Table Of ContentAffective Neuroscience
SERIES IN AFFECTIVE SCIENCE
Series Editors
Richard J. Davidson
Paul Ekman
Klaus Scherer
The Nature of Emotion: Fundamental Questions Persons, Situations, and Emotions: An Ecological
Edited by Paul Ekman and Richard J. Davidson Approach
Edited by Hermann Brandstatter and Andrzej Eliasz
Boo! Culture, Experience, and the Startle Reflex
by Ronald Simons Emotion, Social Relationships, and Health
Edited by Carol D. Ryff and Burton Singer
Emotions in Psychopathology: Theory and Research
Edited by William F. Flack, Jr. and James D. Laird Appraisal Processes in Emotion: Theory, Methods,
Research
What the Face Reveals: Basic and Applied Studies Edited by Klaus R. Scherer, Angela Schorr,
of Spontaneous Expression Using the Facial Action and Tom Johnstone
Coding System (FACS)
Edited by Paul Ekman and Erika Rosenberg Music and Emotion: Theory and Research
Edited by Patrik N. Juslin and John A. Sloboda
Shame: Interpersonal Behavior, Psychopathology,
and Culture Nonverbal Behavior in Clinical Settings
Edited by Paul Gilbert and Bernice Andrews Edited by Pierre Philippot, Robert S. Feldman,
and Erik J. Coats
Affective Neuroscience: The Foundations of Human
and Animal Emotions Memory and Emotion
by Jaak Panksepp Edited by Daniel Reisberg and Paula Hertel
Extreme Fear, Shyness, and Social Phobia: Psychology of Gratitude
Origins, Biological Mechanisms, Edited by Robert A. Emmons
and Clinical Outcomes and Michael E. McCuIlough
Edited by Louis A. Schmidt and Jay Schulkin
Thinking about Feeling: Contemporary Philosophers
Cognitive Neuroscience of Emotion on Emotions
Edited by Richard D. Lane and Lynn Nadel Edited by Robert C. Solomon
The Neuropsychology of Emotion Bodily Sensibility: Intelligent Action
Edited by Joan C. Borod by Jay Schulkin
Anxiety, Depression, and Emotion Who Needs Emotions? The Brain Meets the Robot
Edited by Richard J. Davidson Edited by Jean-Marc Fellous and Michael A. Arbib
affective
neuroscience
The Foundations
of Human and
Animal Emotions
JAAK PANKSEPP
OXPORD
UNIVERSITY PRESS
OXFORD
UNIVERSITY PRESS
Oxford New York
Auckland Bangkok Buenos Aires Cape Town Chennai
Dar es Salaam Delhi Hong Kong Istanbul Karachi Kolkata
Kuala Lumpur Madrid Melbourne Mexico City Mumbai Nairobi
Sao Paulo Singapore Taipei Tokyo Toronto
Copyright © 1998 by Oxford University Press, Inc.
Published by Oxford University Press, Inc.
198 Madison Avenue, New York, New York 10016
www.oup.com
First issued as an Oxford University Press paperback, 2005
Oxford is a registered trademark of Oxford University Press
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording, or otherwise,
without the prior permission of Oxford University Press.
Library of Congress Cataloging-in-Publication Data
Panksepp, Jaak.
Affective neuroscience : the foundations of human and animal
emotions / Jaak Panksepp.
p. cm. — (Series in affective science)
Includes bibliographical references and indexes.
ISBN 0-19-509673-8; 0-19-517805-X (pbk.)
1. Emotions. 2. Emotions and cognition. 3. Emotions—Social
aspects. 4. Psychobiology. 5. Psychology, Comparative. I. Title.
II. Series.
BF531.P35 1998
156'.24—dc21 98-15955
1 3 5 7 9 8 6 42
Printed in the United States of America
on acid-free paper
I dedicate this book to my lost child, Tiina, and my friend Anesa,
who supported me when I was in need.
When the world came down upon me
and the sky closed like a door,
sounds filled my ears from far away.
I lay down on the floor.
And no one near could find me,
and nothing near was mine.
I sank into the floorboards
from the voices, soft and kind.
Until one thought got through to me,
one image filled my mind:
a pencil and a paper lying
close to hand, nearby.
Somehow I took them up and traced
one word and then the next,
until they linked together in a chain
that first perplexed the darkness
in my eyes, then,
rowing on my paper barque,
I soon was far away
and saw the water trail I'd left
rise up into a chain—
a ladder reaching high above
to light and sound and friends.
And that's how I climbed out
of the grief that has no end.
Anesa Miller,
A Road Beyond Loss, 1995
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Preface
The Aims of This Book
This book was written with the student firmly in mind, but it is not a traditional textbook. It
is an attempt to clarify the interrelations between brain and mind as expressed in the funda-
mental emotional processes that all mammals share. I will explore how our knowledge of
the animal brain can help clarify the affective nature of the human mind and how our ability
to appreciate the basic emotions of the human mind helps us understand the functional or-
ganization of the mammalian brain. This borderland of knowledge between the many disci-
plines that study the animal brain and the human mind—the various subareas of neuroscience
and behavioral biology and the many schools of psychology and philosophy—should be
rich in intellectual commerce. Unfortunately, this is not so. At present, these disciplines are
only slowly being introduced to each other, like Europe and the Far East a millennium ago,
and xenophobia prevails. Just as the trade routes between those distant cultures were opened
slowly by explorers and adventurers, interchange between the disciplines that view human
nature from above (i.e., the viewpoint of our recently evolved rational and cultural mind)
and those that view it from below (the ancient networks of our brains) remains tortuous and
unsatisfactory. This book offers one view of how the needed trade routes may be constructed
in the area of emotions and motivations.
A motive of the old explorers was adventure under the guise of economics—the search
for new vistas and knowledge in behalf of what is useful. The motives that underlie the writing
of this book are the same. The aim is to enrich our knowledge about the brain/mind inter
face, but the more primal motive is the adventure of exploring uncharted spaces. At times,
the conceptual paths of the brain/mind charts presented here will be like the routes on an
cient maps that never really depicted the immensity of the journey. At present, ignorance is
more abundant than knowledge in this field, often forcing me to oversimplify in order to
formulate any coherent ideas and explanations at all. It will be a while before we have a true
and lasting science of emotions as opposed to the fragmentary knowledge that now exists.
But instead of just summarizing a patchwork of existing theories that are endemic in the
field, I will aspire to provide a cohesive map to guide future navigations. We are still in the
early stages of analyzing the psychobiology of emotions, and many additional generations
of careful work will be needed before we have precise maps.
This book is dedicated to a new synthetic psychology of the future that will be more
catholic than the present variants—one that will be built jointly on evolutionary, neuro-
scientific, behavioristic, affective, and cognitive foundations. I will attempt to take nature
on its own terms by coming to grips with those intrinsic, ancient processes of the brain/
mind that mediate between environmental events and the natural classes of action tenden
cies that animals spontaneously generate in the real world. The basic premise here is that
the brain is a "symbolic organ" that reflects an evolutionary epistemology encoded in our
genes. The mammalian brain not only represents the outside world in symbolic codes based
on the properties of its sensory-perceptual systems but also has intrinsic operating systems
that govern ingrained psychobehavioral tendencies for coping with those ever-present chal-
lenges that our ancestors confronted in their evolution. Many of these operating systems
arouse emotional states, which are probably internally felt by other animals in ways not that
different from humans. While modern neuroscience has achieved great success in deciphering
viii PREFACE
the anatomical and physiological substrates of many sensory and motor processes at the
neuronal level, it has made only a modest beginning in deciphering the functional charac-
teristics of that "great intermediate net" that intervenes between inputs and outputs. Here, I
advocate the position that many intervening neuropsychological processes, such as the ba-
sic emotions, can now be understood in neuroanatomical, neurophysiological, and neuro-
chemical terms. But that level of understanding does require more theorizing and utiliza-
tion of indirect evidence than is common in the field.
This new, integrative form of psychobiology is still in its early stages. Its development
has been permitted only by the recent growth of neuroscience. The best evidence concern-
ing emotional mechanisms has emerged largely from brain research on kindred animals such
as birds, rats, guinea pigs, dogs, cats, and monkeys. Some comes from work on creatures
even "lower" in the evolutionary "bush," and an ever-increasing amount is also now com-
ing from studies of humans treated with various psychoactive drugs and hormones, as well
as those with brain damage. We must use many sources of knowledge to reveal the nature
of emotionality—what it really means to feel anger, fear, lust, joy, loneliness, happiness,
and the various other desires and vexations of the human "heart."
In my more optimistic moments, I hope that the lines of evidence summarized here could
serve as a foundation for a "new psychology" that recognizes that the discipline must be
grounded on solid neuroscience foundations. Although psychology can continue to deal with
the loftiest human aspirations, it also must become rooted in the evolutionary realities of
the brain if it is to become a true science. But that metamorphosis will be a difficult one,
since most of what psychologists do is not clearly linked to brain issues. Since few psy-
chologists are doing brain research, it is hard to convince them that their thinking should be
premised on a deep respect for and understanding of the organ of the mind. Substantive
neurobiological offerings are still not part of the traditionally mandated curricular require-
ment of many psychology programs. All too often the neural facts are offered in such a dry
manner that students shy away from immersion in such materials. My aim here has been to
provide a treatment with a bit more literary merit than may be typical for a book of this
type. I have aimed to maintain a friendly, readable style, in the hope of attracting the atten-
tion of many readers who truly wish to appreciate the underlying complexities of the human
mind and to understand how our highest aspirations often remain tethered to the values elabo-
rated by ancient parts of our animal brain.
I did choose to cast the present coverage in textbook format, with enough chapters for a
typical semester, divided into convenient thirds. Although this type of course is not yet a
traditional offering in psychology, I hope it will gradually emerge as one for advanced under-
graduates and graduate students studying the psychology of emotions and motivations, as
well as neuroscience and behavioral biology students who wish to have an appreciation of
functional processes shared by all mammals. Also, the work will cover issues that should
be essential for related disciplines such as neurophilosophy and biological psychiatry. In
sum, I have tried to write a book for those interested in psychology who wish to know more
about brain matters and for those interested in neurosciences who wish to know more about
psychological matters. I have tried to write in such a way that little neuroscience or psycho-
logical background is necessary to follow the story lines. At times, the going may be diffi-
cult, especially when it comes to the essential background issues presented in Chapters 4-6
on neuroanatomy, neurophysiology, and neurochemistry. However, I trust that even the nov-
ice will find them reasonably interesting.
I have also sought to keep the lay reader in mind by including enough general material
to make the more detailed materials worthy of continued attention. Many people are inter-
ested in gaining a new and better understanding of the evolutionary sources of the human
mind, which must largely be obtained from brain research. Unfortunately, there are few places
to go to obtain a substantive, yet interesting, coverage of emotional issues. Accordingly, in
this book I have taken a conceptual approach as much as an empirical approach. This is also
reflected in the referencing style, where I have tried to put ideas and carefully selected facts
in the foreground and to leave personalities who have done the hard investigative work, as
well as the myriad research details, in the background. I have used endnotes in preference
to the traditional name-date approach. I mention names only when we encounter the work
of investigators of considerable historical importance. However, the author index should
PREFACE ix
allow effective retrieval of those whose work has been covered. Obviously, there exists much
more raw material to be cited than I have chosen to cover. For instance, I recently received
two reviews on two of the topics covered here, hypothalamic control of feeding and aggres-
sion, and each cited more than 1,000 original pieces of research. It would have been pos-
sible to cite thousands of references for every chapter of the book, but my aim, more than
anything, was to generate a readable text. Thus, my strategy in referencing materials was to
focus equally on review articles and research reports, with a view to maximizing the possi-
bility that students could access related materials reasonably efficiently.
I have also tried to sustain a fairly natural style through the explicit recognition that
we humans are storytelling creatures. Our cultural evolution (and perhaps even our cere-
bral evolution; see Appendix A) has been guided by aeons of sitting around campfires,
sharing our deepest perspectives about the worlds in which we live. The best teaching
must try to rekindle this spirit around the intellectual campfire of the modern classroom.
Thus, one of my main goals is to encourage a renewed interest in the types of experimen-
tal inquiries that can take us toward a substantive understanding of how emotions are
organized in the brain. Unfortunately, in the present scientific climate (where we often
reward knowing more and more about less and less), there is remarkably little integrative
work by active investigators in the field and remarkably little work on the psychobiology
of emotions. Brain scientists are typically unwilling to use mentalistic words in discuss-
ing their empirical findings, and psychologists, because of their lack of training in the
neurosciences, are typically unable to link their psychological concepts to brain functions.
The present effort is based on the assumption that our ability to pursue such linkages,
first verbally and then empirically, is essential for future scientific progress in understand-
ing emotions. Accordingly, I have used the unusual literary device throughout this book
of labeling major emotional systems in folk-psychological terms, using capitalized let-
ters to highlight that I am focusing on certain necessary albeit not sufficient neural sub-
strates for distinct types of emotional processes.
Our stories and our semantic habits have profound consequences for how we proceed in
empirical inquiries, and as I will repeat, perhaps ad nauseum, there is really no other way to
obtain biological knowledge about emotional matters except through arduous brain research
(usually in other species), guided by meaningful psychological concepts. Thus, this book is
written especially for those students who wish to bridge psychological and neurological issues
in scientifically sound ways. For them, I have attempted to mill the abundant factual pep-
percorns into tempting conceptual spices. I have tried to make this difficult journey into the
brain as stimulating as possible without doing injustice to the facts, even though I had to
neglect many important lines of evidence to prevent the book from becoming too cumber-
some. In any event, I hope that there are a number of students of philosophy, psychology,
and the neurosciences who will find my efforts sufficiently refreshing and provocative that
they will eventually set out on their own empirical journey s in search of answers to the many
scientific questions that remain to be asked.
An Overview
This book is divided into three parts: (1) Background issues are discussed in Chapters 1-6,
(2) the primitive emotions and motivations are covered in Chapters 7-11, and (3) the social
emotions form the topic of Chapters 12-16.
In Part I, Chapters 1-3 will elaborate key concepts related to the functional systems in
the brain and taxonomic issues. Chapters 4-6 offer an essential background for understand-
ing the substantive individual topics covered in Parts II and III. Chapter 4 provides a rela-
tively user-friendly summary of neuroanatomy, Chapter 5 touches upon relevant aspects of
neurophysiology, and Chapter 6 addresses key neurochemical issues. These sections may
be tough going for readers who have little knowledge of the brain, but I have tried to make
them sufficiently concise and interesting that, after several readings, even a novice may gain
a sense of mastery of the material. I have also tried to write so that experienced readers will
enjoy them as thumbnail sketches of the enormous fields they are. Other basic background
issues, such as the details of sensory and motor processes, will not be covered here, since
