
unconditionally obeyed by slavelike subalterns incapable of
initiating action on their own (Gallistel 1980; 1981). Responsibil-
ity is distributed among systems and levels of integration, and
there is every reason to suppose that features of conscious
experience reflected in introspection are distributed in these
ways too. The pleasure-pain dimension may be largely an affair
of the caudal brainstem, while social cohesion, may also involve
436 BEHAVIORAL AND BRAIN SCIENCES (1982)3
what Panksepp regards the "command" responsibility of limbic
centers and still more rostral areas of the brain. But the ques-
tion, "Who does it to whom?' should be left to politics and the
analysis of dominance behavior. Especially at these higher
levels of the vertebrate neuraxis (as in open societies), it is
misleading to cast questions about the division of labor among
the components of functional integrations as questions of who
commands whom.
In the organization of the social sentiments, pleasure, as well
as pain, certainly plays an important role, and not one that an
undifferentiated "expectancy" foraging system could explain.
We are not normally inclined to (spiderlike) eat spouses or
(lizardlike) children when we like them very much (although an
aunt used to tell me that she could). And that we do not normally
resort to the swiftest and surest ways of escaping (or perma-
nently terminating) the aversive distress cry of the infant shows
that undifferentiated distress no more than anger, fear, or panic
explains adult behavior - without which it would never have
been adaptive for young mammals to cry out in distress (as
lizards, for good reason, do not!). From similar observations, the
ancient Stoics drew a conclusion about the origins of the social
sentiments, reported by Cicero:
Therefore, just as it is clearly our nature to abhor pain, so it is evident
that from this same nature we receive the impulse toward those we
have begotten. From this develops the general affinity of human
beings for one another; which thus, likewise, finds its source in
nature. (De flnibus bonorum et malorum, bk. 3, sects. 62-63)
But benevolence is often too weak a force to hold the line against
selfish pleasure and pain. We often need to be brought up short
by the threat of isolation (whether moral or physical) from our
fellows, which we internalize as the self-exile of guilt or shame.
(Contrition, confession, and absolution end the sorrow of social
isolation, not the sin.) Our moral and social nature like our
brain, is a tangled skein of pleasure and pain.
NOTES
1. But is the difference between the pain which makes us shrink into
ourselves, and the panic which impels us to seek comfort from others,
only a matter of degree? More probably - and more likely than either of
the overly simple hypotheses considered in this paragraph of the text -
the integration of pain with sorrow-panic and the other negative affects
is a matter of degree. Panksepp, Herman, Vilberg, Bishop, and DeEs-
kinazi (1980) suggest that their social affect system is an evolutionary
development of a system originally regulating only pain. But in the
present paper, based as it is on a discrete command system picture, this
phylogenetic insight into functional organization is obscured.
2. I assume that the hypothalamic area in question is that called Area
3 in Experiment 1 of the referenced original paper (Panksepp 1971a).
Only this group of rats seems to offer a tolerablefitto the present report;
the others seem really to manifest only stimulus-bound attack. But the
original description as near the lateral border of the ventromedial
nucleus, as compared with the current description as merely anterior
basal, leaves some room for doubt.
3. In this connection, see the specific recommendations of Adams, in
response to the commentaries of the Brelands, Rodgers, and Waldbillig
(Adams 1979, p. 232).
Panic, separation anxiety, and endorphins
Donald F. Klein
New York State Psychiatric Institute, New York, N.Y. 10032
Panksepp spends a good deal of time in proclaiming defensively
that introspection is a good source of testable hypotheses with
regard to emotional organization. He is beating a dead horse.
The effective critique of introspectionism was not that it could
not produce testable hypotheses, but rather that it claimed that
it could produce truths that were irreducible and beyond
testing.
We should focus on the validity of a hypothesis rather than its
source. If Panksepp is affirming that we can produce more valid

hypotheses by introspection than by some dry empirical induc-
tion, I tend to agree with him. However, this affirmation, which
we both support, remains yet another testable hypothesis. It is
certainly not clear that any single approach to understanding
emotion has won the prize.
Therefore, I think that Pankscpp should focus more firmly on
the utility of his definitions and the validity of his distinctions
and forget the self-justification.
In his definitional approach, he says, "On the assumption that
emotions arise from the activities of brain circuits, a general
definition of emotion would consist of a list of attributes such
brain circuits possess . . . the list must first be constructed
rationally."
This is just what Panksepp is not doing. What he has done is to
respond to a set of observations and introspections rather than
construct a set of rational a priori definitions. In my view, the
basic observation requiring the emotion construct is that there
are certain classes of behavior consisting of relatively stereo-
typed patterns of activity that respond to life-challenging cir-
cumstances in an innately adaptive fashion, promoting both
individual and species survival. This notion received its paradig-
matic expression in the work of Cannon and was pithily epito-
mized in the belief that anger and fear, respectively, subserved
fight or flight.
Panksepp's assertion that emotions are genetically hard-
wired is not a "rational" assertion but simply a reasonable
inference derived from the observation that these species spe-
cific behaviors are both stereotyped and adaptive to threat. This
seems the major expository stylistic flaw of the paper. A set of
interesting propositions is presented in an assumptive form
rather than in a form that closely relates them to the body of
observation that they hope to explain. For instance, Panksepp
uses his definitional statements to distinguish such states as
"disgust" or "pleasure" from his construct of emotion. Since
definitions are a set of stipulations concerning the use of words,
he is certainly entitled to do this, but if he wishes to persuade us
to accept these definitions, he has to explain why making these
distinctions results in a superior heuristic approach. It does not
seem intuitively obvious to me that disgust does not activate
diverse classes of species-typical behaviors, nor that pleasure
does not generate unique behavioral manifestations. His Aristo-
telian belief that pleasure may be a state that accompanies
stimuli and tends to return organisms towards physiological
homeostasis contradicts his later proposal that "expectancy," a
basic emotion that produces nonhomeostatic exploratory ac-
tivities, is subjectively represented by joyful anticipation (which
I think most people would introspectively affirm to be a pleasur-
able state).
Panksepp's substantive hypotheses are that there are four
basic emotions: (1) fear, elicited by pain and resulting in flight
(with a poorly delineated functional neuroanatomy, although
Panksepp leans toward the importance of anterior lateral hypo-
thalamic sites); (2) expectancy (which Panksepp confusingly uses
as a term for positive expectation only), based on neural circuits
mediating electrical self-stimulation of the lateral hypo-
thalamus; (3) rage, primarily related to the ventro-medi-
alhypothalamus but substantially overlapping with the field for
expectancy self-stimulation; and finally (4) panic.
As Panksepp asserts, this is controversial and innovative. In
most systems, panic is considered to be simply the quantitative
extension of fear, whereas Panksepp affirms that it is a separate
basic motive system related to separation-induced distress.
In my own work with psychiatric patients who exhibit spon-
taneous panic attacks, we have noted a high incidence of exces-
sive childhood separation anxiety. We have also established the
peculiar fact that the so-called antianxiety agents do not block
these spontaneous panic attacks, but antidepressant agents do
(although the patients had not been depressed).
We also hypothesized that the spontaneous panic attack was
quite distinct from the ordinary fearful response. This phar-
Commentory/Panksepp: Psychobiology of emotions
macological approach to affective dissection was reaffirmed
when it was shown that the benzodiazepines were of some value
in lessening expectant anxiety but did not benefit the spon-
taneous panic attack, thus doubly affirming the dissociation of
these states.
Panksepp relates this basic emotional system to endogenous
opiates and, indeed, to the neurochemical dynamics of opiate
addiction and social dependence, and he has reported that
guinea pig separation-distress vocalization increases upon
naloxone injection. As we were in the midst of a study of the
provocation of panics in patients who have spontaneous panics
(panic disorder) by intravenous sodium lactate, we were able at
least partially to test this reasonable hypothesis by infusing four
patients with 20 mgs. of intravenous naloxone. Unfortunately
for the hypothesis, none of these patients demonstrated the
panic attacks typically manifested during sodium lactate
infusions.
To sum up, Ifindthe paper useful as a set of keen observations
and inferences, rather than a set of "rational" definitions. How-
ever, I think that is a positive event.
Generality and specifics in psychobiological
theory of emotions
Eric Klinger and Ernest D. Kemble
Department of Psychology, University of Minnesota, Morris, Minn. 56267
Panksepp's model represents an admirable synthesis of diverse
data and a laudable open-mindedness with regard to sources of
information. The bold sweep of this theory brings together
widely diverse physiological approaches, neurochemical studies
of transmitter substances, behavioral observations of nonhuman
species, and ideas based on human instrospection.
The overall theory is an impressive and quite plausible for-
mulation that is likely to stimulate exciting research. However,
it suffers from two principal difficulties: It makes too few specific
predictions and conceptualizes too loosely to provide unam-
biguous tests with nonhuman organisms; and it treats the avail-
able empirical literature about humans in such broad gener-
alities that it fails to come to grips with some of the important
issues this literature presents. Furthermore, one can argue
about the clinical predictions implicit in the theory. The re-
mainder of this commentary details these problems in order.
First, the means and criteria for unambiguously demonstrat-
ing the proposed executive circuits remain elusive. This is
particularly important in view of the large amount of recent data
showing high apparent specificity of functions that have been
quite resistant to consolidation into broadly general behavioral
categories. How, for example, can we go about demonstrating
the existence of the proposed expectancy circuit? Given the
diversity of species-typical behaviors elicited by brain stimula-
tion, what common features are we to expect from stimulation of
this circuit? If we are limited to increases in forward locomotion,
sniffing behavior, and the life, the model is not likely to be very
useful. Should it not be possible to predict the effects of stimula-
tion in species with novel foraging strategies (e.g., ambush
predators, insectivores, etc.)?
Since the theory presumes functional unity in the executive
circuits that underlie diverse species-typical behaviors, heavy
emphasis is placed on the plasticity of stimulus-bound behavior
demonstrated by Valenstein, Cox, and Kakolewski (1970) as
evidence for the presence of homogeneous psychobehavioral
tendencies. Such plasticity, however, has not always been
found, particularly with smaller electrodes. The theory fails to
specify the common features we may expect to find if these
nonplastic stimulation sites are, in fact, part of a more homoge-
neous system. It is, of course, possible that alternative species-
specific behaviors are nested within the broader "circuits," that
BEHAVIORAL AND BRAIN SCIENCES (1982)3 437
Commentary/Panksepp: Psychobiology of emotions
the selection among them normally depends on stimulus situa-
tions, but that stimulation with small electrodes rigidly elicits
one of these particular lower-order behaviors. This possibility
offers the theory a way out, but it needs to be specified.
The theory similarly needs to specify more clearly how one
might distinguish among different circuits, such as between the
proposed fear and panic circuits. The descriptions of these
circuits and their underlying mechanisms certainly seem plausi-
ble, but the range of test conditions on which this distinction is
made is rather small. The theory would benefit from specifying
test situations that would permit a fuller expression of these
underlying emotional states.
One avowed strength of Panksepp's model is in applying
features of human emotional experience to animal behavior.
However, the theory does not propose specific tests with non-
human species that show rather subtle patterns of emotional
expression or social organization, including several primate and
social carnivore species that have now been described in some
detail. Furthermore, a much larger body of truly comparative
data is needed to evaluate Panksepp's critical assumption of
phylogenetic continuity in the underlying executive functions.
There are a number of issues raised in human self-report data
that bear on Panksepp's theory in important ways. Nearly every
attempt to identify affective dimensions, whether interin-
dividual or intraindividual (Zevon & Tellegen, in press), has
found two or three dimensions that, between them, account for
an overwhelming proportion of variance. The most prominent
two have been variously labeled and interpreted, commonly as
"pleasantness" and "tension" or "activation." However they
might be labeled or interpreted, such dimensions correspond
poorly to the basic command circuits proposed by Panksepp.
The "pleasantness" dimension probably accords well affectively
with Panksepp's expectancy circuit but the "activation" dimen-
sion taps elements of all the other three command circuits.
Panksepp is no doubt right in deploring the interpretive
difficulties in dimensional approaches to emotion and in offering
neurophysiological anchorages as an aid to resolving these
questions. However, a psychobiological theory that takes self-
report data seriously must also deal with the existing data. One
possible implication of these data is that there may be higher-
order hedonic codes that transcend the proposed circuits. Alter-
natively, the "supercodes" may reflect the limitations of the
introspective perceptual process rather than the underlying
circuitry.
A different kind of problem arises out of Panksepp's treatment
of sorrow as a low-amplitude "panic" response (a social re-
sponse), but depression as a disorder of the expectancy circuit,
and both of these as separate from the rage circuit. The difficulty
resides in the relation of this treatment to the following findings:
(1) invigoration, anger, and some form of depression (disap-
pointment, sorrow, grief, etc.) follow the loss of an incentive,
whether social or nonsocial; (2) they follow as part of a usually
orderly sequence dubbed an "incentive-disengagement cycle";
(3) amygdalectomy eliminates both the invigoration and depres-
sion components; and (4) depression is strongly correlated with
irritability (see Barta, Kemble, & Klinger 1975; Klinger 1975;
1977; Klinger, Barta, & Kemble 1974). To parcel out this
seemingly coherent response sequence among three different
command circuits does violence to these findings. At the very
least, they call for an explanation within the framework of the
Panksepp theory.
The treatment of joy in this theory seems unsatisfactory.
Panksepp suggests that "joy may be the fulfillment of expectan-
cies," but also that the expectancy and rage systems reciprocally
inhibit each other. If joy fulfills expectancies, does this mean
reduced activity in the expectancy circuit? And if so, would this
not then disinhibit rage? On the other hand, if joy represents an
increase in activity of the expectancy system, this should in-
crease striving, which does not seem a likely consequence of
438 BEHAVIORAL AND BRAIN SCIENCES (1982)3
fulfilling an expectancy. If we eliminate this possibility, then
just what is joy within Panksepp's system, and where does it
come from? In the case ofjoy and some other affects, the system
seems in need of elaboration and further specification.
Finally, Panksepp's attempt to relate his model to psycho-
pathology in Figure 5, while provocative, is highly questionable
in a number of respects and in some details almost certainly
wrong. For instance, the classification of schizophrenia as aris-
ing from overactive expectancy or panic circuits rides roughshod
over the web of data regarding these disorders. Similarly, the
gathering consensus that psychopathy involves defective fear-
responding is not reflected in Figure 5, except in placing
sociopathy at the underactive end of the panic axis. Psychopathy
(or sociopathy) probably does also involve some kind of attach-
ment disorder, but the fear defect has been found with electric
shocks as well as social deprivation.
In summary then, the Panksepp model summarizes a great
deal of information and has the potential to generate some
exciting research. Its success or failure will depend on how well
the author can elaborate the model further to accommodate
existing behavioral and self-report data and to specify critical
tests and predict outcomes.
Psychobiology without psychosocial
significance
Richard S. Lazarus
Psychology Department, University of California, Berkeley, Calif. 94720
Those in the emotion business are bound to become curious on
seeing this ambitious title, "Toward a general psychobiological
theory of emotions." Yet, given their reductionist predilections,
psychobiological approaches usually have little to say of rele-
vance to those of us with cognitive and social science outlooks.
Cognitivists like myself make a few central assumptions about
emotion. First, we view emotions as the result of evaluative
cognitions (e.g., cognitive appraisals) about the significance of a
transaction for a person's well-being. That is, emotions reflect
the outcomes of encounters with the world as cognized by the
individual, outcomes that may be real, imagined, or anticipated.
Second, each emotional quality (e.g., fear, anger, guilt, sadness)
expresses and includes a somewhat different appraisal of an
outcome; the appraisal involving anger is different from that
involving fear, etc. Third, although primitive animals have
wired-in dispositions to make such evaluative perceptions of
specific environmental configurations, higher mammals gener-
ally learn to interpret the significance of what is happening. This
process is often highly symbolic, as when it is concerned with
definitions of self.
Imagine, therefore, the sudden rush of enthusiasm of a
dedicated cognitive theorist reading Panksepp's early state-
ments in defense of anthropomorphizing about emotions. These
statements suggest Panksepp's conviction that an understand-
ing of emotions cannot be based on reduction of human emo-
tions to the least common denominator of animal life, or to
lower- or midbrain circuits that organize phylogenetically uni-
versal, primitive emotions. The psychobiological and phy-
logenetic orientations traditionally decorticate the emotional
process, metaphorically speaking. Even neurophysiological re-
search as high up as the limbic system omits or understates the
cognitive processes through which the psychological and social
significances of transactions are construed. Reading Panksepp's
account, one's enthusiasm continues for a time with the brief
historical overview of taxonomies of emotion that goes back to
the Greeks and touches on "passions" such as joy, desire,
sorrow, gratitude, resentment, and the like. One thinks, as one
reads on, "Surely here is a psychobiological approach that

disdains reduction and addresses emotions as they are experi-
enced by people in the course of their daily lives."
Alas, it could not be. The bold anthropomorphic beginning is
lost as soon as Panksepp gets to neural circuits, and one wonders
why he thought it fruitful in the first place. We are given a
system based on four basic emotions or "command systems,"
upon which all emotions are presumably built. Panksepp thus
retreats to the time-honored reductionism of research that
never gets beyond the limbic system and is based mainly on
relatively primitive animals. As in the case of other phy-
logenetically oriented systems such as Plutchik's (1980), subtle
and complex human emotions reflecting culturally based mean-
ings and the "feeling rules" of the social structure are treated as
blends of the elemental emotions that are presumably shared by
all animals.
And what an odd and idiosyncratic lot these basic emotions
are on which the richness of human emotions must depend.
"Fear" and "rage" are, of course, traditional and ubiquitous in
theory. "Expectancy" is difficult for me to view as an emotion,
because of its highly cognitive character and its limited arousal
potential, comparable at best to the "orienting response." Final-
ly, it is not clear how "panic" differs from fear. In the typical
social scientist's position, panic depends on a person's sense that
escape routes are closing, and is distinguished from fear through
such a cognitive discrimination and its generation of frantic
escape behavior.
What about Panksepp's treatment of the rich panoply of
emotions in human experience? This central issue is treated so
briefly (one page) that it is mostly begged. His solution consists
of interactions or blends among emotion systems, but they seem
strange and difficult to fathom. For example, it is suggested that
jealousy arises from some "admixture of panic, rage, and expec-
tancy. " But we are not told how panic is involved in jealousy, or
the nature of the expectancy; nor is it obvious what rage has to do
with jealousy, and why rage and not anger. Resentment in
Panksepp's view may arise from "a blend of activities in the rage,
fear, and expectancy systems," a pattern of reasoning which also
escapes me and which is not spelled out in sufficient detail to
moderate my confusion.
I must say with regret that Panksepp offers little help in
bridging the chasm that seems to separate the psychobiological
and the psychosocial sides of emotions. I found tremendously
obscure what was said about physiological circuitry, and after
reading and rereading many of the arguments (for example, on
emotive circuits in "Emotive circuits of the brain: The theory", I
felt keenly the need of a translator. I can't be sure whether my
confusion stemmed from my own modest grasp of neurophysiol-
ogy and its jargon, or from the obscurity of the writing. If
psychophysiologists are able to appreciate the treatment or
resonate with it, all well and good, but I am certain that social
scientists will cry for help, or, more likely, will simply be turned
off.
To return, then, to the theme with which I began, what
cognitively oriented psychologists and other social scientists will
find missing from Panksepp's paper is their own prime concern
in emotion theory, namely, psychosocial significance or mean-
ing. Not only do emotions engender the mobilization for adapta-
tion, which requires reconciling external demands with internal
agendas, as well as the physiological mechanisms for such
mobilization, but they also express and reveal how people
construe what is happening to them with respect to their
individual well-being. If there is any truth in such a view, then
Panksepp's efforts to offer a general psychobiological theory of
emotions fall fatally short, because they completely omit the
psychological and social significance of transactions between
people and their environments and the role of cognitive activity
in sensing this significance. Despite initial signals to the con-
trary, these efforts preserve the unfortunate dichotomy be-
tween psychobiological and social science approaches to emo-
Commentory/Panksepp: Psychobiology of emotions
tion when what is badly needed is some way to bring them
together.
Introspection and cultural knowledge
systems
Catherine Lutz
Department of Anthropology, State University of New York at Binghamton,
Binghamton, N.Y. 13901
Panksepp's call to "approach the brain with realistic experiential
concepts" represents a potential meeting ground for social and
biological research on emotions. His proposal that introspection
be used as a technique for garnering information on human
emotions can, as he notes, balance previous emphases on facial
expression and physiological correlates of emotions. The issue
concerning which of the "experiential concepts" emerging from
introspection should be accepted as "realistic" ones is more
complex, however, than the author indicates. Introspection has
in fact already been in extensive (though rarely acknowledged)
use in the study of emotions as evidenced, for example, in the
ubiquitous reliance on the concepts of emotion as represented
in the English language. Whether our everyday language and
intuitive sense of the nature of emotional experience should
serve as the bases for inferences about brain organization can be
decided only on the basis of data, which Panksepp does not cite.
The nature of the relation between brain activity and verbal
accounts of emotion concepts will not be discovered simply by
searching out those who are judged to be "thoughtful" observers
of their own emotional states. Anthropologists and psychologists
have for years investigated the question of the relationship
between language and cognition, verbal report and internal
experience. They have accumulated several kinds of evidence
that point to the diversity and complex origins of introspective
accounts of emotions and other phenomena. Each of the follow-
ing three areas of research indicates that individual and cultural
knowledge systems concerning internal experience are highly
organized and pervasive, and have profound effects on the
communication and perhaps the nature of that experience.
There has been a long-standing interest in and reliance on
subjects' descriptions of the cognitive processing underlying
their performance on experimental tasks. The relation between
these accounts and actual introspective accessing of those pro-
cesses is problematic, however. Nisbett and Wilson (1977)
review experiments with American subjects that use verbal
reports of cognitive processes and conclude that such reports are
based on the subjects' previously held notions about the relation
between events and responses, not on actual introspection.
D'Andrade (1974) has shown that individuals' reports of their
long-term memory for the observed behavior of others are not
related to that behavior as coded by an observer, but are
correlated with the judged semantic similarity of the behavioral
terms involved. Personal knowledge structures about what
behaviors, responses, and events should go together affect
verbal reports.
A second areas of psychological research has great potential
importance for the understanding of introspective accounts of
emotion. Metacognitive studies are concerned with the way in
which people introspect about, classify, and monitor their men-
tal activity (Brown 1978; Brown & Barclay 1976). Investigators
have also examined, in our own society, the development in
children of knowledge concerning the nature of distinct mental
processes such as reasoning, memory, and imagery (Wellman,
in press). The ontogenetic research strategy that Panksepp sees
as necessary for examining the interaction between cognitive
appraisal and emotive command circuits is also a key to under-
standing the origins of metacognitive skills such as introspection
BEHAVIORAL AND BRAIN SCIENCES (1982)3 439
Commentary/Panksepp: Psychobiology of emotions
and of the concepts and theories of mind and emotion held by
adults.
Third, cultural knowledge structures concerning the mind
and emotion have been documented by anthropologists in the
field of ethnopsychology. Conceptions of the nature of emotion,
its role in the organization of the person and in social interaction,
and typologies of the emotions have been found to vary (Geertz
1959; Hallowell 1955; Levy 1973; Lutz 1981; 1982). The fact that
psychological knowledge systems are culturally variable means
that introspection on brain activity will be mediated by the
categories provided by language and culture. Although the
documentation of variation in emotion-word classification sys-
tems has barely begun, the description of theories and concepts
of emotion held in various societies can provide the basis for
understanding the role of culturally provided knowledge sys-
tems in a person's understanding and experience of emotional
processes.
All this is not to say that introspection is either necessarily
"wrong" about mental activity, or that the cultural mediation of
introspection results in brain activity being hopelessly inaccessi-
ble to the conscious mind. Introspection is simultaneously about
knowledge and process, as these two factors are inseparable.
The suggestion being posed here is that a combination of
developmental and cross-cultural studies can aid in sorting
through the processes which result in account-giving, including
both those which may involve the use of cultural or personal
knowledge structures and those which may not. The brain
activity, which forms at least part of the basis for the initial
construction of knowledge structures about emotions, may yet
be retrieved via the comparison of introspective accounts of
diverse individuals.
Panskepp's theory has the distinct advantage of allowing for
this kind of comparison by going beyond the common focus on
the physiological symptoms of emotions. Emotion, in his
scheme, is characterized by the recognition of diverse environ-
mental events as calling for one class of responses. Introspective
accounts of emotional states and their environmental correlates
may both be more readily comparable. An example can illus-
trate the usefulness of a combined comparative and introspec-
tive perspective for evaluating scientific theories of emotion.
This particular example lends some support to Panksepp's
exclusion of "surprise" and "disgust" from the group of major
emotive circuits. Accounts given by residents of the island of
Ifaluk, in the Western Pacific, of the nature and taxonomy of
emotions explicitly depict "disgust" (the parallel term for which,
in the Ifaluk language, refers exclusively to the response to fetid
or spoiled objects) as being different in kind from other words
descriptive of temporary internal states (Lutz 1982). Also ex-
cluded from this domain is the term for physical startle, which is
seen as a simple behavioral description. There is no separate
term like "surprise" that exclusively describes the reaction to
sudden events. Panksepp's other claims for the centrality of
certain types of emotional circuits should also be compared with
the introspective accounts generated by individuals of other
theoretical and cultural persuasions.
Concerning the alleged four basic emotions
William Lyons
Department of Moral Philosophy, University of Glasgow, Glasgow G12
800, Scotland
1. The structure of Panksepp's psychoblologlcal argu-
ment. Panksepp's basic argument for choosing expectancy, fear,
rage, and panic as the four fundamental emotions, though not as
explicit as it might be, seems to be the following:
i. In the process of evolution, emotions, or at least basic
emotions, have developed as systems for organizing animal and
440 BEHAVIORAL AND BRAIN SCIENCES (1982)3
human behavior in such a way as to cope with emergencies; that
is, emotions are designed to respond unconditionally to stimuli
arising from major life-challenging circumstances and so to
organize behavior . . . which has proved adaptive in just such
circumstances.
ii. There are four clearly delineated and distinct channels -
or four distinct behavioral control systems or psychobehavioral
tendencies - which this process of coping with emergencies can
employ, namely expectancy, fear, rage, and panic.
iii. Therefore, expectancy, fear, rage, and panic are the four
basic emotions.
2. Have emotions been developed by evolution as responses
to life-challenging circumstances? Or, to put the question an-
other way, is the first premise of Panksepp's psychobiological
argument true?
There is a large literature, chiefly in psychology, which claims
that emotions are coping processes or processes developed by
evolution such that they organize behavior to cope with
emergencies. The most famous exponent of this view is probably
Alexander Shand (1914, especially bk. 2, chap. 1), but there
have been many others (more recently, e.g., Arnold & Gasson
1968; Leeper 1948; Young 1961.) Unfortunately, there seems to
be an equal number who claim the opposite, that emotions are
disorganising and disruptive (e.g., Munn 1961; Pradines 1958),
and the truth seems to be that there are undeniable facts to
support both views, even when the alleged basic emotions are
considered. Panic, for example, might speed up responses and
facilitate mass escape by communicating itself to others nearby,
but it might as likely lead to dangerous and unnecessary and
blind flight (Lorenz 1970, vol. 1, pp. 149, 166, 225, 309). Fear
may lead to the frightened animal's running away from the
predator faster than it ever ran before, but it may also lead to
catatonic freezing - and so on. Thus E. J. Murray writes that
"the evidence so far suggests that both emotion and motivation
can positively organize and facilitate behavior, at certain times,
but they may also disorganize and disrupt behavior" (1964, p.
64; see also Lyons 1980, pp. 41-44, 188-92).
Furthermore, the "emotion as coping with emergency" thesis
has always seemed strained once one gets outside the speeded-
up, adrenalin-pumping variety of emotions, and it has seemed
equally strained to limit basic emotions to the class of speeded-
up ones. At any rate, it would be hard to make out a convincing
case that boredom and contentment in animals help cope with
emergencies (and, in humans, that sadness, despondency, en-
nui, depression, shame, or embarrassment are mechanisms for
coping with emergencies). These emotions do not seem to be
on-the-qui-vive emotions; they (or some of them) are backward-
looking responses to the past.
But even within the list of "best candidates" from the class of
speeded-up, adrenalin-pumping emotions, it should be obvious
that such emotions as fear and rage can be positively disadvan-
tageous where calmness and quietness are required. Although
fear may lead an animal to hide, it will also occasion the animal's
loud breathing, increased odor, shaking and quivering, and,
maybe on that account, lead to the predator's discovering its
hiding place. An enraged animal or person will very often fight
less well than a cooler calmer individual. Rage - as well as fear
and panic - can very easily lead to misperceptions and bad
decisions. At the human level, an enraged boxer or street fighter
may be easy game for a cool and calculating opponent.
At a more fundamental level again, why should we assume,
piously, that the process of evolution throws up only useful
behavioral tendencies or patterns of behavior? It may be claim-
ed that, in the different conditions of long ago, the thesis was
true; namely, that fear, panic, and rage were always or most
often useful, though, in both animals and humans, it may now be
the case that these emotions are as much a hindrance as a help.
But will this move save the thesis? For how do we know that long
ago they were or even must have been useful? Is the answer,

"They would not have survived if they were not useful," a good
answer? To simplify the picture, surviving species may have,
say, behavioral tendencies X, Y, and Z. The species may have
survived because of X or Y, or both, but survived with, but not
because of, tendency Z. Z may have no clear survival value,
positive or negative; it may have been, so to speak, carried along
on the backs of the survival-valued tendencies X and Y. Emo-
tions, or some of them (including some of the basic ones) may be
Zs!
3. Are there four distinct and basic "psychobehavloral sys-
tems"? Or, to put this question also another way, is the second
premise of Panksepp's psychobiological argument true?
My first query is whether "foraging-expectancy" is an emo-
tion rather than, say, just an appetitive inclination derived from
"homoeostatic imbalances in the body and their respective
environmental incentives," that is, an appetitive inclination
without any strong somatic or visceral speeding-up or slowing-
down (we must not forget the "calm emotions," Lyons 1980, pp.
9, 30, 116-17). In humans at any rate - and Panksepp urges us to
take such introspective-cum-retrospective evidence seriously -
hunger and thirst do not characteristically give rise to emotions
of expectancy or exploration. When the search for food or drink
is frustrated or proving fruitless, then emotions may begin to
arise, such as, initially, frustration and irritation, then fear and
perhaps panic, and finally despair. Besides, it seems a farfetched
theoretical claim to suggest an emotion is needed to induce "an
animal to move from where it is to where it should be in order to
consume the substances needed for survival." The wants and
desires - or, in the preferred jargon, "incentives" leading to
"motor arousal" - arising from the body's needs in the states of
hunger and thirst seem inducement enough.
Panksepp is more plausible when he later glosses "forag-
ing-expectancy" as "hope, desire - joyful anticipation." But
such a gloss reveals a dilemma. If the basic emotion is left
labeled simply as "foraging-expectancy," and coupled with the
basic behavior of sniffing, it does not seem to be an emotion but a
homeostatically based drive (or instinct or, in general, impulse).
Relabel this basic emotion "hope, desire — joyful anticipation"
and it becomes an emotion all right; it becomes a nonbasic
(because nonsimple) congeries of emotions as suitable to the
anticipation of non-survival-related objects or events as of those
that have something to do with survival.
Panksepp's descriptions and analysis of the function of his
three strongest candidates for basic emotions - fear, rage, and
panic - also seem arbitrarily truncated and unfair to the facts. In
the case of fear, Panksepp proposes that we look to the "hard-
wired circuit" in the hypothalamus from which "flight and
unconditional escape behaviors can be elicited." But Shand,
who long ago attempted a program similar to Panksepp's real-
ized that he had to admit that, even in animals, fear gives rise to
a daunting variety of behavior; namely, behavior expressive of
the "instincts of flight, of concealment, of silence, of clinging
and keeping close to something, of shrinking or starting back, of
immobility or simulation of death, of crying for help " (Shand
1914, bk. 2, chap. 2, p. 207 - and he had read the ethology of his
day, notably Brehms and Hudson). Or is Panksepp's phrase
"unconditional escape behaviors" a catchall phrase that can
accommodate anything? What seems fair to say is that few
ethologists would any longer accept a simple "fight and flight"
view of anger and fear in animals, and so to base a strategy for
discovering the emotion circuits in the brain upon such a view
must produce a very precarious methodology. I do not think that
there is convincing evidence that one can differentiate the
emotions by reference to neatly packaged behavior or, a for-
tiori, to the much more elusive "behavior tendencies." The
behavior that results from a single "behavior tendency" may be
multifarious, and depend very much on the context, as Shand
elaborated. Fear is always fear of something in particular, and -
to take an example relevant only to humans - the fear of getting
fat will give rise to behavior very different from that arising from
Commenton//Panksepp: Psychobiology of emotions
the fear of losing weight! (Re emotions and behavior and behav-
ior tendencies, see Lyons, 1980, pp. 17-25, 41-44.)
I have much to say of a similar nature about Panksepp's
attempts to "home in on" supposedly typical and unique rage-
behavior and panic-behavior, but space and time will allow me
only a few more brief comments. As regards rage, is it plausible
to suggest that one of its "major adaptive function[s] . . . is to
invigorate behavior when activity in the expectancy circuit
diminishes rapidly"? Do animals in the wild fly into a rage when
their "foraging-expectancy" explorations "diminish" for some
reason or another, or should I say when their "hopes and joyful
anticipations" wane? I would need a lot of ethological evidence
to be convinced of this.
Finally, is it plausible to maintain that the "major adaptive
function" of panic is "to sustain social cohesion" when etholo-
gists such as Lorenz (1970) and Lorenz and Leyhausen (1973)
write of "wild" and "blind" and "baseless" panic, of misdirected
and "extremely dangerous panic," and, finally, of "disastrous
panics" in animals? Thorpe (1963) refers to "fatal panic" in
animals, though, admittedly, the evidence was in connection
with animals in zoos. Could "social cohesion" coexist with such
blind, wild, disastrous flight?
Though I have been critical of Panksepp's central psycho-
biological argument, I nevertheless enjoyed the paper. It is
good to see a return to attempts to define and model the
emotions.
Psychobiology needs cognitive psychology
Adam Morton
Department of Philosophy, University of Bristol, Bristol BS8 1RJ, England
Panksepp's real conclusion could be stated somewhat as follows:
There are four interacting but separable limbic neural complex-
es in mammals, each of which is associated with a definite range
of species-specific behavior, which have some pretty basic, but
still not entirely clarified, relation to emotional states of all
mammals, including humans. To put it this way would make it
all seem less interesting and original than it is, though, and so he
formulates a tentative stronger claim, which is more likely to
make the reader read on. This version runs: Distinct limbic
circuits in all mammals can be identified as expectancy, rage,
fear, and panic; and when humans experience emotions, their
experiences (or a large component of them) can be compounded
out of these elements. My comments are addressed to the
relation between the stronger and the weaker version of Pank-
sepp's thesis.
People not only experience emotions, they also have moods
and attitudes, and exhibit character and personality. All these
are related, though it is not clear what the relations are. Some-
times it seems as if Panksepp is talking about this whole class of
states and attributes, and sometimes he seems to be concerned
only with a subclass. That subclass would be not that of the
emotions proper but that of moods and feelings. Attribute
number 4 of his command circuits ("A definition of emotions")
is meant to make the states in question relatively independent of
the environmental factors that occasion them; for the descrip-
tion of jealousy as an admixture of panic, rage, and expectancy,
or resentment as a mixture of rage, fear, and expectancy to have
any plausibility, we must abstract away the typical cognitive
content of jealousy or resentment and concentrate only on the
mood or feeling; and the definitions of sorrow as a diminutive
form of panic and anxiety as a diminutive form of fear seem to
involve a similar decision to ignore the nonfelt aspect of a state.
But, on the other hand, the allusions to the attempts of De-
scartes and others at a grand classification of states of mind, and
Panksepp's speculative extension of his theory to personality
disorders ("Diseases of emotive systems") suggest that he wants
to include all affective states in his theory.
BEHAVIORAL AND BRAIN SCIENCES (1982)3 441
Commentary/Panksepp: Psychobiology of emotions
The main importance of the emotion/feeling contrast con-
cerns cognitive content. Emotions typically have the causes and
effects that they do, in part because of their associations with
complex thoughts. That is why we express them with "that"
clauses. James is afraid that his house may burn down. James's
fear (unlike his anxiety) cannot be identified without reference
to its object, and cannot be accounted for except by reference to
whatever it is (surely essentially neocortical) that allows him to
have such thoughts as "my house may be on fire." Panksepp's
theory tacitly acknowledges this. Fear, for example, is associ-
ated with environmental stimuli of "threat of destruction" (Fig-
ure 1). But a stimulus isn't simply of "threat of destruction "
(contrast it with the stimulus of "bodily irritation" cited for rage);
the stimulus must be conceptualized as representing such a
threat, and unless the mechanisms for such a conceptualization
are included in it, a neural circuit must produce something
other than fear. A similar point could be made for rage, using the
data Panksepp cites on the expression of rage in monkeys: the
effects of rage are not simple behavioral routines, but depend on
cognitively complex representations of the animals' social
group.
I would not conclude from this that Panksepp is confused.
Rather, the moral seems to be that the English terms he has
chosen for the sake of exposition are not quite what he needs. He
does not want to define his subject as feelings, because he wants
to exclude many feelings from primary consideration ("A defini-
tion of emotions") and because his ultimate aim is to cover much
more than just feeling. But the cognitive associations of "emo-
tion" and of English emotional terms aren't wanted either.
Moreover, the terms "expectancy, ' "fear," "rage,' and "panic"
are not quite the right labels for what he has in mind. "Expec-
tancy" is particularly ill-fitting; on reading what Panksepp has
to say about it, one does acquire a feel for the state in question,
but it has no simple relation to expecting (or hoping) that
something will happen.
What should Panksepp say, then? Just that there seem to be
these four neural circuits, and that he hypothesizes that there
are four corresponding psychological states, which are felt
throughout human affective life and are related to the following
states and behaviors in the following ways. . . . But then the
dots have to be filled in. And that's a problem, for one needs a
theory of how the four states affect human behavior and the
affective states we report with our rich, complicated vocabulary.
A purely neuropsychological theory won't do the job, and
neither will pure philosophical analysis. One is too hard and the
other too soft. Somewhere in the middle there will have to be a
cognitive psychology of the emotions, which connects up at one
end with neural and behavioral data and at the other with what
we humanly say and feel. It would be nice if one could get along
just by mixing neuropsychology and philosophical analysis, but
the points at which Panksepp's promising thesis bog down
provide some reason for believing that this combination is not
going to be enough.
Only four command systems for all
emotions?
Robert Plutchik
Department of Psychiatry, Albert Einstein College of Medicine, Bronx, N. Y.
10461
The idea that emotions arise from hard-wired neural circuits and
that they facilitate adaptive responses stems from the Darwinian
evolutionary, ethological tradition and has been described by
many others (e.g., Plutchik 1962; 1970; 1980). What is new and
most controversial about Panksepp's views is his claim that
human introspection may provide direct information concern-
ing operations of emotive circuits. Also debatable is the view
442 BEHAVIORAL AND BRAIN SCIENCES (1982)3
that there are only four emotion control circuits, those which
elaborate states of expectancy, rage, fear, and panic.
There are a number of arguments that can be presented
against the last two assumptions. For one, although introspec-
tions may sometimes be useful in revealing an individual's
emotional states, they are far from infallible guides even to the
subjective aspects of emotion. For example, the psychoanalysts
have made us aware of the fact that subjective reports of affects
cannot always be accepted at face value. Not only are some
emotions totally repressed, and thus unavailable to introspec-
tion, but others are frequently modified or distorted as a result
of partial repressions. Even academic psychologists have ac-
cepted the idea that ego defense mechanisms can transform or
modify the nature of introspective reports (Plutchik, Keller-
man, & Conte 1979).
Other problems with introspection are the following: (a)
Repression may create false negatives; that is, an observer may
erroneously assume that no emotion exists because none has
been reported, (b) Verbal reports of emotions may be deliberate
attempts to deceive another person (as is often the case in so-
called deception experiments), (c) Reports of emotions depend
on individuals' particular conditioning history as well as their
facility with words, (d) Reports of inner emotional states are
retrospective and depend on memory. Remembered events are
notoriously subject to distortions, such as leveling and sharpen-
ing, (e) The inherent ambiguity of language creates the problem
of the "true" meaning of emotion terms. The importance of
context in determining meaning implies that the same verbal
report of an emotion may have a different referent in another
setting, (f) Emotions are rarely if ever experienced in a pure
state. More typically, any given situation creates mixed emo-
tions, which are difficult to describe in any simple or unequivo-
cal way.
For these and other reasons one can conclude that an intro-
spective report of an inner emotional state is only a rough
approximation to whatever that state is. It hardly seems likely
that such crude reflections of emotional states are a royal road to
the nature of underlying brain mechanisms associated with
emotions. Emotions are complex chains of events whose proper-
ties can be inferred only through the use of a wide variety of
sources of evidence.
Incidentally, from an historical point of view, the use of
introspection alone has led to the idea that there are different
numbers of basic emotions. Spinoza proposed three, Descartes,
six, Hobbes, seven, McDougall, twelve, Murray, sixteen, and
others have proposed various intermediate numbers. Introspec-
tion alone is not an adequate guide to the number of basic
emotions we possess, just as factor analysis alone, or physiology
alone, or brain anatomy alone, or stimulation studies alone are
insufficient.
My second set of objections to Panksepp's theory concerns his
assumption that there are only four basic emotions as defined by
transhypothalamic executive circuits. Panksepp prefers to be-
lieve that surprise, disgust, and joy are not basic emotions,
despite the fact that they have clear behavioral referents in
animals as well as humans, and may be controlled by limbic
circuits. He claims that these emotions are "largely time-locked
to the precipitating stimuli, with no sustained regenerative
feedback in the underlying brain systems." This statement
ignores the fact that all these emotions can occur in various kinds
of sustained derivative forms (e.g., revulsion or contempt as
derivatives of disgust, and cheerfulness and friendliness as
derivatives of joy). Contrary to Panksepp's statement, these
states do have unique behavioral manifestations (see, for exam-
ple, Bull 1951; Darwin 1872; MacLean 1975) and do operate
through discrete neurochemical dopamine and endorphin sys-
tems of the brain. Electrical stimulation of the brain produces
not only withdrawal reactions, destruction reactions, and food
intake reactions, but sexual reactions, "disgust" reactions, star-
tle reactions, and watchfulness reactions (Plutchik, McFarland,

& Robinson 1966). The only kind of reactions not produced by
direct brain stimulation are loneliness and grief reactions. It is
highly unlikely that "panic" reactions are defined simul-
taneously by distress vocalizations (how does Panksepp know
the vocalizations reflect "distress"?) and by "explosive behavior,
during which the animal exhibits violent leaping about the test
chamber."
One may also object to the label of "panic" for the emotional
state described by Panksepp as "social loss", or "loneliness." In
my own research, high-intensity fear is typically defined by
subjects as panic, while high-intensity sadness (social loss) is
defined as grief.
The form of the diagrams presented (e.g., target article,
Figure 2) also has certain implications. Expectancy is placed
opposite rage, and fear is placed opposite panic. In contrast, a
number of empirical, multidimensional scaling studies (Plutchik
1980) indicate that grief is opposite joy, fear is opposite anger,
expectancy is opposite surprise, and disgust is opposite accep-
tance. (Incidentally, the concept of expectancy as a basic emo-
tion, along with an evolutionary rationale, was first presented in
detail in my 1962 book.) It is not the case that "rage and
expectancy circuits should strongly inhibit each other." From
both an introspective and observational point of view, sex and
aggression are often found together, and factor analytic studies
have demonstrated that assertive children are often the most
sociable as well.
The idea that basic emotions may be related to psychiatric
disorders is a good one (Figure 5), but it has already been
developed in detail by Schaefer and Plutchik (1966) and by
Plutchik and Platman (1977).
In conclusion, the general ideas presented by Panksepp are
important, but the specific details of the model are debatable,
and his assumptions about the role of introspection probably
wrong. Despite his claims for the importance of introspection,
he presents no insights into brain mechanisms based on this
source of information alone.
On the complexity of emotion
Joseph R. Royce
Center for Advanced Study in Theoretical Psychology, University of
Alberta, Edmonton, Alberta, Canada T6G 2E9
The study of emotion has been refractory to scientific advance,
particularly to theoretical advance. The major deterrent is
complexity. The crux of this complexity is that emotion simul-
taneously involves subjective thoughts and feelings, physiologi-
cal reactions, evolutionary adaptive significance, behavior, and
individual differences. Furthermore, full understanding of a
given emotion requires an alignment at all five levels - and the
alignment includes temporal sequence in addition to consisten-
cy of each of thefivefacets. An inconsistency in the alignment of
any one of thefivefacets would constitute a failure of the theory.
For example, a high rather than a low level of physiological
arousal is the appropriate correlate for the emotive state of anger
or rage.
The problem of complexity in this domain is exacerbated
further by the fact that there is ambiguity as to what the
emotional domain includes. Plutchik (1962), for example, lists
some 21 definitions, and Strongman (1978) concludes that
"emotion defies definition."
Given this degree of complexity and ambiguity, it is not
surprising that the typical theory tends to cover only one or two
of the five facets. Furthermore, since there is no adequate
definition, it is also not surprising to find that the typical theory
fails to cover the full range of possible emotive states.
Panksepp's theory is no exception. Although his approach is
particularly strong in its specification of the biobehavioral and
evolutionary aspects of the problem, it says nothing about
Commentary'/Panksepp: Psychobiology of emotions
individual differences and it is weak in its treatment of the
subjective aspects. Unfortunately, instead of bringing us up to
date on the latter topic, he squanders the available space on a
review of pretwentieth-century philosophical treatments of this
question. While this may be the most difficult of thefivefacets, I
cannot see what Panksepp is trying to achieve by quoting from
seventeenth-century authors such as Willis and Descartes. We
can surely do better than to make lists of emotions, and to talk
about "passions"! But Panksepp (see his Table 1) actually decid-
es to take one of these early authors seriously, namely Cogan
(1802). The point is that contemporary treatments of the subjec-
tive aspect of the problem are of much greater scientific value
than he would have us believe. In fact, they constitute the
widely accepted "cognitive basis of emotion" whose claims are
focused on two related ideas: cognitions as cue-attributions, and
cognitive appraisals as causes of emotional states. Thefirstview,
which is primarily due to the research of Schachter (1964), sees
the detection of the combined internal physiological cues and
external situational cues as the basis for the attribution of an
emotional state. The second view, which is primarily due to the
research of M. B. Arnold (1970) and Lazarus (1966), is ex-
emplified by Lazarus, Averill, and Opton (1970, p. 219) when
they say that "the situation is evaluated as relevant, threatening,
frustrating, . . . etc. . . . This appraisal also includes an eval-
uation of the options for coping and their potential conse-
quences. Cognitive processes thus create the emotional re-
sponse out of the organism-environmental transaction and
shape it into anger, fear, grief, etc." Thus, according to this
view, a cognitive interpretation is not a passive correlate, it is a
cause of the emotive state.
Although the exact role of cognitive processing in emotion is
not clear, it is highly probable that cognition is involved, in at
least the less violent emotive states. This means that the neural
underpinnings of emotion probably include cortical areas of the
brain in addition to Panksepp's four transdiencephalic emotive
circuits. There is also a significant body of evidence (e.g., see
Eysenck 1967; Lindsley 1950; Royce & Diamond 1980) implicat-
ing the reticular activating system, particularly in the case of the
less disruptive emotions.
Of course, these are claims rather than solidly established
facts. However, they are no less established than Panksepp's
biobehavioral emotive circuits, the neural details of which are
not known and, therefore, "can only be surmised." Further-
more, although Panksepp alludes to mixtures of his four emotive
command systems combining with learning in order to account
for the full range of emotive states, this is a very large promissory
note on a problem of central importance to a general theory of
emotions.
Although one is hesitant to be critical of attempts to under-
stand phenomena as complex as emotion, my remarks are
offered in the same heuristic spirit that comes through in
Pankepp's paper. Thus, although I have pointed to some weak
spots, these criticisms are minor when placed in the context of
what Panksepp is reaching for. In short, I am in agreement with
Panksepp's central argument, namely, that his four emotive
neural circuits are a good basis form which to move "toward a
general psychobiological theory of emotions."
On the nature of specific hard-wired brain
circuits
Allan Siegel
Department of Neurosciences, New Jersey Medical School, Newark, N.J.
07103
Panksepp has made a potentially significant contribution to our
research strategies by reminding us of what our priorities should
be in our attempts to discern the brain mechanisms underlying
the various forms of emotional behavior. In particular, he has
BEHAVIORAL AND BRAIN SCIENCES (1982)3 443
Commentary/Panksepp: Psychobiology of emotions
argued effectively that "emotional" constructs achieve signifi-
cance when they are linked to specific circuitry in the brain. The
value of this approach rests in the fact that Panksepp would have
us attend to the details of the neuroanatomical substrates under-
lying the emotional expression of behavior. By pursuing this
course we can then design our experiments specifically to test a
variety of hypotheses about such circuitry. In particular, if we
focus on the anatomical pathways relevant to a given behavioral
process, the data obtained from such experiments may provide
meaningful clues to the sites of interaction at which sensorimo-
tor integration as well as forebrain modulation are most likely to
occur.
Panksepp appears to base his conclusions concerning the
nature of thefixedcircuitry primarily on data from experiments
on rats. It is on this issue that I wish to comment. Panksepp
states that current evidence supports the notion of a "non-
specificity" explanation for a variety of stimulus-bound appe-
titive behaviors, such as feeding, drinking, gnawing, hoarding,
pup-retrieving, predatory aggression, and copulation. His con-
clusions are based on the following lines of evidence: (1) that
through experience one appetitive behavior (elicited by electri-
cal stimulation) can be changed to another (Valenstein, Cox, &
Kakolewski 1970); (2) "that stimulus-bound appetitive behav-
iors do not change when an electrode is passed through the
relevant lateral hypothalamic field (Wise 1971)"; (3) that "The
type of behavior elicited from an electrode does not change
when the tissue under the electrode is damaged (Bachus &
Valenstein 1979)"; and (4) that "the temperament of the test
animal may be more important in determining the stim-
ulus-bound appetitive behavior observed than the exact loca-
tion of the stimulating electrode in the active zone within the
dorsolateral hypothalamus (Panksepp 1971a)."
Over the past 15 years our own observations on the cat, as well
as data reported from the laboratory of the late Dr. John Flynn,
clearly support a different point of view, which corresponds to
what Panksepp calls a "specificity" hypothesis concerning the
nature of the anatomical circuitry underlying various behavioral
processes. This view is that for each behavioral process, there
exists a distinct set of anatomical pathways, which become
activated during the behavioral sequence in question. We have
considered the possibility of trying to replicate the Valenstein
experiment in the cat but have decided against it, because the
length of hypothalamic stimulation per trial, the intertrial inter-
val, and length of experimental session would, in my opinion,
represent a regimen too stressful for application to the cat. Thus,
in order to avoid any discomfort to the animals, data bearing on
this central issue of specificity versus nonspecificity have been
obtained during the course of other experiments performed in
our laboratories. Most notably, the related question of whether
the same site in the hypothalamus could elicit, upon stimula-
tion, two different behavioral responses, was discussed by
Flynn, Vanegas, Foote, and Edwards (1970). They described an
experiment in which horsemeat and an anesthetized rat were
alternately made available to a cat. Following stimulation of the
hypothalamus, most cats displayed attack behavior only, while
both eating and attack were present in a small percentage of the
remaining animals. When both horsemeat and an anesthetized
rat were present in the cage (the horsemeat having been placed
between the cat and the rat), electrical stimulation of the
hypothalamus reliably produced attack behavior in a similarly
high percentage of the animals. Other unpublished observa-
tions revealed that even 24-48 hour, food-deprived cats will
cease eating to attack a rat following electrical stimulation, and
stimulation in the continued presence of horsemeat alone did
not later cause the cat to select food instead of the rat. These
findings clearly point to the fact that most sites in the hypo-
thalamus of the cat are ones from which quiet attack, but not
eating, can be evoked.
Concerning the statement that a "stimulus-bound" appe-
titive behavior does not change when an electrode is passed
444 BEHAVIORAL AND BRAIN SCIENCES (1982)3
through the relevant hypothalamic field in a given cat, we have
observed that the region from which quiet biting attack can be
obtained is quite limited (approximately 0.5-1.0 mm. across),
and that this observation supports the specificity hypothesis
(Fuchs, Dalsass, Siegel, & Siegel 1981). In addition, in studies
dealing with affective display behavior, we have routinely found
that dorsomedial hypothalamic stimulation will evoke flight
behavior and that, as the electrode is passed ventrally for a short
distance into the region of the ventromedial nucleus, the behav-
ior evoked changes to affective display. Thus, in this instance,
the passage of the electrode through the relevant hypothalamic
field does, in fact, produce a change in the nature of the behavior
evoked. Our explanation is simply that stimulation at more
dorsal sites activates a selective set of pathways that mediate
flight responses (Fuchs, Siegel, & Edinger 1981), while stimula-
tion at the more ventral sites activates a somewhat different
group of fibers (Fuchs, Siegel, & Edinger 1980).
Concerning the statement that the behavior is not changed
after a lesion at the tip of the electrode, and that only its
threshold is elevated: this is not, in itself, an argument for a
nonspecificity hypothesis. Such an argument could just as well
be used to support a specificity hypothesis. When a lesion is
placed at the tip of the electrode in the cat, the attack response
may be totally eliminated (Chi, Bandler, & Flynn 1976; Chi &
Flynn 1971). In any event, such an observation has been in-
terpreted by these authors to mean simply that the pathways
mediating attack behavior are highly discrete and were
damaged by the lesion.
The final statement made by Panksepp - that the tempera-
ment of the animal may be more important as a determinant of
the stimulus-bound behavior than the exact placement of the
electrode — is one for which I have not seen support in the
behavior of th,e cat. In fact, some of our cats displaying the most
ferocious attack responses upon stimulation turn out to be
markedly timid and placid in the absence of electrical stimula-
tion. It seems that, in the cat, the most relevant variable is the
placement of the electrode rather than the "personality" of the
animal. Furthermore, our anatomical studies dealing with quiet
biting attack behavior have demonstrated that the pathways
mediating this response are quite specific. Our anatomical
tracing procedures reveal a very consistent pattern of distribu-
tion offibersamong all cats displaying this behavior. In contrast,
when a different response such as affective display or flight
behavior is obtained from a neighboring site, the pathways that
we have traced, and which are associated with these responses,
are clearly different.
Accordingly, on the basis of our observations in the cat, I have
drawn the following conclusions regarding the nature of hard-
wired circuitry in the cat: (1) The sites and pathways underlying
each behavioral response are clearly distinct and discrete, with
only some degree of overlap (or commonality offiberpathways).
(2) The constellation of pathways mediating one response pat-
tern is not the same as that associated with other hypothala-
mically elicited responses (i.e., "specificity" hypothesis). (3)
The "nonspecificity" hypothesis generated from data in the rat
cannot be generalized to the cat. The strategy of utilizing
anatomical tracing procedures is of heuristic value in identifying
the neural substrates underlying various behavioral response
patterns as well as their sites of interaction for achieving sen-
sorimotor integration.
Emotional cookbooks
Robert C. Solomon
Department of Philosophy, University of Texas, Austin, Texas 78712
Panksepp's article seems to me to be a valuable research project
misconceived. The project is not just his, of course; it surveys a

campaign which dates back to the nineteenth, if not to the early
seventeenth century, to explain emotions in terms of their
neurophysiological substrata in the central nervous system.
Unlike many researchers in that tradition, Panksepp does not
pretend to reduce emotions to such neurophysiological (and
associated) disturbances and systems; indeed, his insistence on a
dialogue between neurology and the "subjective awareness" of
our own emotions through "introspection" is an attempt to block
any such reductionist thesis. Panksepp's views, accordingly,
tend to be expressed as a kind of dualism - of method if not of
entities. This will properly arouse some of the philosophers and
cognitive scientists, who would rightly argue that such talk can
and should be replaced by the newer language of functionalism,
in theories of emotion as in other more straightforwardly cogni-
tive fields of study. This presupposes, of course, that emotions
are cognitive phenomena, ways of conceiving and knowing
rather than mere systems of organized behavior, triggered by
neural systems and resulting in a readily identifiable "subjective
experience." This is also what bothers me about Panksepp's
presentation - not just its dualism, but its continued treatment
of emotions as "primitive" ingredients in an emotional cook-
book, raw neurological-behavioral elements that are combined
in a variety of recipes to yield the more complex and subtle
emotions of adult human life. Jealousy is analyzed as a admixture
of panic, rage and expectancy, for example.
The idea that emotions are "primitive" or have primitive
"components," which are more or less instinctual, unlearned,
inherited, and form compounds of other emotions, gives rise to
an intellectual game, which can be traced back through the
middle ages to ancient times. Medieval physiologists conceived
of various lists of basic emotion-components, and then devel-
oped ingenious lists of recipes. Descartes listed his six primitive
emotions in his treatise on "The passions of the soul," and John
Watson, more ontologically frugal as always, reduced his basic
list to three. Robert Plutchik now argues eight, Panksepp four.
But what has changed in the game in Panksepp's hands is that
the classification no longer turns on the rather casual examina-
tion of our ordinary conceptions of emotions and their identifica-
tion (not to be confused with "subjective awareness" or "intro-
spection"); the identification of the primitive emotions turns on
neurological observations. This explains the oddness of his basic
four components - expectancy, rage, fear, and panic. The
inclusion of expectancy, and thefirstjoint appearance of fear and
panic, suggest something more than an eccentric list of ingre-
dients; it is a list of ingredients that, in terms of our ordinary
language of emotions (again, not "subjective awareness" or
"introspection") is simply unintelligible. This becomes obvious,
for example, when we see the ingenious but desperate attempt
Panksepp has to make in order to analyze virtually any human
emotion; his analysis of jealousy, for instance, is not even
plausible, no matter how generous our interpretation. Spinoza
had it better when he suggested that jealousy was a combination
of hatred and envy, but then Spinoza was not saddled with the
frugal neurological list of components Panksepp imposes on
himself.
What is wrong about the cookbook model of emotions, in
which certain emotions are basic ingredients and all of the
others combinations of these, is not the implausibility and
sometimes the absurdity of particular lists of ingredients. Pank-
sepp has it obviously wrong, no matter what the neurological
findings, but Watson and Descartes and Spinoza, too, are
mistaken, and not because of the details of their lists. Emotions
are not made out of basic components, and insofar as a neu-
rological substratum of systems and disturbances can be identi-
fied (as Panksepp and his colleagues are attempting to do), there
is no reason to identify those systems and disturbances with
emotions. Emotions themselves have a structure, which they
may share with other (more primitive) emotions, but it does not
follow that they have components, or that the more primitive
(unlearned) emotions are more fundamental.
Commentary/Panksepp: Psychobiology of emotions
Our paradigm of an emotional reaction has always been one of
those dramatic "outbursts," so easily characterized by rage or
panic, which consists of an emergency reaction to a stimulus, a
more or less stereotyped and perhaps even instinctual sequence
of behavior and the familiar sensations of the physiological
disturbances we now know to identify with certain specific
hormonal secretions. The traditional mistake, canonized by
James (and not uncanonized by Cannon) has been to think of
these reactions as the very essence of emotion, thus ignoring
what, on a moment's reflection, would appear to be the vastly
larger proportion of our emotional life - lifelong love of a brother
or shame about an adolescent crime, envy of the rich or resent-
ment of the powerful. But even within the narrow confines of
that paradigm, the analysis of emotion in terms of basic phys-
iological reactions is inadequate. There may indeed be specific
neural systems to explain the reactions of rage and fear, but
those neural systems, and their products, are not yet rage or
fear. Rage and fear require a cognitive component to distinguish
them, for example, from mere surprise, which Panksepp right-
ly, but for the wrong reasons I am afraid, eliminates from the
usual list of emotions. "Rage,' if it means anything like what we
ordinarily mean by "rage," necessarily includes a component
crudely characterized as "being offended" or at least frustrated;
without some reference to what the rage is "about," we don't
have rage at all. (This raises the question whether dogs and rats
can be angry - there is little doubt that they can feel fear - but
that is not a question I want to raise here.) Panksepp may be
right about his neurological components, but he leaps to conclu-
sions when he assumes that the component itself is rage, rather
than, at most, a causal precondition for that dramatic but
narrowly conceived paradigm of rage which traditionally has
formed the focus of theories of emotion.
There may be distinctive neural systems which in some sense
lie at the base of certain emotional reactions. But these systems
are not themselves emotions, and are not required for the
having of an emotion. What neurological stew can one cook up to
explain David Hume's "calm passions," such as "a sense of
justice," for instance? Emotional phenomena are not easily
distinguished from our more general and less episodic views of
the world and ourselves. Neuroscience has an exciting realm of
its own to investigate, a realm which does indeed explain in part
some of the more dramatic scenes of our emotional life. But even
then, to explain Othello's rage without reference to Desdernona
and Iago will not do, even if our neuroscience is,finally,down to
the precision of a cookbook.
Softening the wires of human emotion
Michael Stocker
Philosophy Department, La Trobe University, Melbourne, Victoria 3083,
Australia
Panksepp wishes to conjoin introspectivist with behaviorist and
neurophysiological approaches to a psychological understand-
ing of emotions. Central to his attempt are first, his claim about
the hardwiring of certain, supposedly basic emotions, and,
second, his suggested understanding of emotions. From the
perspective of a philosopher investigating human emotions, I
found both the first and second themes unhelpful.
First, a process is hard-wired if, at least roughly, it is (at the
relevant time) invariant, automatic, not under conscious or
programmable control; if it is a matter of hardware, not software.
What goes on in a pocket calculator when the square-root button
is pushed is, thus, a good example. And Panksepp writes "each
emotive circuit . . . may . . . be specific (hard-wired), in the
sense that each generates a single, homogeneous psycho-be-
havioral tendency. "
In both classical (e.g., Aristotle 1941) and contemporary (e.g.,
Rorty 1980)1 philosophy, emotions have been seen as involving
BEHAVIORAL AND BRAIN SCIENCES (1982)3 445
Commentary/Panksepp: Psychobiology of emotions
affect, desire, ratiocination, and action. So Aristotle ("Rhet-
oric," II, 2) holds that to be angry I must be pained by a wrong,
seen as such; and I must have a desire to retaliate. Thus, to claim
that human emotions are hard-wired is to hold that the affective,
desiderative, ratiocinative, and action "elements" are hard-
wired - perhaps triggered by some stimulus. Or it is to claim
that these elements, especially the first three, are unimportant
or epiphenomenal. I shall here simply ignore the latter; Pank-
sepp, by his appeal to introspectivism, surely joins me and other
philosophers in rejecting it. I wish here to show why we should
also reject the former.
We at least seem to have some control, for example, through
reflection, over whether we have the beliefs (or thoughts) that
are necessary for an emotion - for example, that we have been
wronged. At least over time, we have some control over our
desires: Education and other forms of socialization depend on
this. Further, whether desire and ratiocination are under our
(present) control, they are often enough not "activated" by an
independently identifiable stimulus. So, someone's stepping on
my foot may arouse my anger and thus the appropriate affects,
desires, and beliefs. But it may not if I do not believe the person
was at fault, or if I do not wish to engage in hostile affrays, or if
my level of affect is too low, or if I was amused, not pained, by
what happened. Further, even if the emotion is aroused, I may
hold my temper and not retaliate. And if I do retaliate, I may do
so in any number of different ways: hit, kick, swear, belittle, call
for help. . . .
There are indications throughout that Panksepp recognizes
these facts (e.g., in "A definition of emotions" at (5) and in
"Learning, reinforcement, and transhypothalamic emotive sys-
tems," last paragraphs) as being true not only of humans, but
also of lower animals. But then, what remains of the hard-wiring
claim? Very little, I suggest, except perhaps that there is some
neurophysiological "pathway" or complex which may be hard-
wired, which may inhibit or enhance affect, desire, and
ratiocination, and which may mediate between those elements
and either or both of, for example, visceral or hormonal goings-
on, and also (just possibly) physical actions, such as hitting,
unless the latter are inhibited. However true this may be, it is
not an especially powerful or novel claim. Indeed, it seems true
if neurophysiology plays any role at all in emotions. It might be
suggested that at least Panksepp has elucidated the nature of
that neurophysiological role. This is outside my competence.
As for the second central point, Panksepp's characterization of
emotions is both unclear and unclearly motivated. Are we really
told why there are just those four emotions? Is it just that they
are the ones so far found? We are not told why the six attributes
in "A definition of emotions" are offered as characterizations of
emotional circuits. We are not told what "life-challenging cir-
cumstances" means or applies to. If "challenging" tends towards
what is threatening, many emotions - for example, curiosity,
amusement, joy - are not captured. If it tends toward what is
simply engaging or what is important to or for life, then what is
not life-challenging? One way to bring this out is to note that in
this second understanding of "life-challenging," the six condi-
tions would fit (at least various) desires and beliefs. Perhaps this
is as it should be. Perhaps they, too, are or involve emotions.
But this is not discussed, although it may be hinted at in
"Learning reinforcement, and transhypothalamic emotive
systems."
Panksepp does not deal with beliefs and desires, perhaps out
of deference to the view that "questions of animal awareness
were [to be] shelved." But without beliefs and desires, how
could one explain or distinguish among (at least various) human
emotions?
Panksepp writes that "the concepts of pleasure/relief and
displeasure/distress are not included in the present analysis."
This might be thought fair enough. Not everything can be
covered at once. But, first, how can one characterize (at least
various) emotions without those concepts? Just consider how
446 BEHAVIORAL AND BRAIN SCIENCES (1982)3
important pain is to anger, or what being pleased or amused by
having one's foot stepped on shows about one's emotion. Sec-
ond, in Panksepp's own characterizations, these concepts figure
importantly: for expectancy, joy; for fear, pain; for rage, irrita-
tion or discomfort; and for panic, distress.
Had Panksepp paid more attention to these factors, he would
have been able to offer us a better understanding of human
emotions. So, for example, he would not have had to speculate
that supposedly incompatible emotions such as positive expec-
tancy and anger (at the same time and/or about the same thing)
involve reciprocally inhibiting "hypothalamic cells." Rather, he
could have considered whether the seeming conflict is due to a
conflict of ideas - note the problems in believing something
both, say, good and bad (in the same respects); or a conflict of
desires - note the problems in both desiring and not desiring
something (in the same respects); or a conflict of affect - note the
problems in being, say, pleased and pained by something (in the
same respects). And, more generally, Panksepp might have
been able to further our understanding of the central and
difficult topic of ambivalence in emotions (on which, see Green-
span 1980).
To draw my comments on the two point together: It must,
however, be recognized that to have pursued affect, desire, and
ratiocination would have been to show even more clearly just
how nonhard-wired, just how soft-wired, human emotions are.
NOTE
1. Rorty (1980), is an especially useful anthology. It includes essays
by both philosophers and social scientists, and it has an extensive
bibliography on emotions under the following headings: general and
historical, physiological, biological, psychological, psychoanalytic, an-
thropological, and philosophical.
The rat as hedonist - A systems approach
Frederick M. Toates
Biology Department, The Open University, Milton Keynes MK7 6AA,
England
First, let's accept that behaviour and subjective experience are
functions of the nervous system. Then, it is reasonable to
attempt a simplified taxonomy of neural circuits and associated
affective states, for example, anxiety. However, I don't like to
think of this as a return to introspection. Surely, verbal reports,
as made to an interested party (e.g., "I feel afraid"), should be
the subject of the taxonomy. Such reports reflect a common
language and cultural heritage involving behaviour patterns and
associated verbal labeling.
My primary interest is in animal behaviour, and here Pank-
sepp considers only one aspect (albeit a vital one) of a complex
system. I am concerned with how the mechanisms governing
behaviour work. Let me start from a perspective somewhat
different from Panksepp's. I see the main task to be the design-
ing of a "model rat," which, in its commerce with the simulated
environment, exhibits some performance characteristics of real
rats (Deutsch 1960; Gallistel 1980; Toates 1980). Even if one
were to attempt a neurophysiologically viable model, the kind of
technique in which Panksepp has faith (lesions, chemical stim-
ulation, etc.) might be of limited use. Given that so many
millions of neurons are available, we might need to implicate
processes and emergent properties for which no obvious identi-
fication technique is available, though we would be constantly
aware of what neurophysiology had to offer. In contrast to
considering whole, behaving rats, looking at only bits of rats can
appear, on close examination, to have left the rat immobile,
buried in "hope, desire - joyful anticipation," fear, or pleasure.
How then might we approach the problem of designing a rat?
Consider what Panksepp calls "expectancy." He writes: "A
fundamental requirement for survival is that an animal move

from where it is to where life-sustaining objects are located."
This is the crux of the matter, but what kind of process does it
implicate? First, it means putting a servomechanism in the
head. It is not enough to have a system that "invigorates motor
behavior" or "modifies sensory acuity." Foraging is not a ran-
dom energization of activity but is purposive and goal-directed.
One might propose a goal-seeking model, in which the rat is
driven in such a way as to "hunt" for maximum activity at a
hypothalamic site. Choice of goal would be dictated by incentive
objects and physiological states (cf. Bindra 1978). A slight worry
with this model is the discontinuity between consummatory and
appetitive states as found by Hamburg (1971). Stein (1964)
designed a theoretical rat that would run a maze or bar-press for
food by ascending a series of predictive cues. Energy depletion
accentuages the incentive value of the cues, in other words, the
hypothalamus would be activated by the cue. Compatible with
Panksepp's argument, a variety of behaviours may maximize
such activity, for example, moving towards incentives or self-
stimulation. In the latter case, the rat isn't left buried in
pleasure, provided that each shock accentuates (reinstates) the
incentive value of the lever, making it the goal for the next press.
Priming may be needed when the rat is "cold."
In the natural environment, things get more complex. Forag-
ing demands a cognitive map; the rat goes to distant sites.
Consider the Amakihi, a species of bird which, in its nervous
system, can label distant food sites as replenished (Kamil 1978).
The map has to be activated by energy depletion, and a given
aspect of it forms the goal of behaviour (Deutsch 1960; Gallistel
1980). If the animal is blown off course, a control system realigns
it (cf. Powers 1978). Similarly, out of fear, animals move from a
source of danger to one of safety (Bolles 1970; Toates & Birke
1982).
Tantalizingly, Panksepp simply notes that his ideas can "easi-
ly handle many of the striking characteristics of animal learn-
ing, " such as autoshaping. But surely the circuits that he reviews
are of very limited usefulness in such an explanation. What kind
of model is relevant to explaining autoshaping? The animal
needs an internal spatial representation of its environment and
superimposed upon this a representation of the causal texture of
that environment, as, for example, light (E,) predicts food (£2).
The system must work by E} evoking a memory of E2 (cf.
Dickinson 1980; Lewis 1979; Toates 1982). It is as if E2 has
occurred; the animal makes an anticipatory response. In the
special case of the pigeon, this involves attempting to eat the
key. This is an enormous amount of information processing,
presumably involving a large part of the brain.
Introspection and science: The problem of
standardizing emotional nomenclature
Holger Ursin
Institute of Physiological Psychology, University of Bergen, 5000 Bergen,
Norway
Panksepp's conviction is that the study of emotions is im-
poverished by our failure to blend human introspection with
objective studies of the physiology and behavioral characteris-
tics of emotions in animals. This is, of course, a rather shocking
statement, but I think Panksepp deserves credit for bringing
this issue into the open. There is little doubt that all of us use
introspection to some extent when we formulate our problems
and discuss the validity of our measurements.
The shock effect may vary among disciplines. In human
psychophysiology and psychoendocrinology, there may be only
mild surprise about his enthusiasm. The most obvious way to
bridge the gap between human introspection and physiological
and behavioral characteristics of emotion is, of course, to study
behavioral and physiological changes in man. There are numer-
ous studies where objective methods have been used, both for
Commentary/Panksepp: Psychobiology of emotions
physiology and behavior, as well as for measuring introspective
elements in the participating subjects. This is no longer any
major source of controversy. However, problems arise when we
try to use introspective terms in our work with animals. Pank-
sepp's remedy is to insist that the terms derived from intro-
spection should be linked to specific behaviors and specific brain
structures. This sounds good, but experience with this approach
is not encouraging. I agree with Panksepp that I should use my
introspection. What worries me is that all you other people may
do the same. Even if my introspection helps me in my work,
does it really help us to communicate and develop a "nonperso-
nal" science?
One important source of variance produced by introspection
is the attribution mechanisms humans use for the experience of
emotional arousal. Panksepp treats this lightly, frowns upon the
James-Lange aspects of emotional experience, and mentions
Schachter and Singer (1962) en passant. Maybe we should not
ignore the attribution mechanisms in psychologists. They may
have even stronger and more varied attribution mechanisms
than the ordinary person. Hence, we may end up with an even
worse variance in terminology and a confusion that introspec-
tion alone cannot solve.
One possible bridge to avoid dangerous phylogenetic jumps is
to compare the instrumental effects of a particular behavior.
There are, after all, a limited number of logical options open to a
logical machine like the brain when it is faced with, say, a danger
or a frustration. When such logical functions have been identi-
fied across species, and found to depend on homologous struc-
tures, it then remains only to put a name on the function in
question. The main criterion for a good name is that it communi-
cate. Flight, freezing, defense and offense are such terms, and
no introspection is required. This is a possible contribution from
comparative psychology that might reduce the confusing and
terminological mess human introspection has left us with.
To some extent this has already happened. The psycho-
analytical data base, to the extent that it exists, is based on
introspection and reports of introspection. The Freudian drive
or "instinct" concept of aggression is not supported by neuro-
psychological or neuroethological research in animals. Aggres-
sion does not appear as a unity. Animal studies show three or
four separate categories (offense, defense, prey-killing, and
male sexual behavior; see Adams 1979). Animal research in this
field has had no benefit from the introspective tradition. Quite
the contrary, this influence brought confusion and erroneous
assumptions about an aggressive "drive." However, due to an
active interface between animal and human research, this drive
and instinct concept now seems to be eliminated, both from the
animal and the human literature. The introspective tradition
may therefore benefit from close contact with behavioral sci-
ence, but does it really work the other way?
This is not a critisism of Panksepp's own work, which is well
within the behavioral science tradition. As a matter of fact, I fail
to see where he has really used introspection in anything but
naming the phenomena he is studying. Also, I fail to see why his
data or his introspection make it necessary to postulate "panic"
(= fear) as a separate emotion. His "expectancy" dimension is an
approach contingency. It seems closely related to response
outcome expectancy (Bolles 1972), but is only a part of the many
expectancy functions described in contemporary learning theo-
ry, at least in its most cognitive formulations. His concept also
has a very interesting and challenging relation to the particular
positive response outcome expectancy, which has been referred
to as coping (Ursin 1980).
Finding a name for operationally defined behaviors is very
important, and it may be fair to use introspection to select names
as long as we are aware that these remain labels for operational
concepts. Failure to realize this may be the most important
source of confusion in our literature. Panksepp has probably put
hisfingeron a very sore and neglected area, but it is difficult to
agree with his prescription for a cure. Our inability to standard-
BEHAVIORAL AND BRAIN SCIENCES (1982)3 447
Commentary/Panksepp: Psychobiology of emotions
ize and communicate when it comes to our labels is overwhelm-
ing. It is remarkable that we still have difficulties with the
taxonomy of aggressive and fear behaviors. This is particularly so
in the fields of animal neuropsychology and neuroethology.
Perhaps an international convention could help us agree on a
nomenclature? This may be necessary to save ourselves, our
students, and our future grants. Anatomists, zoologists, and
psychiatrists have established nomenclature agreements. These
systems do not eliminate all problems, but they identify the
problems and eliminate the unnecessary ones. At least we will
have to show much more determination and will in using each
other's terms and relating our own definitions to definitions
available in the literature. We will also have to rid ourselves of
some of the easy paths to fame. One phenomenon will have to
have one name - and not a reference to the smoothest salesman
and best writer of the pack. Anatomists rid themselves of the
person-names decades ago, at least for most structures. It may
be time we did the same.
Does introspection have a role in
brain-behavior research?
C. H. Vanderwolf and M. A. Goodale
Department of Psychology, University of Western Ontario, London, Ontario,
Canada N6A 5C2
Panksepp's approach to brain-behavior research is based on the
assumption that introspection provides a relatively direct means
of obtaining information about the functional organization of the
brain. Although this assumption is probably false, Panksepp
should be commended for making explicit a tacit assumption
underlying a good deal of physiological psychology and behav-
ioral neuroscience. For years, many researchers have routinely
labeled the behaviors they observe in animals as examples of
fear, pleasure, rage, or anxiety. Whereas we would criticize
them for using a terminology that is derived largely from
introspective experience, Panksepp chides them for not using
introspection enough. Indeed, he asserts that a "single-minded
focus on emotional expression without concurrent consideration
of underlying emotional process has outlived its usefulness in
psychobiology and may be currently hindering progress in
understanding the neural organization of emotion."
The explicit taxonomy that Panksepp proposes for investigat-
ing the neural organization of emotion is derived from an earlier
introspective scheme developed by Cogan (1802). But however
consistent this scheme may be with human self-report, we
contend that introspective systems such as his can tell us
nothing about the neural substrates of behavior. "Mental"
processes cannot be directly examined. We "know our own
minds" largely or entirely as an inference from our own behavior
and not as a result of direct inspection (Hebb 1980). Similarly,
Skinner (1974) concludes that the internal mechanisms that
cause behavior are, in general, not open to direct examination
by introspection. A recent paper by Nisbett and Wilson (1977)
summarizes a body of experiments in the fields of social and
cognitive psychology that supports these views of Hebb and
Skinner. Thus, it is quite unlikely that introspection can provide
decisive information on the functional organization of the inter-
nal processes that give rise to behavior. In fact, it would be
remarkable if introspection were of any assistance in this re-
spect. In this century we have come to believe that neural
activity is the immediate cause of all behavior. However, it is
apparent that introspection reveals nothing of the structure or
function of the brain. If it did, it would not be necessary to
experiment on the brains of animals. An investigator could study
the anatomy and physiology of his own brain by introspection!
A psychological approach of the type advocated by Panksepp
makes further assumptions. In his scheme, emotion and its
components are conceived of as subdivisions or states of the
448 BEHAVIORAL AND BRAIN SCIENCES (1982)3
mind, which, in turn, is assumed to be intimately associated
with the brain. According to Panksepp, the brain contains
circuits corresponding to these subdivisions, and the overt
behaviors comprising emotional expression are assumed to be
initiated or mediated in some way by these primary emotional
networks. That is, it is assumed that brain organization is
subdivided in ways that correspond to major psychological
concepts.
Is such an assumption justified? We think not. In a recent
paper, Vanderwolf and Robinson (1981) summarized a good deal
of evidence showing that cerebral slow wave activity is, to a large
extent, organized in terms of the performance of overt behaviors
such as immobility, head movement, walking, postural changes,
licking, face-washing, etc. This organization appears to be inde-
pendent of emotional state.
A similar lack of correspondence between psychological expe-
rience and neural organization of behavior has been demon-
strated in a very different area of research. Despite the fact that
the unity of our visual perceptions is extremely compelling,
there is increasing evidence that this perceptual organization is
not reflected in any simple way in the organization of our visual
pathways. Instead of a single visual system, there appear to be
five or more independent (albeit interactive) sensorimotor net-
works, each mediating very different visuomotor behaviors
(Goodale, in press; Goodale & Milner 1982). None of this
organization would have been discovered by introspection.
In the same way, it is also possible that brain activity and
neural circuitry are organized, at some level, in terms of feeding
behavior, sexual behavior, parental behavior, thermoregulatory
behavior, grooming behavior, eliminative behavior, territorial
behavior, predatory behavior, and so on. This organization need
not correspond with or conform to any of the neural substrates
underlying human emotional experience. In fact, we would
argue that there are no unitary representations of emotions as
such. Whatever the neural circuits are that permit comment
upon and cognitive processing of our sensory input and motor
output during the performance of behaviors we have come to
label "emotional" or "affective," it is unlikely that such circuits
are related in a one-to-one fashion to the circuits mediating
different kinds of "emotional expression." That is, the brain
mechanisms that generate "emotional" behavior, such as run-
ning away when one sees a bear, may be quite distinct from the
mechanisms that enable one to say, at a later time, "I was really
frightened, so I ran away as soon as I saw the bear."
Suggestions of this type are highly speculative. At present, we
simply do not know how the brain is organized. It seems to us,
however, that an essential part of the analysis of the brain
mechanisms that control behavior must consist of careful de-
scription and study of behavior. Behavior is the end result of the
activity of most of the nervous system and, as in any complex
system, a study of output might reveal something of the internal
organization. Psychological approaches such as Panksepp's,
which consider "mind" to be of primary interest, tend to
discourage systematic study of behavior (Vanderwolf, in press).
For example, Panksepp makes no serious attempt to relate the
effects of localized hypothalamic stimulation to the normal
behavior of the species studied, even though it is widely recog-
nized that a good knowledge of normal behavior is an essential
prerequisite for work in this field. Further, Panksepp gives no
detailed discussion of possible rules by which an "emotional"
behavior might be distinguished from a "nonemotional" behav-
ior, or of how one might identify the presence of any particular
emotion in a normal animal.
In conclusion, we suggest that introspection, in the sense of
direct examination of the mechanisms that control behavior,
probably does not exist and, therefore, cannot be a useful way of
studying the brain. We suggest further that a preoccupation
with psychological concepts actually impedes research, since it
diverts investigators from a thorough study of brain-behavior
relations.

Can arousal be pleasurable?
Marvin Zuckerman
Department of Psychology, University of Delaware, Newark, Del. 19711
Like Gray's (1982a) The Neurophysiology of Anxiety [see BBS
multiple book review, this issue], Panksepp's article represents
a laudable attempt to link emotions with specific neural circuits
in the limbic brain. Panksepp's work is more ambitious, dealing
with four primary emotional states instead of one, and less
insistently behavioral in admitting the value of introspective
data from humans as a source of knowledge, even though such
data are hard to come by in the psychopharmacology and brain
stimulation literature. I am in agreement with these general
approaches to the understanding of emotional states and traits,
and my criticism (as with Gray's work) is addressed at specifics
rather than the comparative approach. My commentary will be
focused on the system of greatest interest to me, "expectancy,"
because of its resemblance to the trait of "sensation seeking"
(Zuckerman 1979).
Although Gray uses the term "anxiety," a term with phe-
nomenological surplus meaning in the human sphere, to de-
scribe one of his three major emotional systems, he uses the
more behavioral term "impulsivity" to describe the adient
system. Similarly, Panksepp uses "fear' to label the avoidant
mechanisms and "expectancy" to describe a system for the
mediation of "exploration-approach-investigation." Why are
these theorists reluctant to use terms like "desire, ' "pleasure,"
or "joy" when describing the emotional system related to the
expectancy or reward, yet willing to use terms like "anxiety" or
"fear" to describe the system related to the expectancy of
punishment?
In the case of Panksepp, his avoidance of the term "pleasure,"
despite his acceptance of human introspective data, is a function
of his drive-reduction view of motivation. Pleasure is conceived
as a state that accompanies stimuli "that tend to return organ-
isms toward physiological homeostasis and emotional equi-
librium." He further claims that such states do not appear to
"generate unique behavioral manifestations." But the system he
describes as "expectancy' (meaning limited to positive expec-
tancy) is one that has been mapped through self-stimulation
studies of the brain as well as externally controlled stimulation.
Self-stimulation or exploration (which may occur in the absence
of specific searches for substances needed to redress "homeosta-
tic imbalances") are activities that create arousal, not reduce it.
In strict drive-reduction terms only the reduction of arousal can
be reinforcing. But tell that to well-fed rats sniffing around a new
environment, or to humans spending large sums of money to
travel and to look at or listen to complex and novel stimuli, or to
sniff or smoke stimulants like cocaine. While drug users may
find pleasure in the effects of depressant and stimulant drugs, an
experimental study of drugs given blind showed that the stim-
ulant D-amphetamine both maintained high levels of arousal
and produced positive affect more consistently than a depres-
sant, diazepam (Carrol & Zuckerman, in press). The latter was
more likely to produce states of anxiety, depression, or hostility,
accompanying decreased arousal.
Are there specific behavioral indicators associated with the
state of "pleasure" other than the tendency to approach the
stimulus? Emotional expressions such as the human smile and
the dog's tail wag seem to provide a behavioral supplement to
the approach-behavior definition. Although I am not sure about
the expression of positive emotion in our more favored subject,
the rat, I believe if we spent as much time studying positive
emotions as we do fear and anger, we wouldfindsuch indicators.
Is there only one arousal system, as Panksepp seems to
suggest? Do norepinephrine and serotonin act equally on all
emotive systems (see Panksepp's Figure 4)? In contrast, Gray
(1982b) and Stein (1978) suggest that serotonin mediates an
anxiety system. Gray also feels that norepinephrine is involved
in part of this system, while Stein believes that this neu-
Response/Panksepp: Psychobiology of emotions
rotransmitter, along with dopamine, mediates reward systems.
Evidence is conflicting on all of these hypotheses, but it should
be noted that there is the possibility of more than one arousal
system. Routtenberg (1968) suggested that the limbic reward
system has its own arousal pathways to the forebrain and that
these constitute a second arousal system, the reticular activating
system being the first. Since his paper, these pathways have
been discovered, lending support to the idea of a type of arousal
generated by, and linked to, reward mechanisms.
Certainly the high levels of arousal associated with sexual
activity are experienced as pleasurable before the drive- reduc-
tion or orgasm. Male sexual behavior is enhanced by brain
dopamine and suppressed by serotonin (Gessa & Tagliamonte
1974). Norepinephrine may also play an excitatory role in sexual
arousal. The question is whether the monoamines simply exert a
nonspecific arousal effect or are specifically effective in sensitiz-
ing reward systems. Is the release of dopamine or nor-
epinephrine associated with the experience of pleasure,
whether from intrinsic reward mechanisms or reduction of
activity in a punishment-anxiety system? Many of these ques-
tions could be resolved by work with humans, in which intro-
spective data are obtainable.
Author's Response
Archaeology of mind
Jaak Panksepp
Department of Psychology, Bowling Green State University, Bowling
Green, Ohio 43403
It is difficult to imagine science without words, but quite
easy to imagine feelings without words. This is under-
standable, for functional neuroanatomy has affirmed that
the brain's ability to elaborate affective representations of
reality evolved much earlier than its ability to generate
scientific-linguistic representations of reality. Further-
more, granted that a major force behind the evolution of
complex human linguistic abilities was the adaptive util-
ity of communicating interrelations among environmen-
tal events (such as would be needed, for example, in
complex forms of group-hunting, defense, foraging, co-
habitation, and migration), the linguistic and scientific
analysis of internal processes such as emotion appears
compromised at the outset. Our imprecision in conveying
emotions verbally (as well as the importance of emotions
in controlling behavior) is captured in the aphorism, "It's
not what you say - it's the way you say it."
The semantic controversies that routinely arise in the
discussion of emotion have long hindered the progress of
research in this area, and understandably, most biolog-
ically oriented investigators have restricted their study of
emotion to behavioral expressions, with little considera-
tion of the underlying internal processes. At least we can
agree verbally on what we have seen, while internal
processes can only be observed through a distorted intro-
spective or theoretical lens.
However, I daresay most people would agree that "the
affective faculties have their origin from within, and are
not acquired by any external circumstances. They cannot
be taught and must be felt to be understood; in them-
selves they are blind and act without understanding;
BEHAVIORAL AND BRAIN SCIENCES (1982)3 449
Response/Fanksepp: Psychobiology of emotions
finally they are partly common to man and animals, partly
proper to man" (Spurzheim 1846, p. 136). My own
position was premised on such seemingly self-evident
truths. Accordingly, my aim was to interrelate the psy-
chology and neurology of emotional processes at a level
primitive enough to permit rather strict empirical evalua-
tion of theoretical propositions. As there is only limited
biological knowledge on the topic, this is a narrow path,
bordered on one side by ungrounded philosophy and art,
and on the other by a radical empiricism which recognizes
only human observation (largely visual) as a credible
source of scientific inference and understanding. I would
argue that to be tempted to one of these sides may be as
detrimental to understanding as to be tempted to the
other. If there is a "royal road," it is one by which
psychobiology and psychophenomenology are interre-
lated at a level that does no severe injustice to either. As
summarized in the commentaries on my target article,
there is little agreement that this can be done. However,
I was encouraged that the arrows of criticism were dis-
tributed rather evenly around the target I provided,
suggesting that I may have struck a realistic, if not a
happy, medium. Who has come closest to the mark
remains to be seen, of course, for the penetration of
future research into the underlying brain mechanisms of
emotion is the metric by which the accuracy of our aim
will be measured.
Two cultures. Two attitudes polarize the debate over
emotions. Commentators commited to understanding
affective experiences in human life are disturbed by
attempts to view emotion in reductionist, neurological
terms. They recommend more conceptual complexity in
discussing emotions than can presently be incorporated
into the psychobiological enterprise (Averill, Jaynes,
Lazarus, Plutchik, Royce, Solomon, Stocker). At the
other pole, several investigators committed to charac-
terizing behavior accurately are skeptical of attempts to
use concepts suggesting that animals experience emo-
tions (Anisman & Zacharko, Ursin, Vanderwolf & Good-
ale) - a stance with which I do not agree because it
arbitrarily closes doors to potential knowledge (also see
Griffin 1976). There is also another road, perhaps most
difficult of all, which I would take with those who suppose
that a realistic account of brain activity underlying emo-
tions will require a compromise between such polar
viewpoints (Fonberg, Globus, Gray, Heath, Izard,
Katz).
What appears too often forgotten by aesthetically,
philosophically, and cognitively oriented contemplators
of emotions is the basic nature of scientific inquiry. In
attempting to simplify complex systems, one necessarily
does some injustice to the manifestations of those systems
in the real world. For instance, in considering the lattice
properties of crystallized water, science provides a cogent
explanation for the six points of a snowflake, but it has not
explained the exact form each snowflake assumes in life.
As emphasized by Katz, even if the rules of nature I
propose are correct, the full emotional richness of human
life will not have been denied, compromised, or deni-
grated: The emotive circuits I outlined may initiate emo-
tions in all mammals, while also interacting with higher
brain functions and cultural processes that distinguish
humans from other animals (as emphasized by Delgado).
450 BEHAVIORAL AND BRAIN SCIENCES (1982)3
Put simply, the emotive circuits I described may be
necessary though not sufficient substrates for the genesis
of emotion in the mammalian brain. Brain software (i.e.,
learning) no doubt operates on the genetically provided
emotive circuits (Averill, Stocker), but such software may
not be possible without the hard-ware I have outlined.
The schizm between the "two cultures" was most
poignantly expressed by Lazarus, whose reaction to my
proposal cascaded from the high hope that something
new and psychosocially significant would be said about
the human condition to the despair of having to deal with
alien neurological facts. I regret that the bridge between
these distant shores of understanding remains to be
completed, but it will require both sides to prepare
common and firm ground. Those with psychosocial and
psychophenomenological interests must appreciate the
nature, importance and limitations of psychobiological
inquiries, and neuroscientists must seek ever more com-
plex functional principles for the brain. Some commenta-
tors steeped in the behaviorist tradition already thought I
had gone too far toward the other side; the despair that
Lazarus felt on reading some facts about the brain was
matched by the concern Ursin and Vanderwolf & Good-
ale felt at the apparent capitulation of an empirically
oriented colleague.
Still, there is a bridge to be built between the cogni-
tive/clinical and the neurobehavioral cultures, and sever-
al commentators agreed that the needed materials may be
at hand (Fonberg, Heath, Izard); and some attempts at
constructing it have already been made (see Willis, 1683,
and see Arnold, 1960, for a more recent admirable
attempt).
Introspection! Why not? Among cognitively oriented
commentators, introspection was generally deemed es-
sential for understanding emotion (Arnold, Averill,
Clynes, de Rivera, Globus, Lazarus, Royce, Stocker,
Zuckerman). On the other hand, Lutz, Plutchik, Ursin,
and Vanderwolf & Goodale were especially cautious
concerning the utility of introspective information in
generating hypotheses about brain function. Everyone
would certainly agree that many mechanisms that con-
trol behavior operate subconsciously, and that introspec-
tion may be directly useful in deciphering only certain
facets of brain organization.
Still, it seems evident that human consciousness ex-
ists, whether it started in 4004 B.C. as the seventeenth-
century Irish divine, James Ussher, would have had our
God-fearing ancestors believe, around 1000 B.C. as
Jaynes suggests, before 10,000 B.C. as the paleolithic
wall painting at Lescaux may indicate, or around the
time transhypothalamic emotive circuits evolved into
basal ganglia in the premammalian brain, as I would
argue. Regardless of when it came about, the concept of
consciousness is as sound as that of gravity, even though
different species are probably endowed with different
degrees of this brain function. Introspection emerges
from this faculty in the human species. Although it will
not help directly to solve all psychobiological questions
(Nisbett & Wilson 1977), I trust that, if carefully used in
the study of emotion, it will deceive us no more than any
other adaptive mechanism evolution has bestowed on
our bodies.
By way of reply to Globus: My perspective on the

utility of introspection in the study of emotions is a
natural outgrowth of my conception of how conscious-
ness evolved in the mammalian brain. I assume that the
most primitive function of consciousness is to facilitate
adaptive response selection from alternative courses of
action: It allows organisms to cope with complex en-
vironmental situations in which several behavioral alter-
natives are competing, with comparable urgency, for a
common output channel in the brain. Such a crisis of
choice (if one can imagine a crisis on an evolutionary
time scale) may have become most urgent to species that
possessed executive brain mechanisms that could con-
currently promote several adaptive behavior patterns to
a single type of environmental challenge. As I have
discussed more fully elsewhere (Panksepp 1981a), emo-
tive command circuits may have such a characteristic.
This flexibility could promote adaptive response-mold-
ing, perhaps by a "reinforcement" mechanism linked to
fluctuating activities in the underlying executive cir-
cuits. In simple animals, such response selection (or
habit formation) might operate at a preconscious level,
but as the trajectory of evolution took species to ever
more complex environments (reinforcing sophisticated
spatial representation abilities), more sophisticated deci-
sion-making processes evolved from the earlier solu-
tions; and today I think we call these processes con-
sciousness. If human consciousness is rooted in the
primitive response selection, response-molding mecha-
nisms that emotive circuits elaborate, introspection
should still have some access to the dynamics of the
underlying command circuitry. This is not to deny that
the introspective view, through the layers of other brain
functions, can be hazy, as suggested by Freud, Lutz, and
Plutchik.
Although my approach could easily be mistaken for
dualism (Solomon), Clobus correctly argues that my the-
sis is not credible from that perspective. True dualism
entails rejecting introspection as a source of scientific
hypotheses about certain brain processes. The implicit
dualism of behaviorism has in fact long been disguised as
monism and should be revealed for what it is - "sham"
monism. If we believe in the continuity of the natural
world, we must acknowledge and use all available
sources of information in order to understand our brains.
Since the human brain must extract external informa-
tion from sensory transduction systems, it seems evident
that all scientific knowledge is ultimately based on intro-
spection (as decoded through language). Accordingly, all
aspects of brain function should be accepted as potential
sources of psychobiological hypotheses. Although in-
terobserver reliabilities may be higher for visual and
auditory information than for other sensory systems, that
is not a rationale for rejecting the other sources of infor-
mation. The subjective experiences generated by taste,
touch, and smell provided fundamental information on
which the physiological analysis of those sensory systems
was based. Why should it be any different with in-
teroreceptive senses or internal brain processes? There
are no good objections; only unverified assumptions that
those sources of information either do not exist or have
little interobserver reliability. I think both skeptical
viewpoints are demonstrably incorrect regarding the
four emotive processes I have discussed, although they
may well be correct regarding certain cognitive pro-
Response/Panksepp: Psychobiology of emotions
cesses. (Thus, de Sousa, Lutz, and Vanderwolf & Good-
ale's citation of Nisbett & Wilson's (1977) critical analysis
for higher order cognitive processes should not be over-
generalized to basic emotive processes.)
As Klein indicates, the classical introspective tradition
in psychology waned because it led to untestable propo-
sitions. When no agreement can be reached with words,
there can be no scientific reward. When mental pro-
cesses are not linked to brain mechanisms, avenues for
arbitrary disagreement will always remain open. Thus,
my point is that introspection (especially as applied to
cross-species analyses of internal brain functions) is best
restricted to brain processes that appear to have a sub-
stantive basis (i.e., which are genetically established
rather than merely experientially derived). Such a re-
striction could have several beneficial effects on the
study of internal events. It may help curb over-imagina-
tive concepts that hinder scientific progress, and it may
encourage cognitively oriented investigators to under-
take reliable, convergent introspective studies rather
than open-ended ones.
I assume that interobserver reliability across different
cultures, in children as well as adults, would be very
high for identifying real-life as well as simulated por-
trayals of the four emotional processes I discussed.1 I
also suspect that most observers could, with little hin-
drance or assistance from culturally conditioned imag-
ination, correctly decipher similar emotional signals in
other mammals. If such experiments yield stable results,
we should accept the reality of the underlying brain
states as readily as if numbers had been read off meters.
The extent to which information derived from human
introspection is pertinent to related species, and the
extent to which information derived from animal brain
research is relevant for understanding human brain pro-
cesses is not known with precision at the present time
(Anisman & Zacharko). Ultimately, the validity of such
generalizations will depend on (1) whether brain net-
works from which human emotions arise are class-typical
or species-typical circuits, and, if the former is true (as it
seems to be), (2) the degree of evolutionary modification
these circuits have undergone. There will surely be
major species differences, but considering the impor-
tance of the adaptive solutions encoded in the executive
functions of such circuits, I suspect that the differences
will be matters of detail rather than of kind. Evolution-
ary molding of these circuits no doubt contributed to
species-characteristic (i) variations in the vigor of the
individual executive circuits, (ii) changes in the degree of
interaction among the circuits, (iii) channeling of differ-
ent sensory and perceptual inputs to these circuits, and
(iv) modification of species-typical expressions that arise
from the activation of these circuits. Of course, our use
of introspective insights to study these processes in other
species need not imply that other animals also have
introspective capacities - only (as so frankly expressed
by Fonberg) that they have emotions.
Introspective analysis of internal processes has ob-
vious limitations. As emphasized by Vanderwolf &
Coodale, it will not provide any information about the
localization of function in the nervous system (except
perhaps in brain-damaged individuals). The mapping of
brain circuits can only be achieved through direct analy-
sis of the nervous system, and for my purposes, well-
BEHAVIORAL AND BRAIN SCIENCES (1982)3 451
Response/Panksepp: Psychobiology of emotions
organized behavior patterns evoked by localized brain
stimulation are especially informative. Thus, the major
use of introspection will be in generating credible tax-
onomies that can be related to brain processes, leading,
one hopes, to incisive behavioral studies using tradi-
tional psychobiological tools. With luck, it may also pro-
vide usable information about the attributes of those
processes, including interrelations among the various
circuits.
The semantics of emotions. Practically all words used to
discuss affective processes are imprecise. Some are free
of undesirable connotations, while others are so heavily
laden with divergent meanings that their communicative
value is compromised. Operational definitions are desir-
able, but the methodological advantage gained is often
offset by the failure of rigid definitions to promote our
thinking about the underlying issues. In defining emo-
tional processes simultaneously with respect to anatom-
ically defined circuits and their many properties, I was
seeking a compromise that might prove useful for the
diverse disciplines that study emotions.
There was substantial disagreement with my selection
of terms. Morton homes in on the problem of undesir-
able semantic connotations. As he notes, I may have
identified four unconditional emotive circuits, which are
necessary for the elaboration of four affective qualities,
with no single word sufficing to label any of these states.
Zuckerman, on the other hand, expressed concern with
the reluctance of psychobiologists to use terms such as
"desire," "pleasure," or "joy." Ursin would retain only
visually descriptive words, which summarize what we
think the animal is doing, such as "flight," "freezing,"
"defense," and "offense." As he suggests, we are in
desperate need of an international congress to standard-
ize terminology, but agreement would seem unlikely.
The problem runs deeper than linguistics - to the
very nature of the emotive neural circuits mammals are
endowed with. Each emotive command system can ap-
parently be activated by diverse stimuli and can control
multiple responses at various levels of the neuraxis (tar-
get article, Figure 3). Since these systems do many
things simultaneously, it is not surprising that there are
multiple semantic representations for the patterns of
neural activity that can be orchestrated by each system.
My prescription is simple: We should not label the
system with respect to its inputs or outputs; rather, we
should seek a label that will do justice to the most
important existential attributes of these circuits, namely,
affective states. By this maneuver, we not only acknowl-
edge our primary source of information about these cir-
cuits, but most people will understand us without having
to learn alien and, at times, content-free vocabularies
(e.g., Type 1 and Type 2 behaviors: Vanderwolf &
Robinson 1981). Also, since the most incisive informa-
tion in this area is bound to come from the study of
children (Izard), diverse cultures (Clynes, Lutz), the
emotionally ailing (Heath, Klein), and the brains of ani-
mals, we should probably select a set of terms that can
be shared across all these disciplines.
Animal behaviorists may shudder and say that the
application of mental-state labels to animals will be disas-
trous - that they will be used as explanations, and peo-
452 BEHAVIORAL AND BRAIN SCIENCES (1982)3
ple will stop analyzing the underlying control mecha-
nisms. I disagree. One could just as easily claim that
such terms as "defense" and "flight" (Ursin) imply that
animals are voluntarily doing something, and hence that
those labels are unacceptably mentalistic. Affective
terms can promote the subtlety of our empirical studies
- increasing rather than decreasing our focus on behav-
ior. And surely, if we insist that psychobiology can be
constituted only of constructs we see with the eye, we
shall have failed to confront nature on her own terms.
Can a consensus be forged? Perhaps - by selecting our
terms according to some overall criterion of utility.
There are two principal reasons, and many subsidiary
ones, for studying emotion in the animal brain. First, it
may provide lasting insights into the common roots of
human and animal nature. Second, it may yield informa-
tion through which human and animal suffering can be
reduced. Thus, it would be most useful if we selected
labels for our emotive circuits that are also applicable to
existential experiences, psychiatry, brain research, and
animal behavior. This was the compromise I was seek-
ing. Perhaps hyphenated designations would be more
acceptable to more disciplines -
exploration-curiosity-foraging-expectation-desire;
flight-caution-anxiety-fear-horror;
offense-irritability-anger-rage-fury;
crying-sadness-sorrow-grief-panic.
The connotative crosstalk between the terms fear and
panic was deemed especially troublesome (Fonberg,
Izard, Plutchik, Solomon). Through cultural condition-
ing, we have learned to use the word "panic" in such a
general way as to convey practically any form of trepida-
tion. Sorrow, sadness, and grief (or perhaps distress, as
suggested by Izard) would have been more understand-
able. However, my choice was aimed at building a
bridge to the relevant data base in psychiatry. As dis-
cussed by Klein, those sudden crises of existential terror
known as "panic attacks" that characterize agoraphobia
seem to be elaborated by brain mechanisms distinct
from those which mediate chronic anxiety neuroses (for
further discussion, see Panksepp on Gray, this issue).
There is reason to believe that "panic attacks" are con-
trolled by brain circuits that mediate separation distress.
Thus, so long as we recognize that the referent here is
sorrow rather than fear, panic may be a useful and
appropriate term.
The label expectancy also caused concern (Fonberg,
Lyons, Morton, Solomon), for it could be taken to apply
either to positive or negative anticipatory states - desire
or foreboding. It could also be taken to be affectively
neutral (Lazarus). I did indicate that the term was ap-
plied only in the positive affective sense, but perhaps
terms such as desire and pleasure advocated by Zucker-
man, would have been clearer. Unfortunately, those
terms do not relate well to the major emotional disorder
presumed to arise from overactivity of this system
(schizophrenia). Perhaps expectancy does not accom-
plish this either,2 but the term does promise to be useful
in animal research. Briefly (since this is discussed more
extensively elsewhere: Panksepp 1981a; 1982a; 1982b),
the idea that lateral hypothalamic self-stimulation cir-
cuits mediate "foraging-expectancy" as opposed to
"pleasure" suggests that we should begin analyzing how

these systems govern an animal's search for food and
knowledge - lines of investigation that remain poorly
developed.
It is unlikely that we can resolve disagreements con-
cerning the meanings of terms such as emotions and
feelings (Fonberg). Fortunately, where animal brain re-
search is concerned, such distinctions are not very
urgent. In suggesting neurally based definitions for
these concepts, I attempted to provide a new perspec-
tive on this old issue. In my scheme, emotive command
circuits are activated by various sensory inputs whose
influence is at least bimodally distributed. The stimuli
that have strong and sustained interactions with these
circuits may be considered life challenging (in answer to
Stocker), and evoke bona fide emotions. Those stimuli
that have weak interactions with these circuits are con-
sidered to have evoked feelings. Accordingly, the num-
ber of emotions is limited to the number of emotive
command systems that exist in the brain, while the
variety of feelings, like that of snowflakes, is infinite.
In summary, the selection of terms for a psychobiol-
ogy of emotions is indeed problematic (Ursin). Many
difficulties arise from the old fear that animal behavior-
ists will begin using mentalistic terms as explanatory
principles (Vanderwolf & Coodale). If such terms are to
be used, they have to be tied to objective mechanisms.
The brain contains convergence pathways that collect
diverse forms of information from the external and inter-
nal worlds; through intrinsic circuit transformations,
these circuits generate coherent psychobehavioral ten-
dencies that are only indirectly measurable as outputs.
Fortunately, many primitive central processes of this
type may have been captured reasonably well by our
introspective abilities as well as in the affective terms
that came to be used during the early evolution of our
human languages. Although the psychobiology of some
future time may find the continued use of such terms to
be counterproductive (Anisman & Zacharko), I think
they serve well for now. Indeed, if such terms do ap-
proximate some essential neurological truths, they will
surely be retained in the future (the way the Newtonian
concepts of "mass" and "force" endure). If, however, we
begin using the human mental states represented by
these words as explanations of animal behavior, (dis-
regarding, for the moment, that evolution appears to
have provided such paths of causality to humans through
the emergence and refinement of brain fantasy mecha-
nisms), then they will be psychobiology's "phlogistons"
and "ethers." Thus, if we ever choose to use plain lan-
guage again to discuss emotions in psychobiology, we
shall also have to pass down to our students the recogni-
tion that words are just words - they help us think about
problems. That done, our job is to unravel the neural
mechanisms subserving those processes (Toates).
On taxonomies of emotions. As Solomon noted, I did
indeed saddle myself with a rather frugal, neurologically
based list of emotions, which does require one to go
through some mental gymnastics to generate the more
subtle human feelings; Solomon concludes that I must
obviously have it wrong, no matter what the neurological
findings (a rather antiscientific opinion). Unless we an-
chor our emotional concepts empirically, we will end up
Response/Panksepp: Psychobiology of emotions
with little more than opinion, and that won't do in the
laboratory.
I am not wedded to the conclusion that there are only
four emotive command circuits in the brain (Stocker),
but each addition to the list should be related to empiri-
cal evidence derived, in part, from an analysis of brain
systems. I have considered including executive circuits
for courting and sexually directed behaviors (lust), since
brain stimulation can evoke genital engorgement and
copulation but, just as with eating and drinking (discrete
behaviors though they are), the common underlying
emotive impulse may arise from a shared executive influ-
ence (one which promotes forward locomotion with in-
vestigation). Indeed, hypothalamic stimulation that
yields sexual behavior in rats can, with time, come to
evoke other consummatory behaviors (Caggiula 1969). I
do not wish to deny by this the existence of distinct
sexual "feelings," but merely to suggest that seeking a
sexual partner is not governed by a separate emotive
circuit (although the aroused appetitive behavior may
well be channeled to an appropriate subject by feelings
arising from genitalia and hormonal interoreceptors). I
am also hesitant to include "defense" and "submission"
as primary systems (see Panksepp 1979). It seems more
likely to me that there exists a distinct emotive circuit for
dominance displays (MacLean 1981), which may elabo-
rate a primitive "will to power" from which assertive
play, human courage, and the so-called "aggression" in
sex (Plutchik) may arise; but the neurological evidence is
presently too sparse to support the cross-species gener-
ality of such a conclusion.
Although more complex models can be generated, it is
to be hoped that such elaborations will not be based
simply on introspective hunches. Plutchik has put for-
ward a geometric model consisting of eight rather unam-
biguous affective processes; but to give surprise, disgust,
and acceptance equal status with fear, rage, panic, and
desire does not seem justified either by subjective expe-
rience or our knowledge of brain mechanisms. Similarly,
the list of primary emotions proposed by Izard (1971) -
surprise, enjoyment, disgust, shame, distress, interest,
fear, contempt, and anger - is longer than animal brain
research can justifiably support at the present time.
Although we can easily get startle/orienting-responses
(surprise?) from a vast number of electrodes in the re-
ticular core of the brainstem, I interpret that to reflect
precipitous brain activity evoked by sudden arousal of
the reticular activating system (target article, Figure 4).
Furthermore, surprise is a transient psychological state
without the sustained affect that characterizes the major
emotions. Disgust has even greater problems. It is diffi-
cult to evoke retching via brain stimulation, and subjec-
tively, nausea is about as affect-laden as stomach cramps
(they both feel bad), but the aversive feeling wanes
rapidly after the precipitating stimulus is gone. With
major emotions, the eliciting stimuli can disappear, but
the emotions boil on. Further, there are no major psy-
chiatric disorders characterized by imbalances of sur-
prise, disgust, and stomach cramps. This suggests that
the neural circuitry differs in kind from that mediating
bona fide emotions. Along similar lines, Lutz indicates
that cross-cultural studies have not found surprise and
disgust to be perceived as emotional states. Solomon also
BEHAVIORAL AND BRAIN SCIENCES (1982)3 453
Response/Panksepp: Psychobiology of emotions
argues against their being emotions, but does not indi-
cate his reasons (though he deems mine incorrect).
Several commentators disagreed with my suggestion
that exectancy should be considered an emotion (Fon-
berg, Lazarus, Lyons), while others did find it compati-
ble with their thinking (Izard, Plutchik). As already
discussed, this disagreement may merely reflect the se-
mantic biases that encumber the field. Considering my
definition of emotion with reference to circuit charac-
teristics, which no commentator seriously criticized, it
should have been clear that the brain system I was
discussing did have properties essential for an emotive
system, regardless of the label used.
My taxonomy of emotions has much in common with
the models of Fonberg and Gray. However, diverging
from Grays conclusion that flight and attack may repre-
sent activity in a single emotive system, Fonberg (in
agreement with my own observations in rats) reports
that distinct fear and rage systems are activated by hypo-
thalamic stimulation in dogs. Basing his conclusion
largely on behavioral studies using electrical foot shock,
Gray suggests that there are separate anxiety-fear and
flight-flight systems. The problem with such experi-
ments is that the convergence of pain inputs onto several
emotive command systems - perhaps all three of the
affectively negative ones in my list - makes it impossible
to distinguish among distinct neural systems. Contrary
to Gray's conclusion, my suggestion that hypothala-
mically elicited flight is a reasonable index of fear is
supported by pharmacological evidence that escape from
such brain stimulation can be reduced by antianxiety
agents (Panksepp, Gandelman, & Trowill 1970).
At a more general level, several commentators were
concerned about the absence of a pleasure/aversion di-
mension in my analysis (Katz, Zuckerman). Although I
may well appreciate the pleasure/aversion dimension at
a personal level, I do not know of any empirical evidence
to indicate that it is elaborated in the brain by distinct
emotive circuits (i.e., ones that bring diverse behavioral
and physiological responses under a common executive
influence). Accordingly, I suggested that pleasure/aver-
sion might be understood as fluctuating activities of the
individual emotive systems. For instance, such affects
may be elaborated by more generalized neural systems
within which the discrete emotive systems are embed-
ded (Katz). The issue of pleasure/aversion is of obvious
importance (and I have considered it elsewhere as a
protopathic background activity in the brain; see Pank-
sepp 1981a), but, except for taking sides on the need for
such concepts, no commentator proposed a cogent em-
pirical way to address the issue in animal research. We
could indeed use simple approach and avoidance as mea-
sures (Katz); but with four systems already in place, one
of which yields approach and the other three avoidance,
the utility of the more general dimension is compro-
mised, save as a class identifier. Thus, in animal re-
search, the concept of pleasure daunts us like the smile
of the Cheshire cat. Of course, in humans we can obtain
the needed verbal reports (Zuckerman), and pleasure
has in fact been evoked by stimulating the human brain
(Heath); many of us believe such feelings may be medi-
ated by brain dopamine and endorphin systems
(Plutchik).
Zuckerman suggests that the dog's tail may be capable
454 BEHAVIORAL AND BRAIN SCIENCES (1982)3
of telling us something about pleasure. In our laborato-
ry, we have become very conversant with this appen-
dage, but we have reached the conclusion that the tail
wags not as a generalized indicator of desire or pleasure,
but as a social, contact-soliciting gesture. As demon-
strated in a recent dissertation by Davis (1980), the dog's
tail does not wag vigorously when it is simply expecting
food or when it is pleasantly surprised, but it wags
frenziedly at the appearance of a desirable social com-
panion (who, in real life, is all too often carrying a bowl of
food, leading to potential ambiguity regarding what the
tail is responding to). It is noteworthy that opioid recep-
tor blockade is very effective in increasing tail wagging.
Perhaps this is because the animal's internal state has
been shifted toward loneliness (i.e., the "panic" system
has become more active). The failure of the same experi-
ment to produce panic attacks in humans (Klein) is in-
teresting, but not inconsistent with our work in the
area.3 Although I would agree that pleasure is a concept
we should continue struggling with in the animal labora-
tory, until it can be measured in a convincing fashion, it
contributes little to our understanding of animal behav-
ior. However, if we do reach a stage where such pro-
cesses can be analyzed in animals, I suspect that plea-
sure will be best understood in the context of
homeostatic principles that help sustain the integrity of
the body as well as the species.
A number of commentators were concerned about my
brevity in describing how complex human emotions may
arise from the more basic ones (Averill, de Sousa,
Lazarus, Solomon). I suggested only that they may be
blended from the activities of the fundamental systems.
This, of course, is the traditional statement of ignorance
about the matter, and no commentator suggested an
alternative view. Still, at a rational level (if Klein will
again permit me to use the term loosely), I would argue
that (contrary to Solomon's opinion) it is reasonable to
consider jealousy (at least from love's perspective) as an
admixture of panic, rage, and expectancy: As one feels
abandoned (panic), and somewhat angry (rage) toward
the person whose behavior threatens the existing bond,
there may still be a longing that things will change
(expectancy). Similarly, shame is a very important
human feeling (Izard, Katz, Jaynes); however, I do not
think it is a fundamental process, but one engendered by
diminished self-esteem (will-to-power), coupled with the
implicit threat of retribution, be it physical punishment
(i.e., fear) or withdrawal of social support (i.e., panic).
Whether my introspective distillation of experience has
any generality can be evaluated empirically, and if it
does, my major point would be that such results may tell
us something about interactions that can exist among the
underlying primary emotive circuits. Presumably the
mysterious glue that will render these mixtures a single
compound (de Sousa) resides in the semantic attribution
faculties of the human brain.
In attempting to simplify matters, my target article
may have portrayed emotive circuits too much as mono-
lithic entities responding to major life emergencies
(Lyons); to understand their role in behavior, we also
need to imagine their role in the organization of every-
day affairs. I suspect that these circuits help control
behavior throughout the day, and modestfluctuationsin
them may evoke diverse affective states with distinct

names. Indeed, some remarkable phenomena could
emerge from the waxing and waning of activity in these
circuits. For instance, Clynes describes the positive af-
fect that can be experienced during music expressing
grief. This may seem paradoxical, but it is quite reason-
able from the viewpoint that music capable of activating
the sorrow-panic system should yield successive in-
creases and decreases in the activity of the underlying
circuit. Thisfluxshould repeatedly evoke opponent neu-
ral processes, and hence may be moving indeed. Al-
though our cognitive appraisal may add to its beauty, I
suspect that it would all sound rather flat if our limbic
circuits did not resonate with the music.
If one attempts to visualize the activities of the under-
lying neural mechanisms on a realistic time scale, the
behavioral and psychological possibilities within these
systems are vast. For instance, perhaps I can allay
Klein's and Klinger & Kemble's concern about positive
affective states existing during both increases and de-
creases in the activity of the expectancy circuit, by spec-
ulating how this and related systems operate in real life.
When a hungry animal is confronted by an enticing odor
(i.e., one that unconditionally evokes activity in the
expectancy system), the aroused dopamine circuits (es-
pecially the mesolimbic ones) would induce the animal
to move forward (by activating appropriate motor sub-
routines), and there would be sequential decreases and
increases of activity in the system as the animal repeat-
edly tracked and mistracked the odor trail (perhaps
yielding minijoys and mini-engagements of an associa-
tive reinforcement network). When the animal has been
brought infinitesimally close to the object that provoked
the foraging, the close interaction with the object would
provide adequate stimulation for consummatory reflexes
to be activated. During the consummatory act, there
would be active inhibition of all emotive systems, includ-
ing rage. Thus, there would be plenty of opportunity for
the generation of "pleasure, " "joy," what have you,
during both the appetitive and the consummatory
phases of behavior, although they may well differ in
kind.
Although this may sound like a simplistic approach to
one of the most important aspects of mental life (Sol-
omon), the existing alternatives are closer to art than to
science. In any case, the analytical view I advocated
should not be considered slighting to the spectrum of
affects. Consider the beautiful paintings created from a
few colors, the music that arises from the vibrations of a
few strings and the diversity of life controlled by the
sequences of four nucleotides. The claim that subcortical
circuits are not essential for having emotions (Solomon)
is contradicted by the vast neuropsychological literature
in humans (Heath; Kelly 1980) and is similar to a belief
that the aesthetic riches of our world can be experienced
without the media in which they are expressed.
There was abundant disagreement among the com-
mentators as to the number of primary emotions that
exist - similar to the disagreements I portrayed in the
historical section of my article and contrary to the in-
ferences of Averill and Royce, my mention of Willis was
not favorable, but an illustration of nonempirical, tax-
onomic nitpicking). So, to shed more light on the issue, I
turn from my desk to my six-year-old daughter playing at
my feet.
Response/Panksepp: Psychobiology of emotions
"Tiina, can you tell me something? [She looks up
agreeably.] How many emotions are there?"
"What's an emotion?" she queries.
"Mm . . . it's the way we feel. How many different
ways can we feel?"
She places a finger to her lips and looks briefly
puzzled, and then rattles off, "Happy, mad, sad. . . . Is
that right, Daddy?"
"I'm not sure - you tell me. Are there any more?"
"Mm, yes, yes, scared! Is that right?"
"You tell me."
"Mm . . . mm . . . mm . . . frowned? Are there any
more?" [She is beginning to look exasperated.]
"Hey, that's really good. Can you show me all those in
faces?"
As I say "happy," she smiles and jumps up and down
clapping her hands; as I say "mad," she frowns, clenches
her jaw, and more or less growls at me; as I say "sad,"
she pantomimes the mask of tragedy; as I say "scared,"
she retracts her torso, balloons her eyes, and shows a
frightened mouth; as I say "frowned," she looks puzzled,
scratches her head a little, and finally crunches her face
up in a way that communicates little to me.
Granted that "happiness" is the positive affective rep-
resentation of desire (expectancies) fulfilled, and that the
labels "mad," "sad," and "scared," can be translated as
"anger-rage," "sorrow-panic," and "anxiety-fear" re-
spectively, a little girl, on the threshold of her age of
reason, with little hesitation has spilled forth some simple
truths about basic human emotions. They were also clear
to Cogan (and I still believe - contrary to the opinion of
Royce - that he deserves as close a reading as modern
writers on the topic). It is also gratifying that this simple
scheme is compatible with some current empirical work
being done with humans (Clynes, Heath, Izard, Klein),
and that it can also be related to metaphysical conceptions
of human emotionality (de Rivera).
Cognitive mechanisms in emotions. Many commenta-
tors with philosophical and cognitive orientations took
issue with the simplicity of my scheme, and pointed to the
many ways in which affective processes are governed by
human thoughts, belief systems, and perceived relation-
ships among events (Arnold, Averill, Delgado, de Sousa,
Jaynes, Lazarus, Lutz, Morton, Solomon, Royce, Stock-
er). I do not disagree, but most of this criticism is
misplaced, arising from a misconstrual of my position on
the utility of introspection in psychobiology or a failure to
appraise realistically the limits inherent in a "general"
(i.e., cross-species) theory of emotions. While on the one
hand I was being congratulated (de Sousa) and, on the
other, criticized (Arnold, Averill) for not having used
introspection adequately myself, a critical point of my
analysis was being overlooked - that introspection has to
be coupled with neuroscience to be scientifically useful.
This self-imposed constraint limited my introspection
largely to matters of taxonomy; but for strict empiricists,
even that was too liberal (Anisman & Zacharko, Ursin,
Vanderwolf & Goodale). However, I regard the higher
order introspective insights provided by some of the less
empirically constrained commentators, whether or not
they correspond to my own personal insights, as scien-
tifically premature until they are accompanied by testable
neurological hypotheses.
BEHAVIORAL AND BRAIN SCIENCES (1982)3 455
Response/Panksepp: Psychobiology of emotions
Many of the cognitive issues raised may also be irrele-
vant to a "general" theory of emotions. They are more
pertinent to "special," nonbiological theories of human
emotion (e.g., the Class II passions of Cogan), which
may be of limited relevance to understanding the behav-
ior of other species. Still, I believe that to understand
those higher emotions, we will have to clarify how the
basic ones are generated. The human brain's ability to
elaborate higher feelings is probably still integrally
linked to basic brainstem mechanisms we share with less
complex creatures, although we may, of course, encoun-
ter totally new organizational principles at higher levels
in various species. For instance, the massive develop-
ment of the human fontal lobe and a paramedian lobe in
cetacean species (Morgane, Jacobs, & McFarland 1980)
may indicate unique principles of organization involved
in human expectancy and in dolphin social-emotional
systems.
There is a major empirical issue embedded in the
perspective that cognitive processes have to be consid-
ered in explaining the genesis of emotions. At the level
of neural organization, the question reduces to: Do emo-
tions arise primarily from brain areas that mediate cogni-
tive processes, or elsewhere? The evidence seems rea-
sonably clear. Cognitive processes appear to be
mediated primarily by neocortex, tissue which is largely
an outgrowth of rapidly-firing neural systems specialized
for processing exteroceptive information (i.e., they arise
from the "somatic" nervous system comprising the
thalamic-neocortical axis). Emotional processes, on the
other hand, appear to be largely an outgrowth of slowly-
firing neural systems designed to collect interoreceptive
information (i.e., they are part of the "visceral" nervous
system comprising the hypothalamic-limbic axis). Al-
though these distinct nervous systems interact at all
levels of the neuraxis, anatomical evidence suggests that
visceral-limbic control of thalamic-neocortical activity
is much greater than the reverse. This is congruent with
common subjective experience, for, as noted by many
observers from the Greeks on, emotions appear to be
more influential in controlling thought than vice versa
(and, hence, this may be a fine example of how intro-
spective experience tells us something concrete about
brain organization).
This is not to deny that cognitive processes can also
influence emotive circuits. People can consciously exert
some control over their emotions. Similarly, the con-
templation of a trying situation can arouse one emo-
tionally even after the precipitating events. Although
these corticofugal controls are much less powerful than
the corticopedal ones, there is room for an updated
James-Lange perspective on the genesis of emotions.
We must remember that, at the turn of the century,
there was no substantive evidence that a visceral ner-
vous system existed in the brain, and James's suggestion
that somatic actions precipitated peripheral visceral
changes which were then perceived as emotions, could
be recast with an eye to current knowledge of brain
organization. Thus, a modern form of the theory could
assert that somatic-thinking processes evoke activity
within visceral-limbic areas of the brain, and this then
leads to affect-laden perceptions in neocortical areas.
Such recurrent controls no doubt exist and probably
contribute to the growth of personality traits and the
456 BEHAVIORAL AND BRAIN SCIENCES (1982)3
development of psychiatric disorders. It is here, at the
juncture of visceral-emotional and cognitive-somatic
processes of the brain that a hard-core reinforcement
principle, so uncongenial to humanists, may be essential
for understanding the sources of behavioral control.
It is noteworthy that another brainstem-cortical pro-
cess has been demonstrated to function by a causal di-
alogue similar to the one described above. While the
executive mechanism for rapid eye movement (REM)
sleep resides in primitive, autonomic areas of the lower
brain stem (Jouvet 1972), it is capable of inducing hallu-
cinatory dreaming activity (typically affect-laden) in fore-
brain areas of higher animals. Humans who have lost
their frontal lobes still exhibit normal physiological man-
ifestations of REM, but the hallucinatory dream content
has been eliminated by the surgery (Jus, Villeneuve,
Pires, Rachance, Fortier, & Villeneuve 1973). In intact
individuals, these cortical consequences of REM sleep
presumably exert a reciprocal organizing effect on the
behavior of the waking organism.
In the same way, executive systems for the various
emotions may arouse certain types of thoughts and feel-
ing states, elaborated in specific zones of cortex and
related basal ganglia, and reciprocal controls from these
higher areas may exert feedback control over subsequent
expression of emotions. Expectancy circuitry may in-
teract in a primary sense with frontal neocortex and basal
forebrain nuclei (e.g., substantia innominata and nucleus
accumbens), fear and rage with temporal cortex and
amygdaloid nuclei, and panic with cingulate cortex, sep-
tal nuclei, and bed nuclei of the stria terminalis. Nearby
basal ganglia (globus pallidus and corpus striatum) may
contain higher order motor subroutines that are acti-
vated by the emotive command systems. By such re-
ciprocal interactions, emotive command circuits may
help precipitate much richer internal states in humans
than might have been apparent in my simple taxonomic
analysis.
If, after all this, one is still inclined to believe that
emotions are basically cognitive, one need but consider
the phylogenetic and ontogenetic evidence. As evolution
provided ever-increasing ability for the brain to think (as
indexed by species-typical expansions of multimodal as-
sociation cortices), the basic affective expressions of or-
ganisms remained relatively stable. To rephrase my re-
sponse's opening remarks, this suggests that affective
representations of reality were built into brain tissue
before complex cognitive representations of reality. Sim-
ilarly, during ontogenetic development of all mammals
(at both neural and behavioral levels), the basic affective
faculties mature earlier than cognitive abilities, indicat-
ing their primacy in the evolution of brain organization.
Although many contemporary thinkers may prefer not to
dichotomize these processes (since they are so thor-
oughly blended in adult subjective experiences), the
neurological data confirm that a dichotomy exists, and
indicate the primacy of the affective rather than the
cognitive faculties.
Yes, psychobiology does need more input from cogni-
tive psychology (Morton), but that discipline has yet to
take up the challenge of dealing effectively with basic
subcortically organized brain processes. It has generally
chosen to focus more on those complexities of human
mind that distinguish us from other animals. Psycho-
 Response/Panksepp: Psychobiology of emotions
biologists might benefit from simple and clear human more tantalizing behavioral options are offered. Similar-
experiments focusing on those basic passions we still ly, rats are more likely to exhibit their own characteristic
share with other animals; and, I suspect, cognitive psy- foraging behaviors - investigation and object carrying -
chology would also profit from such endeavors. rather than specific consummatory responses when this
circuit is activated. I suspect that the consummatory
Facts, fantasies, and predictions. Anisman & Zacharko behaviors evoked by such brain stimulation have long
suggest that my system contains too few equations to misled us about the nature of the underlying circuitry.
solve for too many variables. It is understandable that Thus, I am suggesting that a common denominator may
the simplicity of my proposal might lead to that conclu- underlie the diverse behavioral effects that have been
sion, but such a conclusion overlooks the key element of observed from stimulating homologous neural tissue in
the system — a reinforcement process (in the associative different animals. (As discussed elsewhere, in Panksepp,
rather than the reward sense) that is linked to consum- 1981a, this position does not imply that there are no
matory responses and hence to a rapid lowering of ac- drive-specific homeostatic detector circuits in nearby
tivity in an emotive system. Considering that each emo- areas of the hypothalamus.)
tive system can simultaneously bias a number of flexible The fact that one can get distinct types of behavior
behavioral options, such a reinforcement principle could from different zones of the hypothalamus is not an argu-
mold many behavioral habits from a limited number of ment against my perspective; rather, it is congruent with
command circuits. Behavioral variation under similar it. Transitions from stalking to flight to affective displays
conditions may also be due to differences in species- that Siegel describes are the very evidence on which my
typical behaviors that are sensitized by emotive circuits proposal was based. But these distinct classes of behav-
as well as by variations in the degree and types of in- ior are taken as evidence for general control circuits, as
teraction among the circuits. In addition, as noted be- opposed to rigidly coded neural pathways governing
fore, pain evoked by shock may activate several emotive unitary behavioral acts. Indeed, the experientially in-
systems, yielding confused patterns of activity within duced interchangeability of stimulus-bound behaviors
complexly interrelated circuits. Of course, these logical that Valenstein, Cox, and Kakolewski (1970) originally
alternatives are difficult to evaluate with existing described appears to have been based on the study of a
techniques. single system, the one that is in the present context
Many examples of animal behavior patterns that are labeled the expectancy circuit. My modified non-
difficult for the theory to explain can be derived from specificity hypothesis does not predict that stimulus-
laboratory experiments that constitute ecologically inval- bound flight or rage could be changed to any other type
id experimental paradigms - catch-22's that fail to clearly of emotive behavior, even though considerable response
differentiate underlying control systems. Has an animal molding may transpire within an individual system.
in nature ever been confronted by anything resembling I have previously observed that the specific consum-
lengthy sessions of signaled and unsignaled shock? matory behavior evoked by activation of the expectancy
Moreover, the tradition in experimental psychology of circuit is influenced by the personality of a rat (Panksepp
studying animals socially isolated from conspecifics for 1971); Siegel claims that is not the case in cats. I suspect
prolonged periods in extremely monotonous surround- it is, however, for Hutchinson and Renfrew (1966) re-
ings may be a poor basis for drawing meaningful conclu- ported stimulus-bound feeding from the LH of many of
sions about normal animal behavior. Challenging the their animals exhibiting stalking attack, while Flynn
nervous system with high doses of psychoactive drugs, (1967) and Siegel have noted that it is a rare occurrence.
which drive neurochemical systems beyond levels ever Indeed, Siegel's observation that the most ferocious
experienced in real life, likewise seems a dubious ap- stimulus- bound attack is often obtained from the most
proach for deriving credible conclusions about the oper- placid cats may reflect the operation of a personality
ations of the brain. Perhaps the same could be said about factor (paradoxical though it may seem if one does not
brain stimulation, but as noted by Heath, artificial consider adaptation-level factors). Jouvet (1979) has also
arousal of these subcortical circuits in humans yields noted that rage is an especially striking characteristic of
subjective feelings that are perceived as integral parts of dream-related activity in some normally placid cats.
consciousness rather than as sham responses. Katz questions the interpretation of several stimulus-
Siegel raises several key issues over which there is bound behaviors described in my 1971 paper, specifical-
considerable confusion concerning stimulus-bound be- ly asking why leaping behavior is now taken to indicate
haviors evoked from the hypothalamus. For instance, he the location of a "panic" system, when it was originally
argues that a cat's disregard of food in favor of stalking used to characterize "rage" placements. The confusion
behavior during lateral hypothalamic (LH) stimulation here is simple: I have not changed any interpretations,
argues against the nonspecificity hypothesis I espoused. but merely expanded on observations deemed secondary
This, I believe, reflects a misunderstanding of my defini- to the aim of the original study. That paper described
tion of nonspecificity. It is not that there is unlimited several hypothalamic zones from which aggression could
plasticity to the behaviors evoked by hypothalamic stim- be elicited in rats; at some sites the attack was accom-
ulation, but merely that the stimulus-bound consumma- panied by flight (leaping), probably due to emotive sys-
tory responses typically reported represent epi- tem overlap. However, there was no theoretical discus-
phenomena of more generalized behavioral tendencies. sion of a series of electrode sites above the optic chiasma
Thus, I view the stalking behavior of the cat as a species- that yielded explosive behaviors. I wrote, "At current
typical foraging behavior, arising from the arousal of the onset, these animals would freeze, and after a variable
expectancy command circuit; it is gratifying that cats are period of time that could last for several minutes, they
rarely fooled into exhibiting consummatory acts when would explode (i.e., run around wildly bounding off the

BEHAVIORAL AND BRAIN SCIENCES (1982)3 457

Response/Panksepp: Psychobiology of emotions
sides of the test chamber). Suddenly, they would again
stop all activity and sit quietly, breathing deeply. These
animals never exhibited any aggression" (1971, p. 323;
sites indicated by crosses in histology of Figure 1). The
evoked behavior did not resemble the well-directed
flight obtained from more dorsal sites, and I now enter-
tain the idea that such explosive episodes are the rat
homologue of "panic attacks." Still, I acknowledge that
the anatomical designation of panic is more speculative
than that of the other emotive circuits; and perhaps
distress vocalization sites found in diverse brain areas
may constitute a better estimate of the trajectory of the
panic system (Herman & Panksepp 1981).
Katz also questioned whether existing evidence ne-
cessitated a social cohesion (i.e., panic) system indepen-
dent of pain systems. I have discussed the possible evo-
lutionary links between pain and separation distress
previously (Panksepp, Herman, Conner, Bishop, &
Scott 1978; Panksepp, Herman, Vilberg, Bishop, &
DeEskinazi 1980; Panksepp 1981b), but would re-
emphasize that the induction of separation-distress vo-
calizations does not require any noxious stimulation,
merely the removal of a young animal from its social
environment. The reflexlike nature of this response sug-
gests an intrinsic brain system designed to perceive and
respond to social separation. The fact that this separation
response is exquisitely sensitive to opiates - indeed,
considerably more sensitive than traditional laboratory
measures of analgesia - further suggests that the brain
systems for the mediation of social affect can be dis-
tinguished from pain systems (at least as traditionally
understood). In fact, brain sites from which distress
vocalizations can be elicited do overlap with opioid sys-
tems, but not with classical pain pathways - except in
the lower brain stem. Of course, other emotional sys-
tems also participate in social relationships (de Rivera),
but the evidence suggests that one in particular ("panic")
has evolved specifically for the establishment and main-
tenance of social bonds. Although the affect conveyed by
arousal of this system is aversive, the return journey is
presumably pleasurable. Perhaps the waxing and waning
of activity in this and closely related circuits contribute
to the feeling called love.
I do not claim that emotive systems start in the hypo-
thalamus, as suggested by Delgado; only that the hypo-
thalamus is presently an ideal brain area in which to
conduct systematic analysis of most of these systems.
Indeed, the panic circuit may primarily flow through
dorsomedial reticular areas of thalamus that are linked
with the bed nuclei of the stria terminalis and cingulate
gyms (i.e., areas where distress vocalizations can be
elicited). Surely, the ramification of emotive command
circuits through the neuraxis will yield many surprises.
For instance, while Averill is amused by the introspec-
tively misguided notion of Willis that emotions may be
elaborated by the cerebellum, Heath as well as Bernt-
son, Potolicchio, and Miller (1973) are demonstrating
the participation of deep cerebellar nuclei in the organi-
zation of emotive behaviors. It is unfortunate that the
anatomy and properties of these important brain systems
are still not well understood.4 Although emotive circuits
may not be as hard wired as a radio (Averill, Stocker),
the basic plan appears to be laid down genetically, with
life experiences (especially as manifested in juvenile play
458 BEHAVIORAL AND BRAIN SCIENCES (1982)3
and in neural storms of REM sleep) probably refining
the intrinsic competence of these systems.
Because I related activity in the expectancy command
circuit to homeostatic detector systems of the body,
several commentators (Izard, Lyons, Zuckerman) con-
cluded that I was espousing a drive-reduction approach
to understanding emotional circuitry. This interpreta-
tion is wrong. Although homeostatic detectors probably
modulate activity in these circuits, each system is under
the influence of a variety of incoming stimuli. For exam-
ple, any novel stimulus should be capable of provoking
activity in the expectancy system, thereby precipitating
investigatory activity. The more pertinent the stimulus
is for the animal's survival, the more likely it is to sustain
activity in the emotive system. Also, I believe that the
expectancy system has a resting level of activity (yielding
a baseline level of exploration, curiosity, and interest in
the world), which is increased not only by biologically
important events but also by reafferents from higher
areas of the brain. Invigorated foraging activity could be
reduced in the short term by various consummatory
activities, and quelled for a longer term by complete
satisfaction of the organism's needs. Such home-
ostatically induced circuit stabilization may permit hu-
mans to pursue their cerebral interests. This is not a
simple drive-reduction view, even though implicit in my
argument is a foraging-reduction hypothesis of appe-
titive reinforcement.
Toates need not worry that the approach I have taken
will stagnate at some sterile mentalistic level instead of
proceeding to the analysis of mechanisms of action. The
aim of the target article was to describe the types of
emotive processes that exist in the brain, so as to under-
stand ultimately how the systems operate. I have gone
into more detail on possible mechanisms elsewhere
(Panksepp 1981a; 1982b), but to further extend one tan-
talizing promissory note, I would resketch what the key
mechanism underlying autoshaping may look like: if
there is a foraging-expectancy system, which activates
forward directed motor sequences interspersed with
various species-typical investigatory activities, then au-
toshaping is likely to be mediated by that circuit. Al-
though the rat will need various other psysiological sys-
tems to express itself (including legs and spatial
representation abilities), I would suggest that the essen-
tial process underlying autoshaping is the capacity of
previously neutral stimuli to conditionally evoke activity
in the expectancy command circuit. Thereby foraging
will come to be directed at reward-related stimuli. In
line with that hypothesis, Phillips, McDonald, and
Wilkie (1981) recently reported that DA receptor block-
ade inhibits autoshaping. Although a computer-rat may
help us refine and sharpen our thinking about a prob-
lem, that, I trust Toates agrees, is no substitute for
physiological research (Panksepp 1979b).
Averill and Plutchik question my introspectively de-
rived hypothesis that the "rage" and "expectancy" cir-
cuits inhibit each other. Plutchik bases his skepticism on
the supposition that aggression and sex are often found
together. In fact, sexually related "aggression" is proba-
bly an expression of dominance (will-to-power) rather
than anger ("rage"). Averill's observation that the "expec-
tation" of revenge must be attended by a certain pleasure
highlights once more the connotative difficulties of terms

such as expectation and pleasure, and may indicate how
the fantasy-creating mechanism of the human mind inter-
mittently projects itself forward toward the hope of what
could be, even during the worst of circumstances. Still, at
the moment of pure rage, I believe the feeling of positive
anticipation dwindles toward the vanishing point. Only as
the consuming passion of anger wanes, will opponent
neural processes (Solomon 1980) come into play (e.g.,
target article, Figure 2). In any case, the facts of the
matter strongly suggest that lateral hypothalamic self-
stimulation (i.e., expectancy) and rage circuits mutually
inhibit each other (e.g., Hutchinson & Renfrew 1978).
Klinger & Kemble raise enough issues to fill another
Response. One of their major concerns was the paucity
of specific predictions in the target article. A series of
predictions for the expectancy system are spelled out
elsewhere (Panksepp 1981a; 1982b), so I shall only ad-
dress the key issue of how we might go about determin-
ing whether a neural system constitutes a command
circuit. Although the lesion approach has many short-
comings, it can tell us whether a neural system is neces-
sary for a certain class of behaviors. For instance, Fon-
berg (1972) has demonstrated how corticomedial and
basolateral amygdalectomy in the dog can have opposite
effects on appetitive and friendly behaviors, while a
combined lesion leads to reestablishment of normal be-
havioral balance. This suggests that a modulatory rather
than a command influence was disrupted by those le-
sions. If a complete rebalancing of appropriate target
behaviors can be achieved in animals whose hypothala-
mic emotive circuits are damaged, the command circuit
idea I have espoused would be seriously compromised.
Such data would constitute an especially compelling re-
futation of my position if destruction of the putative
command circuits is accomplished in young animals so
that little prelesion translocation of function could have
occurred. And what might the appropriate target behav-
iors be? For the foraging-expectancy circuit, I assume
that any of a multitude of tasks that required an animal to
search out goods from its environment would suffice.
For the rage circuit, a variety of frustration- and irrita-
tion-induced aggression tasks could be used. For the fear
circuit, I would think unconditional flight behaviors or
many avoidance tasks, where the aversive stimulus is not
present during the test, such as conditioned emotional
response paradigms, would be appropriate. For the pan-
ic circuit, natural separation-induced distress and social
contact maintenance behaviors seem to be reasonable
target behaviors. Furthermore, temporal analysis of cel-
lular activities in appropriate brain sites in relation to the
initiation of natural emotive behaviors (e.g., Adams
1979; Rolls 1980) will be essential for assessing the pri-
macy of these systems in the genesis of emotions.
Lyons raises enough issues for yet another Response,
but his central theme - that emotions can generate
maladaptive behavior patterns - can be handled briefly.
Certainly, every physiological system of the body is
susceptible to deranged functioning - to being over-
stressed and to becoming ill. As sketched in Figure 5 of
the target article, imbalances of emotional systems can
lead to short- and long-term psychological disorders. If
we wish to look at it simply from the "dis-ease" perspec-
tive (represented, perhaps, by the descending limb of
the inverted-U function relating arousal and behavioral
Response/Panksepp: Psychobiology of emotions
organization), the emotional world can be seen as one of
contradiction and confusion, but I would doubt that
evolution has constructed animal behavior on so shaky a
foundation.
As Royce argues, the complexity of emotions seems to
defy comprehensive coverage. As he notes, my paper
had little to say about individual differences and did not
delve into subjective attributes of emotions. Although
the former could be easily incorporated into the theory
by assuming that the vigor of the various emotive sys-
tems can be inherited and modified by various life cir-
cumstances, there is little of substance to be said about
those matters at the present time. Similarly, I was hesi-
tant to undertake anything more than a taxonomic analy-
sis at the subjective level, especially since practically
everything I would have to say on the matter would be
opinion and, at that, largely repetitive of Cogan's in-
sightful analysis. A seemingly endless series of books has
discussed subjective aspects of emotions during the last
several hundred years without promoting major scien-
tific understanding in this field; and, due to the growth
of behaviorism, as well as the impropriety of evoking
strong emotions in the human laboratory, the systematic
experimental analysis of subjective emotional experience
remains in its infancy. Quite independent of what intro-
spection may have had to say about emotion, brain phys-
iologists at the turn of the last century started to unearth
some fundamental mechanisms in ancient areas of the
brain that seemed to be essential substrates for emotion.
I believe that this branch of brain physiology has reached
a stage of development where further progress could be
promoted by modest doses of introspective data, es-
pecially the kind derived from sound empirical analysis.
If there is good alignment between demonstrable phys-
iological systems and introspective categories at a basic
taxonomic level, we may be in a position to proceed to a
deeper analysis.
Summary. We are far from realizing an empirically
grounded understanding of emotional processes in the
brain. The scheme I have presented is incomplete in
many ways, and many of the controversies, which have
paralyzed this field of inquiry, are well summarized in
the foregoing debate. Because of the remarkable diffi-
culty in achieving any consensus in this area, modern
neuroscience has failed to develop a coherent framework
for the study of emotions. The dilemma of human and
animal emotions remains an embarassing skeleton in the
vast cathedral of knowledge being constructed by scien-
tists working on the brain. Some hope that an under-
standing of emotions will be achieved by creeping up-
ward, synapse by synapse, from autonomic, brainstem
reflex circuits toward those higher order brain systems
that dictate the coherent organismic response. Perhaps
that "bottom-up" enterprise will work. However, the
complexity of the reticular neuropil may provide an
insurmountable barrier to the progress of a systematic
empirical march up the neuraxis, and the science of the
brain may be confronted with the need to make some
hard decisions on how intrinsic higher order brain func-
tions are to be categorized and studied.
It is unlikely that traditional animal psychology, in
which the operational referents for terms are based on
external events, will provide concepts adequate for the
BEHAVIORAL AND BRAIN SCIENCES (1982)3 459
Re/erences/Panksepp: Psychobiology of emotions
task. When diverse external and internal stimuli funnel
onto integrative-convergence pathways that elaborate
complex psychobehavioral tendencies in the brain, our
sources of knowledge other than sensory observation
may prove useful. To the degree that those integrative
circuits are class typical and reflected with some degree
of accuracy in human consciousness, we have few rea-
sonable alternatives but to begin systematically applying
our subjective insights to our scientific enterprise. How-
ever, perhaps there are no such convergence pathways,
and emotions are elaborated by a loosely organized al-
gebraic flow of diverse and dynamic influence through a
behaviorally unarticulated reticular net to functionally
fragmented final common pathways, as suggested by
Delgado. If that is the case, the specific outlook I have
recommended is not promising. Still, the fact that we
can evoke a number of distinct, well-organized (not frag-
mented) emotive behaviors from discrete zones of the
visceral brain by the mere application of electrical noise,
suggests that we mammals possess neural systems, pre-
pared through evolution, for achieving behavioral co-
herence at a moment's notice. To understand the func-
tional organization of psychobehavioral processes in the
brain, we must determine how such executive circuits
govern the diverse neurophysiological subroutines that
lead to the objective organismic changes we can see. The
major emotions appear to be elaborated by executive
circuits such as these, and they provide the raw material
from which a lasting understanding of key aspects of
mammalian behavior and the human psyche can be
constructed.
NOTES
1. Izard (1971) summarizes a substantial amount of cross-
cultural data indicating that the identification of affective facial
expressions is quite accurate (for a larger number of emotions
than the four basic ones I discussed). This is remarkable, since
a considerable amount of emotional behavior is contained in
postures and body movements (Clynes) that were not captured
in the still photographs used in those studies. Also, it is impor-
tant to remember that the sounds of emotions are as charac-
teristic as the faces (see Davitz 1964). Thus, I think it is safe to
assume that, if all these dimensions were combined in a emo-
tional recognition task, the "hit" rate for the four emotions I
have discussed would be very high indeed, even for young
children.
2. Klinger & Kemble correctly emphasize that my attempt
to attribute psychiatric disorders to individual emotive systems
(target article, Figure 5) is a great simplification, most notably
in the case of schizophrenia. This is especially true in light of
recent trends to develop substantive subclassifications within
the global diagnosis of this brain disorder. Still, there may be
some utility in the conceptual simplifications I suggest, since
they may be helpful in promoting and guiding further re-
search, especially with animal models. At a general semantic
level, a connection between schizophrenia and overactivity in
an expectancy system suggests that something may have gone
awry in a primitive "reality-creating" mechanism of the brain.
If a fundamental function of the expectancy system is to gener-
ate various search behaviors when an organism is confronted
by environmental stressors or homeostatic imbalances, the
vivid psychological changes that accompany the florid phase of
schizophrenia may become more understandable. When this
system is overactive, heightening of perceptual awareness and
sense of potential meaning in the world, a solution may arise
rather automatically from fluctuating activities of the underly-
ing neural substrate, which is promoting the invigorated search
460 BEHAVIORAL AND BRAIN SCIENCES (1982)3
behavior. Such environmentally inappropriate "solutions" may
crystallize into core delusional systems and henceforth exert
overriding influences on thinking patterns. On the basis of this
kind of reasoning, the widely discussed relationship between
schizophrenia and a form of creativity can be linked to a
common physiological substrate. If the normal function of the
system is to promote exploration and meaning-extraction from
the environment, the pathological expressions of the system
may be solutions that are not restrained by brain reality-testing
principles (reinforcement?), which, under normal circum-
stances, are closely coupled to environmental events. Of
course, a person who has lost touch with consensual reality will
receive diverse forms of social punishment, which will sec-
ondarily bring many other emotional systems into play.
From such a perspective, it is noteworthy that self-stimula-
tion along the trajectory of ascending brain dopamine systems
is our best current animal model for schizophrenia. Not only do
animals appear crazed while self-activating this system, but
practically every drug treatment known to worsen schizo-
phrenic symptoms increases self-stimulation, while those
which ameliorate schizophrenic symptoms, reduce self-stim-
ulation. Thus, keeping in mind some reasonable qualifications,
I do not think it implausible that far lateral hypothalamic self-
stimulation and certain forms of schizophrenia reflect the un-
bridled activity of a brain system whose normal function is to
mediate expectancies - as expressed in foraging at lower levels
of the neuraxis and as a search for meaning at higher levels of
brain organization.
3. We have not been able to provoke panic attacks (as
measured by the induction of separation-induced crying) in
young socially housed dogs, guinea pigs, and chicks with nalox-
one. We can only facilitate the crying process once it has
already been precipitated, and of the species tested, this works
only for guinea pigs and chicks. In dogs, the naloxone increases
another care- or contact-soliciting response - tail wagging. This
suggests that brain opioid systems provide a modulatory rather
than a triggering influence over the relevant circuitry; accord-
ingly, a better test of opioid participation in human panic
attacks would be to determine whether chronic opiate receptor
blockade increases the frequency and intensity of panic attacks
or whether opiate receptor agonists can quell panic attacks in
the same way separation distress is reduced (Panksepp et al.
1980).
4. There is much more evidence concerning trans-
hypothalamic neural systems than I summarized in my target
article. Unfortunately, no anatomically identified circuit has
been firmly related to an emotional process. For instance,
although it is reasonable to hypothesize that one emotive com-
mand system (perhaps fear) is partially isomorphic with long-
axoned amygdalo-tegmental (Hopkins & Holstege 1978) and
hypothalamo-tegmental systems (Swanson & Kuypers 1980),
and their ascending reciprocal circuits, critical functional data
remain unavailable. Although such circuits have anatomical
characteristics that could coordinate diverse physiological and
behavioral processes, only recently has it become feasible to
apply neural and behavioral mapping approaches in the same
animal (Siegel). Only after concurrent behavioral, neuro-
anatomical and neurochemical mapping has been successfully
conducted at brain sites that yield stimulus-bound emotive
behaviors will it be possible to ask precise questions concern-
ing synaptic interactions between circuits (Figure 2).
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Call for Papers
Investigators in
Psychology,
Neuroscience,
Behavioral Biology, and
Cognitive Science
Do you want to:
• draw wide attention to a particularly
important or controversial piece of work?
• solicit reactions, criticism, and feedback
from a large sample of your peers?
• place your ideas in an interdisciplinary,
international context?
and
an extraordinary journal now in its fifth year, provides a
special service called Open Peer Commentary to re-
searchers in any area of psychology, neuroscience,
behavioral biology or cognitive science.
Papers judged appropriate for Commentary are circulated
to a large number of specialists who provide substantive
criticism, interpretation, elaboration, and pertinent com-
plementary and supplementary material from a full cross-
disciplinary perspective.
Article and commentaries then appear simultaneously with
the author's formal response. This BBS "treatment"
provides in print the exciting give and take of an interna-
tional seminar.
The editor of BBS is calling for papers that offer a clear
rationale for Commentary, and also meet high standards of
conceptual rigor, empirical grounding, and clarity of style.
Contributions may be (1) reports and discussions of empiri-
cal research of broader scope and implications than might
be reported in a specialty journal; (2) unusually significant
theoretical articles that formally model or systematize a
body of research; and (3) novel interpretations, syntheses or
critiques of existing theoretical work.
Although the BBS Commentary service is primarily devoted
to original unpublished manuscripts, at times it will be ex-
tended to precis of recent books or previously published
articles.
Published quarterly by Cambridge University Press. Edi-
torial correspondence to: Stevan Hamad, Editor, BBS,
Suite 240, 20 Nassau Street, Princeton, NJ 08540.
" . . . superbly presented . . . the result Is
practically a vade mecum or Who's Who in
each subject. [Articles are] followed by pithy
and often (believe it or not) witty comments
questioning, illuminating, endorsing or just
plain arguing... I urge anyone with an Inter-
est In psychology, neuroscience, and behav-
ioural biology to get access to this jour-
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"Care is taken to ensure that the commentaries
represent a sampling of opinion from scientists
throughout the world Through open peer com-
mentary, the knowledge imparted by the target
article becomes more fully integrated into the
entire field of the behavioral and brain sciences.
This contrasts with the provincialism of special-
ized journals . .."—Eugene Garfield Current
Contents
"The field covered by BBS has often suf-
fered In the past from the drawing of battle
lines between prematurely hardened posi-
tions: nature v. nurture, cognitive v. behav-
iourist, biological v. cultural causation....
[BBS] has often produced important articles
and, of course, fascinating Interchanges
the points of dispute are highlighted if not
always resolved, the styles and positions of
the participants are exposed, hobbyhorses
are sometimes ridden with great vigour, and
mutual incomprehension is occasionally
made very conspicuous.... commentaries
are often Incisive, integratlve or bring highly
relevant new information to bear on the sub-
ject."—Nature
" . . . a high standard of contributions and dis-
cussion, it should serve as one of the major
stimulants of growth in the cognitive sciences
over the next decade."—Howard Gardner
(Education) Harvard
" . . . keep on like this and you will be not
merely good, but essential..."—D.O. Hebb
(Psychology) Dalhousle
" . . . a unique format from which to gain some
appreciation for current topics in the brain sci-
ences . . . [and] by which original hypotheses
may be argued openly and constructively."—
Allen R. Wyler (Neurological Surgery)
Washington
" . . . one of the most distinguished and use-
ful of scientific Journals. It is, Indeed, that
rarity among scientific periodicals: a crea-
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"I think the idea is excellent."—Noam Chomsky
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" . . . open peer commentary . . . allows the
reader to assess the 'state of the art' quickly
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vide a 'who's who' as well as the content of
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ological Structures
"... presents an imaginative approach to learn-
ing which might be adopted by other jour-
nals."—Library Journal
"Neurobiologists are acutely aware that
their subject is in an explosive phase of de-
velopment . . . we frequently wish for a fo-
rum for the exchange of ideas and Interpre-
tations . . . plenty of journals gladly carry the
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promoting ideas. Perhaps even more impor-
tant is the need for opportunities publicly to
criticize traditional and developing concepts
and interpretations. [BBS] is helping to fill
these needs."—Graham Hoyle (Biology)
Oregon