Gruter and Bohannan - 1983 - Law, Biology and Culture - The Evolution of Law

Law, Biology and Culture
The Evolution of Law
MARGARET GRUTER
&
PAUL BOHANNAN
EDITORS
Ross-Erikson, Inc.
SANTA BARBARA
Copyright @1983 Margaret Gruter and Paul Bohannan
Published by
ROSS-ERIKSON, PUBLISHERS, INC.
629 State St., #207
Santa Barbara, California 93101
Printed in the United States of America.
An earlier version was published in The Journal of Social and Biological Structures,
Volume 5, Number 4, October 1982 (Academic Press, London, New York, Toronto,
Sydney, San Francisco), as .'Proceedings of the First Monterey Dunes Conference,"
sponsored by The Goethe Institute, San Francisco, California, and The Hutchins
Center, University of California, Santa Barbara.
Design & typography by Jim Cook
Santa Barbara, California
Cover design by Frederick Usher
Contents
vii List of Participants
ix Foreword
xi Introduction
PART I
1 The foundations in law and morality
2 MARGARET GRUTER—Biologically based behavioral research and the

facts of law
15 RICHARD D. SCHWARTZ—On the prospects of using sociobiology in the

shaping of law: a cautionary note
7 E. ADAMSON HOEBEL—Anthropology, law and genetic inheritance
34 MANFRED REHBINDER—Questions of the legal scholar concerning the

so-called sense of justice
PART II
47 The search for the missing pieces: in biology
50 JANE GOODALL—Order without law
62 JUNICHIRO ITANI—Intraspecific killing among non- human primates
74 PAUL D. MacLEAN—A triangular brief on the evolution of brain and law
90 HUBERT MARKL—Constraints on human behavior and the biological

nature of man.
101 RICHARD D. ALEXANDER—Biology and the moral paradoxes
111 BARTLEY G. HOEBEL—The neural and chemical basis of reward: new

discoveries and theories in brain control of feeding, mating, aggression, self-
stimulation and self-injection
[v]
[vi]
PART III
129 The search for the missing pieces: in social science
134 CHRISTOPHER BOEHM—The evolutionary development of morality as

an effect of dominance behavior and conflict interences
147 PAUL BOHANNAN—Some bases of aggression and their relationship to

law
159 DONALD T. CAMPBELL—Legal and primary-group social controls
171 ROGER D. MASTERS—Evolutionary biology, political theory and the

state
191 Epilog
195 References
List of Participants
Richard D. Alexander- Museum of Zoology, Insect Division, University of

Michigan; Ann Arbor, Michigan 48109/USA
John H, Beckstrom -School of Law, Northwestern University; .357 East

Chicago Avenue, Chicago, IllinoIs 60611/USA
Christopher Boehm –Dept. of Anthropology & Sociology, Northern

Kentucky University; HIghland HeIghts, KY 41076/USA
Paul Bohannan -Dean, Social Sciences and Communication, University of

Southern California; Los Angeles, California 90089/USA
Donald T. Campbell -Maxwell School of Citizenship and Public Affairs,

Syracuse University; Syracuse, New York 13210/USA
William Durham -Department of Anthropology, Stanford University;

Stanford, California 94305/USA
Lawrence M. Friedman -School of Law, Stanford University; Stanford,

California 94305/USA
Jane Goodall- Post Office Box 727, Dar es Salaam, Tanzania
Joachim H. Gruter -158 Goya Road, Portola Valley, California 94025/USA
Margaret Gruter -158 Goya Road, Portola Valley, California 94025/USA
Bartley Hoebel -Department o/Psychology, Princeton University; Princeton,

New Jersey 08540/USA
[viii] LAW, BIOLOGY AND CULTURE
E. Adamson Hoebel- 2273 Folwell Street, St. Paul, Minnesota 55108/USA
Helmut Hofer -Department of Zoology and Comparative Anatomy,

University of Kassel; Heinnch-Plett-Strasse 40, D-3500 Kassel,
Federal Republic of Germany
Jay G. Hook -Department of Psychology, University of Houston; Houston,

Texas 77004/USA
Junichiro ltani -Faculty of Science, Laboratory of Physical Anthropology,

Kyoto University; Sakyo, Kyoto 606, Japan
Paul D. MacLean -Laboratory of Brain Evolution and Behavior; National

Inst. of Mental Health; 900 Rockville Pl., Bethesda, MD 20014/USA
Hubert Markl -Department of Biology, University of Konstanz; Postfach

5560, D-7750 Konstanz 1, Federal Republic of Germany
Roger D. Masters -Department of Government, Dartmouth College; Hanover,

New Hampshire 03755/USA
James Grier Miller -Hutchins Center for the Study of Democratic Institutions;
University of California, Santa Barbara, California 93106/USA
Manfred Rehbinder -School of Law, University of Zurich; CH-8032 Zurich,

Switzerland
Ernst Schuermann -Goethe Institute, San Francisco, California 94100/USA
Richard D. Schwartz -College of Law, Syracuse University; Syracuse, New

York 13210/USA
Harvey Wheeler -Institute of Higher Studies, Carpinteria, California

93013/USA
Foreword
The initiative for this symposium on law and behavioral research came from
Professor Manfred Rehbinder and Dr. Margaret Gruter, scholars of the law
with a strong interest in the life and behavioral sciences. They joined forces
in 1979 after independently realizing the impact this research could have on
their specialty. When the Goethe Institute of San Francisco, was invited to
cosponsor this event, I immediately accepted. The project found its ideal
American Co-sponsor, thanks to the assistance of Dr. Harvey Wheeler and
Professor Paul Bohannan, in the Hutchins Center for the Study of Democratic
Institutions of the University of California, Santa Barbara.
This publication is appearing simultaneously in the United States of
America in English{edited by Margaret Gruter and Paul Bohannan) and in
the Federal Republic of Germany {edited by Margaret Gruter and Manfred
Rehbinder). In the name of the Goethe Institute I would like to thank
everyone whose scientific contributions and organizational assistance made
this undertaking a success. I am certain that everyone involved in the
symposium will agree with me in singling out the hard work and
inexhaustible enthusiasm of Dr. Margaret Gruter for particular recognition.
Without her there would not have been a symposium or this publication.
During the preparation of the symposium I was often asked what the Goethe
Institute, as a general cultural institute of the Federal Republic of Germany,
was doing in such a purely scientific undertaking. While it is true that the
papers here are scientific, the topic under discussion is highly relevant, today
more than ever, to pressing problems in the political and social reality of our
nations. If the Goethe Institute is to live up to its declaration of purpose,
which is "to promote international cultural cooperation," then inquiries like
this book are a fundamental part of our work. I am extremely pleased and
grateful that we were invited to assist in this undertaking.
ERNST SCHUERMAN
The Goethe Institute, San Francisco
Cultural Center of the Federal Republic of Germany
Introduction
MARGARET GRUTER and PAUL BOHANNAN
This first Monterey Dunes conference is an integral part of what is

obviously a major thrust of both biology and behavioral science in the
last third of the twentieth century: a thorough examination of the
biological basis of behavior. We might well ask why such an obvious
area of study is, in many ways, so tardy an arrival.
Why did so many generations of scholars work so hard to create a
biology-free social science? There are, obviously, many reasons. At
least one of them grows out of the Western cultural premise that human
beings are "different" from animals-that we neither "descended"
(Darwin) nor "ascended" (Bronowski) from them. That premise is
manifest at many levels: linguistic, religious, and scientific. At the most
fundamental linguistic level, we find whole sets of words differentiating
the body parts of animals and humans. People have hands, but cats and
dogs have paws. The immediate response of speakers of Indo-European
languages to that point is that hands and paws are structurally quite
different. That is true-but not enough: for many, perhaps most,
languages do not make that distinction. In many African languages, for
example, the same word does for both. Indeed, it even does for "hoof."
The list could continue but that is not the point: the point is a more or
less out of awareness premise that people are so different from other
animals (and usually the word "other" does not appear when this kind of
statement is made) that you cannot even describe their body parts with
the same words. There are further linguistic dimensions to this Western
attitude toward animals: human beings, when they are at their worst,
"behave like animals" in accordance with "the law of the jungle." Thus,
we are likely to ascribe everything that we do not like about ourselves-
our least admired traits-to "the animal in us." The implication is that we
ought to get over being animals. Therefore, at either a conscious level or
an out-of-awareness level, there is a built-in premise that an
achievement as admirable and as “human" as "law" surely does not have
any animal base—indeed, law is, in the popular view, one of the signs
[xi]
that we can transcend our animal beginnings! Except, of course,
"when man, with his monopoly on law:" behaves like an animal. "
Our plea is that, in examining the biological infrastructure of law,
we—even those who obviously do not harbor any of these
prejudices and will tell us that these phrases are "just idioms"—do
not allow ourselves to be tricked by the idioms, as we always are
when we fail to examine "
Premises about the differences between human beings and animals also
lie at the basis of religion. Many a child is confused by the idea that he
or she has a soul (although in some religions, in some perioids of time,
there was some doubt about women) and so do parents and siblings—
but the family dog does not. The dog was banished from heaven just for
being a dog. The dog is of a "lower" order and apparently it is the
presence of the soul that creates the higher order. Of course, in the
process of growing up, those very children soon become involved with
more important issues. They become enculturated and forget the doubts
they felt about the missing soul of the dog. And in the next generation,
they again tell children that dogs and cats do not have souls, but human
beings have.
Although we talked about it very little during the conference, many

participants realized that besides the intellectual interest of the subject
matter, we also had an emotional stake in this matter. In differing
degrees, we all had difficulties in reconsidering some of our cherished
values that seemed to be threatened by the new biological findings and
by the tremendous consequences that might arise from tomorrow's
discoveries. Reality and utopia—law and justice. How do we arrive at
the concept of justice? It was suggested several times that we could not
deal rationally with an idea like justice. It was repeated that nobody can
be completely free of all prejudice or preconceived ideas when it comes
to something so fraught with personal conviction as justice. Our point
here, however, is that early behavioral sciences-despite writings by
Darwin and others on the biology of behavior-seemed always to let such
hidden premises (not to say sensibilities) get in the way of their
thinking. We are still struggling with them today. Westerners
"instinctively" (and much of this kind of "instinct" is, of course,
cultural) read out the biological dimensions from the human behavior
they regard as most uniquely human. Natural scientists do it as much as
social scientists though the arguments used to cover up their premises
may differ. We have tried not to do that-probably without total success,
for premises lie very deep. In this context, we must add another
difficulty that Western
[xiii ]
children sometimes experience-ideas about the concept of justice. In the
United States, school children daily repeat the Pledge of Allegiance —it
contains the words "with liberty and justice for all." What does justice
mean in that context? We do not get much encouragement to ask that
question. Even some scientists and philosophers are adamant that justice
cannot be defined in scientific terms. As a result, the idea of justice, like
most other cultural ideals, is a little bit more circumscribed each time
we do not ask. Of course we cannot turn to science alone for definitions
of justice. Yet, every culture has some view of what is right or wrong, or
of fair play, even if it is not always put into words. So we can ask
whether there are proto-historical precursors to such ideas and concepts.
Also, whether it is possible for a society to function without some more
or less vague sense of justice, no matter, how perverted it may
sometimes become.
Two factors get in the way of understanding our subject: one is the
overzealous extension of the scientific kind of premise. Of course we
must not oversimplify and look for the genes of justice! But it is
important that we not exclude all of biology from our discussions when
we reject part of it. The other factor is more subtle: much of our
culture, like the grammar of our language, is out of awareness. The
premises behind the concepts of justice that our courts dispense are
usually not made overt. That does not make them the less powerful
levers on our thoughts.
We are convinced that, if we can get these points into the overt,
agenda, they will bother us little. But if we do not, then we are truly
beset by our unstated premises.
It will be apparent, as you read the following papers, that the idea of
pairing law and biology still seems daring. All contributors to this
volume have been careful—and a few have been skeptical even about
discussing the two in the same breath. Several of the papers warn us
in no uncertain terms. Markl asks why we should think law even has
a biological dimension. The social scientists are as stern, but make
different points: Schwartz warns us to be very careful in just how we
bring biology into law as it is practiced, and emphasizes that, in his
opinion, lawyers and judges must base their knowledge and decisions
not on science, but on the mores or standards of the community.
Adamson Hoebel urges us to recognize the limits of our own naivete.
Today, the underlying premises—particularly those of the scientific
community—are much more subtle than those of a few years ago.
Hence, in spite of the need for care we all felt, the biology of human
behavior is not as treacherous a topic as it once was. The intellectual
[xiv]
climate of the present is kinder to an examination of this problem than,
that if only a very few years ago. However, such a statement must be
read in context: this introduction is being written during the time that the
"second Scopes trial" is going on in Arkansas. Obviously, there is still a
large segment of the population that denies any biological basis of all
human behavior.
Today's biology and social science are brought ever closer together not
just by their shared interest in behavior but also in trying to overcome
the popular denial of basic scientific facts. it is not just in Arkansas that
people today deny what the most eminent scientists of our time hold as
truth, where the state's attorney argues on behalf of those who believe
that there is a scientific basis to one and only one specific version-the
Judeo-Christian version-of the story of creation. It is not just in
Arkansas where some people close their eyes to the fact that human
beings are biological organisms, subject to biological commands as
much as to religious or cultural influences (these latter differing in
different parts of the world), and that the two must be understood as part
of a single coevolutionary process.
And what about policy? Have people in policy-making positions been
exposed to the biologically based findings of behavioral science- and
have they been exposed at an early enough age to build these findings
into their Weltbild?
The remainder of this introduction will deal with the following topics
that were discussed at the conference:
(1) a history of the relationship among law, biology and anthropology;
(2) the relationship between morality and law, and the evolution of
both;
(3) educating lawyers and fostering law-abiding behavior. .
LAW, BIOLOGY AND ANTHROPOLOGY
Adamson Hoebel's paper gives a sound.view of the relationship among
law, biology and anthropology. Here we add some complementary
points.
Law is just as subject to the premises about the sacred difference of
human beings from the "animals" as is everything else. Thus our
forebearers such as Maine and Ehrlich did not talk about the evolution
of "law" from early hominid behavior or even about the precursors of
law. It also assured that they did not even talk about the fact that human
beings, as legal creatures, were simultaneously biological
[xv]
creatures and that the two dimensions may have some things in
common.
It is true that anthropology has, for well over a century, been committed
to the proposition that the human organism evolved from "lower"
forms. But it is also true that early in this century, cultural -and social
anthropologists banished that idea into a special branch called "physical
anthropology." There specialists could study it separated from day-to-
day cultural activities and nobody was bothered by any kind of
cognitive dissonance.
This is background for today's question: how do we link the human
biological system with the human cultural system? And, in this case,
especially human legal systems? Are there biological imperatives and
how are they to be linked with cultural commandments? Are the two
unrelated, for all that they often work against one another
Law evolved in one biological species, Homo sapiens, a highly
specialized animal. But what are its infracultural roots? Insects deal with
many comparable matters by "hard-wiring." But human beings deal with
them by a special evolutionary step: the introduction of culture, which
includes the capacity-indeed, the necessity-to choose and ultimately to
create both policy and the law that underlies and directs policy.
That proposition takes us instantly to the second point.
LAW AND MORALITY
Richard Alexander, Christopher Boehm and others at our conference
suggested that the evolutionary origins of morality lie in conflict
resolution-conflict management. Indeed, Boehm's idea-that human
ancestors had (to put it into lay terms} more talent for interfering in
dyadic conflict (thus turning it into triadic solutions} than they had for
mere submissive behavior-clears up many difficulties. Both types of
behavior, interfering and submissive, appear in today's non-human
primates. And of course humans too utilize submission in their power
plays—indeed, people like Ghandi thought we underutilize it. Thus if
we are careful not to turn it into too direct an analogy, it is easy to find
"proto-morality" among non-human primates, especially ~n their
characteristic ways of dealing with dominance and submission, and to a
lesser extent their capacity to interfere in disputes (usually on the side of
the weaker animal} in order to regain "peace." It seems to us, that it is
not going too far to suggest that some human ancestors (or
[xvi]
some or all hominoids) had the very qualities we find in some of today's
primates.
Three points are essential here.
(1) The dominance-submission-reassurance sequence is the seedbed of
law and the idea of right and wrong.
(2) Submission-even weakness-brings its own kind of power. The
dominant animal is restricted in what he can do. Watch a humming bird
guarding his food source-it is a full time job. The non-possessing animal
(weaker for the moment) has immense advantages in mobility and choice
to further his personal gain and status. The submissive animal has more
options than the dominant animal.
(3) Primates appear to be better at interfering in conflicts within the
group and restoring peace by reassurance than are most other animals. In
many instances they seem to prefer this method to exclusive use of )
submissive behavior patterns. The desire for balance, harmony and peace
seems to motivate the dominant animal to reassure the weaker.
It seems that in the course of evolution, the triadic solution to dyadic
conflict has been the basis on which more flexible and therefore more
adaptive social systems could be built. Instead of the mere
dominant/submissive dyad, with reassurance, human beings have built a
primary set of institutions on a dominant/submissive/interferer triad,
whereby the reassurance part may be in danger of being lost. This kind of
interference—Boehm calls "the moral community" and' Malinowski
called the "social machinery"—would seem to have gained a central
cultural position.
It was also pointed out at the conference that conceptualization of the self
is involved in this matter, and it was suggested that there is a
development (ascent if you will) self-conceptualization / group morality /
definition of social gains / goals Thus, proto-morality seems to have
evolved into legal behavior and the concept of justice by the working of
two conjoined forces:
(1) the triadic form of interaction, complete with "interferer,"
(2) the growth of self-conceptualization, which involves the realization
that the other person is also a human and merits consideration and that
what we do to him may happen to us-as we have come to see in terms of
a concept of justice.
We can, as a result of positing those changes, see in evolutionary biology
what we once saw only in philosophy. Masters' paper deals - further with
the philosophy and shows something of the overlap. .
[xvii]
Law itself, as we know it, builds on one other distinctly human capacity:
the capacity to state what the law is. Unless such statements of the law
are too divergent from actual behavior or desires, human beings would
seem usually to go along with them, then feel confident because they
know what "the law" is. Undoubtedly, some people know what the law is
and still want to break it, but it is nevertheless important to define the
boundary of behavior that is acceptable to the I group. Therefore, we
have man-made law. In short, specific laws .t work best when they are
overtly stated. However, their effectiveness is not limited to such
"statements." The statement itself allows a rule for behaving to be moved
from one social context to another-a phenomenon that (although, as
Schwartz suggests persuasively, it is not enough) is one of the most
remarkable aspects of human law.
However, we often experience a certain "disjunction" between mores
and the law-which implies that the legal institutions have ,: taken on a
life of their own and perhaps are inadequately connected with the other
institutions of human society, let alone with human biology.
THE EDUCATION OF LAWYERS AND JUDGES
Lawyers and judges, like doctors, have to make decisions on the basis of
inadequate information. Scientific information, including biology .might
well go into and even improve their judgments if it is used wisely-but
judges cannot know everything any more than the rest of us. So where
does one draw the line?
We know that law school curricula need simplifying, not complicating.
Therefore the problem remains: how do we introduce scientific
information (including biology and other behavioral science) as a
simplifying mechanism instead of a complicating one into the
professional training of lawyers and judges? Perhaps it can be done in
high school and college.
During the last 100 years we have paid a lot of attention to the health of
individuals-and have built our social expectations on medical progress.
When medical anthropology joined the effort by studying the links
between disease, medical practice, culture and biology, the result was an
even greater advantage of therapeutic possibilities based on new
technology and scientific findings. Could we make comparable advances
in law? Is not law related to human behavior as much as medicine is
related to physiology and biochemistry and (we are coming to
understand today) behavior?
[xviii]
The emphasis of Western law has long been on the individual social
relationships have been largely reduced to individual rights. But today we
realize that part of being human means having sound relationships with at
least some other individuals. It is for these relationships, rather than
merely for individuals, that we need rules, structures and laws. How far
can we go medically and legally in treating human relationships as we
treat an organism? The social side of this organism should be helped and
sustained in the same way as the individual side. And this can only be
accomplished by the individuals whose Weltbild encompasses the
biological nature of man. Of course, the cultural context in which the
individual acts is also part of human behavior. Human ontogeny is
possible only in a cultural context.
Lawyers have indeed been "treating" individuals, even dyadic
relationships. But too often they do it without realizing that they are
treating only a small area of individual well-being, let alone social well-
being. Individual rights and healthy relationships—these are the two
sides of a coin. To assure their well-being, you need the same basic
sciences—you cannot treat one without affecting the other. You cannot
have the accumulation of knowledge on the one hand and on the other
not allow anything new from the philosophical and scientific -including
behavioral scientific-arena to inform the legal procedure.
We need a broader horizon in legal education. The scholars and
scientists responsible for the professional training of future lawyers and
judges-these are the people who have to be concerned, so that \ training
for the legal profession is built on both a humanistic and scientific
platform. Then you can learn law as a craft in three years of law school.
Education for undergraduates (including those who plan to enter law
schools) and graduate school curricula must be based on he state of
knowledge and the state of the art. What we need are legal scholars who
will reach to the sciences for relevant data to further their own research-
the scientists cannot know what is needed to improve legal processes.
[1]
PART I
The Foundations in Law and Morality

Margaret Gruter and Paul Bohannan
Law is multidimensional. It is, therefore, difficult to define law in
its totality. We greed early at the conference not to spend our time
defining law, for we knew we could get bogged down in it and
never emerge. Definitions of law are influenced not only by the
uses of the law, but by different concepts about the origins of law
and its functions and structure. Several basic theories on the origins
of law reappear throughout recorded history. One of these theories
is that law is given to us by a deity-Moses, for example, got the Ten
Commandments directly from God. Another theory holds that laws
are the product of human reason. Both these theories have
influenced and shaped the changing concepts of "natural law"
during the centuries. (We will not use the confusing term "natural
law," but will speak of "man-made law" on the one hand and
biological laws or "laws of nature" on the other.) Today, ideologies
from Marx to Freud are used to explain the origins of law. Legal
scholars, influenced by various theories of the origins of law, often
provide very narrow definitions of law.
Law is an evolving system that changes under the impact of the
environment, both physical and cultural. It is expressed in the
behavior of the individual and the group. Among the first to
recognize this, at the beginning of the twentieth century, were an
anthropologist of law (Malinwoski, 1926) and a sociologist of law
(Ehrlich, 1913). Ehrlich coined the expression "the living law" to
describe law in action as opposed to law on the books. Today
Friedman (1975) uses the l' term "legal system" for the interaction
of law and behavior. The legal system includes law in action and
law on the books, as well as such subsystems as the courts. The
structures and rules that are part of law have an impact on behavior,
which in turn creates new norms and rules. We may speculate that
some "concept" of law may even have been one of the first abstract
ideas formulated by the human mind
[2]
when, early in the evolutionary history of the human species, the awareness
of rules freed human social organization from genetic commands. Certainly
we can assume that the functions of early law complemented the functions of
human behavior to some degree.
All this necessarily leads to today's questions: how do we link the functions
of human behavior with the functions of the, legal system? The essays in.
this section examine the legal foundations of our efforts to link law with
biology. The authors consider some previous contributions to this topic in the
literature of jurisprudence to see what sort of problems and what solutions
they bring to our task of examining the biological basis of law. They find the
picture of law and justice incomplete and ask the neighboring disciplines to
help legal scholars find the missing pieces.
In Parts II and Ill, several essays deal with morality. Morality, like justice or
ethics, has always been linked with law. It should certainly be of interest to
scholarly inquiries into the relationship between behavior and law. Morality
as a biological phenomenon was the subject of an earlier conference and to
some degree we are building on their published reports (Stent, 1980).
However, we recognize that the concept of morality is an essential part of our
discussions. As part of the introduction to Part Ill, we have published a
segment of the report by Christopher Boehm on the Primates Discussion
Group at the conference, which deals specifically with this topic, but which
makes more sense after the essays on biological research have been added to
those in the first section. All of the essays in the book bear on the search for
the missing pieces.
[2]
Biologically based behavioral research

and the facts of law
MARGARET GRUTER
158 Goya Road, Portola Valley, CA 94025, USA
Can we find precursors of human cultural characteristics among primates? Eugen

Ehrlich in 1913 postulated two primary functions of the living law: organization and
protection of the social order. His theories are compared with observations of
ethologists in order to explore possible new insights into legal behavior. To what
extent do the implicit rules of
Gruter and Bohannan [3]
non-human primate social orders reflect the four behavior patterns that Ehrlich
called "the facts of law"-usage, domination, possession, and" diposition? To what
extent is such legal behavior affected by endorphines and by other other biological
mechanisms?
THE LIVING LAW
If we accept the Darwinian theories that the human species descended from an
ancestor shared by apes and other primates, it follows that we may find precursors of
human characteristics in the animal kingdom, (Edey, 1972: 132-133). Can
ethological observations provide data that point to precursors of legal human
behavior in non-human primates? The theories of Eugen Ehrlich, one of the
founding fathers of the sociology of law, are compared with observations of
ethologists in order to explore possible new insights into legal behavior and the
various factors that contribute to the effectiveness of law.
Ehrlich (1913) formulated his "Fundamental Principles of the Sociology of Law"

approximately 80 years ago, before World War I. One of his major contributions
was to introduce the concept of the "living law" to describe the observed interaction
of people within a legal system-the law in action as opposed to the law on the books.
(Rehbinder, 1977: 10-11; Podgorecki, 1981: 183).
Ehrlich's ideas are based on his observations of the law in action, observations of
what we today would call human social interactions. Observation is the scientific
tool with which ethologists build their theories of animal behavior. Scientists discern
patterns in the social interactions of individuals, whether analyzing the observed
behavior of human beings or other animals. Ehrlich used the term Rechttatsachen or
"facts of law" for those patterns of human behavior which seemed to be basic facts
of everyday social and legal transactions. He stipulated four facts of law: usage,
domination, possession, and disposition
Ehrlich defines law as the inner order of a society. Law has several functions, above
all the function of organization. The law presents the an individual with alternative
choices and points to those choices that the individual's society finds acceptable or
"just." The individual can then choose to act or refrain from acting. By knowing the
moral consensus of the majority, which, in this case, is the same thing as
the"society's concept of justice, the individual can predict with some degree of
accuracy the consequences of his choices.
Ehrhch sees a second functIon of the law: the protectIon of the socIal order. This
function is supported by the "decision-norm," which
[4] LAW, BIOLOGY AND CULTURE
complements or fills in gaps in the legal structure and thus makes law more
effective. The decision-norm within a society is usually expressed through
adjudication, which enables the law to be flexible and to adapt the rigid rules of
legislation to individual cases.
Can we discern functions of organization and protection, a kind of “inner order,” at

work ill non-human societies? Through the millenia, more evolved species have
achieved greater and greater flexibility in their social organization, making them
more adaptive to drastic changes in their environment. It seems certain that this
trend is due to the evolutionary growth of the cortex and the increasing
differentiation of the cortical functions. However, the phylogenetically older parts of
the brain, which developed millions of years ago and served in \ conditions
drastically different from our current environment, still exist and function
simultaneously with our more recently evolved brain structures (MacLean, 1970,
1978a; Hamburg, 1975). Adaptive mutation means that from time to time something
new is added to an existing structure (e.g. human brain) that can correct or
complement those parts of the older structure that have outlived their usefulness and
might drive us in a maladaptive direction.
Perhaps a similar process takes place within the law. The human concept of "law and
justice" reflects the rules, mores and limitations of the social framework within
which each individual grows to social immaturity. Rarely are laws repealed outright
when they no longer reflect the consensus of the group. They usually lead an
existence in the books for many years, even they are not enforced and for all
practical purposes are non-existent in the real world. These outmoded concepts can
be continued or resurrected by people who are strongly bound to a particular
tradition or religion. These traditionalists may react more strongly to the rules of
their sub-society and adhere more rigorously to its laws than does the average law-
abiding citizen of the Western World. Usually, these rigorously obeyed laws are
supported by religious commandments. From an evolutionary point of view,
religions are based on emotions that are nourished by reactions and impulses derived
from phylogenetically older parts of the brain.
Ethics, beliefs, morality, the qualities that make us human, and all the , values we
cherish, are strongly influenced by the regions that already: existed in the reptile
brain millions of years ago. They are "mediated by the old limbic-hypothalamic-
midbrain circuits probably built into the machine because they worked well in its
adaptation over many thousands of years." (Hamburg, 1975: 46)
Although human beings are the only species, capable of conceptual-
izing the rules by which they live, non-human societies also live by implicit rules. In
all societies, a rule by definition is obeyed by the majority and is sometimes broken
by some individuals. The more rigid the code, the greater the adverse consequences
for those who disobey. In the rigid genetically-coded social organization of the bees,
breaking the code of behavior that dictates that a bee must return only to the hive
from which she originated is almost always fatal {Frisch, 1950,1955).
Many human societies have codes and punishments regulating the purveyance of
wrong information, or lying. Distrust of strangers and punishments for deception are
frequently found in simple societies, where misdirected food-gathering expeditions
or misinformation by strangers about fruit-ripening can have damaging
consequences {Kummer, 1980: 5).
The more evolved a social species becomes, the more flexible the rules of social
behavior-and the more difficult it becomes to draw the line between behavior that is
within the norm {compliance) and behavior that is outside the norm {deviance). Of
course, in the biological sense, there is no connotation of good or evil in the concept
of deviance. However, deviance and evil have always been interrelated in human
societies.
Biologists postulate that evolutionary changes {including the continuous

development of the human brain that enabled Homo sapiens to emerge as a species)
had to start with changes within individuals who did not stay within the norm but
changed through mutation. Why then is it necessary for human social organization to
harbor concepts of goodness and evil, justice and fairness, or other value judgments
in coping with those who do not stay within the norms? How does this relate to the
facts of life and the "facts of law ," which are the reality of human interactions?
Is the "sense of justice" {Gruter, 1976, 1980) the individual yardstick for right and
wrong, one of the tools available within the human brain to organize social groups
{the first function of the law) ~ and to protect the group {the second function)?
Ehrlich, like many others, also asked this question. He stated that the scholars of his
day could only provide unsatisfactory explanations, and expressed the hope that "the
jurist and the legislator will gradually become more and more like the modern
scientifically-trained physician in proportion as society is able to trace and present
the laws of the development of human society." {Ehrlich, 1913 [1975: 243-244])
New insights into the evolutionary process show that modern Homo
sapiens could only evolve to the present level because a conceptual framework of
rules evolved with the increasing differentiation of the cortical functions in the
human brain. During this long simultaneous evolution from genetically-controlled
social organization, the rules and norms of behavior took on their own existence as
abstract rules in the minds of individuals through their interaction within and among
social subgroups. The "legal system" is a totality of which written codes form only a
part. Humans express or live legal rules by choosing \ whether to obey or disobey
{Friedman, 1975: 67). They are often \ motivated by emotions while making their
choices.
Emotions can be controlled by certain regions of the cortical part of our brain only to
some extent. Often they originate from, and are supported by, complex mechanisms
in our limbic system. Some of these emotions motivate our responses to rules, which
is our legal behavior. If legal behavior is crucial for our survival as a social species,
it follows that any mechanism that can help in motivating adequate responses to
rules is adaptive. Emotions which are caused by concepts of good and evil, right and
wrong, have been helpful in directing and controlling legal behavior. The ability to
make value judgments and the yardstick by which humans arrive at such value
judgments of right or wrong-their "sense of justice"-have proved adaptive.
An essential part of this mechanism is the "internal reward system" {Routtenberg,

1978; Danielli, 1980) with its ability to release mood- controlling substances in the
brain, such as opioid peptides, also called endorphines. In describing the evolution
of the reward system, Danielli refers to the statement by Marx that "religion is the
opium of the people" and suggests that we might be more accurate in stating,
"ideology is the opium of the people." People who believe in religious or ideological
causes may well be rewarded by a feeling of well-being due to the release of opiates
in their brain. Let us assume for our purposes that the internal reward system is part
of the mechanism that we call the "sense of justice."
THE FACTS OF LAW
Parallels in the structure and function of social organization in different species have
been postulated. Now the four behavior patterns that Ehrlich called the "facts of law"
will be compared with possible precursors in non-human primate societies. Behavior
patterns become evident during observations of social interaction.
Usage
The oldest fact of law, according to Ehrlich's observations, is usage. "Usus" or

"mores" keeps the inner order within societies. Usage is a part of structured social
interaction and can be investigated empirically.
Order can be maintained in human societies by various means among them

constitutional law, legal norms or adjudication. Ehrlich observed that usage is
decisive in individual cases within a given structure, especially in cases where there
are differences of opinion in the interpretation of law or when new developments
have not yet been regulated by formal law.
Usage is the rules obeyed by a majority within a society. It can be abstracted into
formal law but is valid even without official sanctions. What is currently innovative
or even deviant, behavior may become the future norm for the majority as
environmental, biological and technological changes or other developments put
pressure on legal structures and test the present or future adaptiveness of existing
norms.
Examples from non-human societies demonstrate the emergence of new behavioral

techniques, which are slowly adopted by the majority {Marler et al, 1972: 42-43)
and become "normal" behavior. Environmental pressures can force adaptations in
sleeping habits, and even changes in the social structure of non-human primates
{Kummer , , 1971b: 131-135).
In Ehrlich's opinion, usage is determined by economic pressures and the results of

individual competitions for dominance or power within societies. These power
struggles are an integral part of economic changes. For example, the invention of the
railroad, and of motor-driven vehicles in general, required a tremendous number of
new laws and legal instruments to solve the problems brought about by these
developments, often granting new powers to individuals and institutions. In the
United States, the rights of married women had to be extended to permit them to
own real property when predominantly male emigration to the West made many
women "grass" widows. Many commercial practices have existed as "usage" long
before legislation gave official sanction to the rules that were used to facilitate the ~,
exchange of goods and services {Friedman, 1973: 318-322).
Ethological observations document changes in group behavior brought about by

changes in the environment or by the struggle for dominance within non-human
social structures. Goodall's observations of the behavior of a group of chimpanzees
over a period of 20 years illustrate how the struggle for dominance can influence the
social behavior of a chimpanzee community.
During the observation period, there were no drastic natural changes in the
environment affecting the supply and variety of fruits 4 and other desirable goods.
However, observable changes in chimpanzee behavior became apparent early in the
field work when large quantities of bananas were made available to them by field
workers.
Normally chimpanzees have to spend an average of six to eight hours every day
traveling to the food supply, and gathering and eating sufficient amounts. Providing
the chimpanzees with a much-liked food in insufficient quantities for all the
individuals in the group, greatly increased aggression among the chimpanzees, and
between the chimpanzees and baboons that occupy the same territory. Did this
"economic change" affect their ability to control their aggressive drives, the
balancing act that in humans is called "fairness" and is guided by the "sense of
justice"?
Changes in the social organization of the chimpanzees can also be caused by the
personalities of the different chimpanzees, especially the alpha animal, and their
ability to make use of certain changes in the environment. Mike, a relatively low-
ranking and medium-sized adult, made use of empty gasoline cans discarded by the
field workers to become alpha animal by incorporating the gasoline cans into his
display. The resulting noise and commotion he created terrified the other
chimpanzees and allowed him to reach the alpha position within a few months and
maintain the position for almost six years.
To a certain degree, the personality of the alpha animal may influence the behavioral
norms of the group; the absence or presence of the alpha animal certainly does. The
amount of protection for weaker individuals depends on the availability of high-
ranking males. During 1975-78, Goodall observed that two female chimpanzees
were killing infants and eating them. It was observed that the mother of a threatened
infant turned to the males for protection against the potential killers. These males
threatened the wrong-doers and chased them away {Gruter, 1979: 44).
Goodall and others have also observed that the behavior of a mother will influence
her offspring's behavior in many ways. Offspring of high-ranking mothers usually
attain a high rank for themselves when they grow up, an obvious parallel to human
society. The choice of food, the preference for certain fruits, meats or termites is also
learned from the mother {Goodall, personal communication).
All the behavior patterns that make up the norms of a society and give structure to
the group can also be seen as part of the struggle for dominance, which exists in all
primate societies, both human and
non-human. The concept of dominance is related to Ehrlich's second fact of law.
Domination as a ìfact of Lawî

According to Ehrlich, domination results from the fact that weaker members of a
group need protection. Ehrlich postulates that whenever the "dominated" or weaker
individual is unable to protect himself, his protection is taken over by a higher-
ranking individual. In human societies, legal and political definition of human rights
are one method of protecting less powerful individuals.
Lorenz was one of the first to observe that in the so-called pecking order of several
bird species (chickens, ducks, crows, etc.) high-ranking animals may interfere in
fights of lower-ranking birds to protect the weaker of the two fighting individuals. It
has also been recorded for baboons (Wickler, 1971: 138). Goodall's observations
show that a male chimpanzee will come to the defense of mothers whose infants are
endangered, as in the case of Figan, who threatened ; Passion and Pom (the two
marauding female chimpanzees) when they attacked.
Another mechanism in animal societies that serves to protect weaker or "dominated"

individuals is the formation of subgroups such as hierarchal orders for males and
females.
All non-human primates nurse their infants and carry them with t them for several
months, and even for several years. The infant is 1 completely dependent on, or
"dominated" by, the mother. This relationship is nourished and strengthened by
many biological, physiological and psychological mechanisms. The close bond
evokes a feeling of well-being in both mother and infant.
Hormonal processes during pregnancy, childbirth and the nursing stages strongly
influence maternal behavior. Reciprocity and feedback are an important part of the
first social interactions ensuring survival for the infant and providing satisfaction to
the mother.
Even in non-human primates, this behavior is not dictated absolutely by biological

mechanisms or genetics. Students of chimpanzees and other primates have observed
that females are capable of learning maternal behavior to some degree. Female
juveniles learn from observing their mothers caring for younger siblings. There is a
likelihood that the female child of a successful mother will also become a successful
parent. Chimpanzee mothers are capable of becoming "better," more efficient and
caring mothers with their second child.
It is possible that a sense of well-being similar to that originally

generated by hormonal changes during the childbearing period can be experienced

again when the mother shares food with offspring, or even with other group
members. This tendency towards altruistic behavior based on interactions that
generate reciprocity and mutual feelings of well-being can perhaps expand and
develop in other interpersonal relationships, even if these feelings are no longer
triggered by' " hormonal processes but by other stimuli to the internal reward
system.
Ehrlich postulates that domination is a fact of law in all human societies, and that it
provides mechanisms that protect weaker members of the group. Does this imply
some altruistic motivations in the stronger individual? If altruism is a factor in
dominance, is there a biological basis for altruistic tendencies that exist in different
social species (Dawkins, 1976)?
The readiness to act altruistically could only develop as the result of many different
stimuli. An important discovery is that a reward system does exist as a cerebral
mechanism that produces a sense of well-being , and can be triggered by various
actions and stimuli. These various stimuli cause the production of substances in
certain centers of the brain, similar to but more potent than morphine, which cause a
feeling of well-being in the individual (Routtenberg, 1978; Danielli, 1980; B.
Hoebel, this volume).
People have long been accustomed to the fact that emotions are states of feeling that
can be affected or triggered by the intake of drugs such as opium, marijuana,
cocaine, LSD and alcohol. Recent experiments with the salts of lithium demonstrate
the potential for chemical , substances to alter feelings and behavior. One researcher
posits that "the.physiochemical simplicity of lithium arouses the hope that it will
provlde a light to clarify the neuronal basis of moods (Tosteson, 1981:4). This type
of research may help to discover when and why obeying the law makes humans feel
good, even when obedience involves self-sacrifice or so-called "altruistic" behavior.
As new research in neuropharmacology elucidates the sites and causes of these

chemical processes in the brain, methods are being explored to locate and measure
the chemical compounds and to localize the receptors and analyze their functions.
What research can discover about motivations towards legal behavior due to
endorphine production is especially relevant to legal scholars. Research may also
show ways to measure environmental influences on these processes.
Domination as a "fact of life" seems to be related to the evolutionary growth of the

human brain and the chemical substance produced in the brain that gives individuals
a feeling of well-being. Human society
could only have evolved because the human species has been capable of forming
systems of ordered interpersonal relationships, or social orders, which are not
genetically transmitted in a rigid code but are flexible and adaptive to the demands
of changing environments. One of the facts that makes social order possible is the
feeling of well-being produced in individuals who feel part of their social order, are
comfortable with their status within the hierarchy, and derive protection,
companionship and security from it. Ostracism is one of the severest punishments in
many social groups, because of the devastating effect on the individual deprived of
his "place" in the social order.
The ability to accept and tolerate "domination" is also necessitated by the lengthy
period of social immaturity for the young in human and non-human primates.
Without some form of domination that produces well-being, the young in these
societies could not survive. This leads us to the third "fact of law," possession,
insofar as the mother-child domination may well have resulted in the concept that if
one person "dominates" exclusively, he or she thereby "possesses."
Possession
Possession, Ehrlich's third "fact of law," can be seen from an ethological point of
view as the result of the more differentiated social organization that evolved
simultaneously with the brain. The concept of possession may have partly developed
from the mother-child relationship. Human and non-human primate societies alike
give the nursing mother the "right" to actually hold and possess her infant, as long as
she performs her maternal duties. In non-human primates and other animals, the
mother-child relationship is usually respected (but see Fossey, 1981). Respect for
possession has also been observed in pair-bonding situations among the hamadrayas
baboons (Kummer , 1980: 100; Gruter, 1977).
However, bonding is only one of the possible mechanisms that can evoke respect for
possession in others. Generally, it seems that this feeling or attitude results when
others perceive the physical closeness of individuals (such as mother and child) or of
an individual and an object (a person using a tool or a hunter holding his prey) as
one unity or gestalt (Gruter, 1977, 1979) in which the dominant one possesses the
other. Llewellyn (1977) concludes from this physiological phenomenon that people
in the abstract sense also accept the unity of certain concepts like "sale," "lease" and
"corporation" as one concept, but do not pay attention to the attributes or legal
conditions that make up or :":, are part of the legal definition of "sale." He concludes
that people
not recognize norms by perceiving details, but by perceiving the entire picture.
Perception of a familiar gestalt evokes impressions of rightness in the viewer,

producing feelings of well-being. Unfamiliar or "wrong" impressions trigger feelings
of fear (a possible explanation for the prevalence of xenophobia ill many social
groups} that can turn into aggression accompanied by a change ill hormonal
production. Until a balance of positive feelings of "rightness," rather than negative
feelings, can be reached, the imbalance and impressions of "wrongness" may cause
feelings of depression, despair, and destruction (Gruter, 1977: 217}.
The concept of possession may have evolved from the "right" (in the eyes of the
observer} of an individual to possess exclusively what he carries with him,
supported by the production of endorphines. The visual stimuli of perceiving
individuals together, or an individual and an object (container, tool, weapon or prey}
as a gestalt, may trigger, endorphine production. Even chimpanzees defend what
they can carry with them, perhaps expecting that it is their "right" to possess ,
whatever they can physically carry.
During a period of field observation of the chimpanzees in Gombe, field workers fed
the chimpanzees considerable quantities of bananas. Some animals consistently tried
to hold on to more and more bananas, which repeatedly slipped from their grasp.
Although the animals appeared to be frustrated and almost bewildered by the
situation, they persisted in their attempt to carry off or "possess" more than they
could hold. The fact of "possession" also led to aggression. Goodall reports that the
only time in 20 years of observation that she was seriously attacked by a chimpanzee
was when she attempted to take a banana dropped by a female who held dozens in
order to give one to a young chimpanzee who possessed none. The greedy
chimpanzee can be said to have regarded the bananas she was carrying as her
possession or "property" (Goodall, personal communication}.
The respect engendered by the possessor's determination to defend his property may
account for situations in chimpanzee societies where higher-ranking animals beg
food from lower-ranking animals during meat-sharing after the prey has been
caught.
Respect for the possessor and his possession may have developed as a behavioral
trait in a biological sense, as well as culturally-supported behavior, over a period of
hundreds of thousands of years. A framework of conceptualized behavior patterns
has evolved that includes domination over objects,and criteria for possession similar
to Ehrlich's "facts of law."
This development can be seen as part of human phylogeny, as well as a part of

human development in early childhood and through life (ontogeny).
Disposition
Disposition, Ehrlich's fourth and final "fact of law," involves contracts. In the legal
sense, contract requires two dispositions-offer and, acceptance-and a third essential,
agreement. Once contract is accepted as a fact of law, the concepts of breach of
promise, breach of contract, guilt, responsibility, liability, damages and punishment ,
follow. Some form of contract has probably existed in all human societies
(Malinowski, 1926 ).
Ehrlich claimed that all norms, whether legal or extralegal rules, serve an
organizational function in society. All norms are prescriptive or proscriptive,
according to Ehrlich, demanding certain actions or non-actions. All norms that
demand actions within a society must be both heteronomous and autonomous-out-
directed-because they derive their effectiveness from the group's concept of justice,
and inner- directed because each individual must recognize and obey these norms
("sense of justice") for the society to function. Recognition of and obedience to the
rules by the majority of the members of a group, constitute a valid norm. These
norms then represent the structure of the society, or group organization.
"Group organization" evolves into a system founded on the interactions between

individuals (domination), between individuals and objects (possession) and their
reciprocal actions (dispositions or contracts).
NEW INSIGHTS-NEW QUESTIONS
How do Ehrlich's theses compare with Darwinian and Mendelian theories and
findings, and with the accepted laws of nature by which the human species evolved?
One problem posed by Darwin's theory .has been to make it compatible with the
tendency towards altruism that exists at least partially in all human societies.
According to Darwinian theory, "altruistic genes" would die out naturally since their
existence would lessen an individual's fitness to reproduce. Those individuals who
sacrifice themselves or their reproductive success in favor of altruistic actions
endanger the propagation of their qualities, more than individuals who do not act
altruistically.
Darwinian theory, however, refers strictly to individuals rather than families, groups,
races or species. Darwin did not use the term
"survival of the fittest" in the sense that a certain class or race within a species will
survive, or one species compared to another species. His statement was merely that
the individual who is capable of having more reproductive success than others in the
same environment, and who is capable of raising his offspring to sexual maturity,
will transmit more of his genes to posterity. Hamilton, Trivers and others linked
Mendelian theories and Darwin's to demonstrate that altruistic behavior that helps
close relatives to have reproductive success can indeed transmit more genetic
material of the altruistic individual to following generations (inclusive fitness)
(Trivers, 1971; Hamilton, 1964; Axelrod & Hamilton, 1981). The "altruistic"
individual, of course, does not have to be aware of this.
Human tendencies towards reciprocity, postponement of gratification for future

good, and the ability to act on cost-benefit calculations, play an important role in the
development of modern social organizations. Precursors of these attitudes can also
be observed in the behavior of some non-human species.
Chimpanzees are capable of thinking and can project to the degree that they will
forego instant gratification for a better return in the future. Chimpanzees carefully
select only ripe fruit, leaving unripe fruit for the future after careful touching and
testing. Chimpanzees have also made rudimentary tools to fish termites out of a heap
without destroying the entire heap, choosing a lengthy and laborious method of
fishing for small insects rather than destroy their entire termite supply.
There are examples in other species where individual animals use foresight and
planning to achieve a goal (Gruter, 1979). Certain rituals of courtship and mating in
many species depend on reciprocal actions, as do maturity rites and other rituals.
These are behaviors that require timing, some form of planning, or postponement of
gratification.
Reciprocity in the interaction of non-humans can be seen as the precursor of contract

in human society. Perhaps the link between all these different aspects of reciprocal
behavior is that the individual gains a sense of well-being when he acts in
accordance with partly innate and partly learned rules. Due to the production of
endorphins in the brain of the law-abiding citizen, can an individual become
dependent on, or at least accept, the rules of his society? If obedience to the law and
the acceptance of one's place in society can trigger the production of endorphines,
"the opium of the people" may have a beneficial effect after all by preserving the
continuity of social structure. This does not mean the perpetuation of the status quo
at all
times since many different religions or ideologies with different goals can trigger the
same mechanism. This interpretation also does not imply that the individual must
blindly follow innate commands. Obviously, the individual still has a wide choice;
he can refrain from behaviors that cause both pleasure and pain, and can choose
different stimuli to gain the same or similar, effect. Whether this is a wide or a
limited choice, "free will" or a narrow spectrum of alternatives may well be ill the
eye of the beholder.
On the prospects of using sociobiology in shaping the law:
A cautionary note
RICHARD D. SCHWARTZ
College of Law, Syracuse University, Syracuse, NY I32IO, USA
Caution must be exercised in promoting the use of biological knowledge for

guidance in legal policy decisions. Scientific information is frequently needed to
heighten rationality, but judges often limit the place of such information; in so
doing, they are not necessarily being parochial. A crucial problem for legal use of
scientific knowledge concerns the boundary between scientific and normative
judgment. In the Durham v. US (1954) decision, Judge Bazelon opened federal
courts in the District of Columbia to fuller participation by psychiatrists in
determining insanity than had been provided by the M'Naghten (1843) rule. Two
decades later, the same court rejected the Durham rule because it led juries to
abdicate judgment concerning defendants' moral responsibility.
The tendency for law to seek significant guidance on normative questions from the
population, rather than from technical experts, is well grounded in our legal
tradition. Even where normative agreement exists, science does not necessarily
contribute to the resolution of normative questions. In a heterogeneous society,
biological knowledge can be of greater use in analyzing the processes by which
normative consensus can be generated through social interaction. Legal thinking has
already begun to examine the ways in which law can affect the norm-generating
process, but much remains to be learned concerning the capacity of human beings to
control conflict, optimize satisfactions, develop standards and satisfy the sense of
justice
The task of law in reinforcing and helping to create normative order, cannot be
successfully delegated to science. If scientific knowledge is to assume a place in
affecting specific legal decisions and forming specific laws, it must pass close
scrutiny and compete successfully with traditional sources of law. It is more likely to
contribute, I suspect, at the level of general legal principles and processes than in the
direct shaping of specific laws.
Guessing about future interdisciplinary contributions is necessarily a speculative

activity. Whether probable utility can be determined in , advance or must await
evolutionary selection varies with the field in question. It is not clear whether we
know enough at this stage about what biology can offer law to guide future efforts.
The problem might better be approached by asking first about the absorptive
capacities of law. What does law take from other fields and what does it reject?
What internal processes in the development of law bear on the relevance of
biological science to law?
Some light may be cast on these questions by looking at the sources of law in
societies such as ours. To do so, I shall draw unsystematically on some judicial
decisions and jurisprudential ideas. These are intended to illustrate (certainly not to
prove) some selective propensities of our legal system.
(1) Law limits the scope it accords to other disciplines in matters of normative
judgment.
(2) Law seeks instead to rely on moral judgments of the population, where these are
available, to shape legal standards.
(3) Science does not necessarily provide persuasive guidance for or against a law
which accords with the mores.
After illustrating these propositions, I shall suggest that biological contributions to

law might usefully focus on the processes of interaction which generate normative
consensus. Developments in several disciplines indicate that concepts of justice,
possibly inherent in the species, contribute to the formation of norms. In open
societies, which tend toward anomie and normative conflict, there is a particular
need for arrangements which facilitate normative consensus. In the effort to
construct such opportunities, law can greatly benefit from the advice
of all of the human sciences, including sociobiology.
The capacity of sociobiology to make contributions to specific laws depends not

only on what it has to offer but also on who is listening. There are a number of
discreditable reasons for legal resistance to scientific knowledge, such as ignorance
and professional rigidity. These are often difficult to distinguish from certain more
justifiable bases for legal caution in modifying law to accommodate current
scientific thinking.
I have in mind especially the ambiguity of the borderline between is and ought and
the related question of who should participate in deciding the ought. The record of
inter-disciplinary contributions to legal policy indicates that is- ought problems
frequently arise after initial enthusiasm over the prospective assistance which
science can offer. A familiar pattern has been. repeated in several instances. The
sequence begins with a recognition that some knowledge from another field can be
of great value in resolving legal questions. On this premise, basic legal decisions are
made which rest on the findings of the other field. In applying these basic decisions,
law draws on experts from the other field to provide factual information relevant to
specific cases. As this is done, it turns out that the experts disagree on factual
matters of specific relevance to the decision at hand. Their differences not only
reflect the ambiguity and limitations of scientific knowledge; they also point up the
relevance of normative considerations to the resolution of the issue. When scientists
are brought into a specific case, neither law nor science can reliably draw the line
between fact and value, between is and ought.
A fine example is found in the use of psychiatrists during the past three decades in
the Washington, D.C. Circuit Court of Appeals to testify concerning the insanity of
criminal defendants. The history of the insanity defense, so well summarized
elsewhere (e.g. Goldstein, 1967), need not be described in detail here. Suffice it to
say that in the Durham case, Judge Bazelon replaced the traditional M'Naghten rule
with the so-called product test in the hope that psychiatric testimony could be
introduced more freely and could playa larger part in determining whether insanity
would excuse what otherwise would have been a criminal act. M'Naghten had
limited the insanity defense originally to the question of whether the defendant knew
the nature and quality of his act or, if he did, that it was wrong. Durham broadened
this test conceptually to include any instance in which the
act in question was the product of a mental disease or defect.
The Durham rule appreciably increased the incidence of successful use of the
insanity defense. It led to decisions by jurors which were dominated by the
testimony of psychiatrists. As a result, verdict inconsistency (always present) could
be attributed to the happenstance of forensic testimony and psychiatric attitude.
When St. Elizabeth's psychiatric hospital shifted over the weekend to the view that
psychopathic persons were mentally ill, Friday's finding to guilt became not guilty
on Monday morning (in re Rosenfield, 1957). The major problem seemed to be the
transfer from jury to expert psychiatric witness of the basic normative judgment.
Through a series of successive cases, the Court tried to specify and narrow the rule
so that the jury would have more specific judicial guidance. The purpose of these
developments, sometimes explicitly stated, was to prevent the determination of
blame-worthiness from being dominated by psychiatrists. The psychiatric
profession, concerned primarily with therapy, leans toward a guilt-free conception of
human affairs. It strives to understand all and to forgive all, in order to cure as many
as possible. This conception does not readily fit with the legal approach which, for
purposes of standard setting and enforcement, seeks to fix moral responsibility, i.e.
culpability. Ultimately, the court moved in Brawner 'V. US (1972) toward a revision
of the rule that gave much more explicit criteria to the jury, in the expectation that
they would thereby be enabled to decide the normative issue of responsibility
themselves. Judge Bazelon, in a partial dissent, joined in reversing Durham while
expressing the view that the court should be even more explicit in limiting the role
of the psychiatrist.
II
More is involved in such examples than a simple competition between the

professions. The courts reflect here and in many other areas an inclination to take
into consideration the views of the public. In the example of the insanity defense, the
courts explicitly express the concern that the jury will be deprived of its normative
function. This concern is also manifested by the courts in the traditional deference to
the legislature and in the use of changing community standards in judicial review to
determine constitutionality.
Some court opinions reveal with particular clarity the importance of community
standards. In the first of two major capital punishment cases, Chief Justice Burger
gave this account of the Court's accepted way of construing the Eighth Amendment's
ban against cruel and unusual punishment.
Gruter and Bohannan [19 ]
The Eighth Amendment prohibition cannot fairly be limited to those punishments

thought excessively cruel and barbarous at the time of adoption of the Eighth
Amendment. A punishment is inordinately cruel, in the sense we must deal with it in
these cases, chiefly as perceived .by the society so characterizing it. The standard of
extreme cruelty is not merely descriptive, but necessarily embodies a moral
judgment. The standard itself remains the same, but its applicability must change as
the basic mores of society change (Furman v. Georgia, 1972: 382).
Differences of opinion are found among the judges not as to whether the mores are
important in interpreting the "cruel and unusual" clause, but as to how public
sentiment should be measured. Some would rely on the legislature to reflect public
opinion (except in instances where there existed "unambiguous and compelling
evidence of legislative default" [Furman 'V. GeOrgIa, 1972: 384]). Others would
supplement legislative action with information derived from referenda ! (presumably
a more direct measure) and public opinion polls. For J Justice Marshall, the criterion
to be applied was whether public (opinion, if enlightened by the kind of information
available to the Court, would have favored abolition. And Justice Brennan urged a :
shift from the level of specific norm, existing or potential, to a broader principle.
Capital punishment, he said, is unconstitutional because it does not accord with "the
evolving standards of decency which mark 1~, the progress of a maturing society a
criterion initially enunciated for , the Court by Justice Warren in the earlier Eighth
Amendment case of Trop 'V. Dulles (1958: 101). The standard of decency in this
instance involved the concept of human dignity. Capital punishment, said Brennan,
was unconstitutional because our moral evolution as a society has brought us to the
point of valuing the dignity of each individual human being. In our value system, we
are ready to believe that "even the vilest criminal remains a human being possessed
of common human dignity" (Furman 'V. GeOrgIa, 1972: 273). That being the case
the discard of any human being by execution is for Brennan unconstitutlonally cruel
and unusual.
Each.of these approaches indicates an effort by the judges to shape law of societal
norms. The variations have to do with how the normative position of the society is
best registered, whether it is ; current or potential, whether it is pervasive or limited
to a (particularly significant) segment of the population, and whether it is derived
from and consistent with a general set of values.
In using capital punishment as an example, I have chosen a line of

l
decisions in which the courts most explicitly consider public opinion. In other
spheres, this tendency is more obscure or absent. We need to know more about its
distribution in the universe of court cases. It seems to me, though, that recourse to
public opinion (however it may be ascertained) constitutes a systemic tendency
which is particularly marked where societal norms are strongest.
The tendency to seek consistency between the mores of society and the law is an
important characteristic of our legal system. The theory of representative
government, as carefully described by Dahl (1956) and others, rests on the premise
that law and government will receive greatest support in our type of society if the
interests and standards of the population influence the making of the laws. For this
reason, the courts regularly defer in form at least to the legislature by according to
its statutory acts the "presumption of constitutionality. When the Courts declare a
statute to be unconstitutional (as formulated or implemented) they seek justification
under a broader set of values embodied or inherent in the Constitution which are
assume to express deeply anchored, general value commitments of the society
concerning the rights of individuals, social categories, organizations, or
governments.
III
The propensity of law to be guided by public opinion does not mean that the mores
do in fact playa large part in shaping the law. While that relationship may be
fundamental in primitive societies, as suggested by Bohannan ( 1965) in his concept
of double institutionalization, public opinion offers much less guidance in urban
societies because such societies are so normatively heterogeneous. As Dicey (1905)
pointed out concerning England in the late nineteenth century, there may not be a
public opinion on many issues, and if there is it may not be transmitted, or, if
transmitted, not be taken seriously in the shaping of laws. If anything, our situation
shows even greater diversity than did nineteenth century England.
Recourse to science as a basis for normative judgment may reflect the need for some
sort of consensus which is so lacking in our pluralistic, anomic society. But I am
skeptical at the prospect that science can support particular laws or mores in a
sustained and effective manner. It seems to me far more likely that it can contribute
to belief in some general principles and procedures which may eventually influence
mores and laws.
Science seems poorly fitted to shape laws because laws are so heavily normative in
content. They are expressions, that is, of what people feel to be proper behavior. The
societal sense of what is proper is derived largely from socialization and experience,
rather than from scientific observation. If anything, scientific studies of the range of
human societies demonstrate the enormous diversity in normative beliefs found in
human cultures. “The mores,” Sumner remarked, “can make anything right and
prevent the condemnation of anything." In the face of such diversity, the search for
universals succeeds if at all only at a highly abstract level. We may find that all
societies have some form of family, for example, but we must also acknowledge that
the family can be polygynous, monogamous or polyandrous; that it can be stable
over a lifetime or highly unstable; that it can be rigorously exclusive or co-existent
with extensive philandering; and so forth.
How then might science help to provide guidance to law, for example in the matter
of family structure? It might (1) call for toleration, (2) provide technical (functional)
information or (3) aid in the development of societal norms.
(1) The first possible use of biology is to reinforce the norm of toleration. Given the
diversity of human societies and of primate behavior, scientists might urge that laws
seeking to constrain freedom of behavior should be regularly avoided. Of course
such a position must specify limits. Law must presumably constrain freedom where
freedom is used to harm other individuals or the society. This classic position,
tracing back through J. S. Mill to John Locke, sets terms of inquiry but does not
answer the question. What behavior must be constrained to avoid harm to the
individual and the society?
Law frequently wrestles with such questions. An interesting example is found in
the instance of polygyny. In the early test of this issue, presented by the Mormon
religion, the Supreme Court upheld the proposition that freedom of religion did not
protect this practice from statutory prohibition even though the practice was deeply
imbedded in a system of religious belief. In doing so, the Court drew r on the
prevailing mores:
Polygamy has always been odious among the northern and western nations of
Europe and, until the establishment of the Mormon Church, was almost exclusively
a feature of the life of Asiatic and of African people. (Reynolds v. US, 1878: 164).
Later the Court condemned polygyny as a "return to barbarism"

(Church of Jesus Chnst of Latter Day Saints 'V. US, 1890: 49) and a "notorious
example of promiscuity" (Cleveland 'V. US, 1946: 19).
The Court thus expressed the prevailing mores and supported a" legislative policy
which enforced those mores. In order to override the . First Amendment guarantee of
freedom of religion, the Court was bound by its own doctrines to find that the
legislation had a "valid “secular purpose." Such a purpose was found in the effort to
preserve monogamy. The Court affirmed the legitimacy of that purpose primarily by
reasserting the monogamous mores of this society, calling to mind their
pervasiveness in the north-west European cultural area, and ethnocentrically
condemning the alternative family patterns as barbarian and alien.
Could the case be made scientifically that preserving monogamy is not a valid
purpose? All societies have mores of one kind or another The mere fact of
intersocial variations in mores, while true, does not seem to obviate the need for a
particular set of mores in any given society. Survival as a society may well depend
on some degree of normative consensus. Within wide limits, the fact of agreement
may be more important than the content or substance of the agreement. Law may be
seen as one device (perhaps the major device) by which a complex society can
achieve and express necessary normative agreement. Thus, a scientific argument for
a norm of toleration would not necessarily carry the day.
(2) If science is to attack laws which express the mores, it might have a better
chance if it questioned the instrumental effects achieved by particular laws of this
type. The courts are doctrinally prepared for such functional arguments. Secular
purposes, justifying limitations of First Amendment freedoms, are linked to effects.
An expressive purpose may be enough, but the courts are also concerned with "
consequences. If a law produces the opposite of its intended result, it may fail to
meet the test of a rational relationship.
In the polygyny cases, the Court used language of both purpose and effect. Writing
for the Court in Cleveland 'V. US (1946: 19), Justice Douglas not only condemned
polygyny on moral grounds, but also spoke of "the sharp repercussions that
[polygynous practices] have in the community." What repercussions he had in mind,
beyond the moral concern, is not clear.
One possibility worth considering concerns the economic significance of differing

family structures. It is the essence of anthropological functionalism that the
consequences of a particular practice must be understood in terms of the conditions
in which the practice
occurs. Polygyny, which is functional in an agricultural economy, (especially with a

high female/male ratio), does not retain its economic, social or reproductive
functions in industrial societies. This functional difference between types of
societies is supported not only by correlational observation but also by studies of
economic development. Those societies which undergo rapid modernizatlon tend to
move toward monogamy and those which are monogamous move more rapidly
toward modernization (Goode, 1.963). On that basis, one might propound the
argument that polygyny is dysfunctional for a modernizing or modernized society.
Would that kind of analysis lead the scientist inevitably to support laws against
polygyny? Even at the technical level, for example, what ab out post-industrial
society?
Another value in the polygyny case is the role of religious belief. A functional
analysis produces indeterminate results here as well. Sociobiology may tell us
something about the universality of religious belief, manifested in primate "sun-
worshipping ceremonies" or in prehistoric hominid burial practices. If every primate
society manifests religious belief, what significance does that carry for polygynous
religion? Does it mean that (1) every society must have a single, unifying religious
belief which should be supported by social and legal controls, or (2) that every
member of society should be free to practice non-intrusive religion together with
like- minded others? Depending on the choice, the religious factor also could weigh
for or against the prohibition of polygyny.
(3) There is a third way in which science might contribute to the resolution of legal
policy questions by aiding in the development of social norms. Scientific efforts
might be best employed if engaged in trying to understand the ways ill which
normative systems develop and work in societies. We know that viable societies
always maintain some minimal degree of normative consensus. Failing to do so, a
society loses its stability, its capacity to maintain the functions which it must
perform to hold together. In such circumstances, societal collapse can lead to
immediate misery and uncertain results as the process of reconstruction begins. In
general, societies strive in various ways to maintain their equilibrium and' avoid
collapse, anarchy or revolution.
Scientific knowledge may affect these outcomes by helping us to understand

minimal conditions of normative order. Contemporary work in the social sciences,
jurisprudence and philosophy has given us
some interesting leads which suggest that a useful empirical inquiry is possible.
Sociobiology can, I believe, play a valuable role in such work.
IV
All human societies (and many other species) have standards of behavior which
regulate interaction. Social control provides a basis for transmitting and enforcing
these behavioral codes. These standards are in addition reinforced by the reciprocal
rewards or reciprocities which are associated with the interactions. Controls and
reciprocities constitute the mutually reinforcing mechanisms which maintain these
conditions of normative order.
In every human society, these codes are verbally formulated as norms (though not
necessarily as laws) used to guide and evaluate individual behavior. As noted, they
vary widely in content from one, society to another. Yet they seem to have in
common the following characteristics:
(1) they are generated by and manifested in social interaction;

(2) they are enforced against those who violate them;
(3) they lead to reward for those who comply with them;
(4) they are supported by a substantial segment of the society, directly or
vicariously;
(5) support for them is enhanced by the belief that they are fair or just.
Even a casual sampling of anthropological literature illustrates each of these. Their

universality can for the present be assumed.
My hunch is that these standards are a product of some intrinsic property of the
human species, quite possibly shared with many other species. These codes may
vary widely, like language, from one society to another. But like language they may
reflect common species characteristics. If so, it is important that we understand the
deep structure of these characteristics.
By putting together the ideas of several students of the concept of justice, we can get
some idea of what might be involved. Piaget (1932) tells us that rules are adopted by
groups of children after they reach a certain age (seven to nine years) and have had a
chance to play together. Prior to that age, they have believed that rules come from
some powerful, inflexible authority (e.g. God, father, the mayor) outside of
themselves by whom they are preserved and enforced. This
"heteronomous" orientation toward authority gives way as they mature. It is replaced

by an "autonomous" orientation such that the children realize that rules are made by
the group for its convenience and that they may be changed to enhance the
satisfaction of the group.
An element emphasized by Rawls—in potential conflict with Piaget—is that the

rules, once accepted, must be stated in advance of knowing to whom they will
apply, that they cannot be perceived as fair if it is known that they change to suit the
advantage of a single person (Rawls, 1971: 11-17). This accords with Lon Fuller's
idea that warning, stability, and even-handed administration are essential elements
of "the morality that makes law possible."
The idea of standards does not mean equality, however Piaget notes that a big
winner in children's games may be required to return some of his winnings to the
game to keep it going, but that he is never deprived of all gains. Barrington Moore
reflects at length on the (for ~ him) puzzling phenomenon that people accept a very
small share ~ rather than rebelling. His eventual explanation-that they come to
accept the very deprivation as a normatively good thing-he illustrates with a story
from India. When a harijan retainer is invited into the home of a newly equalitarian
Brahmin youth, recently returned from school, the invitation is vigorously but
courteously rejected by the servant with the words, "you may have given up your
religion, young master, but we have not given up ours" (Moore, 1978: 61). While
such standards tend to be fixed (more so perhaps than Piaget suggests) in stable
cultures, they are regularly being given new normative content in open, rapidly-
changing societies such as ours. Walster points out that a perception of imbalance
between the value of work contribution and compensation for the work leads to a
tension. This is resolved, her research suggests, in one of two mutually incompatible
ways: by increasing the compensation or by derogating the quality of the work or
worker .
These ideas (and many more like them) indicate the prospect that scholarship will
help us increasingly to understand the social bases of behavioral standards. I believe
that sociobiology can make its most important contribution in this area.
To learn more about norm formation does not imply any particular or immediate
effects on questions of legal policy. In the example of 4 family structure, the
prohibition against polygyny may be so deeply rooted in the structure of our society
that any analysis would simply .i confirm the strength of the taboo and its
inevitability under existing social conditions. Even so, it would be interesting to
learn what might
be discovered if, for example, advocates and opponents of polygamy (let us include
polyandry as well as polygyny) were able to consider the consequences of
permitting these forms. Why should we not develop procedures for discussing, even
simulating, variations which might have societal value? Such deliberation should be
informed by scientific \ knowledge, not displaced by it.
There are many areas in which the society seems burdened because of difficulties in
discovering latent normative dispositions. Better understanding of such dispositions
is, I believe, desirable not only for the sake of the knowledge itself but also for the
uses to which it could be put. Like any knowledge, it has the potential for misuse.
But a program aimed at facilitating normative consensus by structuring voluntary
interactions and encouraging mutually satisfying resolutions seems to me worth
developing. Can sociobiology help in this by informing those charged with
responsibility for legal procedures how to arrange them more effectively?
Better understanding of norm-forming processes is of great importance for

maintaining minimum conditions of normative order. In open, heterogeneous
societies such as this one, the evidence of anomie (normlessness) is found on every
side. Some of this normative diversity provides valuable liberties for the individual
and variations for the society. When anomie passes a certain limit (which we cannot
yet identify), however, it can destroy mutual trust, confidence, legitimacy and a
willingness to adhere to and support the code of the society.
Open societies particularly need to cultivate congruence between law and the norms
of society. Where these do not match, law tends to be viewed as alien to one's
concerns if not an oppressive instrument imposed in the interest of a dominant class.
But the absence of pervasive norms means that law will often fail to accord with
someone's (if not everyone's) conception of what is right. The remedy for this state
of affairs is not necessarily to be found in a renewed effort to use scientific expertise
to shape and support particular legal decisions. Sociobiology and other sciences of
human behavior may contribute much more effectively and appropriately to legal
policy formation by explaining norm-forming processes. With the help of such
knowledge law may provide the best opportunities for normative coalescence. It can
promote interactions between opposing parties which facilitate dispute resolution
and norm formation. Labor-management relations illustrate that possibility (Fuller,
1971). Law can facilitate the orderly termination of relations where these need to be
maintained. This is illustrated by property
division and custody arrangements in marital dissolution cases [C; (Mnookin and
Kornhauser, 1979). Law can provide a setting in which latent mores can be
collectively expressed and examined. This is illustrated by the use of aggregate jury
determinations in declaring the death penalty for rape unconstitutional (because
disproportionate, excessIve and random) (Schwartz, 1979: 319-325).
These instances are intended only to illustrate the potentialities for law, by arranging
interaction, to facilitate dispute resolution, mutual satisfaction and norm formation.
Much more remains to be done in this direction. Any knowledge that can be found
in sociobiology concerning how disputes are resolved, reciprocities generated and
norms formed will be welcomed as a contribution, where it is most needed, to the
reconciliation of law and the mores in an open, stable, democratic society.
CASES CITED
Brawner v. US, 471 F.2d969 (D,C.Cir.1972).
Cleveland v. US, 329 U.S. 14 (1946).
Church of Jesus Christ of Latter Day Saints v. US 49 (1890).
Durham v. US, 214 F.2d 862 (1954).
Furman v Georgia, 408 US 238 ( 1972 ).
M'naghten-Daniel M'Naghten's Case 8 Eng. Rep. 718, 10 CI. & Fin 200
(1843).
Reynolds v. US, 98 U.S. 145 (1878).
Rosenfield, In re, 157 F; Supp. 18 (D.D.C. 1957).
Trop v. Dulles, 356 US 86 (1958).
Anthropology, law and genetic inheritance

E. ADAMSON HOEBEL
2273 Folwell Street, St. Paul, MN 55/08, USA
Although anthropology has accepted organic evolution as established fact for

more than a century, anthropologists have made few serious attempts to anchor
human institutions in the specific social behavior of animal aggregations.
Throughout the first half of this century, American and
British social and cultural anthropologists with few exceptions rejected cultural
evolutionary theory in favor of structural-functional study of directly
observable societies. Contemporary anthropologists tend to reject or downgrade
the significance of panhuman genetic factors in favor of the dogma of culture as
sui generis. However, law as a phenomenon of social structure can probably be
linked fruitfully to behavioral research with genetic implications—with
cautious stretching of the limits of naivety, through examination of the so-
called imperatives of social maintenance, and the 'law jobs'.
In approaching the problem of whether the ordering of social behavior in human

populations may in part be rooted in our genetic inheritance, we accept as
established fact that all human beings are basically the product of organic evolution.
We acknowledge with Dobzhansky (1962:115) that 'many features of human
ontogeny make no sense at all except on the assumption that they are retentions of
developmental patterns of remote ancestors. This is clear enough and has been
accepted by anthropologists for more than a century-but only with respect to
anatomy and certain physiological processes.
In the early years of post-Darwinian enthusiasm, Western anthropologists such as
Tylor, Lubbock, Frazer, Westermarck, Bachofen and Morgan built theoretical
systems of social and cultural evolution enunciating developmental sequences for
the emergence of human subsistence techniques, mating, kinship, property,
government and religion. In a more modest way, Sir Henry Maine did the same for
the evolution of law. In no instance, however, was any serious attempt made to
anchor human institutions in the specific social behavior of animal aggregations,
although hypothetical assumptions concerning prehistoric forms of mating, kinship
and 'horde' organization were advanced.
Early in the present century, as ethnology became self-consciously empirical,
anthropologists, except archaeologists, became increasingly, hostile to theories of
cultural evolution. In the United States, Robert Lowie (1920:107) proclaimed, 'There
is no royal road to the comprehension of cultural phenomena and painstakingly
demonstrated that the social history of a particular people cannot be reconstructed
from any generally valid scheme of evolution but only in the light of its known and
probably cultural relations with neighboring peoples’. Morgan's and Frazer's
deductive interpretations of the sororate and levirate forms of marriage as cultural
relics of group marriage were dismissed as 'empty guesses, which may be
disregarded. They are real
institutions intelligible in their context; they are not rendered one ~ whit more
intelligible by conceiving of them as a survival of a condition which has never been
observed' (Lowie, 1920: 185).
From 1910 to 1930 American anthropology limited itself to intensive ethnographic
work in field studies and in limited historical reconstructions of culture growth
among Indian tribes. In Britain, both Malinowski and Radcliffe-Brown rejected
cultural evolution and historical reconstruction as scientifically bootless because
they dealt with empirically unverifiable data. Radcliffe-Brown turned British, and
part of American, social anthropology into a search for regularities ~ in social
structure based largely on Durkheim's proposition that social reality is external and
sui genens, and that behavior and personality are the product of individual
internalization of the consscience collective the cultural ordering of norms and
values. Although Radcliffe-Brown acknowledged both organic and social
evolution, in practice he evinced no interest in either, holding that social
anthropology is concerned only with persons in reciprocal relations based on
mutual interests and values socially determined.
In sum, social anthropologists in the United States and England, except for
Malinowski's naive and highly generalized use of bio-genetic 'needs' and 'drives' as
culture-generating forces, tended to restrict their researches to social structure and
social facts. They either dogmatically reject the possible significance of panhuman
genetic factors or more modestly assume that the question is one of such low priority
in research strategies that it is better for them not to devote limited energies and
resources to pursuit of the question.
The logical foundations of this position have been spelled out in a symposium
volume, Closed Systems and Open Minds: the Limits of Naivity in Social
Anthropology {1964) edited by Max Gluckman. It deals with two issues of
importance to our fundamental problem here: can the assumptions and findings of
biophysical behavioral research on the one hand and those of the sociology and
anthropology of law on the other, be fruitfully linked? These two issues are (1)
limiting one's field of study and (2) making naive assumptions outside one's
special field. He identifies five procedures used in scholarly formulations of
comprehensive endeavors: incorporating complex facts without analysis;
circumscribing a field; abridging the conclusions of other sciences; making naive
assumptions about aspects of reality other than those under investigation; and
simplifying events within the field of investigation (Gluckman, 1964: 17).
We all do these things. Only by narrowing the field of inquiry can
we achieve expertise and advance empirical knowledge of reality. We all
incorporate certain facts as given-an anthropologist will accept the fact of a rain
forest as a tribal habitat without attempting a meterological explanation of it. We
abridge the findings of atmospheric—and astrophysics when we attempt to deal with
important long range climatic cycles in the Pleistocene era. We assume a need for
individual identity, which is outside the area of our competence. And of course we
simplify the facts within our field of investigation-any ethnographic report is a more
or less generalized construct of what is going on in tribal society.
The problem is; how tightly can we maintain our closed systems if without
stultifying development of a grand theory of human beings, and how much openness
and naivete can we accept in cross-disciplincary incorporation without muddying
our perceptions with dubious trash?
Sociobiology is in bad repute. Setting aside widespread emotional prejudice
emanating from political bias, repugnance to sociobiology" rests on its proclivity
to reject circumscription of its field while making , innumerable assumptions about
aspects of reality outside the fields of biology and ethology of insects and lower
orders of animals. Worse, it excessively simplifies organic phenomena within, its
own field.
Lumsden & Wilson’s recent volume (1981) strives to rectify this last failing and
they have made some progress in linking genes and culture. However, Washburn,
one of our most innovative physical anthropologists and a pioneer in studies of
primate behavior, warns that modern genetics is radically different from the kind
described by Wilson (1978) and that sociobiology omits any reference to the
complications of modern evolutionary theory. We are told that over the past 20
years molecular biology has revolutionized evolutionary theory. Dobzhansky et al.
(1977: 156) state that recent research shows a seeming deluge of genetic
variability' that 'proves the classical theory is clearly invalid', and that
determination of the relative importance of selection and chance or neutral facts
stands as 'the most important unsolved[emphasis added problem in our
understanding of the mechanisms that bring about 'biological evolution'. Yet
Chagnon and Irons's Evolutionary biology and Human Social behavior (1979) in
Washburn's view rests on 'the synthetic theory of the 1940s plus inclusive fitness',
while its 78 pages of bibliography contain 'almost no ~ references to even the most
important developments in evolutionary theory (Washburn, 1982).
We must tread cautiously, for few anthropologists (including this
author) know anything about contemporary scientific evolutionary theory.
Although Stent (1980: 4-10) devotes respectful attention to social anthropologist
Sahlins' polemic attack on sociobiology, physical anthropologist Konner (who
does know the field) dismisses Sahlins' critique on the ground that 'his grasp of the
basic scientific issues is so poor that discussion of his critique is really impossible'
(Konner , ~ 1982).
It is imperative that we all recognize and respect the limits of our naivete.
Now, can law fruitfully be linked to behavioral research with genetic implications?
I would say: only with a cautious stretching of the limits of naivete. This, I, for one,
am willing to do with open interest and limited expectations.
Much has been achieved in behavioral science research in comparative
jurisprudence in the last five decades. 'Law is one of the best-studied subdisciplines
of anthropology; the literature is small but of high quality' (Bohannan, 1963: 284). It
has a strong empirical base ,"7 emerging from a series of carefully executed field
studies of a variety of .,~; tribal systems in North and Central America, Africa, and
Oceania, ranging from simple to complex in subsistence economy and social
organization. It has a common approach rooted in the so-called trouble case
method—the observation and recording of issues of dispute and conflict, and how
they are perceived, conceived and resolved by the members of a given society. It has
at the same time generated considerable healthy dispute among its theoreticians over
such questions as to how law may best be conceived and identified, to what extent it
is a real phenomenon in its own right or a mental construct of the observing analyst.
The criteria for identification of law-the nature of norms, dispute handling processes,
sanctions, authority and levels of hierarchy within a social system to which any fj
body of law may apply-have effects that differentially narrow or broaden the domain
of law, according to the preferences of the researcher.
Here I offer a compact abridgement of my own views of what legal anthropology
has to offer on the nature and function of law in human societies and the degree to
which it may be rooted in the organic nature of man and subhuman species. Law is
a cultural phenomenon that cannot be studied apart from the entire social entity of
which it is a part. Law is a human invention developed as an adaptive mechanism
for the maintenance (effective survival) of the individuals, subgroups and the entity
that constitute a society. It is that aspect of social
structure that serves to standardize behavior and to regulate relations between
individuals and groups. It can be focused on as a loosely discrete phenomenon but
never to the neglect of its cultural matrix and never without linkage to all other
aspects of the productive and social system of which it is but a small part.
Its attributes, for purpose of identification are:
(1) Regularity, Law consists of social norms identified a modal behavior
characteristics ill defined situations. Regularity IS rarely absolute; behavior is
variable around a clustered mode, and most legal systems make allowance for
'permissible leeway'. Regularity introduces the element of predictability.
(2) Sanction. All social norms are sanctioned. Conformance evokes rewarding
(positive) responses, internal and/or social; deviance evokes punishing (negative)
social and, perhaps, internal individual responses. Legal norms are sanctioned by
the forceful application of physical injury or the confiscation of material '0'0 ~
goods-restitution or the payment of damages or fines.
(3) Legitimate ('official') authority. The legal process is initiated and carried
through by either the wronged person and members of his subgroup or by
designated officials (headmen, chiefs, priests, judges, councils) explicitly
representing the societal interest as a whole. Due process means that the
procedures used are socially recognized and accepted as right and proper and that
the sanctions are appropriate and acceptable.
There are indeed functional prerequisites for survival: organically and genetically
determined and culturally conditioned for the individual human organism.,
culturally adapted in the main for human populations as societies. Whether there
are genome presets that underlie a few cultural adaptations remains an open
question. Certainly, biology imposes limitations on what may be culturally
achieved; more than that, it mechanically controls much of the essentially organic
.0.." field of individual behavior (neurological, metabolic, reproductive etc.).
A socio-anthropological formulation of the functional prerequisites for societal
maintenance, which builds on the earlier work of Malinowski and Talcott Parsons,
sets out the following:
(1) to maintain the biologic functioning of the group members;
(2) to reproduce new members of the group;
(3) to socialize new members into functioning adults;

(4) to produce and distribute goods and services necessary to life;
(5) to maintain order within the group and between it and outsiders;
(6) to define the 'meaning of life' and maintain the motivation to survive and engage
in the activities necessary for survival, (Bennett & Tumin, 1948),
Legal institutions are directed to function (5) above. Four specific maintenance tasks
of law may be identified.
(1) To define relationships among the members of a society, to identify what
activities are permitted and what are ruled out, so as to maintain at least minimal
integration between the activities of individuals and groups within the society: the
ordering of the fundamentals of living together.
(2) To allocate authority and determine who may legitimately exercise physical
coercion as a socially exercised privilege/ right-and how. This point is derived from
the necessity of channeling aggression, of taming naked force and directing ~ force
to the maintenance of order within the social system.
(3) To dispose of trouble cases as they arise; to heal social breaches when breaches
of defined relations are alleged to have occurred.
(4) Explicitly, to redefine relations between individuals and groups as the conditions
of life change; to maintain adaptability.
The specific substantive contents of primitive law systems have been found to be
highly variable in accordance with the numerous manifestations of social structure
around the world. But they are not infinite. Social evolutionism is once again in
vogue among American anthropologists. As for myself, I have long held that it can
be r, discerned in broad and general terms in the area of law (Hoebel, 1954: ~
Chapter 12). I believe that the postulate of coevolution in organic and cultural
development holds good. How much of social ordering is genetically predicted for
human beings is no moot question. My mind is open to the search for epigenetic
patterns in the multifarious formation of legal culturgens, but let us not each in our
separate disciplines fail to hold a tight rein on our respective limits of naivete.
[34]
Questions of the legal scholar
concerning the so-called sense of justice
MANFRED REHBINDER
School of Law, University of Zurich, CH-8032 Zurich, Switzerland
(translated by Ulrike E. Lieder)
With laypersons the sense of justice is one of the determinants of legal

behavior. With lawyers it is a source of law. The origin and nature of this
phenomenon are uncertain. For almost 100 years the debate in German
jurisprudence marked time: is the sense of justice innate or acquired? The
question sprang to life in 1979 when two suggestions were made: ( I) that
the sense of justice is related to what Freud called identification, and (2) that
it is associated with a norm filter of the sort that Swiss neurologist von
Monahow described in 1927 as the biological conscience. Extending these
excursions of jurisprudence into neighboring disciplines, we ask: (a) is the
sense of justice related to the internal reward system? (b) does the sense of
justice show different levels of maturation? and (3) is the sense of justice a
biologically regulated iterative process that leads to self-conditioning? If we
lawyers could have empirical answers to such questions, our understanding
of the working of the law would be enhanced.
For almost 100 years legal scholars have looked to the neighboring disciplines for
help in their controversy on the nature of the sense of justice' and its role in assuring
the righteousness of the law. This controversy which continues undiminished to this
day started in Europe with a lecture by the eminent German scholar Rudolf von
Jhering (1884) entitled ‘On the Origins of the Sense of Justice', at a meeting of the
Law Association in Vienna. Jhering spoke against nativism, a term coined by the
physicist Helmholtz (Bihler, 1979: 3). Nativism claims that man has innate value
concepts responsible for his legal behavior. Jhering countered with the maxim, ‘It is
not the sense of justice which created law, but rather, it is law which created the
sense of justice.' (Jhering, 1877: xiii). Why this controversy developed among legal
scholars needs same explanati6n. Laypersons probably
think that the sense of justice equals the common man's concept of right and wrong,
and that it is something that lawyers do not need. After all, lawyers should know
their law-generally speaking, at least or they should know where to find it. Then
why do they need a sense of justice?
THE SENSE OF JUSTICE IN LEGAL SCIENCE
Although law is a rational instrument of social control, lawyers can-unfortunately-

not do without emotion or a sense of justice, <cc, their own as well as that of others.
Creating norms
This becomes evident when law as a device of social control does not provide a
legal norm for the solution of a given problem. In this case, the lawyer as legislator
must create a new norm. This is also true for judges if there are gaps in the law: like
legislators, they must create a new norm. For example, Article I, Paragraph 2 of the
Swiss civil code ~ provides that if the code does not contain an applicable rule, the
judge .is to rule in accordance with common law. If there is no common law, he is
to decide according to the rules which he would establish if he were the legislator.
In this case, judges will either 'find' spontaneously a just solution based on their
intuition, and later give a rational explanation for their decision; or, if they initially
do not prefer a specific solution, they will rationally examine the problem and find
a solution based on certain rational aspects, and will then (more or less satisfied)
accept this solution as 'just' and attempt to convince others. In the case of very
'technical' legal regulations the judge may not have any intuitive feelings at all, yet
he must still consider the feeling of others as to what constitutes a fair and just
decision. The sense of justice thus plays a significant role in the creation of new
norms. It is important in assuring the effectiveness of these norms, since the degree
of compliance will be low if the norms are not in accord with the concept of justice
within the social group.
Application of the norms
The sense of justice within a society also is instrumental in determining the degree
of compliance with a norm, and for this reason is a decisive factor in the
application of existing legal norms. The consideration of the consequences of the
application of legal norms is based on the fourth method of interpretation of law
which—according to
[36] AW, BIOLOGY AND CULTURE
prevailing opinion-is the most important of the four classical methods of

interpretation. Existing legal norms can be interpreted grammatically, historically,
systematically and teleologically. The teleological interpretation attempts to optimize
the intent of the law. To this end, the judge must know which interpretation of an
abstract rule will achieve the effect on society intended by that rule. This requires an
awareness of the moral consensus of the particular social group in which the rule
functions, and to what degree it fulfills their expectations of justice.
However, legal scholars do not agree on which of the four techniques of

argumentation is the most persuasive and therefore most decisive for the solution
of practical legal problems within the framework of existing rules. That is why, in
the eyes of laypersons, the application of law is very unpredictable and thus
'unscientific'. According to today's prevailing view the choice of methods depends
on the preconceived ideas of the judge and on whether those to whom the law is
addressed share his preconceptions and will therefore accept his interpretation
(Esser, 1972). The mechanism which was described as important in the creation of
norms also plays a role in these preconceptions, that is the sense of justice of the
judge and of the 'law consumers' plays an important role indeed, along with
rational argumentation. Anglo- American case law, which accepts court decisions-
concrete application of abstract rules-as new law (Geiger, 1970: 253-261 is the
best explanation) may well have been aware of this situation.
Judging a normative error
There are numerous problems which the judge can only solve with the help of his
sense of justice. This is always the case when a rule refers to normative standards
as, e.g., when referring to public policy (the definition of the Supreme Court of the
Federal republic of Germany is the 'sense of justice of all those who think in terms
of fairness and justice' BGHZ 52/17,20), morality or other undefined legal
concepts such as human dignity or pornography. To illustrate this point, let us
examine how law deals with a normative error. The proliferation of laws in the
welfare state makes it impossible to know all the laws. To an ever increasing
extent, the 'law consumer' can claim with good reason that he/she did not know the
law. In earlier times, the Roman legal principle 'error iuris nocet' was acceptable.
Today, however, we must look for more sophisticated solutions. In establishing an
accused's guilt or innocence, the law now demands that the court inquires whether
the accused had made a
reasonable effort to consider the legality of his actions. It is not expected that they
know the exact terms of the law but that they must make an effort, within a
layperson's ability, to understand the legal situation. This awareness of the legal
consequences is called 'parallel evaluation in the layperson's sphere' in current
criminal law (Jeschek, 1978: 236f). Because the layperson lacks professional
expertise, their evaluation has to be based on emotions.
THE CONCEPT OF THE SENSE OF JUSTICE

IN GERMAN LEGAL LITERATURE
Lawyers, of course, do not wish to give the impression that they work with the help
of emotions. The legal scholars' professional arguments about the nature and
significance of the sense of justice, therefore, have not focused much on the
principal application of law and the creation of legal norms, the fields of legal logic
(Ehrlich, 1918) and legal policy. Rather, the subject has been discussed among legal
philosophers who have focused on the justice of positive law. It is interesting to
note that-with a few exceptions to be discussed later the basic ideas have hardly
changed since Jhering's time, in spite of progress which has since been made in
sociology, psychology, the behavioral sciences and-most recently-in biology.
Is the sense of justice innate or acquired?
In his time, Jhering fought against a school of philosophy whose most prominent
representative was Gustav Ruemelin. Following the political unification of
Germany in the Bismarck Reich, Ruemelin, in his 1871 speech 'On the Sense of
Justice' (Ueber das Rechtsgefuehl], had stated that law had its basic root in an
innate sense of order, 'which aims at the harmony of our lives and of the world'.
This speech reflected the spirit of anew nationalism which supported the idea of a
national codification of the existing law. Ruemelin felt that law should create an
ideal order of life conditions which corresponds to the people's concept of morality,
expressed in the sense of justice and the conscience of the individual. Tensions
would arise when the positive law no longer corresponds to society's basic moral
ideas which are, of course, subject to historical changes. These tensions could be
resolved by adjusting the law to the prevailing concept of justice. Opposed to this
interpretation, von Jhering advocated a theory of the empirical, 'historical' evolution
of the sense of justice. According to his theory, the experiences of the prevailing
social conditions are 'mentally
inhaled', that is, internalized by the individual, and become part of their personality.
However, each of these theories on the origins of the sense of justice had to make
considerable concessions to the other. Ruemelin could not claim that there exists a
constant, invariable sense of justice rather, he assumes that the moral evolution of
man outpaces existing law, and: as a consequence, existing law will no longer
satisfy societies sense of Justice. Von Jhenng could not claim that the sense of
Justice builds only on acquired concepts, but attributes the progress of law to the
insights of 'great minds', the intuition of geniuses.
To this day, these two explanations of the origins of the sense of justice—innate
(nativistic) as opposed to acquired (historical)—are still at the center of the
controversy. In his Psychoanalytical Jurisprudence, Albert A. Ehrenzweig (1973:
219) does not call the sense of justice inherent in all men an instinct, because—as
he states—its biological origins cannot be traced. He postulates, however, that in its
function, the sense of justice determines the law just as hunger controls the intake
of food, or the sex drive controls procreation. Nature is too wise to rely solely on
man's intellectual capacities. rather, due to his genetic heritage and as a result of
long periods of acculturation, man was endowed with a sense .of justice, just as he
inherited other drives, like hunger and sex. In this function, the sense of Justice had
always been recognized both by the advocates of natural law (Naturrechtler] and
those of positive law (Positivisten].
Erwin Riezler (1969: 40-46), however, emphasizes in what was until recently the
leading study on the subject that the sense of justice depends on cognitive
(intellectual) concepts which are not innate, but historically conditioned and
empirically acquired, yet law and the sense of Justice are interdependent.
Is the sense of justice a source of law?
Innate and acquired—these are possible answers to the question of whether the
emotional response of right or wrong IS part of our genetic endowment or whether
it has been learned in the process of socialization. Innate and acquired—these are
also the answers to the socio-psychological question of priorities: law before the
sense of justice or the sense of justice before law. To Jhering, the sense of justice
is only a diagnostic tool for finding the existing positive law. To most other
authors, the sense of justice, in addition to positive law, is also a source of law.
This applies in particular to all those who believe in a natural law,

superior to positive law. The modern-day advocates of natural law have introduced
Max Scheler's and Nicolai Hartmann's concept of material ethics into the discussion
(Bihler, 1979:10). They consider the sense of justice a value-finding tool, a genuine
source of knowledge revealing absolute objective values (Hubmann, 1977), even
though they might be subject to historical changes. There is also a consensus among
the majority of legal scholars who do not recognize any a priori, standards, that the
sense of justice works as a source of law in those areas we mentioned in the
beginning—legislation and adjudication. Here, an important. contribution was made
by Erwin Riezler (1969)," who developed a tripartition of the sense of Justice in
1923 (see also Weimar, 1969; Geiger, 1970: 412-415; Venzlaff, 1973). According to
Riezler, the sense of justice can be:
(a) a feeling of what law is at a given time (positive sense of justice)
(b) a feeling of what the law should be (ideal sense of justice) and
(c) a feeling that law should be obeyed (general sense of justice, respect for the legal
Order).
Therefore only in the sense of (b) can the sense of justice be regarded as an
independent source of law.
Is the sense of justice rational?
As an independent source of law, the sense of justice requires rational explanations

in order to be accepted by others. This is as true for a constant natural law as for a
natural law with changing contents. Rational criteria must be available to explain
contradictory statements on a given concept of justice and to provide reasons
documenting the error and supporting the chosen alternative (Bihler, 1979: 20).
However, this also applies to the creation of norms by an ideal sense of justice as
Riezler sees it, because the legal norm thus created requires a rational basis if it is to
be accepted as binding. Therefore, it is not surprising that legal scholars agree that
the sense of justice has, besides its emotional component, also a strong rational one
with a clear pre-ponderance toward the intellectual (Bihler, 1979: 22).
However, the contradictio in adjecto of an intellectual emotion should make legal

scholars aware that something must be wrong. Before we direct questions to the
neighboring sciences that are competent in this problem area, let us consider two
examples of recently published German legal literature that document new insights
developed in the fields of psychology and neurophysiology.
Sense of justice without law? .
The first example is Michael Bihler's Psychologie der Rechtsgewinnung

(Psychology of Acquiring Legal Norms) which appeared in 1979. It makes Riezler's
work, which was until then the leading work on the sense of justice, obsolete. Bihler
postulates a model in which the emotional and rational elements of the sense of
justice are separate. He does not see emotion as a product of the sense of justice:
rather, he suggests that the feeling of Justice is a psychological identification (in
Freud's sense) with a one-sided bias. The sense of justice spontaneously; favors one
party over the other in a legal conflict. The person who takes sides is motivated by
empathy. The sense of justice as an act of solidarity in this sense is 'the result of a
process of identification which is triggered when a predetermined partial identity
between the person involved in the conflict and the person taking sides is affectively
cathected, i.e. when the case is provocative' (Bihler, 1979: 101).
Does the sense of justice have to go through a normative filter?
The second example of how non-legal insights have shed new light on old legal
problems concerns the application of the so-called biological normative filter to the
consideration of normative errors in law. In his 1979 study, Zur juristischen
Dimension des Gewissens (On the Legal Dimension of Conscience), Ernst E. Hirsch
utilizes the theory of a biological conscience which had been developed by
Constantin von Monakow in 1927. Hirsch applies this theory to the legal system and
especially to the assessment of lawyers' behavior during the Nazi regime. Should
hard-core National Socialists have been told by their sense of justice to reject the
then valid law if they had made what the German Supreme Court requires them to
do an adequate effort to assess the legality of their actions'? Hirsch says that,
according to von Monakow, man has innate behavioral programs, located in the
limbic system. These programs are later internalized and appear to him as clearly
evident commandments that must be obeyed. This so-called biological conscience
determines his moral awareness. Every person thus has an innate biological
normative filter. This filter is supplemented by another normative filter, acquired
through learning, which reflects the moral norms prevalent in his social
environment. 'Because " man, due to his lack of instincts, is by nature a cultural
being and therefore vulnerable and exposed to suggestive influences of all kinds, his
biological normative filter can be restricted or even totally blocked by the cultural
one. As a result, the individual's moral behavior is primarily or exclusively
influenced by (the conditions of the cultural
environment in which he lives' (Hirsch, 1979: 82). 'Conscience' is a biologically pre-

programmed 'sense of justice' whose contents, however, can be modified and
manipulated by the social environment.
QUESTIONS TO THE NEIGHBORING SCIENCES ON
THE 'NATURE' OF THE SENSE OF JUSTICE
In the case of the two examples mentioned above, it took 50 years to introduce the
insights of neighboring sciences into legal reasoning. This is by no means an
exception; rather it is symptomatic for jurisprudence. The conservatism of this
discipline should certainly not be seen merely in a negative light, for the latest fad
is not always the best. Yet we wish that questions would be addressed to the
neighboring sciences on a more updated basis to explore the relevance of new
discoveries for law and that we would devote more comprehensive studies to this
subject. In the following, I will concentrate on some approaches that might help to
explain the sense of justice.
Sense of justice as social interest?
In the legal literature, the sense of justice has a dual character. Emotional and
rational elements are inseparable intermingled, albeit with a clear emphasis on the
intellect (Bihler, 1979:22). This epistomological duality of an intellectual emotion
IS probably due to the tendency to compromise inherent in legal reasoning. If the
sense of justice is a source of law, then, according to the famous Article 1,
Paragraph 2 of the Swiss Civil Code, the creation of law would have to follow
Kant's categorical imperative. However, Kant considers this action to be in the
world of the a priori. Thus, the spontaneity of reason and not emotions, as proposed
by Adam Smith in his theory of moral sentiment, determine such judgments.
However, legal scholars are unable to choose between the idealistic and the
naturalistic ethic (a controversy that goes back to Plato and Aristotle), and they
therefore attempt to combine the two. But a scientific approach to law has to
include the social sciences. Therefore it seems appropriate to disregard the world of
the a priori for the time being and to ask empirical questions about 'the sense of
justice'. The sense of justice is thus a psychological process. As was mentioned
before, Bihler characterizes it, with Freud, as an emotional process of identification.
I wonder if it might not be I better to characterize it-with Alfred Adler (1928: 267-
272)-as social 'interest (Gemeinschaftsgefuehl—'feeling of community').
Adler's 'Gemeinschaftsgefuehl' is, in principle, also an identification

with one point of view or another. Adler describes it as the ability 'to see with the
eyes of another person, to hear with the ears of another person, to feel with the heart
of another person' (Adler, 1928: 267). However, if we rely on Freud, as Ehrenzweig
did in his psychoanalytical jurisprudence, we would focus too much on the
individual and his psychological problems. By contrast, as he says himself, Adler's
individual psychology is, in reality, a social psychology because it focuses
primarily on the relations between the individual and the community. This
interpretation comes close to the concept of law as a device of social control. It
seems to be more adequate to explain legal processes using the concepts and
theorems of individual psychology than using those of psychoanalysis. It has
already been emphasized elsewhere that the sense of justice is not so much the
identification with one's own self, but that it also means empathizing with someone
else's situation according to the principle 'one man's words are no man's words, you
have to listen to both sides' (Rehfeldt & Rehbinder, 1978: 142).
If one looks at the sense of justice as social interest, the question remains why a
given legal solution, i.e. a certain point of view, is considered to be right and just.
Bihler believes that here the intellect is not actively involved, but that we can see a
transference of emotional "evaluation at work, which is familiar from studies of
early child development. The child does not make a distinction between internal
feelings and external objects; rather, he/she perceives his/her feelings as
characteristics of the object. The adult experiences much the same way during the
psychological process of making value judgments. In the learning process, he/she
emotionally internalizes the values prevalent in society and considers them part of
the object [Haftwertkonzept] (Bihler, 1979: 135-145). A solution is considered just
because the criterion 'just' is perceived as apart of the object being evaluated. Since
Theodor Geiger's detailed discussion of the practical nihilism of values, the
sociology of law knows that speaking of a just solution is theoretically unfounded
and merely an illusive justification of an ;i emotional judgment. But the
Haftwertkonzept explains at the same time why this illusion is a psychological
reality.
Can the sense of justice trigger or express a sense of well-being?
If the sense of justice is no more than social interest and if there are no reasonable
grounds for calling the resulting decisions 'just', the question arises as to why this
statement is made always and everywhere. Let us say that we put ourselves in
someone else's position and
decide whether he is right or wrong. Then we would like to see the world
organized according to these evaluations. In extreme cases, this desire goes to the
point of self-destruction (as is shown, for instance, by Heinrich Kleist in his
novella Michael Kohlhaas). Could this desire for ‘justice’ possibly be connected to
a psychological gratification, to a sense of well-being? This idea already appears in
Adam Smith's theory .of moral sentiment: Smith called the sensual feeling of lust
the root of morality (Cassirer, 1918: 250). From the rat experiments of Olds and
Routtenberg we have learned that this sense of well-being is located in the so-
called pleasure centers of the brain. In the legal literature, Gruter. (1980: 108) was
the first to print to this as the possible motivation and reward for compliance with
norms.
Until now, sociology of law only considered part of the problem, namely social
security in the sense of freedom from risk (Rehbinder, 1977: 149). Compliance
with the norms would thus result in diminishing fear, for conscience, according to
Freud, means social fear. However, if we consider the decrease of social fear as
the only motivation for compliance, then we focus too much on the negative
sanctions of law, i.e. the various disadvantages, e.g. lower social prestige as a
consequence of deviance. But there are positive sanctions as well-tax advantages,
subsidies-which motivate compliance and provide expectations of rewards. And,
finally, there are cases when people tend toward an ideal of justice. Certainly, law
must rely for its enforcement on the 'law consumers' self-interest and their fear of
punishment. However, law succeeds best by triggering the sense of justice. In that
case, complying with the law generates a sense of well-being that is independent
of the feelings of security and self- interest.
Sense o f justice as criterion of Socialization ?
Compliance with the law is thus motivated not only by social fear, but also by
moral conviction. A stance such as Martin Luther's —'Here I stand, I cannot help
myself—shows a standpoint resulting from the conquest of social fear. Freud
and, following him, Hans J. Eysenck thus defined the concept of conscience too
narrowly. According to David Riesman's famous distinction between internally
and externally controlled people (Riesman, Denney & Glazer, 1953), people are
controlled by external motivations. Their sense of Justice IS thus dictated by
social fear. However, there are also people who are
directed by motivations that are not controlled heteronomously, but, rather

autonomously. This group complies with the law not because of social
conditions and social fear, but rather because of their own insights
Jean Piaget (1932[1965: 395]) also makes a distinction between

heteronomous rules which exercise authoritative control, and autonomous
rules, which are based on the acceptance of the reciprocity of social relations
in the sense of indispensible cooperation. He sees the development of a child
as a shifting from authoritatively controlled behavior to autonomous
behavior. On this basis, Lawrence Kohlberg (Lickona, 1976: 31-53)
distinguishes six steps in the development of moral concepts. These steps
are stages in a person's maturing process and indicate to which point his
socialization has succeeded (Lickona, 1976: 219-40). To be sure, the various
stages of development are reached by cognitive, not emotional abilities.
however, a rational attitude can be internalized to such an extent that it
becomes an emotional compass. Sociologist Robert K. Merton's model
(1957) of anomy also demonstrates that a deficient sense of justice is due to
insufficient socialization. To comply with a norm without accepting its
intent is anomic behavior. Merton calls this behavior ritualism. It might
better be called pseudo-conformity or anomic conformity. This behavior
characterizes an authoritarian personality and leads to social tensions
because it leaves room for alternative value judgements which can cause
group conflicts (Rehbinder, 1977: 162).
The sense of justice as a biological mechanism .
We can prove empirically that the sense of justice enables us to embrace and live by
certain concepts of justice. However, we do not know anything about its
determinants—that is, what determines its contents. Gruter (1980: 97) calls legal
ideals the results of an interaction between society's concepts of justice and the
individual's sense of justice. Since society's concepts of justice can only influence a
person's emotions if they are internalized due to fear of sanctions, self-interest, or
love for a certain ideal, we must imagine this interaction as follows: pre-existing and
new emotional reactions are balanced on a case-by-case basis. This is possible
because man, being a deficient being (Gehlen, 1962), can make up for missing or
inappropriate genetic dispositions by learning. His memory retains what he has
learned (Dawkins, 1976); and the newly acquired knowledge influences his
genetically directed behavior and can, if it is adaptive, become part of his genetic
material. The sense of justice should thus
be seen not statically only as the result of an interaction between the individuals'
expectations and the demands of society, but rather dynamically as biological
iterative process. In this process, a primary evaluation {Geiger, 1970: 318) passes
through Innate and acquired normative filters until a feeling of satisfaction {internal
reward) sets in, which then conditions the primary evaluation of the future.
For theoretical as well as practical reasons, it would be most important for the jurist
to find out how the psychosomatic interactions between hereditary dispositions and
individually acquired experience take place, to what extent this affects the
individual's primary evaluation and how much room there is for individual decision
making. Even if the biological theories are presently of only heuristic value, it. will
be important for legal scholars to formulate them in more detail so ( that legal
scholars could address relevant questions to the biological sciences.
PART II
The search for the missing pieces:
in biology
MARGARET GRUTER AND PAUL BOHANNAN
Repeating what we said in the introduction, we want to underscore at the beginning

of this section one important point that we all agreed upon. It is stated forthrightly in
the paper by Markl below: we are not limiting our inquiries to the relevance of
sociobiology for law—indeed, we aim to go far beyond sociobiology into the rest of
biology and the : behavioral sciences when we consider the relationship between law
and human behavior, especially when we analyze the behavioral s responses to law.
Biology encompasses many specialized and diverse scientific approaches to the still
unsolved mystery of life. Genetics is just one branch of biology in which a lot of
new territory was explored during the last 30 years; another branch is ethology;
apart of ethology again is primatology.
To learn about biologically based behavioral traits in human beings, we have to
work with all the different branches of biology involved in today's pioneering
research. In some areas biologists have come up with data which the most
prominent ones among them hold for truth. " We can build our own hypotheses on
these findings and test their validity.
Behavior can be influenced and sometimes determined by phylogenetic factors or
by experiences during ontogeny. In most cases both factors shape the behavior of
any living organism. The broader the base on which we build our studies the more
likely we are to arrive at a 7 scientifically sound appraisal of the behavioral
functions essential for individual and group survival.
Biological research, of course, has been the foundation of modern medicine which
uses the findings of biologists to treat the individual.
[47]
Animal experiments as a basis for the study of causes and possible treatments of
individual health problems have been universally accepted. A major attempt to apply
findings gained in observing animals in their interaction with con-specifics (in this
case their mothers and peers) was started in the 1950s by Harlow, whose well-
known deprivation experiments with monkeys led to insights in mother-child
relationships. Ethologists, physiologists, neurologists and other scientists agree that
by studying a number of different non-human primates, general principles have
emerged, in light of which we can examine human child-mother relationships. It
means that with only one or a few more steps in the same direction, we can look at "
other relationships (including those among more than two individuals) and
investigate sensory inputs and reactions in the brain as they relate to behavior like
aggression or depression and their effect on legal behavior. In doing this, we can
arrive at behavioral responses to rules. It will illuminate moral concepts and
motivation, the function of specific laws, the effectiveness of law in general. We
will learn about the role of law in the phylogenetic and ontogenetic development of
the individual as well as in human social organization.
The major areas to be considered here are the closely linked ones of behavior and
brain. We start w1th the primate record. Goodall— whom we sorely missed at the
conference, but she was in the field in Gombe—emphasizes the distinction between
order among the chimpanzees at Gombe (sustained partly by the patterns of
dominance interactions) and law as humans intentionally practice it. Goodall then
analyzes field data and gives us examples of social interaction among Gombe
chimpanzees. She notes importantly that, although chimpanzees sometimes interfere
in the disputes of others, they have nothing resembling sanctions.
Among chimpanzees certain behavior sequences seem to serve very important
functions in maintaining social order: not only the appropriate submissive patterns,
so necessary for acceptance in society, but also the reassurance gesture and the
seemingly strong need to re-establish "normal relationships." This need seems to
prevail in the individual even if terrible injuries are inflicted or a baby, vehemently
defended by his mother, is k1lled by a group member. The reassurance gesture—the
outstretched hand, the touch that appeases both parties involved in conflict (in
human terms, forgiveness) is one of the moving episodes conveyed on film taken in
the field in Gombe. Boehm's comments below emphasize that we have ceased to
value or even to make sufficient use of submissive gestures in the human behavioral
repertoire. It would seem that when dominance is no longer complemented by

submission and reassurance, the evolution of traits for morality may be favored in
a species needing complex modes of social organization for its survival.
Itani's paper summarizes what is currently known about intra-specific killing
among non-human primates; its importance stems not only from his effective
summary of the data, but from its place in the context of on-going debate about
murder in human origin myths (Girard, 1980). Itani emphasizes the distinction
between analogy (in which similar functions result from different mechanisms) and
homology (in which functional similarity results from similar mechanisms). On the
whole, we will adopt the view held by many prominent ethologists that moral traits
or precursors of morality in non-human primates are analogs of some types of
human behavior. With the papers by Goodall and Itani, even the myth that human
beings are the only creatures to murder their own kind has been proved wrong-our
vanity makes us not only the best animal, but also the most evil one. Some human
beings appear to have a deep need to underscore again and again the farthest
reaches of human depravity, just as others (or are they the same ones?) have the
need constantly to reassert human superiority over the "beasts."
With MacLean's paper, we turn to the important matter of the brain-that least
explored human organ: the way it has developed during evolution, and the way it
works as one of the necessary, foundations of human social control and law. No
one doubts that the institutionalization of law is primarily a product of the
neocortex, but surely we can also no longer doubt that all the rest of the brain is
involved, and that some structures of the brain are far older but still appear to playa
vital role in our total behavior, including our disputing behavior and dispute-
settling behavior.
Richard Alexander takes an evolutionary biologist's look at the “central paradoxes
of moral philosophy" and warns us that "approaches that deny biology are
potentially deadly." His essay should be closely compared to that of Masters in the
next section. What both have to say about biology, law and philosophy seems to us
to be putting the philosophical basis of law into a genuinely new key—a key that
includes consideration of the human biological heritage.
Bartley Hoebel's piece is rooted firmly in biology but begins the processes of bridge-
building toward law. He asks whether there are neural and chemical bases of
rewards that make human beings feel good about being law-abiding and living in the
kind of predictable
society that an effective legal system assures. In order to do that, obviously, he has
to make some assumptions that can be put as questions to behavioral science, an
essential step if we are concerned about what Hoebel calls "a jump from genetics to
motivation." Hoebel's rather daring position is that "accomplishment of law goals"
may be {there is, of course, as yet no proof) rewarded by endogenous opiates, just as
sex, drinking when thirsty and eating when hungry are rewarded by such opiates (for
which there is proof). "The 'gravitational force' that gives anthropological outcome
an appearance of purposefulness could be a genetically programmed brain circuitry
that releases rewarding chemicals when law rules are followed—which is to say,
when the perception matches the standard. Why else do we feel "good" when we
behave "well"? As Hoebel says, " According to this theory, when we pick a rule, any
rule, and obey it, perhaps the brain releases some opiate. That much could be
innately programmed. The rules themselves could either be innate (like swallowing)
or learned {washing food). He suggests that there is one master "law": "Thou shalt
match behavior to behavior model, and if you fail, adapt either the behavior or the
model. It is what Powers was saying in 1973 when he entitled his book Behavior:
the Control of Perception. Hoebel adds: "Model matching is the law of laws." In
these circumstances, we must discover "the evolutionary, neural and social
principIes by which people redefine their relationships as the conditions of life
change.
[50]
Order without law

JANE GOODALL
P.O. Box 727, Dar es Salaam, Tanzania
Codes that regulate behavior of non-human animals depend on more

learning as the brain of the animal becomes more complex. Chimpanzee
brains are more similar to human brains than those of any other known
animals. Chimpanzees maintain social order by postures, gestures and calls;
among the most important of which is a sequence of aggression, submission
and reassurance. At puberty young males find a place in the dominance
hierarchy by relying on strength and intelligence, specifically with charging
displays and challenges directed first to females, then to the lower ranking
males, then rising through the ranks. Some are
seriously wounded. There are three types of "possession": territories;
females and young; and things gathered, found or made. The “owner” must
be prepared to defend all. Examples of killing and cannibalism of infants are
given; males responded to the mothers calls for help, but no sanctions
against the killers were observed. "
All human groups have certain traditional codes of behavior, deviance from which
may result in punishment. These codes are passed down from generation to
generation, and it is on such traditional behavior patterns that many of the rules, or
laws, of that society are based.
Non-human animals also have codes that regulate their behavior within their social
groups. In birds and lower mammals, these are usually very rigid and highly
predictable, often conforming to the stimulus-release models of classical ethology-
those governing court- J' ship, competition for mates or territory, and the raising of
young. As we progress up the evolutionary ladder of increasing brain complexity, we
find that learning, within the individual life cycle, plays an increasingly important
role in the acquisition of adult behavior.
When behavior is, to a large extent, "instinctive" there is no need to impose rules or
"laws" from without—they are built into the animal c' from its birth and it relies on
them for its survival. But as the individual becomes ever more significant in shaping
his society, the more may arise the need to regulate his activities.
Chimpanzees are man's closest living relatives. This has been shown
biochemically—such as in the structure of the blood proteins and the number and
form of chromosomes. In particular, the structure and circuitry of the chimpanzee
brain is more similar to the human brain than is that of any other creature.
Chimpanzees live 40 to 50 / years and the period of childhood and development is
prolonged-an ~ individual is not fully socially mature until 14 or 16 years of age.
Thus, it may be of interest to examine the way in which order (without law) / is
maintained in chimpanzee society, searching for elements which might be useful in
exploring the biological basis of human legal behavior.
GENERAL DESCRIPTION OF THE SOCIAL ORDER

Chimpanzees live in communities that vary in size. There are between 30 and 60
individuals, including infants, at Gombe. Chimpanzees comprising a community
recognize one a»other and may travel in peaceful association, although they do not
move about as a socially
cohesive unit. Rather, individuals form and re-form in constantly changing

temporary associations. The only unit that is stable over long periods of time is a
mother and her dependent (up to eight years) young. Bonds between mothers and
their grown offspring and between siblings are strong and may persist throughout
life. Pairs of non-related adult males may also form very strong supportive
relationships lasting for years.
Adult males are dominant over females and youngsters. The males themselves
are ordered into a hierarchy so that one male usually emerges as the top ranking
or alpha male. Of course, the hierarchy is seldom static for long periods of time
since some individuals are becoming older and weaker while youngsters are
becoming stronger and more aggressive. However, provided the alpha male is in
a strong position, social order within the community is maintained with relatively
little aggression. Most of the time each individual knows his (or her) status and
disputes can be settled by means of mere threatening gestures without recourse to
physical fights.
Females have a much less clear-cut hierarchy. There is almost always a top-
ranking female, and there are a number of very obviously highly-ranked females,
as well as a number who rank very low. Females spend much less time in each
other's company than do males moreover their status, relative to each other, is
always changing depending on their reproductive state and the presence or
absence of grown offspring-who will support them.
MAINTENANCE OF SOCIAL ORDER

The maintenance of the social order is dependent on a rich repertoire of postures and
gestures and calls. One most important sequence is "aggression, submission and
reassurance." After an aggressive incident the victim may approach the aggressor
and make appeasing gestures, such as holding out the hand or crouching low on the
ground, while whimpering or screaming. The aggressor usually reaches out to touch,
pat, kiss or embrace the supplicant. This serves to calm, or reassure, the latter. It is
this kind of behavior which makes it possible for the chimpanzees to enjoy relaxed
and friendly relations, even in a society where violence may erupt suddenly. This
sequence, also indicates the importance, in chimpanzee social life of friendly
physical contact. Threatening gestures, rather than physical fights, serve to regulate
most disputes between chimpanzees. The most significant and the most impressive
is the male "charging display"
when he hurtles along, throwing branches or rocks, swaying vegetation, making

himself look more dangerous than he may actually be. It is a supreme example of
bluff.
In chimpanzee society, almost always, "might is right" and it is the lower ranking of
the pair, whatever the context, who must make appeasing gestures. The participants
are then able to resume a relaxed relationship, at least to outward appearance. In
fact, the subordinate may not be feeling as unruffled as it seems.
High ranking Fifi and low ranking Gilka squabble over food. Fifi attacks Gllka
who screams, withdraws, then holds out her hand, which Fifi touches. Both
resume feeding. Suddenly Gilka screams and flies at Fifi, hitting her. This
coincides with the arrival of Gilka's eldest brother who stands, hair bristling.
Now it is Fifi who retreats, screaming.
This kind of incident, where the stronger of a pair of relatives or friends protects the
weaker is, common. But do strong chimpanzees ever protect the weak irrespective
of social relationships?
Adult males are, in general, protective of all infants (and may discipline them too).
Infant Freud pesters a male baboon, kicking at him playfully. The baboon
loses patience, pulls Freud from the branch and bites him. As Freud
screams a young male chimpanzee, Goblin, charges over and hits the
aggressor. Freud soon resumes his pestering of the baboon. Almost at once,
Goblin seizes Freud, slaps him hard and leaves. Freud, subdued, goes to his
mother!
Since there are no permanent pair bonds between non-related adult males and
females, any male can, theoretically, impregnate any female, so that the general
protective attitude of adult males to infants can be likened to a father-child
relationship. This kind of relationship may not be extended towards infants of other
communities. Three times, groups of adult males severely attacked females from
neighboring social groups (encountered near the territorial boundary), seized, killed
and partially ate their infants.
Sometimes, even within the community, the protective attitude of a male towards an
infant breaks down. If an infant gets in the way of a .charging display, it may be
used as a display object, picked up and flailed, dragged or thrown.
: Young adolescent females may transfer, temporarily or permanently,
into a neighboring community. Initially these young females run the risk of
persecution by resident females and usually keep close to the big males for
protection.
A young male, Figan, is traveling with a transfer female, Sparrow. Figan's
mother and sister joined them and began to threaten Sparrow, chasing her
as she fled, screaming. Figan, unwilling to attack members of his own
family, displayed vigorously between the squabbling females and stopped
the aggression.
Sometimes a high ranking male will break up a fight between a lower ranking male
and a female. This, however, is not necessarily directed toward "rescuing" the
victim: sometimes his display is aimed at re-asserting his rank relative to that of the
aggressor. And there are even times when a male may rush over and join in such an
attack against the female victim. As many as four males may join in an attack in this
way. All this suggests that the sense of chivalry is not very far advanced in our
chimpanzee relatives.
THE DEVELOPMENT OF INTER-INDIVIDUAL

RELATIONSHIPS WITHIN SOCIETY
The infant chimpanzee is born with a repertoire of inherited species- specific
behavior patterns. But experiments conducted in captivity, in which infants were
raised in social isolation for the first months of life, suggest that they must learn, in
social context, when and how to use these communication signals.
In the wild the behavior of the infant is initially molded by the mother. She prevents
him from doing harmful things and may physically remove him from the potential
trouble or start to play or groom him, temporarily distracting his attention. As the
infant grows older, inappropriate behavior is corrected by his mother by more active
punishment. For example, she may gently bite him. If he then throws a tantrum she
may punish him more vigorously-but she will reassure him, usually be embracing, at
the same time.
From the moment that he begins to leave the protection of his mother's body, the
youngster starts to learn what he can and cannot do in his society. He learns this
mainly by trial and error and by observing and imitating others. He learns quickly
that other individuals may not be as tolerant as his own mother. During play with
peers he will find that he is stronger than some, yet he must learn to hold his
punches if the youngster's mother is dominant over his own. He "' learns, too, that
while he can approach most adult males when they are relaxed, he may not be
tolerated when they are socially roused. And he finds that behavior which is
acceptable to some adults may provoke irritable responses from others.
As he grows older, the lessons are harder. Juveniles learn that they must be more
cautious during feeding and keep their distance from others—even their own
mothers—to avoid aggression. In the early days, some youngsters respond to
reprimands with tantrums, but gradually they adjust to the new state of things and
show the appropriate submissive patterns that are necessary for their acceptance in
society.
It is during adolescence that some of the hardest lessons must be learned,
particularly by the male. Adolescence is not too difficult for females. Although they
leave their mothers during periods of oestrus {and may move into neighboring
communities) they usually return. They can learn most of what they need to know to
function appropriately as adults within the family context. Females learn submissive
patterns during infancy and will continue to use them, at least towards the males, for
the rest of their lives.
The young male, however, must leave his family in order to learn about such
primarily male activities as hunting for meat, patrolling the territorial boundaries,
repelling incursions by neighbors and recruiting young females. The young male,
from late infancy onwards, becomes increasingly fascinated by the big males. His
first excursions away .from his mother are almost always in the company of one or
more of the adult males. At the same time as his fascination for the males is
increasing, so too is his fear of them. Increasingly he is liable to threats :" and attacks
from them. Moreover, he is the ideal 'scapegoat' on to whom adult males can redirect
their aggression.
Meanwhile, the adolescent male is growing strong. Around puberty, the testosterone
levels in his blood increase rapidly, and he shows a I strongly correlated increase in
aggressive behavior. Initially he directs -" this aggression towards the females of his
community. He displays at them again and again {provided the big males are not
present) until eventually he is able to dominate all of them by the time he is 13 to 15
years old. The one really stable relationship of the adolescent male is that with his
mother. He does not try to dominate her and he continues to show respect for her.
Again and again, after spending time with the big males, particularly after times of
social tension, the adolescent male leaves the group and seeks out his mother and
family.
Finally, when the young male is between 14 and 15 years old, he begins to try to
establish a place for himself in the hierarchy of the socially mature males. How does
he manage this? In most human societies there are some sort of initiation rites to
establish the fact that a young man has 'come of age' and has the right to enter the
world of men. The chimpanzee male, however, is not automatically treated as an
adult just because he has reached full body weight and developed the strong canine
teeth of the mature male. He must rely entirely on his own strength or intelligence to
make the transition to adulthood though an elder brother may support him during
this difficult time. Above all he makes use of the charging display when challenging
the lowest ranking of the mature males. The more frequent and the more impressive
the charging display of the young male, the quicker he is likely to rise through the
ranks of his elders. It is probably the occasional avoidance response of an older male
to the tempestuous arrival of a late adolescent male performing such a display
(during which inhibitions are, to some extent, lost) that gives him the confidence to
actually direct his display towards his erstwhile heroes. The struggle is not easy,
and young males may become very badly wounded in their attempts to better their
social position. As a result they may drop right back and remain in low ranking
positions for several more years.
The transition from adolescence to maturity is easier for the female than for the male.
When she is about 10 years old she develops an adult sexual swelling and is accorded
mature reproductive status by the big males—who mate her for the first time. One or
two years later she gives birth to an infant. Both these events occur without the
female having to do anything about it and both, in themselves, are signals to other
chimpanzees of her new mature status.
INCEST TABOO
At Gombe, no physically mature male has been seen to mate his mother. Mating
between siblings, while it does occur, is at a very low frequency, and three out of
four young females tried vigorously to avoid the courtship displays of their elder
brothers. As mentioned, young females have a tendency to mingle with neighboring
males during periods of oestrus so even if they subsequently return to their home
communities their first infants are likely to be sired elsewhere. Young females also
show a tendency to avoid the courtship of old males and this provides a mechanism
whereby they may avoid incestuous relationships with their fathers.
THE CONCEPT OF 'POSSESSION'

In general, possessions are of three major types.
(1) The oldest kind of possessions, in the evolutionary sense, are undoubtedly those
involving the occupation of certain areas of the animal’s environment-territories.
Some species have individual territories which they protect from invasion .even by a
member of the opposite sex except during the breeding season; others maintain a
joint territory for family or larger social group.
Those species which roam over large areas of country-hunters and nomads-often
select one area within the overall range, which changes from time to time, as a
territory to be defended from conspecifics with whom they may share the rest of the
range.
(2) One individual may show possessive behavior towards another individual or
individuals of the same species. A male may fight and defeat a rival male in order to
take possession of a female. Both males and females may show possessive behavior
towards their offspring while these are still dependent. This kind of possession is, of
course, widespread in the animal kingdom.
(3) Finally we come to possessions that have been found, gathered, or made. The
winter food store of a mouse, a nest which has been constructed for sleeping or
breeding, an object used as a r tool, clothes, a herd of cattle, or a dog.
In all three categories, possession cannot be inferred unless the 'owner' is prepared to
defend what is his or at least shows fear or distress if he is dispossessed by a stronger
rival. There may be occasions I when he shares his possessions with others, but we
must have evidence that, at least on some occasions, he maintains or tries to maintain
exclusive possession. There will, of course, be joint-ownership—a " pride of lions
driving competitors from their kill, mother and father fighting together for the lives
of their offspring.
Let us briefly discuss these three kinds of possession in chimpanzee society. (I)
Chimpanzees are territorial: adult males regularly patrol their boundaries, chasing off
or even killing strangers. The main study community at Gombe divided in 1972: over
a four-year period the J males of the larger, northern group systematically attacked
the "."' individuals of the southern splinter group. Eventually the entire
break-away group was eliminated and the victorious males, and their females and
young, once again had access to the area which they had lost after the division.
(2) Male chimpanzees do not form exclusive, lasting pair-bonds with adult females.
In some situations a female in oestrus may travel with a group of males and be
mated by all with little overt competition. At other times the highest ranking male in
such a group may show possessive behavior towards a female in oestrus and, to
some extent, can then inhibit other males from mating with 'his' female. If however,
there is a diversion, the other males in the group will quickly take the opportunity of
stealing a quick copulation. If the possessive male notices he charges towards the
mating pair and attacks one of them-usually the female since if he fights the male the
female may be mated by a third male.
The most successful reproductive strategy for a male chimpanzee is to form an
exclusive consortship with a female during which he leads her away from the center
of the community range and tries to keep her away from other males for the duration
of her oestrus period. If she tries to escape from him during the early stages of such
a consortship, he may attack her quite severely.
Mother chimpanzees are possessive towards their infants, and this is unchallenged
by other individuals. Others may look closely at a newborn but seldom try even to
touch it during the first few weeks.
Older siblings, usually the first to be allowed to touch the infant, will also show
possessive behavior towards the new family member and chase away other
youngsters who try to play. If this is not possible, the older child may join in the play
session and unobtrusively take the place of the small sibling, thus preventing the
undesired contact.
When an infant dies, the mother will continue to show possessive behavior towards
the body, usually for three or four days—until it begins to decompose. While she
may treat the corpse quite differently from alive baby—slinging it over her shoulder,
letting it drop to the ground and so on-nevertheless she does not move without it and
she will not allow others to take it away.
(3) Things which have been found, gathered or made including sleeping platforms or
nests that are constructed each night (ownership of which is very seldom disputed),
food items (including meat) which have been picked up or gathered together, objects
such as tools, and objects used as toys in play. These things are very temporary
possessions-a nest may be occupied for 12 hours or so and a portion of meat may
take five hours or more to consume, but the other things are
usually discarded after minutes rather than hours. This is hardly surprising since
the objects referred to are either food, in which case it is eaten, or sticks, leaves, or
bunches of fruit that can be found .whenever they are needed. It is significant that
the 'toy' which was carried about for the longest time was the remains of a pillow
that young Figan took from my tent one morning, and was still in possession of,
after many tussles with his peers, the following afternoon. Pillows are not part of
the typical Gombe environment!
Possession of some of the things listed may be very hotly disputed—in particular,
meat. Meat is much less easy to acquire than the more usual vegetable and fruit
food items of the chimpanzee diet and the prey killed is seldom large enough to
satisfy all those who are present during its consumption.
Immediately following a kill there is much excitement during which those present
try to seize a share. After this division individuals who have obtained a share move
off to feed. Those who have not been successful usually gather round the lucky
ones and beg. They reach out to touch the meat, requesting permission to remove
apiece or to join the possessor in feeding. Or they hold out their hands, palm up.
The success or failure of a begging individual depends on a variety of factors: the
personalities concerned; the nature of the relationship between possessor and
seeker; the amount of meat available; and the degree of satiation of the possessor .
Meat is not only acquired by begging. A male will unhesitatingly, take the prey
from a female or juvenile. Yet he may beg, patiently, from a possessor who
normally ranks below him in the hierarchy. This does not mean that he
acknowledges the other's 'right' to his meat. Rather it suggests that the usually
higher ranking male knows that the other will not easily relinquish such spoils:
meat is worth fighting for and he is likely to hang on tightly and crouch over it.
Moreover the possessor, who may usually show submissive behavior towards the
begger, is concentrated on feeding and is not, perhaps, emitting the appeasing
signals which the other has come to expect this could produce feelings of anxiety in
the begger.
It is of interest to note that a chimpanzee recognizes that an object is 'mine' or 'his',
even when the object is spatially separated from its 'owner'. This is shown very
clearly during termite feeding when the chimpanzees push grass stems into tunnels
in termite nests, then pick off the clinging insects with their lips. A feeding
individual must leave his tunnel from time to time to select new tools. At such
times a lower ranking individual may move over and push his own stem into
the temporarily freed passage, but will repeatedly glance towards the 'real' owner as
he does so, and will hastily move away when the other heads back. When a
chimpanzee is sitting eating a pile of fruits he has gathered, such as bananas, he may
discard the skins. A subordinate may reach out towards a skin very cautiously
indeed, watching the ‘owner's' face carefully. Only if he sees no sign of threat will
he actually take a skin. A toy, especially a valued toy such as a piece of cloth, may
be laid down by a youngster who is feeding. After awhile another youngster may
creep towards the cherished object, keeping a wary eye on the owner. In such
situations, the owner is usually well aware of what is going on and will instantly
race over to retrieve his or her toy, even if this is out of his sight. One mother,
Passion, was feeding on driver ants. She got bitten, dropped her long, peeled stick
and retreated to remove the ants from her person. Her adult daughter Pom glanced at
her mother, then exchanged her own tool, which was shorter, for the abandoned one.
Immediately Passion reached out and touched 'her' stick, Pom handed it over and
continued with her own.
In 1975, the adult female, Passion, watched by her adolescent daughter, Pom, seized
the newborn infant of a community female, Gilka, killed it and ate it. She shared the
flesh with her family. The following year Glika again gave birth. Passion and Pom,
working as a team, jointly attacked Gilka, seized the baby and ate it. It was Pom who
killed this victim—biting into the forehead as her mother had done the year before.
Two months later, Passion and Pom were watched as they jointly attacked another
female, Melissa. After a fierce struggle, mother and daughter managed to seize her
newborn infant, which they ate. In mid-1977 and early 1978 three more attempts to
kill infants were observed, all of which failed. Subsequently no further incidents of
this sort have been seen.
The behavior, more than any other recorded at Gombe, could be regarded (in human
terms of reference) as criminal deviance. What was the response of other members
of the community? Was any form of punishment meted out to Passion and Pom?
It is Important to realize that all the observed attacks by this mother and daughter
were made when no chimpanzees, except the victims and their families, were
present. The extent to which other members of the community knew about the
killings was not clear. One mother, Melissa, was found with a newly dead baby,
killed by a bite on its forehead. She was with a number of males and we suspect that
this baby had been one of Passion’s victims, but that the arrival of male chimpanzees
(alerted by Melissa's screams) had prevented the feast. If
so, these males knew about Passion's behavior. Both Gilka and Melissa, after losing
at least one infant each to Passion and Pom, were fearful when the killers
approached subsequent babies. When adult males were nearby they responded
quickly to the screams of the mothers. Twice they displayed towards the killers, and
once attacked Passion.
Five different mothers were involved in the observed killings or attempted killings.
Two of them (one Melissa) seemed, quite quickly, to lose their initial fear of the
killer females. One continued to show anxiety in Passion's presence for over a year
after an attempt had been made on her infant's life and another, Gilka, remained
very fearful of Passion for the remaining three years of her life.
The behaviour of the fifth mother, Miff is of particular interest. Her first observed
encounter with Passion was when her infant was a week old. Miff at once screamed
loudly and ran from Passion until she encountered two big males. Then, her
screams changing to fierce, threatening barks, Miff turned and headed back towards
Passion. The two males followed and when they arrived, both displayed at Passion
and Pom who fled. At that time, Miff, clearly, was aware of Passion's murderous
tendencies. Three weeks later Passion made a determined attempt to seize Miff's
baby. However, Porn, heavily pregnant, did not help, and Passion was hampered by
her own four-month-old infant. Miff was able to protect her baby during a very
fierce fight.
Three months after this, another encounter between Miff and Passion was observed.
It was Miff's juvenile son who saw the killers first he gave threat barks and the
whole family ran off screaming. When they encountered an adult male, Miff again
turned and led him back to Passion: he displayed vigorously and Passion and her
family retreated. Miff and her family's fear of Passion and Pom continued, unabated
for the next two years.
Thus, while it is clear that the adult males, when available, promptly respond to
appeals for help from threatened mothers, only mild forms of aggression have been
observed in response to threatening behavior by Passion and Pom. Even some of
the victims appear to quickly overlook (or forget) their grudges against the deviant
females.
SUMMARY
There is order in a chimpanzee society—an order which is maintained without 'laws'
in the human sense of the term. Because so many chimpanzee behavior patterns are
so similar to some of our own, we
may see codes of behavior similar to those which have led to laws that are enforced
in order to try to maintain 'law and order' in human society. A better understanding
of evolutionary factors that have molded human social behavior might suggest why
some human laws a are widely accepted, while others are more difficult to enforce.
This is a fruitful field for future research, but it can only be meaningfully undertaken
when a trained legal mind along with a student of chimpanzee behavior co-operate
to tackle the problem involved.
Intraspecific killing among non-human primates
ONICHIRO ITANI
Kyoto University,
Faculty of Science, Laboratory of Physical Anthropology,
Sakyo, Kyoto 606, Japan
Intraspecific killing episodes among wild non-human primates are summarized
from an evolutionary viewpoint, with regard to the social structures of the
species. The summary is restricted to the higher primates, the Old World
monkeys and anthropoids, which have well- developed brains and more
complex social structures. This phenomenon is found only in societies with
other than pair-type social units; all have one-male or multi-male groups. Eight
aspects of intraspecific killing among non-human primates are discussed:
structure of basic social units, intolerance among males, male precourtship
aggression, tendency toward incest avoidance, population density,
developmental cycle of unit groups, identity of the victim and cannabilism.
Episodes of intraspecific killing have been reported for 132 species in nine
genera. It is notable that, for all these species, infanticide has been observed.
Therefore, the episodes were classified into infanticide and killing among
adults; characteristics of each category were indicated. It is apparent that
pongids have a wider variety of killing than cercopithecoids. One of the
characteristics of intraspecific killing among non-human primates is that there
are few counteractions from a third person or from the society toward the killer.
Although an integral theory to interpret this problem is not easy to formulate
and we have to wait for future studies for detailed analysis, it
must be emphasized that these phenomena are related to the origin of institutions
and the evolution of society.
I. INTRODUCTION
Several scholars-Angust & Thommen (1977), Poirier (1974), Hrdy (1979) and
Goodall (1977)-have reviewed the topic of infanticide among non-human primates
for comparison with their own data. The present article deals not only with
infanticide, but also with other aspects of intraspecific killing; it assumes the
evolutionary viewpoint, and focuses on the phylogeny and social structure of the
species among which the events were observed.
How are we to interpret intraspecific killing among primates? Such events are fairly
common in human society. The recent accumulation of field data on non-human
primates has revealed the presence of counterparts among them as well. However
this issue does not involve the whole primate order. Although the studies on
prosimians are not conclusive, no intraspecific killing has been reported among
them in the wild. Among ceboids, there is only one report of infanticide- among
howler monkeys (Collias & Southwick, 1952). Therefore, the problem of
intraspecific killing is for all practical purposes restricted to cercopithecids and
pongids. Indeed, thus, intraspecific killing is seen primarily in the presumably
phylogenetically advanced species. This fact suggests that the phenomenon is
related to well-developed brains and complex social structures.
II. PROBLEMS CONCERNING INTRASPECIFIC KILLING
(1) Basic social unit
All species of anthropoidea, with the sole exception of orangutans, have bisexual
social groups. For the orangutan, no intraspecific killing has been reported.
Cercopithecids have matrilineal social units, except for one species mentioned
below, while pongid societies are not matrilineal (Itani, 1977a,b, 1980a,b). Males
among the former and both sexes or at least females in the latter transfer between
unit groups. All species of gibbons have unit groups of male-female pairs, and there
is no report of intraspecific killing. Recently, mentawai langurs (Presbytis
potenziani) have been found to be the only cercopithecid species having pair-type
units (Watanabe, 1981), and no intraspecific killing has been observed for this
species. Therefore, species with pair-type societies can be excluded from the
present discussion.
'The societies of cercopithecids and pongids with other than pair- type social units'
are thus the subjects of this essay. they all have one-male groups of 10 to 100
animals. It should be noted that many social exchanges, including intraspecific
killing, occur between social units or between a unit group and non-group
individuals.
(2) Intolerance among males

Conflicts between males are usually avoided by two means: one is to maintain
interindividual distances and thus to refrain from direct contacts. The other is the
dominance order among males. Conflicts are avoided by inhibition behavior of
subordinate individuals toward dominant individuals (Kummer, 1971), and by
interindividual recognition of dominance. However, these avoidance mechanisms
are not perfect. Examples of conflicts due to their breakdown are numerous.
Intolerance between males is often related to domination and possesion of females,
Solitary orangutan males maintain larger distances between one another
(MacKinnon, 1974), and in a sense, they may be considered vagabonds who have
abandoned possession of females.
(3) Male precourtship aggresstion

Accounts of aggression by males is commonly observed in multi-male groups, such
as among macaques (Itani, 1954; Southwick, 1965). This behavior is thought to
promote the courting relationship between a particular male and a particular female.
The aggression toward the female may intensify her sexual arousal. However, cases
in which such precourtship aggression results in infanticide are widely seen among
cercopithecids and the gorilla. Although it is unknown whether the killings are
intentional, it appears that the males may know that the loss of the infant is
followed by physiological changes in the female who had been nursing her infant,
causing her return to sexual arousal.
(4) Tendency toward incest avoidance

Higher primate societies universally tend to avoid incest. Although analytical data
on this problem are inadequate for most species, the on-going study by Takahata
has clearly revealed this tendency for the Japanese monkey. For 54 dyadic
relationships of the first and second degree of consanguinity in a troop of Japanese
monkeys, no copulation at all was observed. The semi-closed structure of unit
groups allowing selective individual transfers, can also be interpreted in terms of
incest avoidance. The transfer of either males or females from their natal
.
unit-group to another prevents the occurrence of incest within the unit-group. If each
society emphasizes incest avoidance, a third safety valve may be hypothesized: the
elimination of individuals born through incest. The one-male group structure has an
inherent possibility of father-daughter incest. Not all, but some of the infanticide
commonly seen among societies with one-male unit groups undeniably eliminates
individuals born of incestuous unions. However, whether such a context is
recognized by them, or what might precipitate such behavior, is unknown.
(5) Population density
Yoshiba ( 1968), studying Hanuman langurs (Presbytis entellus), eco- logically and
sociologically compared a population of basically one- male groups in south-west
India with a population of basically multi- male groups (Jay, 1965) in north India.
The primary factor affecting the difference in grouping patterns was suggested to be
populatipn densities of 87-136 animals/km2, in the south-west in contrast to 2.7
animals/km2 in the north. Infanticide was observed only in the south-west.
Rudran (1973), who studied two subspecies of Presbytis senex in Sri Lanka,
reported that the P. senex senex population, with a density of 92.6 animals/ km2,
had relatively infrequent male replacements and low infant mortality. It is
undeniable that high population density complicates social relationships. It seems
likely that the high population density gives rise to the male-intolerant one-male
group structure, in which infanticide is inevitable.
(6) Developmental cycle of unit groups

In some species, the developmental cycle of unit groups is recognizable by the size
and composition of a unit group (Itani, 1980b ). For the one-male groups of
Hanuman langurs, the interval between male replacements, through the initial stage,
growth stage, and mature stage, is estimated to be about three years (Hrdy, 1974 ).
Sugiyama ( 1965) estimated the cycle of the group rejuvenation to be 4.5-5 years.
For these species, the size and composition of a group indicates the stage of its
development, and signals to out-group males whether the group is ready for male
replacement. For out-group males, the mature stage must trigger male replacement,
which involves infanticide.
(7) Identity of the 'Victim
Of course, the identity of the victim of a killing differs from one episode to another.
In the context of intolerance among males, males are victims, while in the context of
male precourtship aggression, females and their infants become victims. When sex
and age have ethological significance, then such stimuli as the silverback of male
gorillas, swollen sex skins of female chimpanzees and macaques, the Gestalt of
mother with an infant, or the neonatal pelage, can be releases that trigger killing. Not
only these physical traits, but also the group membership or personal history of each
individual may be distinguished and become the trigger. The mere presence may be
distinguished and become the trigger. The presence or absence of acquaintance or
personal recognition may differentiate the unfolding of the episodes. A particular
individual, for some idiosyncratic reason, may become the target. In addition,
characteristic attitudes, facial expressions, behavior and vocal sounds must be
involved. The study of the personality and identity of the victim is an important
theme in analyzing such data.
(8) Cannabilism
The episodes of intraspecific killing in primates include some cases of cannibalism. It

is quite problematical whether the significance of such an episode lies in the killing
or in the eating of the victim. In the case reported by Suzuki ( 1971 ), in which the
victim (an infant chimpanzee ) was still alive while one of its legs had been eaten,
cannibalism appears primary. However, it is not easy to determine which is the
intent-to kill or to eat. Here, I take the viewpoint that killing is the primary, and
eating the secondary, motivation. Although there are not yet enough data for this
proposition, the occurrence of cannibalism closely coincides with the dietary habits
of the species. The food habits of higher primates can be grouped into the generalized
diet, which is unspecialized or omnivorous, and the specialized diet, which is based
on some particular food-a species may, for example, be folivorous (Itani, 1979).
Cannibalism has been reported so far for the red-tailed monkey (Cercopithecus
ascanius) Japanese monkey (Macaca fuscata), chamba baboon (Papio ursinus);alld
chimpanzee ((Pan troglodytes). All of them have generalized dietary habits. Among
the red-tailed monkeys, cannibalism occurred after an infant was killed following a
male replacement in a one-male group. More than ten episodes of similar infanticide
following male replacements have been reported for three species of Presbytis, but in
none of these was cannibalism
observed. It should be noted here that the Presbytis species have specialized food
habits. Although cannibalism may be regarded as an extreme expression of
generalized food habits, the possibility that the appetite precedes the incident should
also be considered. In addition, chimpanzees and gorillas, for which unquestionably
deviant cases have been reported (Goodall, 1977; Goodall etal., 1979; Fossey, 1981)
require us to reserve the category of individual pathology.
III. TYPES OF INTRASPECIFIC KILLING

Up until now, intraspecific killing has been reported for 13 species in nine genera of
non-human primates in the wild. It is notable that, for all of these species, infanticide
has been observed. Therefore, it seems reasonable to classify the episodes into those
involving and those not involving infanticide.
(1) Infanticide
The approximately ten cases in the two species of Presbytis and the two cases in red-
tailed monkeys all occurred after male replacement in one-male groups, and share
the following common context. The male of a one-male group was replaced by an
out-group male, who then fatally injured the infants nursed by the group females.
The females came into oestrus, copulated with the new male, and bore new infants
in about a year (Sugiyama, 1965; Mohnot, 1971; Rudran, 1973; Hrdy, 1974;
Parthatharathy & Rahman, 1974; Struhsaker, 1977). In P. cnstata, after a change of
dominance order in a multi-male group, the new leader expelled the other males to
form a one-male group, in the process of which infanticide was observed
(Brotoisworo, 1982). The two cases in red-tailed monkeys were accompanied by
cannibalism (Struhsaker, 1977). The maintenance of the matrilineal one-male group
seems to require male replacements, which are accompanied by infanticide.
This type of infanticide is apparently related to the type of social unit. In this type of
social unit, male replacement is an outcome of the intolerance between males, and
precourtship aggression results in infanticide. This series of social changes
eliminates individuals born by incest in consequence, and also is related to the local
population density and to the development of unit groups.
Colobus guereza (Oates, 1978) and Theropithecus gelada (Nieuergelt, cited in
Angust & Thommen, 1977) also seem to exhibit this type of infanticide. It has also
been reported in Macaca sylvana (Burton, 1972).
The one-male unit of Papio hamadryas is an infrastructure of the basic social unit of
this species. Once a one-male unit is formed, no male replacement usually occurs for
that unit. But in an experiment Kummer ( 1968) removed the male of a one-male
unit, and observed that the out-unit male took over the unit and infanticide followed.
Similar phenomena have been reported for many captive colonies (Angust &
Thommen, 1977). It seems probable that such changes also occur after the death of a
unit male in the wild.
For Japanese monkeys, which havemulti-male groups, infanticide has been reported
from two troops. In one case, a lone male approached Shiga (A troop ), and from
June to November, attacked nine out of the 11 infants born that year. Two infants
were killed and three were 1 injured (Tokida, 1976 ). This series of attacks is
peculiar in that it 1 occurred in the non-mating season. This lone male joined the
troop in December of the same year, and did not commit any more infanticide until
he moved to C troop five years later. The second case was observed in Hakone. A
mother and an infant not with their troop were attacked by a lone male, who grabbed
the infant and ate it (Editorial Board of The Nihonzaru, 1974).
The following can be deduced from these two cases. These episodes were not intra-
group phenomena, they cannot be interpreted in terms of male intolerance, and they
are probably related to precourtship aggression, although they occurred in the non-
mating season. These cases were recorded for a particular species that, compared
with other species, has been observed for long periods. Although it is difficult to
generalize from these two cases, they indicate that societies with multi- male groups
can also have episodes of killing similar to those found in societies with one-male
groups.
Fossey (1979, 1981) reported for the gorilla that six of 38 infants with births
confirmed during 13 years of observation became victims of infanticide. These also
occurred either during male replacement or antagonistic encounters of two groups,
or as a stage preceding the kidnapping of a female by an outsider male. Fossey
stated that it is the means of transmitting their genes, and that infanticide is
necessary for maintaining a healthy degree of outbreeding. Although there is a
fundamental difference in social structure between the matrilineal one-male groups
of cercopithecids and the non-matrilineal social unit of gorilla, it is notable that both
have one-male group composition and both are accompanied by infanticide.
For chimpanzees, records of infanticide have increased to 12 since the first incident
was observed in the Budongo Forest by Suzuki in
1971. Seven cases come from Gombe (Bygott, 1972; Goodall, 1977; Goodall et al.,
1979) and four cases from Mahale (Nishida, Uehara & Nyundo, 1979; Norikoshi,
1978; Norikoshi & Kitahara, 1979; Nishida, 1980; Kawanaka, 1981). Nine of the
cases were accompanied by cannibalism.
Kawanaka ( 1981) reported in detail on the case of infanticide he observed, and

thoroughly traced the personal life history of the victim's mother, an important
method of analyzing such data. He also compared the 12 cannibalistic episodes. All
the victims were within the infant stage, between newborn and three years of age
(Nishida, 1968); all were entirely dependent on their mothers. Among them, the
four cases reported from Gombe in which a female harmed infants ( category-b in
Goodall, 1977) were restricted to newborn infants. The victim's sex was male in
seven cases, female in three cases and unknown in two cases. If the four category-b
cases are excluded, the victims were five males, one female and two unknown,
suggesting that male infants were being selectively killed. This may be one reason
for the chimpanzee's biased sex ratio (fewer males).
The contents of the episodes are diverse and difficult to categorize. However, except
for the four category-b cases at Gombe, which can be assumed to be intra-unit-group
phenomena, it seems undeniable that all the cases were related to the complicated
movements of females between unit-groups. In the other three cases at Gombe
(category-a in Goodall, 1977), the infants of stranger females were killed, after
which these females became members of the captors' unit group. Each of the four
cases at Mahale occurred in its own complicated context. Kawanaka pointed out that
since in some cases there were time lags of two to five years between a female's
immigration and the infanticide, they cannot be considered 'inter-unit-group
phenomena' as Nishida et al., (1979) had concluded.
The category-b cases differ from the episodes found among gorillas or
cercopithecids, and it should be noted that they all involved only one particular
female named Passion and her children. An infant disappeared from a group of
gorillas, and bone splinters were found in the feces of a female belonging to the
same group, suggesting cannibalism (Fossey, 1981 ). This indirect evidence of
cannibalism and Passion's cases were probably of similar contexts, and they might
have been infanticide for the purpose of cannibalism.
(2) Killing among adults
As a rule, interactions that may lead to killing among males are avoided by them.
Even among macaques, which have multi-male groups, intra-group males may have
severe fights and injuries but such behavior seldom results in death. However,
between troop males and solitary males, aggression may be more fierce, with the
possibility of killing (Itani, 1954). Aggression between one-male groups and all-
male groups of Presbytis is reported to be very fierce. Many researchers have
reported that males are sometimes severely injured. Rudran (1973) inferred that a
young adult male's death was due to an antagonistic interaction between a lone-male
group and a predominantly male group. Such interaction can be regarded as
revolving about the male position in a one-male group.
Among chimpanzees, the interolerance between males of different unit groups

sometimes results in killing. The case reported by Goodall et al., (1979), in which
three males of the Kahama community were killed one after another by the males of
the Kasakela community, indicates that males within a unit group have a strong
bond among themselves, and they take a rather ruthless attitude toward males of
other groups with whom they do not share the bond. In the chimpanzee population at
Mahale, six adult males and one young adult male have disappeared from M-group
(Nishida, 1980; Itani, 1980b ). When the balance of dominance gets out of order,
aggressive inter-group encounters increase. In this context, the cases of Gombe and
Mahale are in agreement. Since 1979, only one old male remains in K-group, but the
group has not yet dissolved.
These aggressive interactions are different from those involving the position of the
male in a one-male group, as in Presbytis. Perhaps these killings are inevitable,
rooted in their social structure, and are the phenomena most difficult to interpret. If a
chimpanzee unit group is maintained by the exchange of nulliparous females with
neighboring groups, killing might result in the destruction of a neighboring group
that is a source of new females. Or do they attack to eliminate a weakened group that
is not a good source of females? Goodall et aL, ( 1979) reported that the males of the
Kasakela community often patrolled the boundary of the home range of the Kahama
community. Similar behavior has been observed at Mahale. Although the en-
counter of M- and K-groups that Nishida and I observed did not result in any killing,
it had the atmosphere of a skirmish in a war. This behavior was even strategic: they
looked as if they were aiming for the best chance of encountering another group
(Itani, 1977 b ). There has
Gruter and Bohannan [7 I]
been no direct observation of such killing among male gorillas, but they must have
similar antagonistic interactions that may lead to killing (Baumgartel, 1976).
However, antagonistic interaction of a group versus an individual, or a group versus
another group, with the intent to kill, is peculiar to chimpanzee society.
An adult male killing an adult female has been reported twice for chimpanzees at
Gombe and once for gorillas. Such an episode is rare. In precourtship aggression,
infants not females often become the victims. In one of the chimpanzee cases, the
males of Kahama community were found inspecting the body of a very old female;
it was inferred that she was a victim of chimpanzee aggression. (Wrangham cited in
Goodall et aL , 1979). This female was probably a stranger, and her death cannot be
interpreted in the context of an intra-group episode. Moreover, since the victim was
very old, there is little possibility of precourtship aggression. In the other case, the
males of the Kasakela community attacked a female of the Kahama community who
died four days later (Goodall et aL, 1979). This was also an inter-unit-group
episode. The gorilla case was an intra-group episode, the background of which was
subtle balance between an aged silverbacked male and his mature son. The son
killed an old female who had been one of his father's partners for a long time
(Fossey, 1981 ).
None of the cases in which adults were the victims was accompanied by
cannibalism. Cannibalism among non-human primates is restricted to infant-
eating.
IV. DISCUSSION
Through the foregoing review of intraspecific killing among primates in various
contexts, it has become apparent that an integral theory to interpret this problem is
not easy to formulate. Many researchers have attempted to explain this issue by
sexual selection or egoistic behavioral strategies to maximize individual
reproductive success (Hrdy, 1974; Angust & Thommen, 1977; Struhsaker 1977;
Hrdy, 1979; Nishida 1979,1980; Fossey 1981). However, these theories encounter
difficulties in explaining the male killing of a female or intra-group killing (Goodall,
1977). In the infanticide following a male replacement, as in Presbytis, the male
who has succeeded in taking over the leader's status is supplanted in three or four
years, and the new leader will kill some of the infants he sired. How do the
authorities explain this reproductive equality? Their theories seem to lose
significance
unless winners continue to be winners and reproductive inequality persists.

For some categories of intraspecific killing, data are not sufficient, and owing to
the difficulty of observations in the wild, we have to wait for future studies for
detailed analysis, with the personal histories of the individuals involved. It would
be regrettable if only such dramatic episodes receive attention, while the reality of
each society is neglected. Social episodes that are not as dramatic seem to be the
clue to explaining these events. Episode analysis, while taking the factors in
section II into consideration, is the only way to solve the present problem. These
factors are general and basic, discovered through the long-term studies of primate
societies, but they were not framed with the intention of explaining the adaptive
significance or purposefulness of intraspecific killing (e.g. classes of infanticide in
Hrdy, 1979).
It can be said that the higher the species phylogenetically, the more frequent and
varied is intraspecific killing. Table 1 shows the combination of the attacker and
victim for pongids and cercopithecids. It is apparent that pongids have a wider
variety of killing than cercopithecids. these episodes are unquestionably more
frequent in the inter- unit-group context than in the intra-unit-group context. For
the inter- unit-group episodes, Table 2 further differentiates pongids and
cercopithecids.
Finally, there is one important problem ignored by most researchers. One of the
characteristics of intraspecific killing among non-human primates is that there are
few counteractions from a third person or from the society toward the killer. This
may be related to the fact that most episodes are in the inter-unit-group context.
But, in the case of P. entel/us at Dharwar, females just watched males fight, no
behavior by the females to hinder infanticide was recorded, and their attitude
toward the injured infants was unhelpful (Sugiyama, 1965).
On the other hand, for the cases at Abu, Hrdy ( 1974) reported. In every case,
females other than the mother intervened. Sol and Pawless repeatedly played the
most daring roles in defense of the infant'. Despite that, the infants were killed. In
the case of an intra-group episode, in which an adult male gorilla killed an aged
female, the observe(r reported, 'Other group members looked on but did not
interfere' (Fossey, 1981 ). At least six out of the 12 reported cases of infanticide
among chimpanzees were followed by an attack by the captor on the victim's
mother (Kawanaka, 1981). Such an attack may have been a kind of precourtship
aggression, but it is likely that the mothers were protecting their young from the
captors. The killed
Table 1. Combinations of attacker and victtm in intraspeCtjic killing among

non-human pnmates
VICTIM
ATTACKER Adult Male Adult female Infant
Pongoids
Adult Male Common Rare Common
Adult Female Nil Nil Rare
Cercopithecids
Adult male Rare Nil Common
Adult Female Nil Nil Nil
Table 2. lntraspeciftc killing among nonhuman pnmates, sorted by the

Vtctim’s identity and context
VICTIM
Adult male Adult Female Infant
Intra-unit-group-episodes
Pongoids Nil Gorilla gorilla Pan troglodytes
Gonjlla gorilla
Cercopithecids Nil Nil Presbytis cristata
Inter-unit-group-episodes
Pongoids Pan troglodytes Pan troglodytes Pan troglodytes
Gorilla gorilla? Gorilla gorilla
Cercopithecids Papio ursinus* Nil Cercopithecus ascanius
Presbytis senex Macaca fuscata
Macaca sylvana?
Papio liamadryas
Papio ursinus*
Prebytis entellus
Prebytis entellus
Presbytis senex
Theropithecus gelada
*Saayman (1971).
infants were consumed by many group members. Were they simply having a little
taste of meat without regard to the victim's identity or species? Or did it signify the
acceptance of the fact of killing? Or was it a communal feast to eliminate the fact of
the act from the group memory? About these points, there must be some differences
at least between the case of Passion's cannibalism and tbe other cases. This
raises the question of whether their action of killing is within the norms or outside
them. Probably, higher in the evolutionary scale, intraspecific killing has more
complicated episodes and increases in frequency (Roper, 1969).
In this process the questions are how the counteractions towards such actions have
emerged and how they have been established in societies. , These questions are
related to the origin of institutions, involving the " rules of female possession, the
resolution of inter-group intolerance, and the evolution of society.
.
A triangular brief on the evolution

of brain and law
PAUL D. MacLEAN
Laboratory of Brain Evolution and Behavior,
National Institute of Mental Health,
9000 Rockville Pike, Bethesda, MD 20205, USA
A comparison of the brains of existing vertebrates, together with an

examination of the fossil record, provides evidence that the human brain has
evolved and expanded to its great size while retaining the anatomical and
chemical features of three basic formations that reflect an ancestral
relationship to reptiles, early mammals, and late mammals. Such factors
must be taken into account when considering the origins and evolution of
the law. The present paper focuses on recent evidence that the proto-
reptilian part of the forebrain plays an essential role in the display of power
involved in establishing and defending territory. Such findings are of basic
significance in regard to the underpinnings of law which depends on the
collective show of power and the collective use of power in protecting both
the rights of an individual and those of a society. The concluding discussion
deals with the questions relating to the roles of the two newer evolutionary
formations of the brain in (i) adherence to the primitive principles of
'precedent' in deciding the law; (ii) providing through reification a pervasive
respect for the sovereignty of the law; and finally (iii) generating a concern
for a merciful application of the law and merciful justice.
Somewhat like citizens of Great Britain, the lives of all of us are regulated by an
unwritten constitution, the basis of which we must look for in the human brain. The
argument in my triangular brief on the evolution of the brain and law hinges on
evidence about the brain that can be quite easily demonstrated to anyone. Since time
permits only a cursory examination of this immense subject, I will focus mainly on
the primitive brain roots of the 'will-to-power' because, in essence, the existence of
law depends on a collective show of power and the collective use of power. In
conclusion, I will make two comments about functions of evolutionary newer parts
of the brain that are relevant to two important aspects of the law. The first comment
will pertain to brain mechanisms underlying reification with respect to the
sovereignty of the law (i.e. the supreme power of the law) that inculcates in a society
a pervasive respect for the law without which a judicial system could not exist.
Finally, the second comment will pertain to most recently evolved parts of the brain
that have made possible a concern for a merciful application of the law and merciful
justice.
THE TRIUNE BRAIN

A comparative study of the brains of existing terrestrial vertebrates, together with an
examination of the fossil record, indicates that the human brain has evolved to its
great size while retaining anatomical and chemical features reflecting an ancestral
relationship to reptiles, early mammals and late mammals (Fig. 1). We have, so to
speak, a hierarchy of three brains in one, a triune brain (MacLean, 1970, 1973a ). As
we shall see, there exists a period of time between us and our reptilian ancestors that
amounts to more than 10 million human generations. Apropos of the just mentioned,
unwritten constitution, it is of utmost significance that the two older evolutionary
formations of the brain lack the neural capacity for verbal communication.
As one of Chaucer's characters was fond of saying, 'Quid juris est?' What is the law?
If traced to its earliest roots, we would have to go back to the unwritten law of the
sea, which in a later era creeps ashore and carries over as the law of the land. For our
purposes, it will suffice to introduce the history of the law with our reptilian
ancestors living more than 250 million years ago. We will have to base our
inferences on what can be learned from the fossil record and from neuro-behavioral
findings on existing reptiles that belong to a different reptilian stock from ours.
Given this angle of our triangular brief, we
will then be in a position to consider the two remaining angles representing

psychological functions of the paleo-and neomammalian brains.
Figure I. THE TRIUNE BRAIN. In its evolution the primate forebrain expands along
the lines of three basic formations which anatomically and biochemically reflect an
ancestral relationship, respectively, to reptiles, early mammals and late mammals.
The three formations are labeled at the level of the forebrain which may be regarded
as the psychenecephalon (From MacLean, 1967)
THE PROTOREPTILIAN BRAIN

In recent years developments in histochemistry have been of invaluable help in
identifying corresponding parts of the forebrain in reptiles, birds and mammals. A
stain for cholinesterase intensely colors ganglionic structures representing the
protoreptilian forebrain. In clinical neurology, these structures are listed as
belonging to the so-called basal ganglia. Since there is no term that applies generally
to the ganglia in the telencephalon, I will simply refer to them as the striatal complex
or for short, as the R-complex (MacLean, 1973a,b). In reptiles and birds most of the
R-complex is found in the paleostriatum located below the dorsal medullary lamina.
In mammals this complex
comprises the olfactostriatum {including the olfactory tubercle and nucleus
accumbens), the corpus striatum {consisting of the caudate nucleus and putamen),
globus pallidus and surrounding collections of gray matter. In the last 15 years it has
become well known that parts of the R-complex are rich in dopamine and in opiate
receptors. Dopa- mine is a neural sap that derives from arborizing nerve fibers
climbing upwards from cells in the substantia nigra of the midbrain. If these vines
wither and die from whatever cause {age, disease, experimental manipulation) the
organism becomes almost immobilized.
Because of disorders of motility in patients with disease of structures C connected

with the R-comlex, neurologists long ago concluded that " J the complex itself is
part of the motor system under the control of the motor cortex of the neocortex. This
view has prevailed ill spite of the recognition that destruction of large parts of the R-
complex may result in no apparent motor deficits. Moreover, electrical stimulation
within extensive parts of the R-complex results in no apparent motor responses.
It was because of such considerations that several years ago we designed our present
Laboratory for taking a new approach to investigations on the functions of the R-
complex. In the past it had been customary to make neurobehavioral observations on
animals living isolated in cages and subjected to psychological tests in which they
performed within some artificial apparatus or manipulated inanimate objects. In
designing our present laboratory, it was the purpose to learn whether or not
experiments on animals living under seminatural conditions, and interacting with
other animals, might reveal functions of the R-complex that would otherwise not be
apparent.
ReptIlian behavior
As in other fields of science, progress in neurobehavioral research depends partly
upon the recognition of similarities and differences of things. Otherwise known as
the therapsids, the mammal-like reptiles are of gerat human interest because they are
so close to the roots of our family tree. Long before the dinosaurs they covered the
face of the earth when there was but one giant continent now known as Pangaea. On
the basis of what is known about their limb bones and body carriage, head-shape,
dentition, articulation of the jaw and other features, the advanced therapsids closely
approached the mammalian condition {Romer, 1966; Colbert, 1969).
Here are a few questions that further paleobiological research might

[78] LAW , BIOLOGY AND CULTURE
help to resolve: did the therapsids lay eggs? Is there fossil evidence that they were
accompanied by young? Were they capable of audiovocal communication, or is it
likely that they were mute like most of today's lizards? Will more detailed study of
sites of muscular insertions yield clues as to whether or not the therapsids engaged in
communicative displays such as those to be described shortly? As may be revealed
by cranial endocasts, what was the morphology of the brian in the most \, advanced
therapsids?
For comparative purposes, it is unfortunate that no existing reptiles are directly in
line with mammals. One variety of the stem reptiles was sufficiently lizard-like as to
merit the name Varanosaurus (Romer , 1966; Colbert, 1969), the same name
identified with today's monitor lizard, of which the Komodo dragon is one example
(Auffenberg, 1978). Of existing reptiles, lizards would probably have the closest
resemblance to the therapsids.
Analysis of reptilian behavior

In analyzing the behavior of lizards one can identify more than 25 forms of behavior
that are also characteristic of mammals (MacLean, 1975). The upper part of the list
Table 1 incudes forms of behavior that primarily involve self-preservation or
preservation of a social group, while the lower part lists kinds of behavior seen in
social groups and in procreation. As will be discussed, the first three items
pertaining to behavior involved in the selection of homesite, establishment of
territory, and use of domain are particularly pertinent to our present topic.
The behavioral profile
The total of all the building blocks of behavior constitute the behavioral profile or
ethogram of an animal. Exclusive of verbal behavior, one recognizes in human
beings and other terrestrial vertebrates two main aspects of the behavioral profile.
For descriptive purposes, the two aspects might be likened to the profiles of two
mountain ranges seen from a distance. In one range are the distinctive peaks and
subpeaks representing the chain of activities in an animal's (1) daily master routine
and (2) subroutines. In the other range are four main peaks and subpeaks
corresponding to four primary types of behavioral patterns, otherwise called
displays, used in prosematic communication. Prosematic, meaning rudimentary
signaling, applies to any non-verbal signal-vocal, bodily, or chemical-used in com-
munication (MacLean, 1975). In lizards, the four main types of
Table 1. Special forms of basic behavior

1 Selection and preparation of homesite
2 Establishment of territory Domain
3 Use of home range
4 Showing place-preferences
5 Trail making
6 Marking of territory
7 Patrolling territory
8 Ritualistic display in defense of territory, commonly involving the use of coloration
and adornments
9 Formalized intraspecific fighting in defense of territory
10 Triumphal display in successful defense
11 Assumption of distinctive postures and coloration in signaling surrender
12 Use of defecation posts
13 Foraging
14 Hunting
15 Homing
16 Hoarding
17 Formation of social groups
18 Establishment of social hierarchy by ritualistic display and other means
19 Greeting
20 Grooming
21 Courtship, with displays using coloration and adornments
22 Mating
23 Breeding and, in isolated instances, attending offspring
24 Flocking
25 Migration
communicative displays are referred to as (I) signature; (2) challenge; (3) courtship;
and (4) submissive displays.
At our laboratory we are testing the hypothesis that the R-complex plays a basic
role inthe regulation of the daily master routine and subroutines, as well as in the
different kinds of communicative displays that have been mentioned. In deference to
our topic, I will briefly describe the four main kinds of displays and then summarize
our neurobehavioral findings on the challenge display.
Four main displays. The common green anolis lizard serves to illustrate the four
main kinds of displays which variously incorporate dynamic and
static modifiers: ( 1) The signature display consists simply of three to five combined
head nods and pushups, together with a brief extension of a crimson throat fan
occurring after the second head nod. The signature display reflects a kind of self-
assertion that finds expression in both non-social and social contexts. It seems to
afford individual recognition among members of a group. (2) Challenge displays are
of two types (distant and close in) and are used mainly by territorial males (i) in
establishing territory; (ii) maintaining dominance within a social group; and (iii)
fending off invaders. The challenge display includes dynamic components of the
signature display, combined with a number of static modifiers that make the subject
appear larger in size. (3) The courtship display begins with a signature display
followed by a number of head nods and an approach towards the female with a
prancing gait. (4) The anolian submissive display is characterized by four slight up
and down motions of the head and is performed by lizards of either sex and of all
ages.
Neurobehaviorat findings
For present purposes it is enough to say that birds and mammals also engage in some
variation of the four main kinds of displays just mentioned. Here I will summarize
the outcome of our neurobehavioral investigations relevant to challenge displays
seen in lizards and monkeys. In our inital work on lizards, we focused on the
challenge display of territorial males because it is the easiest to manipulate and
quantify. Utilizing an experimental approach that does not interfere with
thermoregulation, we found that lesions of the R-complex, but not of other parts of
the hemisphere, abolish the challenge display of territorial males ( Greenberg et at.,
1979).
In a study continuing since 1961, I have been investigating the role of different brain
structures in the display behavior of squirrel monkeys. In both the challenge and
courtship displays, the assertive monkey vocalizes, spreads one thigh, and directs
the erect phallus towards the other animal. The 'signature' displays of one subspecies
incorporates the features of the challenge and courtship displays. This display may
be used as a form of greeting, and, interestingly, monkeys of this particular
subspecies will regularly display to their reflections in a mirror. We informally refer
to the mirror displaying monkey as the
Gruter and Bohannan [8 I]
gothic-type because its white ocular patch comes to a peak over the eye like a gothic
arch. We identify the monkeys of subspecies that do not perform the mirror display
as 'Romans' because the supraocular part of the patch is round like a roman arch.
Since it allows control of several variables, I have used the mirror display test as a
means for detecting what parts of the brain are implicated in prosematic
communication. In experiments on more than 120 monkeys, I have found that the
medial pallidal segment of the R-complex is a site of convergence of neural systems
involved in the species-typical mirror display of adult squirrel monkeys. Electro-
coagulations made within this region (MacLean, 1978a) or its projecting pathways
either eliminate or result in a fragmentation of the display.
Comment on neurobehavioral findings

The results of these experiments indicate that in animals as diverse as lizards and
monkeys, the R-complex plays a basic role in displays used in social
communication. They also provide indirect evidence that the R-complex is essential
for conspecific recognition through the performance of like kinds of behavior. The
performance of like kinds of behavior (for which I use the term 'isopraxis') is what
typifies a species (MacLean,1975).
Isopraxis is one of six important interoperative behaviors seen in reptiles, birds and
mammals (the others are routinizing, tropistic, repetitious, reenactment and
deceptive behaviors). Without going into detail, I shall simply point out that in
human activities these behaviors find expression in such human activities as the
performance of daily routines and subroutines; responding to partial representations,
whether alive or inanimate; adherence to fashions, both social and scientific;
repetitious and obsessive-compulsive acts; obeisance to precedent as in legal and
other matters; ceremonial reenactments; and all kinds of deception.
I mention these proclivities because one finds it frequently stated that all human
behavior is learned. One anthropologist has stated, 'The evidence indicates quite
clearly that everything that human beings do as human beings, they have had to
learn from other human beings' (Montagu, 1956). If that is so, one might ask why is
it that in spite of the high degree of human intelligence and culturally learned
behavior, human beings continue to do all the ordinary things that animals do? Take,
for example, the close-in challenge display. Here, there appear to be some
carryovers from animal to human displays that
[82] LAW , BIOLOGY AND CULTURE
are so subtle that they have escaped even the notice of expert ethologists, making it
all the more remarkable that, if as claimed, everything human beings do as human
beings, they have had to learn from other human beings. In the close-in display,
lacertilians rise up on all fours and present themselves sideways while stepping in a
stilted, staccato manner that makes them appear off balance. Some rodents perform a
similar broadside display, but it happens so rapidly that observers may fail to notice
it. I had been unaware that the challenge display of two adult, rival gorillas
incorporated lacertilian features until I saw Dian Fossey at one of our Laboratory
seminars perform what she refers to as the 'parallel display' of two silverbacks.
When she mimicked their sideways presentation and their walking with stilted,
awkward steps, one was immediately reminded of the close-in display of certain
lizards. In the case of chimpanzees, Jane Goodall (1968) has described a bipedal
swagger which appears to correspond to the strutting display of the gorilla. Her
description reminds one of the posture and movements of a Japanese wrestler.
In all of the above examples, the stilted, staccato steps seen in the displays seem to
carry the message of a series of exclamation marks, calling to mind the goose step of
a military parade and its similarity in profile to the Schriigstellung gait of a Komodo
dragon (Auffenberg, 1978). Among different species of animals, the sideways
presentation and stilted staccato steps have such an uncanny resemblance that it
would almost seem that the challenge display had been genetically packaged and
handed up the phylogenetic tree of mammals.
The focus on the challenge display might give the impression that it was more
important than the other displays. One might argue, however that the submissive
display is the most important of all displays, because without it no one could ever
survive! As ethologists have made it popularly known (e.g. Lorenz, 1966), a
submissive display serves the role of toning down an aggressor which might
otherwise destroy the weaker individual.
The question of power
These foregoing remarks call attention again to the question of the nature of power,
the origin of power, and the use of power. Biologically, we find power exerting itself
primarily in connection with an animal's territory and with mating and breeding. To
judge by some writings, there continues to be heated debate as to whether or not
human beings are naturally territorial. In the strict ethological sense,
Gruter and B()hannan [ 83 ]
erritoriality refers to an animal's demonstrated determination to protect a particular
piece of ground. The discussion of territory becomes more manageable if it is dealt
with in terms of an animal's domain, which in addition to territory, encompasses a
home site and an ill-defined activity range (Table 1). In an operational sense, the
domain represents a place of refuge, a place to hunt for food, and a place to mate and
to breed. Reduced to first principles, it represents a"' space required for an organism
for both self-preservation and the preservation of the species. In his memorable
study of bird life, Eliot Howard ( 1920) concluded that in many species of birds, the
male must show its ability to establish and protect a certain area before it will be
recognized as a suitable mate by the female.
Whether or not a particular species of animal appears to be territorial may depend
upon the availability of food. Nevertheless, even when the need for food requires a
roving existence, animals have an ill-defined domain, and somewhat like a traveling
circus, the homesite and territory travel along with them. Monkeys such as the red
howler or apes such as chimpanzees and gorillas may confine themselves to a
domain (Goodall, 1968; Fossey, 1976) in which there exists a moving social space
which other animals are not allowed to enter. It is usual for strictly territorial animals
to patrol their territory, whereas those with a moving territorial space may patrol
their domain
(Goodalletal.,1979).
For animals that are not strictly territorial, there may exist artificial territories at
the time of mating. The U ganda kob, for example, return to breeding grounds where
they must compete for, and hold onto, leks in order to earn the attraction of the
female and the opportunity to mate with her (Buechner, 1961). In human affairs,
there perhaps exists a counterpart for this in the seasonal games that occur at the
local, national and international levels.
The will-to-power
The striving for territory is but one manifestation of the struggle for dominance
everywhere manifest in nature. What is the origin of this life force? Why does it find
more forceful expression in some individuals than in others?
As is well known, the will-to-power eventually became the driving force of
Nietzsche's philosophy. The idea of the will-to-power and the superman was an
indissociable part of Nietzsche's revelation in August 1881, about the doctrine of the
eternal recurrence (Kauffman, 1968).
He concluded that the will-to-power is the basic life force of the entire universe.
'Thus life taught me," he wrote (Nietzsche, 1888). In Nietzsche's superman, we hear
echos of Aristotle's 'great-souled man', who being far superior to other human
beings, 'is justified in despising other people' (Nichomachean Ethics). Similarly,
Nietzsche's superman has the draconian right of riding roughshod over other
people. As one interpreter explained, , All that proceeds from power is good, all that
springs from weakness is bad' (Forster-Nietzsche, 1954).
In the world of animals, one will hardly find the will-to-power more dramatically
expressed than in the behavior of some lizards. To see two rainbow lizards in their
resplendent colors striving for dominance is like returning to the days of King
Arthur. In a contest, once the gauntlet is thrown down, the challenge display gives
way to violent combat and the struggle is unrelenting. Twice we have seen dominant
rainbow lizards humiliated in defeat. They lost their majestic colors, lapsed into a
depression, and died two weeks later.
It is one thing to describe the will-to-power as it occurs in nature, but quite another
to offer any explanation for it. In our laboratory, we have a large holding terrarium
for new shipments of the common green anolis lizard. In this habitat it invariably
happens that some anolians rise up from the ranks and strive to achieve dominance
over the animals perched on one or the other of two main branches. Reminiscent of a
top sergeant barking orders, once a lizard achieves control of a branch, he is
generally able to maintain order and discipline among the others by repeatedly
engaging in his signature display. In rsponse to this display, the other lizards may be
seen to nod their assentive display in unison.
Marking territory
As familiarly illustrated by the house dog and cat, animals with a large olfactory
apparatus mark their territories with urine. The message, so-to-speak, is 'stay away'.
We have just seen, on the other hand, how a visually dominant animal such as a
squirrel monkey thrusts the erect genital as part of an aggressive visual display.
Wickler has described sentinel monkeys in tribes of baboons and green monkeys
which sit at lookout sites with their thighs spread and a display of partial erection
while the rest of the animals feed or take a siesta (Wickler, 1967). He regards this
display as an 'optical marker' of boundaries, warning other animals not to intrude. In
mythology we see the erect phallus superstitiously endowed with the power of
protection. In primitive cultures in different parts of the world, house guards (stone
monu-
Gruter and Bohannan [ 85 ]
ments showing an erect phallus) are used to mark territorial boundaries. It was as
though a visual, urogenital symbol were used as a substitute for the olfactory,
urinary, territorial markings of macrosmatic animals {MacLean, 1973a).
The human use of symbols affords unlimited boundaries to protected ‘conceptual
space'. In addition to our personal space and domestic space about which we feel
possessive and protective, we incorporate into the scheme of belongingness the
boundaries of a town, city, county, off-shore areas, and now in modern times, the
territory of outer space. We can add to this listing the space we assign to schools,
churches, clubs and the like. Administrators in the same business firm ; try to avoid
overlap in areas assigned to salesmen. Friends in the same profession shun
overlapping competition. Many teachers and scientists have the reputation of
establishing intellectual and research territories and protecting them with all their
might. If human beings are not born with some degree of territorial itch, it is
remarkable that there is so much preoccupation with trespass and non-trespass and
that in every advanced culture complex legal systems and a whole body of law have
evolved for settling disputes regarding ownership of lands and possessions. If all
human behavior is learned, then human beings seem naturally to have learned very
well to be territorial! Esser has described institutionalized boys with severe mental
retardation and learning handicaps who will, in anew setting, establish
miniterritories and readopt them after a year of separation {Esser, 1968,1973).
The matter of size and power

Among lizards the spoils usually go to the larger animal. Lizards have an uncanny
way of recognizing a larger animal. A Komodo dragon will immediately size up an
approaching stranger and step aside if it is larger {Auffenberg, 1978). Evans has
shown, however, that size is not the only factor {Evans, 1936 ). The territorial lizard
on its home ground appears to hold an advantage over an intruder. There are other
factors that do not involve power and size per se. Recalling what takes place in the
political arena, it may be the number of displays that decides a winner in a contest
{cf. Masters, forthcoming). As Oesmond Morris has commented upon in discussing
'typical intensity', it may be the number of telephone rings, rather than the loudness
of the ring, that brings the party to the phone {Morris, 1957). As we as human
beings have learned so well, there may be other modifiers such as color, various
trappings, size of retinue, that help to make up for what
particular individual lacks in size. Ethologists have made it popularly known that
ritualized aggressive displays and their passive counterpart -submissive displays-
make it possible under most circumstances to preserve an individual's rights and
avoid unnecessary, and sometimes mortal, conflict. In this country and elsewhere in
the world there is currently much debate about the desirability of using a display of
police officers for maintaining law and order at home and of displaying military
power for preserving international peace.
Having delved into some of the primitive brain roots of power, we will next consider
briefly the two other main evolutionary formations of the brain and indicate how
they influence the collective power of the law and elaborate upon such primitive
principles as 'precedent' in deciding the law.
THE PALEOMAMMALIAN BRAIN (LIMBIC SYSTEM)

In the evolution from reptiles to mammals, three key behavioral changes were the
development of (1) nursing, (2) parental care and (3) play (Maclean, 1978b). Further
research on the mammal-like reptiles mentioned earlier may help to give clues about
transitional changes leading up to typical mammalian behavior. One might say in
essence that the history of the evolution of mammals unfolds as the history of the
evolution of the family. Taking care of the family, not only at the time of birth, but
also as in the case of human beings, for a prolonged period, amounts to a progressive
evolution of the sense of responsibility and what we call conscience. Many species
of existing lizards prey upon their young. Will further paleontological field research
reveal whether or not the mammal-like reptiles were accompanied by young? Or is it
possible that like the offspring of today's giant Komodo dragons, the young had to
escape to the trees to avoid being cannabilized? With the evolution of mammals
there appears to have come into being the primal law and commandment: 'Thou shalt
not eat thy young or other flesh of thine own kind'. The later commandment, 'Thou
shalt not kill', is inherent in the primal commandment.
In the evolution of mammals vocalization and hearing become of utmost importance
for maintaining parent.-offspring relationships. Presumably vocal communication
helped to insure contact among the diminutive early mammals living in the dark
floor of the forest. The so-called 'isolation call' is probably the most basic
mammalian vocalization, serving to maintain maternal-offspring contact and group
affiliation.
.
Most of the old cortex identified with early mammals is found in the great limbic
lobe which is found as a common denominator in the brains of all mammals. In
1952, I suggested the term limbic system as a designation for this cortex and the
structures of the brain stem with which it is primarily connected {MacLean, 1952).
Research of the past 40 years has provided evidence that the limbic system derives
information in terms of emotional feelings that guide behavior required for self-
preservation and the preservation of the species. Like a great metropolis, the nerve
cells of the limbic system fall into three main districts. Two of the districts are
closely connected to the olfactory apparatus. The population of nerve cells in district
1 are concerned with behavior involved in self-perservation-namely, feeding,
fighting and self-protection {MacLean, 1952); while the population in district 2
participates in primal sexual behavior and socio-sexual expression {MacLean,
1973a).
In reptiles, there is no counterpart of the third district that expands to a great size in
primates and culminates in the human brain. The main communication line for this
district bypasses the olfactory apparatus. We have found that hamsters deprived of
all the neocortex at the time of birth are capable of all forms of hamster-typical
behavior, including mating and rearing young {MacLean, 1978b; Murphy, MacLean
& Hamilton, 1981). If, however, they are deprived of the limbic cortex of this third
district, there are deficits in maternal behavior. Moreover, such animals do not
develop play behavior-a mammalian trait so fundamental for harmony in the nest
and social affiliation later on. It was as though these animals had regressed towards a
reptilian condition. In the evolution of this great third subdivision of the limbic
system, we discern the roots of responsibility to other members of the same species-
a responsibility that is so fundamental to the even-handed administration of the law
and Portia's merciful justice.
The clinical study of limbic epilepsy provides the best evidence that the limbic
system is basically involved in emotional experience and expression. During the
aura occurring at the beginning of an epileptic storm, the patient's mind lights up
with vivid emotional feelings that range all the way from intense fear to ecstacy.
Significantly, such a storm may spark feelings like those associated with belief and
conviction regarding what is real, true and important. Afterwards, so to speak, the
lights go out in the limbic system and during such times an individual may carry out
very complicated behavior dependent on the neocortex and yet have no memory of it
afterwards. They behave like
disembodied spirits. Such considerations indicate that without our primitive limbic
system, we would have no gut feeling for the reality of our existence, as well as no
belief or conviction regarding what is true and important. Such considerations not
only have far reaching implications in regard to epistemology, but also in regard to
the law (see below).
THE NEOMAMMALIAN BRAIN

On the basis of what we have reviewed so far, it would appear that the R-complex
and limbic system are sufficient in themselves for giving expression to the basic
animality. To make this generalization, however, is not to downplay the importance
of the neocortex which mushrooms progressively in higher mammals and reaches its
greatest development in human beings. Nothing is more certain neurologically than
that the neocortex is necessary for language and speech, and that we owe to it the
infinite variety of ways in which we can think and express ourselves.
The constitutional union of minds
Prefatory to a few concluding comments, two of which were mentioned in the
introduction, I will further qualify what was just said by noting that in the case of
law or anything else, it is evident that the new brain is the handmaiden for
rationalizing, justifying and giving verbal expression to the protoreptilian and limbic
parts of our brains that appear incapable of communicating their wants and needs by
the use of words. Take, for example, the matter of precedent. In observing reptiles,
one finds that they are slaves to routine, precedent and ritual. Obeisance to precedent
often has survival value. If, for example, a particular crevice served as an escape
from a predator on one occasion, it may do so again. Hediger has provided
numerous examples of the role of precedent among animals and human beings-the
sum and substance being that if something worked for you once, it may do so again
(Hediger, 1955).
For someone outside the legal profession it may not seem clear why a particular case
cannot be decided on its own merits. Instead, endless time and money are spent in
searching for precedents. Why should it be that one must uncover the case of some
individual living years ago and in some remote place in order to prove that one's
own case has merit? Lawyers will explain that 'the law' likes to be even-handed, to
be as fair as possible to all parties. This, they claim, is best assured by
trying to find a similar case that might have been decided by a particularly renowned
judge or by one of the higher courts. The greater the authority, the greater the weight
of the judgment. What they fail to emphasize is that whoever sits in judgment
derives great reassurance if it can be shown that the ruling on a similar case survived
an appeal. Consider next how the neocortex cooperates with the limbic part of us in
the matter of the reification of the law. In helping to draft the Constitution of
Massachusetts, John Adams injected the contention that we are 'a government of
laws, and not of men'. The reification would have been complete, had he added
women. I call this a reification because it was as though the body of law had been
made flesh through the agency of some external power. It was mentioned earlier that
the limbic system appears to account for our affective feelings associated with belief
and conviction regarding what is real, true and important. This primitive mind
without the capacity to read, write or speak, seems to be responsible not only for
sizing up the reality and suitability of food, mate and the like, but also provides the
feeling of conviction and belief that we attach to our ideas and theories, regardless of
whether they are true or false. Similarly, it must be presumed that it plays a
fundamental role in hypostatizing the sovereignty {the supreme power) of the law.
Without such a reification, the law could hardly be the powerful, pervasive force that
is able to keep a society in line and to do so, for the most part, without even being
seen.
Finally, one last comment. For solutions of situations that arise in the external world,
nature designs the neocortex so that it receives signals primarily from the eyes, ears
and body wall. With its focus on the external world, the neocortex develops at first
somewhat like a coldly reasoning, giant computer. It is the kind of computer that has
the capacity to devise the cruelest of cruel ways of destroying our own kind-
examples of which we have seen in our recent wars and crimes against humanity. As
though foreseeing long ago that a terrible genie was in the making, nature began to
enlarge that part of the neocortex -the prefrontal area-which for the first time in the
world brings a sense of concern for the welfare of all living things. In the progress
from Neanderthal to Cro-Magnon man one sees the human forehead develop from a
low brow to a high brow. Underneath the heightened brow is the prefrontal cortex.
The prefrontal cortex appears to be the only neocortex that looks inward to the
inside world. Significantly, it establishes connections
.
with the third great subdivision of the limbic system concerned with parental care
and play. Clinically, there is evidence that the prefrontal cortex by looking inward,
so to speak, obtains the gut feeling required for identifying with another individual.
It is this new development that makes possible the insight required for the foresight
to plan for the needs of others as well as the self-to use our knowledge to alleviate
suffering everywhere.
Constraints on human behavior and

the biological nature of man
HUBERT MARKL
Department of Biology, University of Konstanz,
Postfach 5560, D-7750 Konstanz I, FRG
The contribution of biology to a better understanding of human legal behavior seems

severely limited for three reasons: (I) Since laws are cognitive constructs of the
human mind which must be verbalized to become effective, man is the only species
in which legal behavior can be studied. All inferences from animal behavior studies
and from evolutionary considerations are highly speculative with respect to human
legal behavior. (2) In the ontogenetic development of human behavior there is
adaptation of the behavior to the environment, including culture. There seems no
reliable procedure to factor out their relative contribution, particularly since genetic
adaptation can be easily phenocopied. Therefore it is only rarely possible to separate
a 'biological' component of human behavior from a 'cultural' one. (3) Most theories
pertaining to the evolution of behavior in animals (and more so in man) are 'weak'
theories with some retrodictive but little predictive power; they allow us to define
probable modes for behavioral averages but say little about the behavior of
individuals, which is at issue in legal considerations.
Rules are constraints on the behavior of systems. Rules are descriptive if they
separate the more probable behavior states from the less probable ones. They are
prescriptive or proscriptive rules or norms if they differentiate between admissible
behaviors and forbidden ones. Thus, in applying rules, behavior is first classified and
then evaluated with respect either to the probability or the desirability of realization.
Norms or laws are cognitive constructs according to which different actions are
classified and evaluated. Therefore, they can be reconstructed from the overt
behavior of an organism only with very limited reliability-unless, of course, we can
find ways to get access to the mental representations that guide the organism's
behavioral decisions. Inasmuch as the characteristics or even the existence of mental
representations in non-human beings remain in doubt (Griffin, 1976, 1978, 1982), it
seems anything between difficult and futile to address the question of normative
behavioral rules in animals. Analyzing the evolution of behavioral rules is, thus,
particularly difficult.
Every effort to trace human behavioral evolution must be guided by careful
consideration of the different processes through which rules can be imposed on
behavior. The final outcome-a behavioral action -is the result of a multi-stage
constraining process through which a particular activity is selected out of the total
universe of behavioral abilities. In the course of evolution to 'higher' forms, there has
been an amplification of levels at which constraining or selective influences can be
exerted. Man exhibits a high degree of freedom of behavioral' choice-human
behavior can be influenced, modified and manipulated throughout life more than
that of any other known organism. In parallel with this seemingly limitless capacity
for behavioral variability, we find a plentitude of normative rules that constrain
behavior: rules that may go to such extremes that nothing is left to be decided by
individual discretion. Thus, the human being is every bit as much a slave to its
normative representations as any insect is to its genetic program.
This is the first point I want to make: the propensity to legal behavior seems clearly
correlated with demolition of such constraints on behavior as we find typically
represented in animals. This point is not new (for example, Bonner, 1980), but it
needs re-emphasis each time one tries to understand the biological bases of human
legal behavior. This correlation points to a crucial fact in the evolution of 'man the
rule-maker'. The laborious and precarious effort necessary to constrain man's
behavior by norms must have been outweighed by gains in behavioral flexibility and
freedom of choice.
However, when one speculates on how 'human nature' constrains moral and legal
conduct, one might just be looking at the matter from the wrong angle. It may be
that the emphasis would be better placed on the adaptive benefits of being
biologically underdetermined and dependent on guiding principles-poorly adapted,
but highly adapt- able to, and in need of becoming adapted to, normative rules. The
pre-eminent task of human ethology and social biology would then be to find the
causes of unleashing man's behavior from genetic constraints rather than to track
down any last remnants of biological fetters. It might, in fact, be misguided to look
primarily for the biological determinants of legal behavior when our main interest
should be the explanation of that remarkable evolutionary process that " leaves, by
its very nature, a helpless creature at the mercy of the inventiveness of his fellow
beings and his forebears and their capacity to adapt their behavior. Such a process
obviously has costs in cultural failures and maladaptations, but those costs may be
overridden by the benefits of cultural malleability.
But even if we accept it as our task to search for the biological determinants of
norm-related behavior, there is a methodological problem that seriously limits the
usefulness of comparative socio-biological models in trying to understand human
behavior. If sociobiological reasoning about the biological nature of man means
anything at all, it refers to genetic constraints on and specific predispositions toward
behavior. More specifically, it must refer to constraints on the behavioral
development of the newborn: to epigenetic rules (Lumsden & Wilson, 1981) which
owe their existence, if not directly to the individual's genes, then at least to the
interaction of genes with external influences on prenatal ontogenesis. In an organism
without learning capacity, such an innate endowment would fully specify the
possible behavior potentials. But in humans we can (at least theoretically)
differentiate between a whole series of constraining and directing influences. Such
constraints are, first of all, the blueprint for development contained in the genome of
the zygote. The directing influences are represented, second, by prenatal
environmental influences on the developing embryo. Post-natally, at a third level,
the child is subject to the directing influences of behavior modification in culture-
dependent learning situations. At that stage, an immense richness of information is
incorporated into the child's behavioral system through imitation learning,
conditioning, cognitive self-structuring, and by formal teaching. Through these
devices the individual can partake in the traditions of its culture. Thus, within the
range of the constraints imposed on the individual by its genes and its prenatal
epigenesis, learning cultural traditions further narrows behavioral options. Each
culture makes specific selections from the gamut of behavioral possibilities open to
a human neonate.
Looking at this process in more detail, we find that no individual can experience the
whole range of information at the disposal of any
given culture. Rather, a further selection occurs: the specific culture taken in is
constrained by earlier development decisions and also by cultural ideas about what
is suitable for one sex or the other, by family structure, social status and parental
predilections. Throughout this process" there is an additional source of behavior
modification-the individual's personal experiences that are different from those of
other members of the same society.
Thus, in a continuous interactive process, there is a complex mixture of genetic,
foetal, cultural and personal individualization; the outcome is a unique person with
unique patterns of species-typical, culture-typical, and personal capacities,
constraints and inclinations.
Finally, during each moment of its life, while actual decisions about particular
behavioral actions are being made, the individual decides on the basis of preparation
by its genes, its ontogeny and its various learning experiences. The decision also
depends on how much time for deliberation is available, on which particular fraction
of all this stored information is available, on evaluation of available options and on
the risks and benefits to be expected from alternative actions as they are perceived
by the deciding ego.
All of these behavior-constraining factors result from selection processes. The
specific genetic blueprint has been arranged by means of natural selection during the
evolution of a species. The contents of the cultural tradition derive from a history of
cultural selections. An individual's personal learning history makes him select
particular behaviors and suppress others. And, in a situation that demands immediate
decision, the actor selects the most promising alternative.
Ultimately, reproductive (Darwinian) fitness is the major criterion of selection.
However, most behavioral actions cannot be immediately evaluated by the
individual in terms of Darwinian fitness. Therefore, the individual's behavioral
machinery is equipped, either through the genetic program or through traditional and
personal experience, with intervening goals and accounting procedures. In the long
run, of course, no intervening goals should be retained unless they are positively (or,
at least, not negatively) correlated with Darwinian fitness.
In fact, the relationship between intervening goals and ultimate fitness soon
becomes very complicated and unpredictable. Even in animals, it is often difficult to
assign "adaptive values" (that is, positive fitness correlations) to particular
behavioral traits. A large part of sociobiological modeling aims to explain how
seemingly unfit, "altruistic", behavior could indirectly enhance the genetic fitness of
the performer (Markl, 1980).
In man, conditions are given for all possible indirect ways of gaining fitness in
addition to the classical direct one. Therefore, describing the relation between
intervening goals and ultimate fitness for many behavioral traits and motivations
becomes a prolific exercise in plausible speculation rather than one in presenting
convincing evidence. The Panglossian trap is always set: explaining everything that
exists as somehow secretly and miraculously profitable to inclusive fitness in the
long run. This propensity is even more disconcerting in light of the argument that
behavioral mal adaptations in humans are due to their biological past. It should be
emphasized that maladaptability in a trait (in the biological sense of the word) has to
be quantitatively measured in terms of the bearer's inclusive fitness as compared to
competitors who do not bear the trait. Only so can maladaptability be regarded as
proven. However, man's ways of gaining indirect fitness seem limited only by the
fertility of the actors' (or the investigators') imagination. Many allegedly
maladaptive behaviors seem to have little direct detrimental effect on man's
Darwinian fitness, especially under advanced conditions of medical care, which is
the natural environment of humankind in an evolved civilization. Therefore, we are
still waiting for a demonstration of a biologically imposed behavioral characteristic
that is maladaptive because of the changed living conditions of modem civilization.
We still cannot point to a trait that makes its bearers suffer from lowered inclusive
fitness because of the evolutionary load on their behavior. There seems to be far
more convincing evidence of cultural maladaptations than of maladaptation in
biological programming.
If it is so difficult for an assiduous investigator plausibly to relate intervening
goals to ultimate fitness, how much more difficult must it be for the average
individual! This may mean that these intervening goals have, by genetic selection or
cultural selection or both, been carefully programmed to be positively correlated
with ultimate fitness. This is a functionalist credo to which many biologists may
want to subscribe (Lumsden & Wilson, 1981). But many social scientists
(Luckmann, 1979) may regard it as utterly useless for explaining man's cultural
diversity. This state of affairs can, however, also mean that an individual human
being's Darwinian fitness counts so little in cultural evolution and in determining the
historical success of competing cultures, that the intervening goals of everyday
behavior are scarcely under ultimate fitness control at all. Therefore, they are not
actually "intervening", but rather have become cultural ends in their own right,
pursued for their own benefit, and evaluated in accordance
with standards of cultural fitness that are only remotely kept in check by ultimate
Darwinian fitness limitations.
From that perspective, the amazing freedom of action in man can be seen as the
result of behavioral organization in which non-cultural causes of fitness are of far
less importance than cultural ones. That is to say, postnatal experience rather than
prenatal endowment may set the frame for cultural performance. If "biological
fitness" differentials are of little influence on "cultural fitness" differentials among
members of a society, and if cultural fitness rather than reproductive fertility of its
members is what determines a society's capacity for coping with the challenges of
life, then we would not expect to find much isomorphic gene-culture matching at all!
If that is so, then models from sociobiology or population genetics lose their
explanatory power at that stage in human evolution at which cultural fitness began to
be predominant over biological fitness. Biological gene pools, which are selected to
outre produce competing gene pools, would gradually have become subordinate to,
perhaps even superseded by, cultures that compete by out producing each other in
goods, knowledge, ideas and skills. Of course, the gene pools are still there and have
to reproduce. But their fitness could become less and less dependent on the primary
biological phenotypes of their members and more and more so on the store and
tradition of cultural experience of their particular societies.
The direct way to test the limits of biological nature on human behavior would be to
discover the constraints on and predispositions for behavior in the human neonate.
Raising animals under conditions of stimulus deprivation or stimulus substitution
has proved to be the most effective method for animal ethologists to tackle the
question of the conditions under which behavioral modification by experience is
possible or necessary. For ethical reasons we cannot systematically manipulate the
environment of a human child in a similar way. I do not belittle the imaginative
efforts of human ethologists in exploring human nature when I add that what we
have actually learned about biological constraints on human behavior from studying
sensory discrimination, spontaneous or conditioned responsive behavior, vocal or
other expressive behavior in normal babies or babies with defined congenital defects
(Eibl-Eibesfeldt, 1973, 1979; von Cranach, Foppa, Lepenies & Ploog, 1979) is
soberingly little. Such studies have indeed demonstrated that the newly born human
is far from being a tabula raJa on which experience inscribes the person's future
character. The baby is prepared by nature to communicate in an adaptive way with
its social environment, especially with its caretakers. It is prepared to
respond to language and to produce human vocal sounds. Above all, it is prepared to
construct from its limited experiences, in an orderly step-wise fashion, its world of
cognitions and its cognition of the world, as Piaget has demonstrated (Piatelli-
Palmarini, 1980).
However, it is just here that the trouble sets in. By constructing this subjective and
interior world of cognitive representations, by actively interacting with and probing
the outside world, and by developing an ability to perform a variety of rational and
emotional operations on these mental representations, the child escapes from the
grip of our ethological methods. The older it grows, the more irrevocable its escape-
and the more interesting it would be to see how much biological (genetic) program
lies behind the different aspects of its behavior. This is not a matter of "not yet
having found the right techniques to disentangle nature and nurture." If we want to
talk at all about real human beings, we have to talk about the constructors and
creators of their cognitive selves as an intricate tapestry of interactions between
genetic nature and empirical experience. To try to separate the contributions of
nature and of learning would be about as sensible as to ask whether the human
species is male or female. It is the pattern of the tapestry and not the origin of the
threads that makes the person! Therefore, the question of how inborn nature
influences norm-related behavior of the adult becomes almost devoid of reasonable
meaning.
There will be strong objections to these ideas from many human ethologists and
sociobiologists. Don't we have the inferential methods for searching for behavioral
universals such as communications rituals (Eibl-Eibesfeldt, 1979) or for
demonstrating by cross-cultural and cross-species comparisons that social
organization follows sociobiological model predictions, e.g., with respect to the
avunculate (Alexander, 1979)? Alas, these methods will not solve the problem.
Neither is it so that genetically programmed behavior must be species-universal.
Evolutionary game theory has taught us to see the adaptive potentials of mixing
strategies and behavior-genetic polymorphism. Nor is it so that species-universal
behavior must be genetically programmed: the ontogeny of bird song, among other
studies, has taught us how firm the grip of learned programs can be on a whole
population. Nor would the proof of a particular behavior or behavioral organization
favoring its producers' inclusive fitness-that is, being demonstrably biologically
adaptive-tell us anything about the causal origins of such organization. Who said
that cultural tradition or individual experience should result in fitness-detrimental
behavior? The potential of phenocopy looms over all inferences from
function to cause. There is no doubt that culture can phenocopy, nature, since all
these behavior-guiding mechanisms are selection, processes according to functional
outcome criteria.
Pursuing all this, we end with the quaint conclusion that we can recognize a
biological adaptation only from a maladaptation under changed living conditions. I
have already considered how difficult it seems to prove maladaptiveness of behavior
in man-and, after all, biological (genetic) maladaptations too can be nicely
phenocopied by culture-traditional maladaptations.
Even if we grant that demonstration of universality- or model- conforming fitness-
adaptiveness of a behavioral trait gives us, if not convincing proof, at least plausible
indications of agenetic hangover from our animal past, would that be a strong,
predictive piece of behavioral theory? Or would it be one that sometimes fits the
data and at other times not-that is, a suggestion of low predictive value? I am afraid
the latter would be true. The funny thing about most alleged human behavioral
universals or adaptive dispositions seems to be that social scientists have little
difficulty in coming up with exceptions to the rule. We slide easily from universal
and preprogrammed to probable and inclined.
Now, if someone claims that aggressiveness is a human universal, and if this is to
mean that some aggressive behavior is highly probable in man under appropriate
circumstances, then that is no news. What is really interesting about this universal
propensity is what makes different people so differently aggressive. The human
nature argument states that something can occur. But what we want to know is when
it will occur and when not, and how its occurrence can be modified. And that is
surely less a matter of preprogrammed nature which sets the range of modifiability
than of specific education, individual experience, and previous success-all of which
determine specifically where the individual ends up within the total range.
The comparative method of Darwin and Lorenz is undoubtedly a powerful tool of
evolutionary investigation for tracing relations between organisms, if the causative
mechanisms behind the traits compared are principally the same and if we can
assume historical continuity between these traits. In other words, if we can regard
them as homologous. For many strictly genetically controlled traits in plants and
animals, the comparative method is a resounding success. It is no less so for some
purely cultural traditions such as those of concern to comparative linguistics. But
where there is a complex mix of causative influences which can replace (phenocopy)
each other, the method loses grip.
Another pitfall of the comparative method that one must carefully avoid is the
assumption that those traits that are common {"universal") to different systems are
somehow more fundamental in the sense of important than more recently acquired
or restricted characters. The opposite may very well be true if we talk about
behavioral adaptations. The universals may just as well be old-fashioned remnants
of little importance for the adaptation of a system to its particular environmental
niche, while specialties may make all the difference for the goodness of this fit and
for competitive success. The comparative method, almost by definition, tends to
emphasize generalities and to neglect specifics in precise reverse of their actual
adaptational importance.
This is not the place to go through the different human behavioral traits in order to
weigh the plausible evidence for more or less contribution of “biological nature." I
find it difficult to draw strongly nativist conclusions for those human behaviors that
involve the participation of our cognitive capacities. Cognitive processes are
involved at every step in the evolution of man as the animal with culture. The ability
to reify conceptual representations {beginning with the concept of self) and the
ability to value such reified representations motivationally in widely varying forms
and degrees according to one's cultural socialization, individual experience and
insights, are the most interesting behavioral characteristics of man. At the same time,
this capacity to create a world of facts and goals of one's own- derived from
experience but able to detach itself from outer reality to such a degree as to become
virtually a world of independent subjective reality of its own-seems most loosely
connected with the "biological nature" from which it undeniably originally derived.
When one tries to define the minimal set of plausible biological dispositions that
preform such a cognitive system sufficiently that its imprints can be found in human
behavior, one may find only very few of them. Primary among them must be a
disposition to define and to value a concept of personal identity and to guard its
integrity {Luckmann, 1979). There is the inevitable disposition to construct this
identity as one of asexual being-male or female, or even both, but never neuter.
Further, there may be a natural inclination always to organize the social surrounding
into three circles around the ego: one that can be labeled the kin group and often, but
not always, comprises the closest biological kin; a second that may be designated the
ingroup {the set of kin-groups that together make one's band, tribe or nation, one's
cultural allies) and beyond that the circle of strangers,
.
the foreigners, the out-group people. This three-circled, self-centered organization

would not make much sense, unless it were connected with specified behavioral
tendencies to treat kin-group and in-group and out-group people differently in
various subsistence-relevant aspects. What these aspects and tendencies are, and
how they are enacted in behavior, can be subject to great cultural diversity. But from
sociobiological theory we would always expect to find a polarity gradient from
assistance/cooperation to discrimination/aggression- from the inside to the outside of
the circles. This may be part of our natural endowment: to be nepotistic, to make a
difference between kin and non-kin, between group-members and group-strangers.
Incest avoidance and exogamy could be derived from or grafted upon such a
behavioral gradient. I am not sure whether this list of biological minima has to be
made very much longer. None of these surmised propensities will functionally
develop without environmental, especially social, interactions. They must not be
taken as causally determinant tracks, but as facilitated valleys of developmental
flow. Nature seems rather to suggest than to prescribe human behavioral
development.
I am not convinced that the concept of property (beyond the idea of something being
presently under one's control as a kind of extension of one's personal identity) must
be based on human nature as these other dispositions have been assumed to be. The
same holds for the notions of reciprocity and distributive justice, although it is clear
that no human culture can do without them. They could just as well be cultural
phenocopies (if the genetical model ever existed) or new inventions that put human
social groups on a different level of organization from that of social animals. This is,
they may be based on cognitive representations of time, of cause-effect, of rules of
conduct and of relations and obligations between oneself and group members.
Whether, on the motivational level, we have to assume anything beyond aversive
and rewarding feelings with respect to the basic physiological homeostasis of the
body (which by necessity for the baby includes social interactions) and the sexual
urge as natural emotional endowment, again seems doubtful to me. It seems possible
to derive all other emotional capacities and motivational goals from the action of
experience on these basic motivations. However, this may be an oversimplified
picture-it may be that more specific basic, unlearned tendencies have to be
postulated if we are to arrive at a parsimonious description of such behavioral facts
as our inclination to rest undis- turbed, to explore, to dominate, to care for offspring,
to communicate
feelings and ideas and skills, to teach and conversely to learn from and imitate other
group members.
What can all these deliberations mean for the natural bases of human legal behavior?
First, there seems to be very little evidence of specific biological determinants for
norm-controlled behavior except for the strong tendency to invent, to expect, to
conform to and enforce norms. Since norms are cognitive concepts they can only
derive from the interactionary social construction of the self and of interindividual
relations. Whether there are any other specific biological constraints on norm-related
behavior than the mere fact that nature provides us with a brain that is capable of
these cognitive constructions, can be disputed.
Second, one could expect to find some limits to entirely free malleability of norm-
related behavior in man where natural dispositions are most probably involved. Thus
the value given to one's own personal integrity, the value given to kin-relations
(nepotism), the value given to in-group versus out-group people and sex-related
propensities should be 'natural' friction areas for legal behavior. Whether property-
related or contract-related behavior, or behavior that involves domination, could be
expected to be under comparable biological influence, with similar consequences in
legal behavior, remains to be seen.
Beyond that, I see little that I can suggest with much confidence as an
evolutionary biologist's contribution to the understanding of legal behavior in man.
There are a number of more or less suggestive, plausible or frankly speculative ideas
that could be contributed from a general sociobiological perspective. However, I
doubt that these can marshall any more evidence in their favor than a number of
other, competing, theoretical explanations.
Legal behavior has to postulate freedom of behavioral choice. Freedom of
behavioral choice demands cognitive representation of the world and selective
operations (like thinking) on these representations. This limits the explanatory
possibilities of sociobiological, Darwinian modeling of extant human behavior
decisively, in spite of the fact that it is certainly called upon to explain how thinking
man could have evolved.
.
[101]
Biology and the moral paradoxes
RICHARD D. ALEXANDER
Museum of Zoology, Insect Division,
University of Michigan, Ann Arbor, MI 48I09, USA
Considerations from biology suggest (1) that human interests can be

generalized as reproductive, involving activities by individuals that tend to
promote the survival of their individualized sets of genes; (2) that ethical,
moral and legal questions arise out of conflicts of interest that exist because
of our history of genetic differences; (3) that human behavior probably
always involves egoistic tendencies and moral inconsistency; 4) that the
stages of moral development described by social scientists correspond to
the patterns of life effort discussed by biologists; (5) that the idealized moral
systems of philosophy and religion have been developed as models that are
promoted in others but not (or more than) in one's self; and (6) that what are
usually seen as the closest approaches to these idealized models are the
sources of our most severe problems because they involve between-group
competition and strife.
Ethical, moral and legal questions arise out of conflicts of interest among human
individuals and groups. Although this assertion seems to be accepted universally,
those who write on ethics, morality and law rarely emphasize it (Pound, 1941; Perry,
1954; Kelsen, 1957, represent the exceptions). Evidently, no student of human
behavior has undertaken the obvious challenge of explicitly identifying human
interests and quantifying their conflicts. Equity theory from psychology, net- work
and exchange theory from sociology and anthropology, and theories of interest from
law, political science and economics are partial attempts. These theories, however,
are all restricted to a superficial level, involving only reciprocal transfers of good or
beneficent acts. They neither identify the ultimate significance of goods and
beneficent acts, nor deal satisfactorily with the all-important class of interactions
that frustrated equity theorists have termed "deep and intimate" (Walster, Walster &
Berscheid, 1978). III other words, these theories provide no means of defining
human interests in a general or complete sense, therefore, no means of dealing
generally with the intensities and directions of individual efforts (see Alexander,
1979, and references therein).
A theory of interests is a theory of lifetimes: what they are about and
how their goals are achieved. A growing body of information and theory from
biology now provides a reasonable and testable answer: lifetimes have been molded
by natural selection to yield the greatest likelihood of survival of the individual's
genetic materials. This likelihood is maximized by success in reproduction, which
includes producing offspring, and assisting both descendant and non-descendant
relatives. The "deep and intimate" interactions causing difficulty to equity theorists
are actually those most directly involving reproduction-those occurring between
mates, potential mates and relatives. The currencies that mold the proximate
mechanisms of altruism in these interactions are genetic, not a matter of returned
goods or services, and this is the reason the payoffs have not been apparent to
investigators outside biology. Even the investments and returns of reciprocity
(exchange, equity) are ultimately comprehensible only in terms of their eventual
effects on the "deep and intimate" interactions of mates and relatives. Included are
wealth, status, good will and innumerable other items.
Biologists divide lifetimes into somatic and reproductive effort: use of calories and
taking of risks in (1) building the body or soma ( = amassing resources) and (2)
using the soma to reproduce ( = redistributing resources in the interests of one's own
genetic materials). Reproductive effort is in turn divisible into mating effort (on
behalf of gametes), parental effort (on behalf of offspring) and extraparental
nepotistic effort (on behalf of all relatives other than offspring). There are good
reasons for supposing that normal lifetimes include no other kind of effort
(Alexander, 1979).
I would regard the central paradoxes of moral philosophy to be those of (1) the
incompatibility of egoism and utilitarianism (seeking the greatest benefits to one's
self versus seeking some version of the greatest benefits to the greatest number) and
(2) the associated problem of duality in human nature. These paradoxes have been
developed and discussed in many forms, but always independently of the current
biological view of interests and lifetimes. I shall argue that they remain paradoxes
not because of some inherent irresolvability but because those concerned with them
have not adequately discussed the costs and benefits of either egoism or altruism.
Kalin (1968), for example, speaks of "winning" and "coming out on top," and
Frankena (1973, 1981) of getting "the best score," but neither describes the actual
currency involved. Some authors speak of survival, but it is unlikely that humans or
any other organism have evolved to survive (Alexander, 1979, 1982b ), and it is easy
to show that they all do
Gruterand Bohannan [103]
things that reduce their likelihood of survival. Essentially all authors consider
pleasure or happiness as reward {benefit) and pain and suffering as punishment
{cost), but none can explain in egoistic terms either the voluntary acceptance of pain
or the pleasure of helping others. Because the indisputable prevalence of egoistic
behavior eliminates any likelihood of a purely altruistic or utilitarian society, except
as an unattained {and as yet unexplained) pursuit or ideal, the problem of duality,
and of moral inconsistency as normal behavior, persists.
Biological theories of interests and lifetimes have the power to resolve these
paradoxes, at least in terms of the natural history of moral systems {the "why" of
behavior in respect to morality). Thus, an organism whose interests are in its own
genetic survival must first develop a soma {be a wholly or largely egoistic juvenile),
then reproduce {show the "altruism" of parenthood and nepotism) while maintaining
the soma by which it continues to reproduce {thus retaining egoistic tendencies
during adulthood). Direct and indirect reciprocity {Alexander, 1979) are distinctive
human overlays that add to the complexity, but they create no special problems.
They may be seen as indirect somatic or nepotistic efforts routed through pseudo- or
temporarily-altruistic investments in the welfare of others who are expected to
reciprocate with interest.
The stages of moral development in the individual, as interpreted by Kohlberg
{1981) and others, are remarkably supportive of this biological view. Represented,
first, is a purely somatic {selfish, "amoral") stage. This is followed by the
introduction of reciprocity through a system of rewards and punishment, usually by
the parent. The individual gradually foregoes immediate rewards in favor, I would
argue, of larger later ones {reciprocity). Acceptable rewards may be both
increasingly later and increasingly less direct {in the senses of involving diverse
currencies, and of coming from society at large rather than the person or persons
directly involved in the original social act). Eventually the individual also begins to
forego personal {somatic) rewards in favor of unreciprocated rewards to others
{nepotism). And he becomes increasingly able to assess the profitability of social
acts without outside help.
From these arguments about interests it follows that conflicts of interest arise out of
the history of genetic differences. This hypothesis is strongly supported by the
absence of observed conflicts among non- human individuals in clones and other
cases of long-standing genetic identity, and by the general diminution of altruism
with decreasing
relatedness within human societies the world over. It explains human individuality,
and bears upon powerful human issues, such as what Wallace called "the
impossibility, despite all the labor of God, Freud and the Devil, of one man fully
understanding another, or the loneliness of existence" which he regarded as "a pan-
human theme." It explains the unique cooperativeness of unrelated pairs pledged to
lifetime monogamy, and of genetically different workers in the colonies of social
wasps, bees, ants and termites. In both cases the genetically different individuals
involved share interests because they reproduce through the same third parties: the
offspring produced jointly by the monogamous pair and the siblings of worker
insects produced by their common mother. It is significant that Kohlberg's final
stage of moral development is that in which the individual has learned for himselj
how best to assess his personal costs and benefits in following (and using) whatever
social rules prevail.
Viewing humans and their moral behavior in terms of natural selection provides
stark and dramatic answers to some serious and very general questions: the
incompleteness of justice; the persistence of conflicts of interest; the failure of
idealized moral systems; and the absence of universal happiness and satisfaction.
Part of the answers lie in the relative nature of success in evolutionary terms:
In natural selection the likelihood of a genetic element persisting depends
entirely on its rate of change in frequency in relation to its alternatives;
changes in absolute numbers are irrelevant. Among the I attributes of .living
creatures, whatever can be shown to have resulted from the action of natural
selection may be expected to bear this same relationship to its alternatives.
Thus, we should not be surprised to discover that the behavioral striving of
individual humans during history has been explicitly formed in terms of
relative success in reproductive competition, that justice is necessarily
incomplete, that happiness is not easily made universal, and that ethical
questions continue to plague us, and can even become more severe when
everything else seems to be going well {Alexander, 1979: 240).
I stress that our interests are not individual because of genetic differences per se, or
current genetic differences, because such information has never been directly
available to humans. Relatives are known through circumstantial evidence, and only
recently have geneticists learned what the average relatedness actually is for
relatives whose learned assumptions about relatedness from genealogical
connections
.
and kinds of social interactions are nevertheless usually correct. The individualized
genetic constitutions of the successions of our ancestors caused natural selection to
save and mold proximate mechanisms whereby appropriate efforts could be
mounted by individuals in each successive generation to realize their separate and
individualized interests. We learn who our relatives and friends are, and how to treat
them; but our learning responses are themselves evolved, and often very specific and
channeled.
The hypothesis that conflicts of interest derive from the history of genetic
differences also generates new and sometimes startling questions: What are the
benefits of the group to the selfishly reproducing individual? Why does one kind of
ultrasocial group (eusocial insects) achieve its greatest numbers and unity (up to 22
million) as a single nuclear family in which one individual does all of the
reproduction while the other (humans) achieves its greatest numbers and unity (now
approaching one billion in China) by leveling the reproductive success and
opportunities of its members (through socially-imposed monogamy, graduated
income taxes, gradations of negative correlations of government support with family
size, restrictions on "free" enterprise etc)? How do these questions relate to the
morality of individuals and the idealized moral systems discussed by moral
philosophers?
The altruism of human nepotism and reciprocity is discriminative: Different
relatives, and relatives of different needs, are distinguished. Friends are treated
individually. As yet, no evidence of truly indiscriminate, species- or population-wide
altruism has been reported for any organism, and there is no undisputed evidence for
unlearned recognition of relatives in any species (Alexander, 1982a ). These facts
are crucial to understanding moral paradoxes and the rise of moral systems.
Indiscriminate altruism requires no special proximate mechanisms—no social
learning. I would venture that without genetic individuality, and the consequent
discriminative altruism in nepotism and reciprocity, social learning would have
remained simple, and human society as we know it could not have evolved. The
very concepts of ethical, moral and legal would be unknown.
To think of humans existing without conflicts of interest is to assume situations
involving or mimicking group selection, in away explicitly opposing the notion of
individuals striving to maximize their separate reproductive successes. It seems to
me that this is the ideal state of morality postulated by philosophers and social
scientists. If so,
perhaps biology gives us the reason for understanding interpretations such as that of
Perry (1954: 100).
Morality is like a cultivated field in the midst of the desert. It is a partial and
precarious conquest. Ground that is conquered has to be protected against
the resurgence of original divisive forces. The moralized life is never immune
against demoralization. At the same time that morality gains ground in one
direction it may lose ground in another. Changes in if; the natural and
historical environment and the development of man " himself are perpetually
introducing new factors and requiring amoral reorganization to embrace
them. In the last analysis all depends on the energy, perseverance, and
perpetual vigilance of the human person.
Numerous philosophers have suggested that morality, at least as expressed in the
behavior of individuals, is in fact only an ideal, or a pursuit, and not something that
is actually realized. This idea seems consistent with the approach from evolutionary
biology that I have been describing. Thus, it is common, if not universal, to regard
morality in the behavior of an individual as consisting of a kind of altruism that
yields the altruist less than he gives. In a utilitarian system (defining utilitarianism as
promoting the greatest good to the greatest number) morality would not always
require that complete and indiscriminate altruism cause individual losses. This
would not, for example, be the case when the interests of the group and the interests
of the individuals comprising the group are the same. Such a confluence of interests
would happen each time the group was threatened externally in such fashion that
complete cooperation by its members would be necessary to dissipate the threat, and
when failure of the group to dissipate the threat would more severely penalize any
remaining individuals than would the use of all the individuals' effort to
(successfully) support them {this is the true, but in these times of nuclear threats
forgotten, meaning of the term "national security"). In other circumstances, as when
some competitiveness has a likelihood of benefiting individuals in the group {i.e. the
individuals' interests are not all completely tied up in the survival of the group or its
success in dissipating some external threat), morality of an ideal sort would require
the kind of genetic altruism, unlikely in evolutionary terms, in which the altruist
truly gets back less than he gives. Of course, if an external threat came from another
group of humans, the definition of morality as indiscriminate altruism would again
be in jeopardy.
Reflecting on these circumstances, we see that if approaches to
morality are expressed consistently, and to the degree usually achieved in society,
because there is continual pressure to bring about a condition of morality, this
pressure is likely to be applied by each individual so as to cause his neighbors, if
possible, to be a little more moral than himself. To say it another way, it would be to
the advantage of each individual that other individuals in his society- especially
those not closely related to him-actually achieve the ideal of completely moral
behavior. Any ideally moral person would incidentally "help" every other person in
the society, however slightly, to achieve the goals that evolutionists believe have
driven evolution by natural selection, because he would hurt himself {a competitor)
by dispensing his beneficence indiscriminately. Accordingly, one might expect that
every individual in a society would gain from exerting at least a little effort toward
encouraging other individuals to be a little more moral {altruistic) than they
otherwise might have been. Among the many ways of furthering this aim is included
the setting up of an idealized model of morality and the encouragement of everyone
{else) to become like that. One way of promoting this outcome is to designate as
heroes {i.e. as appropriate targets for special rewards) those who most closely
approach the ideal moral condition. This line of reasoning predicts that sainthood
will be awarded to individuals who spend their lives on explicitly anti-reproductive
behavior. The prevalence among saints of asceticism, self-denial, isolation from
relatives, devotion to the welfare of strangers, and otherwise indiscriminate
tendencies to be altruistic supports this hypothesis. So does the fact that sainthood is
generally awarded {long) after the death of the awardee.
So we are provided with the general hypothesis that the concept of morality, and the
establishment of systems promoting ideal moralism, at least appear to have as their
aim the support of the goals of society as a whole. For this reason, within society,
each and every individual may be expected to promote in his associates tendencies
to be moral. Because of continuing possibilities of differential success within
groups, though, we can also understand that each and every individual may also be
expected to promote a slightly greater degree of " morality" {altruism) in his
neighbor than in himself. And we can understand why the idealized morality of the
philosophers is never a reality in society as a whole and occurs only as an accident, a
manipulation or in special circumstances.
The question may be raised, why anyone should be susceptible to
being manipulated unduly far in the direction of morality, given that we have been
subjected to such manipulations for so long? Why, in other words should moralizing
ever be effective?
I think there are at least four contributing factors. First, the degrees of morality that
are actually reproductively appropriate will vary dramatically as societies move
between periods of extreme danger and relative security, making it difficult to know
how to behave. When will a specified degree of failure to accede to exhortations to
be altruistic cost more than it yields, because of (1) failure of the group on which
one depends for success, or (2) responses within the group to one's failure to be
altruistic?
Second, individuals may be expected to take advantage of the dramatic shifts in
most profitable degrees of altruism to deceive others about costs or dangers so as to
induce in them unduly altruistic behavior. It is obvious that aspiring leaders often
use such deception to promote their own leadership, as an antidote to the supposed
threat and as a promoter of unity.
Third, we may expect that the individuals in society such as we have been describing
will evolve to deceive others about the degree of altruism they themselves are
exhibiting: Everyone will wish to appear more altruistic than he is. There are two
reasons: This appearance, if credible, is more likely to lead to direct social rewards
than its alternatives. It is also more likely to encourage others to be more altruistic. If
one's associates are altruistic, then he can afford to be more altruistic than if they
were not. We may expect everyone to be concerned that everyone else appear
altruistic so that people in general will feel comfortable with a higher degree of
altruism than would otherwise be the case.
Fourth, if kin recognition is learned (Alexander, 1979, 19820), mistakes are likely in
this context, and one may insinuate himself into the role of relative so as to receive
inappropriate nepotism, or even to pretend to be nepotistically altruistic so as to
receive the appropriate altruistic responses.
Playing upon the tendency of everyone to strive to appear more altruistic than one's
self, and using the other ploys just described, may produce a considerable amount of
successful social manipulation. These various factors seem to be the elements
necessary to produce and maintain what we commonly call moral systems, and
moral behavior in individuals. They represent the means of resolving the
philosophers' paradoxes with respect to morality, and for understanding why moral
systems have always fallen short of our ideals, and
why we establish and maintain such ideals. If accurate, these arguments may also
clarify the routes by which we can most closely approach what are seen as idealized
moral systems, and perhaps most confidently avert moral disasters.
The introduction of indirect reciprocity, whereby society as a whole or some large
part of it provides the reward for altruism and the punishment for selfishness,
simultaneously served both society and the individuals comprising it, and provided
the vehicle for socially manipulating individuals to levels and kinds of altruism
detrimental to them (or their reproductive success). It is somewhat paradoxical that
the tendencies and pressures in the direction of idealized moral systems should serve
everyone up to a point, but then be transformed by the same forces that molded them
into manipulations of the behavior of individuals that are explicitly against their
interests and in the interests of those ostensibly promoting everyone's interests by
promoting trends toward morality in the system.
The concept of a single just God for all people, however, it is believed to have
originated, implies social unity. I would regard this concept as one representation of
an idealized moral system arising out of religion; and it is just as difficult to follow
as those generated from moral philosophy. It is not trivial that the concept of a single
God for all people differs from that of a 'tribal' God looking out for the interests of
only one group or society. Adhering to this concept requires denial of practices like
slavery, caste systems, and other within-group discrimination. Despite its
prominence and use during times when groups are threatened externally, the concept
in some sense fails whenever such external threats involve ( or are) other groups of
people. This failure is, of course, denied by the invention of anti- Gods, or Devils,
and the ascription of others' motivations to their control. As a US Christian picketing
over the arrival of some Russians put it, 'I could love them if they were my enemies,
but they are the enemies of God!'
The concept of God also implies continuity of social unity—a long-lasting,
intergenerational social contract. If nepotism is our evolved function, then God (in
the sense of vox Populi, vox Dei) really can guarantee a reward 'in Heaven', or after
our individual deaths-or a kind of 'everlasting life' (for our genetic materials)-as a
reward for moral behavior during life. This guarantee is in the form of a renew- able
contract in reciprocity which occurs when those who remain after our death use our
own life of' morality' to judge our children (and other relatives who remain) as
suitable risks to continue receiving (and
giving, and receiving and giving, and receiving and giving the benefits of social
reciprocity. The guarantee actually exists because, t unless those in a position to
honor it do so for us, the same possibility r will not exist for them. The ceremonies
associated with death, and the reverence given to the dead, are surely, in part,
ritually related to this I guarantee.
If morality tends to mimic the effects of group selection, if moralizing seeks to
promote this mimicry, and if tendencies for people to be altruistic are self-
reinforcing within societies, then it is not remarkable that sincere, knowledgeable
and well-meaning people sometimes resent the arguments that natural selection is
not powerful at group levels, and that humans, as individuals, have evolved to be
interested in furthering their own reproduction. Such persons may well believe ( or
sense) that publicizing or stressing such arguments, even if they are correct, will
diminish altruism and morality by providing an anti- moral model. The indications
that humans have regarded moral models as extremely important in achieving
societal goals cause such a belief to be completely understandable. Nevertheless, this
attitude runs counter to the goal of diminishing human problems through improving
self-knowledge.
Our truly serious problems of morality and law stem, not from the behavior of
individuals, but from the behavior of groups that may show most dramatically
within themselves the indiscriminate altruism that represents approaches to
the idealized morality of philosophy and religion. Indeed, loyalty and
patriotism are revered as the highest forms of morality and virtue within
groups. But this same level and kind of within-group morality has also
created our most devastating problems-those involving intergroup conflict-
that we must somehow supercede. What we seek, when we think of world
peace and world law, has no precedent in the history of life, not to say that of
humankind. There seems to be no evidence that humans or any other
organism have achieved the species-wide indiscriminate altruism
represented in the idealized moral models of philosophy and religion.
I offer only one conclusion in this brief and perhaps unsettling essay: that, in the
effort to solve humanity's most profound problems, there is potentially great value in
adding a perspective from modern evolutionary biology to those developing out of
philosophy, the social sciences, religion, history and the humanities. This biological
perspective must be added, not as an argument for determinism, but precisely to the
contrary, as a possible way to greater freedom, deriving from greater knowledge of
the cause-effect patterns that underlie our
history and our nature. Some of my colleagues in biology, and many people outside
biology, deny that humans can be understood in biological terms. Others cling to the
notion that we evolved by an innocuous (and hypothetical) form of group selection
and can somehow return to it. Or they argue that if this is not the case, we should
deny the truth and pretend ourselves toward world peace and human justice; or that
it is better to be ignorant with an idealized moral model before us than to know
about an immoral history. I believe that people who think in these fashions are
wrong. Worse, because of the enormity of the problems that face us, I regard
approaches that deny biology, and sometimes deny reality, as potentially deadly.
Essentially, everyone thinks of himself as well-meaning, but from my viewpoint a
society of well-meaning people who understand themselves and their history very
well is a better milieu than a society of well-meaning who do not.
The neural and chemical basis of reward:

new discoveries and theories in brain
control of feeding, mating, aggression,
self-stimulation and self -injection
BARTLEY G. HOEBEL
Department of Psychology. Princeton
University. Princeton, NJ 08540, USA
Discovery of endogenous opiates raises the possibility that they serve

as a reward for behavior, including law-abiding behavior. This paper
begins with a discussion of homeostatic behavior, drinking, feeding, and
then social behavior, mating and aggression. It is possible that peptide
fragments of longer protein chains serve as "drive peptides" in the sense
that each is evolutionarily adapted to serve a battery of body and brain
function. Dual function, gut-brain peptides have recently been
discovered for digestion and satiety, blood volume and thirst, and
ovulation and sexuality. Aggression-control pep tides are a logical
possibility.
Studies are described in which feeding, mating and killing responses are
induced or blocked by electrical or chemical stimulation of the
hypothalamus in the rat. Animals will also work to stimulate their own
brain electrically or with the catecholaminergic drug, amphetamine, or

with the opiates morphine and enkephalin. These reward systems could
conceivably reward not only homeostatic behavior and self-centered
social behavior, but also altruistic behavior. A hypothetical model of
brain function is proposed in which endogenous catecholamine and/ or
opiate rewards are released by behavior that matches a memory or
"rule" of prior behavior. Thus complying with internalized laws or
expectations is in itself rewarding. Altruism as such is not necessarily
innately rewarding, but matching performance to expectation probably
is. Behavioristic, physiological psychology can explore the role of
opiates in legal expectations; behavioristic cultural anthropology can
explore the legal expectations that people depend on for their reward. In
summary, a theory is presented in which innately coded peptides prime
physiological and behavioral patterns for feeding, mating and
aggression; catechola mines modulate arousal and activation; and
opiates reward successful compliance with rules, most of which are
learned, some of which are laws.
INTRODUCTION
Gruter (1979) and Danielli (1980) have suggested that endorphins in the brain could
serve as opiate rewards for altruism and law-abiding behavior. This is an interesting
idea. Endogenous opiates could provide all kinds of satisfaction including the
satisfaction of eating, drinking, mating, child rearing, aggression, defense and other
forms of individual and social responsibility. Neuroscientists and physiological
psychologists have an unusually clean and elegant way to test these theories. If the
opiate antagonist, naloxone, blocks the opiate- supported behavior, then the
behavior is mediated by endogenous opiates. For example, naloxone blocks
morphine analgesia, acupuncture analgesia, morphine euphoria and morphine-
induced feeding. Therefore parts of the systems for pain, pleasure and hunger
motivation involve opiate receptors. Similarly, endogenous opiates have been
shown to playa role in pain suppression by placebos and pain anticipation after a
warning (Fanselow, 1979); thus opiate release can be conditioned. Reacting to a
warning could be taken as a model of law-abiding behavior. Even more to the point
of this conference, certain social behavior patterns are innately reinforced by
opiates. Chicks display cries of distress if they wander off from their mother and
siblings. The same reaction occurs if they are removed to a "jail" or given naloxone.
The reaction of distress and anxiety is unmistakable at the appropriate age. Obeying
innate law to stay by mommy is
rewarded by opiates; breaking nature's law is punished by opiate withdrawal
symptoms. If someone develops a clear, simple model of altruism in a laboratory
animal such as the rat, then I could explore ~ the role of endogenous opiates in
altruism as well. At present I can tell r you more about opiates in feeding behavior .
Opiates were discovered in the brain when Hughes et aL (1975) recognized a
morphine-like compound within the structure of a long '\ protein isolated from the
pituitary gland. It was named endorphin (short for "endogenous morphine"). We now
know the parent compound is split by enzymes to produce a variety of important
peptides, including endorphin, that control both physiological processes and pain.
Clearly an animal's genetic codes for producing the long parent compound and the
enzymes that split it up will influence the animal's physiology, its reaction to pain
and perhaps certain behavior patterns.
There are strains of mice and rats that have agenetic tendency to be obese. According
to Mendelian laws a predictable fraction of the offspring will be fat. It was
discovered that the fat ones had more beta-endorphin in their pituitaries than the lean
litter mates (Margules, 1979). If it could be shown that endorphin affects feeding
behavior, then we would have a clear link between genetics, brain peptides, and a
life-long motivated behavior pattern. Such a jump from genetics to motivation is a
primary concern of this conference.
MOTIVATION
Investigators in several laboratories tried injecting morphine and beta-endorphin
into a region of the hypothalamus, just above the pituitary where norepinephrine
(i.e. noradrenalin) is known to induce feeding. Morphine or beta-endorphin induced
feeding after some delay. More- over the effect was reversed by naloxone which
proved that there are opiate receptors for feeding in that brain region (Sanger, 1981).
Meanwhile, relatively short opiate peptides had been discovered inside neurons of
the brain and spinal cord. Made up of a string of amino acids, these potent little
peptides are cracked off of larger proteins made by the genetic apparatus in the
nerve cell bodies. These short, opiate peptides, called enkephalins, are released
directly into the nerve synapses instead of into the blood circulation. Thus they are
more like neurotransmitters than hormones. In the spinal cord enkephalin neurons
can close the gate on incoming pain signals. Apparently specific local stimuli such
as pricks to acupuncture haku
points in the skin can release enkephalin in various segments of the spinal cord.
There the en kephalin may suppress the painful stimulus and even pain from other
sources. Generalized stimuli, such as stress and exertion, on the other hand, can
release endorphin in the general r circulation. Given that opiate receptors for feeding
had been found in the hypothalamus the next step was to try injecting en kephalin to
see if feeding was a generalized endorphin effect or perhaps a more specific
enkephalin-mediated effect.
We found that enkephalin caused rats to eat just as norepinephrine and beta-
endorphin had done. The effect was partially reversed by naloxone. One thing is
strange. The enkephalin-like compound we used acted very slowly. Norepinephrine
injected in the hypothalamus can induce eating in a minute; the enkephalin analogue
took half an hour. Norepinephrine seems suited to controlling meal size; enkephalin
is slower and may have more to do with the urge to eat, rhythms of eating or other
slow cycles.
We eat because food tastes good, and because we feel good after a meal. There may
be two separable systems for these two pleasurable aspects of eating. Belluzzi &
Stein (1977) proposed that brain opiates give the satisfaction that comes after
successful consummatory behavior. According to this view, opiates may be the
chemical basis of pleasure, the goal of drive reduction. Our task now is to figure out
which neurochemicals and which brain systems cause "drive induction" such as
appetite for food, and which give "drive reduction" such as satiation.
Another question is why we stop eating at all. Something must stop us from eating
all day so we can do something else such as make love, play with the children,
compete, create or what have you. Again, glandular pep tides have been discovered
first in the circulation and then in brain neurons. Peptides secreted by glands in the
gut are thought to act as satiety signals. They have been discovered not only in the
general circulation, but also within neurons of the brain. Neuroscientists surmise that
these pep tides might be released at synapses to act as neurotransmitters, or they may
be released to modulate the effects of other faster-acting neurotransmitters. It looks
as if the nervous system, which derives from ectoderm, may have incorporated these
chemicals into neurons to control feeding, and the endodermal organs may have
adapted the same chemicals as signals about fuel availability and digestion. Peptides
from the gut could conceivably diffuse out of the bloodstream into the brain
synapses and have modulatory effects
concordant with the brain's neural transmitter. The end result would be a neural
circuit for feeding that could be biased by the same peptide from either source,
either the brain or the body. There may be a .general principle here. It is as if a
given peptide is used by the brain and the body to control the same overall need in
all its aspects from modulating organs to modulating behavior. This hypothesis
remains to be proven for feeding systems. Studies of drinking are further along.
Drinking is more essential than eating. People on self-imposed hunger fasts have
been known to live for 40 days and 40 nights, but the punishment for not drinking
is swifter. Without the right amount of water, chemical reactions are disrupted
because the fluid concentrations are wrong; circulation fails for lack of blood
volume; the body overheats for lack of coolant, and food sticks in the craw.
Regulation of water balance depends on two stimuli, osmotic concentration of the
body fluid as signaled mainly by osmoreceptors in the hypothalamus, and by blood
volume as measured by receptors in the kidney. In the first case of low osmotic
concentration many things happen at once. Hypothalamic receptors send a
hormone (ADH) telling the kidneys to reabsorb water that would otherwise
become urine; the same hormone contracts blood vessels (Guillemin, 1980). A
neural signal peps up the heart; and a message goes out to the rest of the brain to
prime reflexes for drinking. The triple result is a yellow, concentrated urine,
elevated blood pressure and "thirst." This is all a reaction to too much salt relative
to water in the body fluids. The concentration must be close to that of the sea water
in which we evolved. The points to notice are the physiological redundancy, the
chemical feedback loops, the neural orchestration and the role of behavior in
homeostasis.
Let us look at it again, this time focusing on blood volume instead of concentration.
Suppose an animal has been in a fight and is bleeding. The kidney sends a
hormonal message (angiotensin) that does the same three things as ADH. Water is
reabsorbed, blood vessels constrict and the sensory-motor system is "primed" to
drink. In most cases of "priming" the threshold for detection is not lowered, but
instead the sensitivity to particular stimuli is raised; more sensory neurons fire and
the motor output is more vigorous. In addition, there may be a strong sensation. In
the present example there is an awful thirst.
The motivation called "thirst" translates into something measurable in three ways.
People can say it in words; this is the verbal variable. Animals prove it by
overcoming obstacles between themselves and water. For example, rats run faster,
pull more weight, or
press a water pump more frequently. This is the "drive," "motivation," or

performance variable. Third, animals repeat the performance that paid off. This is
the operational definition of "positive reinforcement," the reward variable."
These same principles seen in feeding and drinking also seem to hold true for the
complex social behavior known as mating. Again, a newly discovered peptide seems
to choreograph the beautiful interplay of brain and behavior. Actually I should say
brains and behaviors, because mating is an emergent property of the back and forth
interaction of stimuli from two animals acting as one system.
A peptide, leutinizing-hormone-releasing-hormone (LHRH) is produced in cells of
the hypothalamus and released to trigger leutinizing hormone (LH) from the
pituitary (Adler, 1981). LH is the hormone that causes development of the egg,
ovulation and copulation. It is released after estrogen causes courting and before
progesterone causes embryonic development and nesting. In each case a pituitary
sex hormone adjusts the physiology of the sex organs for a particular phase of
reproduction. Everyone knows that the primary sex organs develop during this
process; vocal chords and hair or feathers take on signal functions. It is less well
known that brain areas for receiving sex signals also grow, and the erogenous zones
change their sensitivity. For example, estrogen actually increases the size of the
pubic area that will generate sensory nerve impulses when caressed. Estrogen also
potentiates the spinal output for copulatory postures. Of course the sex equipment,
sex signals, sexual sensitivity and mating reflexes are not enough. Complex
behavioral output is also primed. LHRH has just been discovered in long neurons
reaching out from the hypothalamus to all the brain nuclei involved in sexual
behavior patterns. LHRH is probably an overall organizer of orgasmic sex. It
commands the master gland to secrete the LH surge, and it stimulates a million
synaptic contacts in sensory and motor relays for sexuality. I would not be surprised
if it someday proved to modulate the rewarding incentives we feel during
intercourse and the satisfaction of orgasm. LHRH, in both men and women, has all
the earmarks of a "drive peptide" (Olds, 1977) for mating and reproduction.
Aggression plays a major role in survival. It starts when a pup must struggle for a
nipple, and the behavior grows into the struggle for food and living territory. The
link between aggression and feeding is unmistakable in carnivores, but there has
always been some controversy as to whether feeding and aggression have clearly
separable brain mechanisms, or whether they are just two manifestations of the
same basic system. We have done studies in which brain stimulation with electricity
or chemicals produced mouse killing in totally naïve rats that had never killed a
mouse before {Smith, King & Hoebel, 1970). They had never even seen a mouse
killed until we stimulated their hypothalamus, and then they proceeded to commit
muricide. This suggests to me that some types of aggression have separate neural
components which are innate. Prior learning also plays a major role {Cools, 1982).
The murderous behavior may never be performed unless triggered by appropriate
environmental stimuli or brain stimulation. In one study rats worked for mice to kill
even though they were never allowed to eat the mice. Such killing satisfies the
laboratory definition of a reward.
There are many stimuli for aggression. They are so varied that they define many
types of aggression; for example, prey aggression, male-male aggression, aggression
for infant protection, and aggression that is triggered by pain {Valzelli & Morgese,
1981). The link between sex hormones and aggression is very clear in many species;
thus one cannot study aggression without being aware of the underlying hormonal
state. A "drive peptide" for aggression has not been found yet, although we can
predict it on the basis of the foregoing discussions of feeding, thirst and mating.
If a specific aggression peptide were discovered it would finally prove that

aggression is a distinct behavioral category, extricable from feeding and mating.
Such a peptide should have the same general properties as angiotensin for inducing
thirst, cholecystokinin for satiety and LHRH for mating. It should have an
organizing effect on several physiological and behavioral systems to favor an
aggressive response at the expense of feeding, drinking or mating responses.
In the last five years scientists have learned to intervene in these motivational
processes. For example, the new drug Captopril blocks the angiotensin signal and
thereby lowers blood pressure. Now people can lower their blood pressure by
chemical intervention to prevent heart attacks. Our laboratory is looking for a drug
that could block the physiological and behavioral manifestations of hunger. Then
people who eat and have a body weight suitable for a cold climate or a famine, could
have a normal body weight instead. Even though feeding seems to be organized in
much the same way as drinking, it is more complicated because there are more
stimuli. Taste, stomach fullness, intestinal contents, liver store of glycogen, blood
sugar, circulating fat, insulin, and even the outdoor temperature can all control food
intake. For thirst there were two important stimuli, blood concentration and
volume, for hunger there may be a dozen. Mating is more fascinating, for here there
are crucial stimuli that must be properly sequenced between two individuals.
Nevertheless, the newly discovered peptides provide the missing link between the
classic hormones and behavior. Manipulating the enzymes that make or break these
peptides is one of the new hopes of the drug industry. Another is manipulating the
genes that make the enzymes.
In summary, any given motivated behavior seems to consist of subsystems which
can be brought into action over the course of a few hours by hormones, driven in
minutes by peptides, and triggered in a split second by the appropriately coded
sensory signal or direct electrical stimulation. The next step in our analysis is to go
beyond motivation, beyond "reflexes," "priming," "drives" and "sign stimuli," all of
which create behavior tendencies. These propensities to behave in certain ways are
the essence of sociobiology, but to understand law we have to go further and study
brain systems that are necessary for reward and punishment.
AROUSAL
Various motivation systems have to compete with each other for access to the
animal's attention and response mechanisms. Superimposed on all this is a flip-flop
mechanism in which the hierarchy of priorities can be switched by the
parasympathetic and sympathetic nervous system. Under the calm dominance of the
parasympathetic system, breathing comes before temperature regulation, which in
turn takes precedence over drinking which comes before eating, which comes
before mating. Under the emergency conditions that bring the sympathetic system
into action, fight or flight comes first, sex may dominate eating, feeding may erupt
with regard to energy level and a drink of water is the last thing on the animal's
mind. The animal may even disregard its temperature, or for a minute forget to
breathe.
Adrenaline and noradrenaline are the physiological signals from the adrenal gland
that prime our bodies for emergency action. In the brain, the same chemicals are
released from neurons that modulate all the motivation systems discussed above.
The well-known reticular activating system in the hindbrain contains a system of
widely ramifying neurons that modulate most of the brain. These neuro-hormones
arouse us to pay attention. They also inhibit processes that can wait. For example,
some of the neural pathways for inhibition of feeding have been discovered (Hoebel
& Leibowitz, 1981). Interestingly,
they inhibit both the hypothalamic systems for eating and for not eating. It is as if
they were evolved to control meal quantity and quality in the parasympathetic
relaxed state, and to inhibit the whole works during emergencies. Apparently
specific neurons inhibit food intake after a meal when the input coming from the
gut signals a full stomach or full intestines. Other adrenergic neurons may inhibit
mating after copulation. Note that the same adrenergic synapses would inhibit both
feeding and mating in an emergency when adrenaline from the adrenal gland pours
into the bloodstream and diffuses into these synapses. This theory that adrenaline
leaking into the hypothalamus blocks feeding and mating is unproven, but it makes
sense.
It is curious that a relaxed state is usually necessary for eating or courting; whereas
ejaculation involves a switch to the sympathetic aroused state. Thus feeding and
courting occur in a common physiological state, and ejaculation shares a state with
some kinds of aggression. Animals, even humans, may display some confusion in
their response to the excited state. If a person is made excited with a shot of
adrenaline in the absence of any natural internal code to guide their behavior, then
they base their response largely on environmental cues. They may be either morose
or joyful, murderous or ecstatic depending on the social context. This is one way in
which excited animals can be entrained to exhibit a socially cohesive response. An
emergency, such as an earthquake, will set up the sympathetic state in everyone.
The individuals who see the problem, or think they do, will tip the first domino that
pushes everyone into the same emotion. Thus, in emergencies, feeding and courting
will stop. Whether the person then has an orgasm, commits a murder, laughs or
cries depends on the internal factors such as the pep tides that set the stage for
appropriate behavior. In the absence of such preparation, environmental stimuli,
real or imagined, play a bigger role. Environmental stimuli include the law. Internal
factors have a "law" of their own. The brain somehow synthesizes the two and
makes active choices.
ACTIVATION
After motivation and arousal the next step is activation. The story goes that bed-
ridden Parkinson patients could only activate themselves if there was afire in the
hospital. Such a life and death emergency could activate them to run out to the
sidewalk where they would collapse again. The greatest breakthrough in modern
psychopharmacology
.
was the discovery that Parkinson Disease patients needed I-DOP A to make the
neurotransmitter, dopamine. The dopaminergic system has its neural cell bodies in
the midbrain. The nerve axons project up through the hypothalamus to the forebrain.
Part of this dopamine system is absolutely necessary for initiating movement.
Dopamine gives us the willingness to work {Hoebel & Novin, 1982). Without it, we
only sit and dream. "
IMAGINATION
There is a very primitive set of serotonin-containing neurons in the hindbrain that
projects up, down and all around the brain like the adrenergic arousal type discussed
earlier. It has been possible in the last few years to record from these big, old cells in
awake cats. The faster they fire, the more alert, active and tense the cat becomes.
During sleep they gradually slow down. When they stop firing, postural muscles
relax, the head drops and dreams start. Imagination runs wild. Perhaps that is why
some discoveries are made during sleep. Part of this serotonin system clearly inhibits
imagery when it is active. It seems to ready the animal for concentrating on a single
problem and a quick motor response. When this readiness is missing the animal can
either lose muscle tone and dream, or maintain muscle tone and hallucinate. Normal,
everyday creativity and imagination probably require an in-between state of medium
tone and free flowing imagery. The hallucinating schizophrenic overdoes it. The LSD
freak loves it. The peyote cult worships with it. In this unfocussed state the stream of
imagery can be influenced by either internal or external stimuli. Alarm bells remind
us of phone bells, rain reminds of urine, crickets can sound like railroad trains and
eagles become gods. Freud seemed to think that dreams got him closer to the laws of
sociobiology unfettered by the lessons of later life. I know of no hard evidence that
dreams are a representation of the innate or childlike any more than a representation
of socialization and culture. However, it is fairly clear that when serotonin cells shut
up, suppressed ideas can speak up.
In sum, the animal, be it cat or person, seems to have its thoughts "loosely coupled"
or disinhibited when serotonin turns off. We do not know the functional significance
of this. Is it time for regeneration of chemicals for arousal, activation or reward? Is
this a state for new learning? A time for novel associations? Or just an
epiphenomenon of the age-old need to lie low when our defenses are weakest?
ARTIFICIAL STIMULATION OF INNATE BEHAVIOR

The sociobiologists would have us believe that there are genetically
preprogrammed properties lurking in the brain. If so, it is conceivable that one
could use electrical stimulation to induce behavior patterns that the animal never
learned. I would not make such a bold statement if we had not already done it.
Brugger and Hess long ago showed that hypothalamic stimulation in cats could
induce voracious feeding (Hess, 1957). Flynn (Adams, Bandler, Sheard & Siegel,
1981) showed that cats would kill rats during hypothalamic stimulation and that the
stimulation sensitized the cat's snout and enlarged the area that would trigger a
snapping response. As mentioned earlier, we found that hypothalamic stimulation
could induce mouse killing, "muricide," in rats that had never even seen a mouse
killed. Chemicals injected into the hypothalamus could do the same thing, and other
drugs to block the endogenous neurochemicals could block mouse-killing (Smith,
King & Hoebel, 1970). Apparently non-killers had the latent propensity to become
killers, and muricidal rats could be inhibited with drugs. Incidently, the tendency to
kill varies from strain to strain, and it can be modified by crowding, food
competition and other environmental variables (Karli, 1982).
Even though brain stimulation can induce any of several behavior patterns, its main
effect seems to be general arousal and activation. The precise behavior which
emerges depends a great deal on the physiological state of the animal (Hoebel,
1976, 1979) and on the environmental stimuli (Valenstein, 1973 ). Stimulation-
bound feeding can be blocked by a full stomach or a lack of edible objects in the
cage. In such cases the stimulated animal will search around and then probably
drink water or copulate or shred wood, or hoard food pellets, or jog in a running
wheel, depending on what is available.
We have already discussed motivation in terms of innate and learned shifts in an
animal's hormonal, peptide and catecholamine chemistry. These shifts, we saw,
biased neural development and neural activation. This approach is relatively new
because it depends on modern neuroscientific procedures to measure the relevant
chemicals in the relevant parts of the brain. The older approach was to define
motivation entirely in terms of stimulus deprivation or behavior output. Behavioral
biologists such as Dethier (1970) thought that directed activity, such as a fly flying
up wind, was sufficient to apply the label "motivated." Physiological psychologists
such as Teitelbaum
(1967) insisted that learning of an arbitrarily chosen behavior (i.e., operant

behavior) was the key to a successful definition. The two sides rested their cases
and the jury has been out ever since. Is drive directional activity where the direction
can be innate? Or is it goal- directed activity where the response is dictated by the
environment? Olds (1977) was one of the first to attribute drives to peptides,
essentially a chemical definition of drive. The latest breakthrough in neuroscience
has been the discovery of peptides inside of catecholamine neurons. Can it be a
coincidence that a gut peptide has been discovered inside of nerves of the dopamine
activation system? We are now working on the possibility that parts of the
dopamme system are coded for various behaviors such as feeding. They may be
coded not only in their anatomical arrangement, but also in their dual-transmitter
system. One transmitter could be for activation and one for a particular drive or
motive. In this way a circuit for feeding, for example, could be inhibited by either
the neural input that releases CCK, or by chemicals in the circulation that influence
CCK receptors. This is hypothetical, but it goes a long way toward explaining how
an activated animal can be biased towards eating, or mating or aggressing. It also
suggests that the genetic code for manufacturing various drive- controlling
chemicals is in some sense the animal's code of behavior. Time and research will
tell whether the chemical code is detailed enough to be in any way analogous to a
legal or ethical code. From what we are discovering about feeding, I would predict
that pep tides lay down the detailed rules of motivation. We have seen that there are
two peptides for different types of thirst, and peptides for satiety. We have hints
that there may be different peptides for different types of hunger, specifically, salt
hunger, sugar hunger, fat hunger and protein hunger. We also know there are
different hormones promoting courting, copulation, nesting and nursing with
peptides for at least some of them. Therefore logic suggests, although the evidence
is missing that there will be different chemicals promoting each of the dozen
different forms of aggression. However, there is no evidence whatever to suggest
that peptides could innately code any law as detailed as incest taboo, nor all the
rules of fair fighting.
Let us suppose that an animal is primed for action by the motivation system,
aroused, willing to work and awake enough to deal with reality, the next step for the
animal is to learn from experience.
LEARNING
In classical conditioning one stimulus is substituted for another; in instrumental
conditioning one response leads to another. In classical conditioning the
experimenter more or less arbitrarily picks any neutral stimulus which, with
training, takes on the power to elicit a relatively fixed response. It is amusing from
a sociopolitical point of view that Russia seems to place particular stock in this
approach. America has championed instrumental conditioning which is response
training instead of stimulus training. The response to be learned can be picked by
the trainer within limits set by the animal's response capabilities and its level of
motivation, arousal, activation and rationality.
Research suggests the short-term "reward" for learning is partly catecholaminergic,

dopamine for the incentive to work (Mogenson & Yim, 1980) and norepinephrine
for stamping in a memory of what pays off ( Belluzzi & Stein, 1977). Long term
satisfaction is probably an opiate-like state involving the brain's own opiates. These
chemicals have been shown to be necessary for each of the functions named. They
are not sufficient by themselves because many neural systems interact to produce
learned behavior. As necessary chemicals, we can use them to talk about the
functions they serve. Something subtle, but important has been going on in this
paper. I have been trying to tell you about brain chemistry as we know it and then
to attach concepts that seem to fit the chemical function. The object is not to get
trapped into looking for chemicals to fit psychological or ethological concepts.
Once we find important chemicals and circuits we can define the concepts to fit.
Some of our classic concepts may not be discoverable at the chemical or simple
circuit level; they may be emergent properties of large systems. In a sense we are
trying to find the minimum neurochemical standards for to-be-named components
of operant behavior. These neurochemical and neuroanatomical functions will be
the internal rules of willful behavior; the rules which legislate how we learn. This
information is not necessary to lawyers, because to control behavior one needs to
know laws of stimulus and response, not laws of brain function. This information
is, however, very interesting to those who want to know how laws emerge from
brain.
How does "reward," i.e. positive reinforcement, emerge from the chaining together
of many sensory-motor reflexes in the motivated, aroused, activated animals?
Teitelbaum (1981) has recently been struck by the fact that seemingly complicated
acts like walking over to a
lever, orienting, pressing it and eating the food reward are all made up of many tiny
actions each triggered by an environmental stimulus. The floor triggers an action in
the foot; that foot triggers the other; the smell turns the head; the turning head turns
the eyes, the eyes the whiskers, the whiskers the mouth, the mouth the tongue, the
jaw the swallow, and all this proceeds in a beautifully choreographed sequence of
reflexes leading to food in the stomach. If any step is interrupted the chain may be
broken. The willful act when seen under a behavioral microscope is "mere
reflexes"; each spontaneous voluntary behavior had its source. Just as a ballet which
exhibits the pinnacle of athletic control and deep emotions can be reduced to an
orchestra score and a motion score, so also any skillful act is seen as a program of
linked reflexes. What we need to understand is the energy source that welds the
links in the chain. The simplest view is that each link, once forged, leads to another,
and no part of the brain needs to know the whole pattern. The common sense view,
on the other hand, is that some part of the brain has a goal in mind. Fortunately we
have neurochemicals and neural circuits necessary for both. Both link-by- link,
action-by-action, catecholaminergic reward that could weld operant behavior, and
the grand finale of opiate intoxication that could signal success in hedonistic terms.
Let us assume that the homeostasic mechanisms we spoke of at the beginning

utilize hormones and pep tides for specific drives as described, and that these
chemically-defined drives do two things. They increase the available opiate for
feeding, for mating, for aggression, or whatever, and they also open the gates for
appropriate classes of reflexes which are triggered in proper order by the natural
sequence of stimuli in the environment. When the environment changes, then the
behavior will change almost randomly until the reward system links one new
successful act to the next. The program of reinforced synapses which led nerve
impulses from one reflex to the next will then lead the animal through the same
routine again and again. Each time, I suppose, opiate is released. It must last for
minutes, hours, or even days. Each biteful of food, each genital stimulus, each prey
killed, hypothetically releases more opiate until tolerance builds up or an anti-opiate
(opiate receptor blocker) takes effect. The behavior will stop when any of the
necessary chemical substrates shut down or are inhibited. A drive peptide like
angiotensin may diminish and thus fail
to cause thirst. A satiety peptide like CCK may inhibit feeding, adrenergic arousal
may fail and the animal gets lazy. When opiate tolerance sets in it is time to try a
new behavior with which reward is still accessible. Heaven help the animal that can
find no reward sufficient to satisfy one of these basic systems. Then a state like
opiate withdrawal sets in. These may be the animals that become sick with hunger,
sick with grief, dying of thirst, and as for aggression the urge to beat someone at
something is all too well known. If these are really withdrawal states, then heroin or
morphine should cure them. The proper experiments have not been done, but opiate
sales lend some credence to the idea.
By putting electrodes or hollow cannulas into the brain we can short-cut some of
these processes. We can artificially drive the animal to eat, copulate or kill. We can
also reward almost any sequence of behavior electrically or chemically.
All mammals, and some of our more distant relatives too, can learn to stimulate their
own brains electrically. Rats, cats, dogs, dolphins, monkeys and humans all appear
to delight in pressing a switch to turn on a stimulator that is connected to an
electrode in an appropriate part of the brain. We use rats to study self-stimulation in
our laboratory. They respond at a rate of 3000 times per hour. This well-known
phenomenon was discovered by Olds & Milner in 1954. In the next 20 years it was
shown that the rewards of self-stimulation are similar to the natural rewards of
feeding and mating (Hoebel, 1976, 1979).
Recently, rats have been trained to stimulate their own brains chemically instead of,
electrically. This will allow us to break the chemical code for brain reward. As you
could predict, the chemicals that animals will self-inject into their brain are the
drugs that people abuse. Rats will press a lever to self-inject morphine or
amphetamine (Hoebel & Novin, 1982) through hollow needles implanted in care-
fully chosen brain regions. Soon we will be able to describe the neural pathways and
the neurochemicals that are primarily responsible for generating rewards.
Skinner (1938) defined a reinforcing stimulus as one that changes the frequency of a
foregoing response. Examples are palatable food, sexy stimuli and painful stimuli. A
given response followed by a reinforcing stimulus usually causes the stimulus.
Reaching into afire causes pain. Taking morphine causes the pain to go away. Thus
animals learn to be careful of fire and learn to use morphine. Similarly, some
[126] LAW, BIOLOGY ANDCULTURE
responses are socially painful and some are socially rewarding; thus animals adapt
to social usages. E. A. Hoebel (1954) has concluded that one of the four main
functions of law is "to redefine relations between individuals and groups as the
conditions of life change. It is to maintain adaptability." Thus law fosters learning.
There is one sure-fire way to tell whether or not a stimulus is a reinforcer. If you, the
experimenter, can arbitrarily pick a response and have it learned merely by
following it with the stimulus, then the stimulus satisfies the definition of a
reinforcer. In other words, if the animal adapts its behavior, if it learns, then we have
reinforcement. Similarly if a pair of animals, or a group, alters its behavior then we
may find some cause and effect relation between the group behavior and a stimulus
that reinforces the behavior. According to E.A. Hoebel's analysis we are looking for
law. I will squeeze Hoebel's description of the function of law into my own
Skinnerian jargon by proposing that law functions as a reinforcer. That is how it
"maintains adaptability." Law as a reinforcer is in the same conceptual realm as food
or sex. Therefore, if I can find the neural basis of feeding and sex reward, it may be
like the neural basis for law reward.
Skinner, the experimenter, can write a law defining a response- stimulus relationship
and then try to teach it to a pigeon. E.A. Hoebel, the anthropologist, works the other
way around. He can observe the behavior of primitive tribes and then deduce the
functions of their laws. Both stress two facts. First individuals or groups often
mistake fortuitous response-stimulus relations for cause and effect relations. this is
superstitious behavior in Skinner's terms, and legalized magico-religious belief in
superhuman spirit beings in Hoebel's terms. Second, both stress that individuals or
groups have regularities in their behavior. If there is a cause and effect relation for
Skinner's pigeons to find, then the pigeons not only learn it, but almost all pigeons
react alike. For example, "variable interval schedules of reward" produce a perfectly
standard pattern of behavior. The behavior is basically just a rational approach to
maximizing the reward. Skinner sees no need to postulate any innate laws except to
say that the animal is equipped to learn that effects follow causes. Similarly, Hoebel
assumes that a group's law ways are rational, acquired solutions to law-jobs. "When
the law-jobs get done, these "norms inevitably become the common denominator of
legal culture {Hoebel, 1954-: 287). It is not how the job gets done, but the outcome
that counts. There are no reflexes, no innate givens, no motoric programs to be
played out. What is interesting is the function that the behavior fulfills. The function
is to
get the rewarding stimulus, i.e. to get law. The strict behaviorist always picks a
response for the animal to learn, and thus the behaviorist himself learns relatively
little about goals of behavior. The culturist tries to find out what the people are
getting done, and thereby learns what is rewarding to them. The excitement that the
behaviorist Skinner generated came from the demonstration that a rule (a schedule
of reinforcement), led rats '" and pigeons to respond rationally.
The excitement the culturist Hoebel offers is that different cultures have similar
goals. The response topography may be drastically different from one culture to the
next, even weird or superstitious, but if the responses are doing the law-job, then
there is purposefulness. Hoebel discerns common purposes. Just as food, sex and
killing are rewards for most rats, issues of food sharing, wife stealing and murder
seem to be paramount in achieving law. Perhaps the accomplishment of such law
goals is rewarded in part by endogenous opiates. The pathway to the law goal is
probably not innate, but the capacity for rules, per se, to be reinforcers could well be
innate. It is a small step from eating food to feeding babies, and from there to
feeding refugees; small steps from sex to sexuality to love, or from opiate supported
affiliation to social altruism; small steps also from muricide to homocide to
genocide, and an even smaller step from neurochemical rewards to drug addiction.
The path of behavior as it seeks reward is as varied as the path of water trickling
down a hillside. The "gravitational force" that gives anthropological outcome an
appearance of purposefulness could be a genetically programmed brain circuitry that
releases rewarding chemicals when law rules are followed.
According to this theory, when we pick a rule, any rule, and obey it, perhaps the
brain will compare the rule model with the actual behavior. If there is a match, then
the brain releases some opiate. That much could be innately programmed.
Where do the rule models come from? Some of the neural circuits for basic reflex
rules could be built in, for example the "rules" for swallowing. Some rules could be
learned, such as rules for washing food, or washing someone else's food, or altruism.
Other rules could be created fresh, funny and different every time, like the sanctions
of colorful, spirit beings. After anyone of these rules for model behavior is in the
brain, then a match, mismatch decision would determine opiate release. A match-up
would feel good. A mismatch might cause "cognitive dissonance" that leads to
another attempt at either achieving the cognitive model or at altering the model to fit
one's behavior. My social psychologist friends call that attitude change.
As a physiological psychologist, I am simply suggesting that if Gruter ( 1979) and

Danielli ( 1980) are right about opiates playing a role in law abiding behavior, then
perhaps the mechanism includes the elements discussed above: (1) hormones to
program cellular development and change sensory sensitivity and motor capability;
(2) "drive peptides" to create behavior tendencies in concert with physiological
needs for water, energy and hormone regulation; (3) catecholamines such as
norepinephrine and dopamine for arousal and activation; ( 4 ) monoamines such as
serotonin for imagining and dreaming; (5) a match, mismatch mechanism for
comparing rules and reality; (6) chemical rewards for reinforcing a match-up
achieved through either superstition or correct logic; and (7) a chemical punishment
for failure to match up.
The above mechanism is largely genetic. Genes program the proteins, fats,
carbohydrates, neurotransmitters, hormones, enzymes to split off peptides,
monoamines, opiates and maybe opiate antagonists. The brain theoretically develops
a wiring diagram in which hedonism comes about by matching behavior patterns to
internal models of behavior patterns. This could be the key to releasing opiates. Thus
the genes program relatively few specific laws, as long as one general law is there:
"Thou shalt match behavior to behavior model, and if you fail, adapt either the
behavior or the model." Model matching is the law of laws. Model matching based
on homeostatic hedonism, the pleasure principle, is a relatively simple matter for the
genetic code. The sociobiological fun starts when we do the law-job. That is
discovering the evolutionary, neural and social principles by which people redefine
their relationships as the conditions of life change. The mischief starts when we use
artificial sweeteners, aphrodisiacs, ritualized aggression, amphetamine, exogenous
opiates, and explanatory fictions to bypass the process or impose conformity. I hope
and trust our research into homeostatic hedonism, self-stimulation and self-injection
will create more happiness than mischief.
PART III
The search for the missing pieces:
in social science
MARGARET GRUTER AND PAUL BOHANNAN
We noted above that ideas about morality like those in Alexander's and B.G.
Hoebel's papers begin the immensely interesting task of building bridges between
biology and the law. This section contains a selection of papers in which building
bridges between the biological and cultural foundations of law is the major purpose.
First, two anthropologists speak. Boehm's suggestion about the place of the moral
community in human evolution makes an original statement about the possible
origins of human law. Bohannan discusses aggression-one of the topics that lie at
the basis of law. If aggression is to be understood, all the biological and the social
sciences must provide input or else we will be defeated by its complexity.
Quite another sort of bridge was essayed by the third anthropologist, William H.
Durham. He started from ethnographic observations and reached toward biology
.He found that regularities-and laws-of many tribal peoples of West Africa
concerning the consumption of yams during the height of the malarial season favor
(apparently without the people's being overtly aware of it) their relative good
physical health. Because he has found some of the evidence in the medical
literature questionable, he withdrew his paper at the last moment. We were
fortunate to have it at the Conference and look I forward to its revision and
publication.
Roger Masters' bridge is a bold one: the suggestion that the principles of
evolutionary biology form a framework for examining the ideas of the great
political philosophers on which our very governments and laws are built.
It is at this stage that we wish to turn again to the papers on morality and those on
the non-human primates to set them into the context of the entire symposium. The
best introduction to this section is to quote
[129]
a comment made by one of our participants, Markl, at an earlier conference, and

then to go on to a report prepared by Boehm of the discussion in the group
concerned with primate studies at Monterey Dunes.
First, Markl's comment:

"[ a] sensible sociobiological interpretation would, of course, be to suppose
that morals can (along with having other effects) also increase evolutionary
fitness but that they do not do so inevitably by force of genetical
determination. It is entirely within the frame of biological evolutionary
reasoning to assume that there is no genetic disposition toward specific
moral norms, and that man is free to select moral goals, while granting that
only those moral systems that give their adherents higher fitness than the
followers of competing moral systems could and will prevail" (Markl in Stent,
1980: 215).
Boehm's report provides a valuable insight into studies of primatology and
morality. Some of the information is relevant to the papers in Part II above, on
biological research, but all of it is relevant to this section in which the question of
morality becomes central-there is, indeed, no way to discuss the biology underlying
human legal behavior without dealing with the areas of morals and the primate
analogies. The other participants in the discussion group were Itani, Hofer, and
Hook.
GROUP REPORT: PRIMATE STUDIES

Christopher Boehm
Law and morality both minimally involve a sophisticated assessment by individuals
of primary group social dynamics, and a calculating, inventive capacity to condition
other individuals through a variety of sanctions. In one sense, all dominance which
is calculating (i.e. involves a good understanding of the situation and an inventive
approach) can be considered proto-legal.
In their further development, law and morality also involve consensual values
concerning social behavior which, because they are translated into social action,
may be viewed as rules. These rules are attuned to the perceived needs of the group,
as opposed to individual needs and impulses. Because both social calculation and
intent to control are present in non-human primates, along with something like an
individual sense of responsibility for others, we may find here precursors of human
morality. However, it is only when the formulation of goals and the implementation
of sanctions become collectivized that we can speak of morality in the human
sense.
Social dominance behavior in non-human primates has been studied
chiefly by looking at dyads. But triadic relations are more interesting if we want
to compare them to human morality and law.
(1) A hamadryas baboon female, already a member of a harem, copulates behind a
rock with a non-harem male. Several times she interrupts copulation to check
circumspectly on her possessive harem master, who generally imposes his rules
(that she stay close by) by glaring, eyebrow-raising, chasing, and if necessary,
biting. Kummer (1971) has demonstrated experimentally that these females are
conditioned to be herded, so something similar to rules and punishment would
appear to be present.
(2) Hofer relates a similar incident among chimpanzees, in which subtle and
presumably deceptive non-verbal communication was used between a female and
a younger male to set up a tryst, this exchange taking place in the presence of two
possessive larger males.
(3) Itani related a frequently occurring macaque example, in which a subdominant
male and a female sit just outside the group in a consort relation. The alpha male
notices them and comes over to attack the female, who flees. But she returns to
her consort when the coast is clear.
In all three cases, there would seem to be a clear understanding of both rules and
punishment for breaking the rules. An inventive attempt was made to maximize
hedonic gratification by avoiding detection. There appears to be a relatively
sophisticated understanding of the intentions of the animal wielding superior
power, and a capacity for long-range anticipation combined with flexible problem
solving at the social level.
Another area of triadic interaction is conflict interference. Goodall reports that
high-ranking males may interfere in conflicts. Sometimes the motive seems to be
protection; sometimes the male merely wishes to reassert his own rank. Either
helps the victims, but only the protective impulse can be compared to human
conflict intervention as an altruistic act.
Sometimes interference involves more than three parties. With macaques, group
attacks may involve large numbers, still with only one "scapegoat." Analogy to
ostracism or expulsion in humans is suggestive, but ethologists do not understand
such behavior at present, and it cannot be stated right now that anything similar to
collective sanctioning takes place. Indeed, both Itani's and Goodalf's papers' show
that even when homicide occurs within the non-human primate group, the group
never responds as a whole.
In macaques, triadic interference is clearly different from alliancing behavior in
which one animal helps another to win a fight. Boehm
suggests that some active appreciation of social harmony could be present, fostered
by a genetically prepared sense of aesthetics favoring pattern consistency.
One form of proto-moral behavior would be a concern for social harmony that
overrides concerns for individuals when a third party breaks up fights without
showing preference to either combatant. Another is altruistic behavior in which an
individual displays a sense of social responsibility and the altruist takes personal
risks.
Goodall gives an example of control exercised for someone else's good: Goblin
physically removes little Freud from a branch, where he's about to tease a large male
baboon to the point of a second biting attack. The first time, Goblin had interfered
protectively. Another example, described by Lindburg {1971) for Indian rhesus
monkeys, involves a group encountering humans in an area where humans
sometimes hunt monkeys. The leading male serially threatened various members of
his group to "herd" them in a safe direction, rather than withdrawing immediately to
maximize his own safety and trusting them to follow.
Possession of objects is prominent in morality and law. In many human societies,
sexual possessiveness heads this list, and parallel behaviors have already been
pointed out in baboons, macaques and chimpanzees. Goodall provides a dramatic
example in chimpanzees, in which Passion temporarily abandons a long termite stick
and later discovers her daughter Pom has discarded a shorter one to use Passion's
longer stick. Passion simply touches her stick, and it is returned immediately. This
may be considered to be a {non-verbal) symbolic expression of right of prior
possession, presumably accompanied by an intention to punish.
In a similar way, many fights are controlled by a mere gesture on the part of a
dominant animal, particularly with chimpanzees and gorillas. It is this capacity to
send and receive messages about the consequences of breaking the rules that makes
such protomoral social systems highly effective in terms of both individual and
group selection. Such subtle communication curtails risks of physical damage, but
also reduces investments of time and energy.
As labile social sensitivity develops, the costs of dominance-based systems of social
control are decreased, and with humans public opinion took over as the chief
sanctioning mechanism within the group. However, such flexible systems also
permit abuses.
Further parallels might be discerned by giving close attention to the ontogeny of the
human capacity to respond to and to generate rules,
using Piaget's basic work. These parallels seem not merely ones of analogy, but could
go back to some common genetic traits. Goodall is correct in asserting that much of
the order in primate societies is created by dominance orders seemingly operating as
self-organizing systems. It is for this reason that only protomorality and protolaw are
present-the true moral community with its strong, conscious and collectively
communicated "rules" is confined to human beings. "
We would like to add something to Boehm's report. Turning back to B.G. Hoebel's
paper, rooted firmly in biology, we find him discussing the internal reward system
as it applies to human behavior and drives (hunger, thirst, sex, etc.). His studies are
done on rats. When it comes to developing new methods to treat the individual's
health, such experiments have long been accepted by biologists as well as by
lawyers. In medical research, nobody seems to doubt the possibility of finding
similar reactions in the brains of humans as are found in other animals.
Could it be that our brain as well as that of chimpanzees is the seat for the drive
towards balance in social organization, towards structure and the obedience to rules?
That, like chimpanzees, we develop patterns of social structure, helped by the
formation of subgroups, be they peer or dominance oriented; that behaviors as
expressed in the dominance-submission-reassurance sequence are rewarded by the
internal reward system? This system might not be so highly differentiated and
developed in chimpanzees as in humans; it might not be activated by observations of
misbehaviors (criminal or evil actions in human terms) to the degree that it would
strongly stimulate third party interference in dispute settlement. But it might be the
same system nevertheless, only less differentiated and adapted to a different
economic niche in the overall scenario of living creatures. In this case we might be
able to learn much more from long-term field studies of our close relatives;
especially if these observations are complemented and further investigated by
experiments in primate facilities. There is no reason why we should not be able to
learn about fundamental biological rules of group relationships just as we have
learned and are continuing to learn about the regularity and predictability of
biological processes in the individual living organisms.
[134]
The evolutionary development of
morality as an effect of dominance behavior
and conflict interference
CHRISTOPHER BOEHM
Department of Anthropology, Sociology and Philosophy,
Northern Kentucky University, Highland Heights, KY 41076, USA
Morality may be defined as the problem solving activities of a moral

community, a primary group which uses a wide range of sanctions directly to
reduce conflict, which also sanctions perceived causes of conflict, and
defines and controls other deviances judged to be antisocial. So defined,
morality is a precondition for law. In comparing human with non-human
primates, conflict management is one of the most impressive parallels. This
empirical parallel is built upon, to construct an evolutionary scenario for the
development of morality and law in their proto-forms.
Selfish dominance consisting of pure physical coercion has nothing to do with law.
But when culturally patterned understandings about what is socially desirable
develop in a context tempered by dominance or authority, and in being informally
sanctioned they help to regulate social behavior, we speak of customary law. Such
sanctioning is practiced by all human groups, acting as moral communities which
agree on a code of prescribed and proscribed behavior. Moral communities
deliberately condition their members in directions they believe will promote social
harmony, and the more this purposive regulation of social conduct comes to be
formalized and institutionalized, the closer it approaches modern law based on
written legal codes, with formal judicial bodies, policing specialists, and a penal
system.
The continuum between non-literate and modern legal systems is well recognized
among legal anthropologists and others. However, extension of this continuum to
other highly social species or to hypothesized behaviors of our direct precursors has
been tentative. Gruter's ( 1977, 1979) general analysis of analogs to legal behavior in
non-human primates has been one important attempt to blaze a trail in this direction,
while Lorenz (1966) and Bischof (1980) point to a number of analogs to moral
behavior in other animals. By contrast,
Kummer's (1980) assessment of moral homologs in non-human primates remains
negative. But this may be due to the rather restricted definition of morality he works
with.
PRELIMINARY SCENARIO FOR EVOLUT1ONARY

DEVELOPMENT OF THE MORAL COMMUNITY
I shall explore dominance, coercion, submission, flight, conflict resolution and
leadership as probable loci for the evolutionary development of protoforms of moral
and legal behavior in earlier stages of human evolution. I take for granted the
arguments of Tiger (1969) and Eibl-Eibesfeldt (1974), that bonding and positive
affect are also of great functional importance, but have not received their due in
sociobiological analyses of human behavior. My reasons for concentrating on
politics rather than love will very quickly become clear. Given the excellent data on
social dominance and conflict interference in extant primates, it is possible to
develop relatively specific, homology-based hypotheses on the origin of morality
and law, hypotheses specific enough to be tested for relative plausibility against
other such hypotheses.
It is difficult to conceive of the early evolutionary development of law, without
speaking first of the development of proto-morality and morality. Elsewhere (Boehm
1981c, 1981d) I have suggested that the origin of morality came through deliberate
interference in conflicts within co-operating groups of primates. Such interference
occurs among adults in many extant monkeys which exhibit dominance rankings,
and in great apes as well. Because such efforts apparently involve an intention to
restore social harmony, I have called them "prot0--moral."
These animals display less cooperation at the group level than is generally assumed
for protohominids, yet they are sufficiently sophisticated, socially, to manipulate
many exacerbated conflicts. It is assumed that protohominids also recognized the
value of social harmony and controlled obvious social problems such as severe
fights with still more insight. However, monkeys and apes must not be underrated.
They employ an impressive range of deliberate techniques to interfere in conflicts so
as to terminate them (Boehm, 1981d).
Certain of these extant primates, notably chimpanzees and orangutans, also are able
to recognize themselves in mirrors (Suarez & Gallup, 1981). In early humans, it is
assumed a similar but more developed capacity for individual self-conceptualization
developed into
a sense of group self-interest, which by far transcended anything developed by
extant monkeys or apes.
The assumption, then, is that morality developed as a form of rational problem
solving. The hypothesis is that morality had its earliest roots in conflict
management. It later came to encompass group sanctioning of individual deviance in
other areas, including eventually deliberate limitation of raw power used by
dominant individuals acting as leaders.
Once all this happened and verbal symbolic communication arrived, restoration (or
maintenance) of social harmony by collective sanctioning, and subsequent
intentional regulation of dominance itself, were developments which provided
important preconditions for the evolutionary development of protolegal behavior,
and eventually of law. Minimally, this must have involved a set of agreed upon rules
of conduct, and some collectively espoused means for applying and enforcing such
rules.
This brief scenario provides a bare outline of the evolutionary sequence I shall try to
reconstruct. I now treat various aspects of the argument separately, offering a series
of rather specific hypotheses about the relation of social dominance to the
development of morality. Later, I shall integrate these hypotheses into a more
definitive scenario for the origin of morality and law in their proto-forms.
EIGHT HYPOTHESES
Hypothesis 1:
Prehominids exhibited behavioral lability.
Our precursors were capable of flexible adaptive modification, in Kummer's (1971a)
sense. In other words, the balance between genetic preparation and learning was
such that considerable flexibility in social organization was present. This means
under conditions of environmental change, that relatively major adaptive
modifications at the phenotypic level could be made rapidly. For example, upon
becoming more terrestrial this ape may have escalated its predator defenses very
quickly, through development of cooperative bluffing and fighting.
Hypothesis 2:
In prehominids, there may have been relatively stronger preparations for
dominance, and weaker ones for submission.
I hypothesize that genetic preparations for social dominance mechanisms in early
hominids were comparable to those of great apes, baboons or macaques. It is
assumed that as largish arboreal primates our direct precursors interacted at group
levels above the nuclear family level, and such groups were adaptively dependent on
genetic dominance and upon the resulting social dominance hierarchies. More
basically, such groups consisted of bonded animals capable of recognizing one
another individually.
The potentially controversial assumption is that submission was less prepared
genetically, compared to dominance. This is based on the following reasoning. It is
widely agreed that our precursors lived arboreally. In such niches, many extant
species tend to display flight behavior in the face of dominant aggression rather than
submissive behavior which neutralizes the aggression. However, flight is not easily
substitutable for submission among ground dwelling primates because fleeing
animals are very exposed to predators.
To summarize this hypothesis, our precursors, adapting to terrestrial niches as
lakeside savannahs opened up, already had well-developed tendencies toward
dominance behavior previous to this adjustment. They had resorted mainly to flight
as an effective individual response to aggression, and this worked perfectly well up
in trees. On the ground, a social dominance hierarchy became still more critical for
group survival. But something other than flight was needed, if dominance
interactions were not to seriously damage reproductive success.
Hypothesis 3:
In protohominids, adverse effects of dominance were inhibited through
deliberate interference by third parties in conflicts within the group.
In making their adaptation to lakeside savannahs these apes developed largely
group-traditional controls on dominance behavior, rather than depending heavily on
flight or submission, as genetically well-prepared mechanisms. These controls were
built upon a previously less developed capacity for deliberately interfering in
internecine conflicts through a variety of strategies, the aim being to stop an episode
of exacerbated agonism.
Such behavior is exhibited regularly in langurs, macaques, baboons, chimpanzees

and a number of other primates, conflict interference constitutes only a secondary
control on aggression. The primary controls are genetically well-prepared individual
submission, while interference takes over when such submission is ineffective and
an agonistic episode becomes exacerbated. However, among protohominids I am
hypothesizing that conflict interference, as a form of deliberate problem solving, had
to account for a much greater proportion of social control on dominance, compared
with any extant non-human primate. The resulting cohesive group was able to ward
off terrestrial predators, and eventually was able to hunt large game cooperatively.
Hypothesis 4:
Terrestrrtzl adaptation created selection pressures
favortng more socially sensitive individuals.
In macaques, much dyadic and triadic agonism is rather fully acted out through
physical contact and even fighting. In baboons, a greater proportion of such
communication takes place through posturing, gestures and vocalization. In
chimpanzees and gorillas, the latter tendency is so great that subordinates anticipate
aggression much of the time, or are able to pick up on very subtle cues because of
their greater social sensitivity. This is true of dyadic dominance interactions, but also
of triadic alliancing or interference situations.
As protohominids developed a greater dependence on interference, selection at both
individual and group levels favored a social sensitivity which permitted individuals
to anticipate the outcomes of dominance or interference episodes, and to modify
their behavior in advance. This brought selective advantages to individuals because
time, energy and physical risk were reduced, as costs accompanying the gains of
social dominance hierarchy. It also increased reproductive successes of groups as
semi-isolated breeding populations, as did increased cohesiveness and flexibility
resulting from such sensitivity. One result was an animal that was becoming
increasingly receptive, through learning, to social control; another was a tradition of
social manipulation far subtler than that of chimpanzees and gorillas.
Hypothesis 5:
Tool use may have exacerbated dangers of intragroup agonism.
It seems probable that the homicidal potential of hominids was increased by tool
inventions which made predation on larger game possible. Particularly since canine
teeth suitable for ripping or slashing were absent, it would appear that even use of
large stones as hand-held " weapons or projectiles would have greatly escalated the
probability that intragroup conflicts might seriously diminish reproductive success
of a group. As a result of such inventions, selection favoring conflict interference
may have escalated concomitantly, through a combination of individual and
interdemic selection effects.
Hypothesis 6:
The moral community was a necessary development in the human line.
At some point, advanced development of self-conceptualization and sy.mbolic
verbal communication permitted verbal labeling and generalization concerning the
same obvious social problems which had been intuitively but deliberately
"managed" through conflict interference. This was the earliest manifestation of the
moral community which exists in every contemporary human society. Most likely,
the first problems discussed were ones of a violent nature: quarrels injurious to a
consensually appreciated group-interest in maintaining well-recognized forms of
cooperation. This involved conceptualization of collective interests as being more
important than individual interests. As agreed-upon rules of conduct were built out
of the experience of the moral community in social problem solving, proto- legal
behavior emerged.
The moral community may be defined technically as a primary group having a
conscious and collective sense of its own self-interest, which generalizes its
preference for social behavior into idealized models and restrictive rules, and
knowingly sanctions these preferences so as to condition individual behavior in
desired direction. As a precondition, there must exist the individual social sensitivity
mentioned in Hypothesis 5. But also individuals in the group must understand social
dynamics well enough to harness this sensitivity in manipulating the behavior of
individuals judged to be deviant.
Hypothesis 7:
Dominance in leadership becomes regulated by the moral community.
As moral communities emerged, the already diminishing coercive dominance of
individual leaders was further limited. In part this dimunition was due to genetic
selection, since increased social sensitivity made coercive methods less efficient.
But it was also due to a growing individual sensitivity to public opinion, on the part
of followers and leaders alike. Thus, there came to be deliberate collective
curtailment of individual power, as a matter of right and wrong. This was the origin
of the egalitarian ethos so regularly exhibited by smaller extant human societies.
Followers began to decide exactly how strongly they wished their leaders to express
dominance, for which increasingly moderate genetic preparations still persisted.
Dominance patterns became heavily shaped by group tradition, since leaders had
become sensitive to subtler social pressures.
As groups came to manipulate their leaders through social pressure, an enormous
gap grew between protohominids or early humans and other primates, in the
expression of dominance behavior. This was significant both in the acting out of
dominance, and in its inhibition as well. The result was a less genetically dominant
species, but one which continued to have relatively strong genetic preparations for
dominance, compared with those for submission. Interference, and a more thorough
elaboration of bonding, did much of the job performed by submission gestures in
other terrestrial primates.
Hypothesis 8:
Timing was essential; when did the moral community and
proto-legal behavior arrive?
I have made these developments contingent upon the arrival of spoken verbal
language. However, it is emphasized that linguistic development need not have been
very elaborate, to permit conceptualization of group self-interest and generalization
about desirable versus undesirable varieties of behavior. The crucial factors are
capacity for self- conceptualization and an intelligent ability to generalize, not full
development of verbal symbolic language. Chimpanzees and orangutans (but not
baboons or macaques) are well along the road to self- conceptualization already
(Suarez & Gallup, 1981). And chimpanzees (with gorillas) have shown themselves
to be capable of learning and manipulating symbols inventively, in spite of limited
capacity for verbalization using phonemes invented by humans.
Because there is no good evidence which limits the articulatory potential of our
precursors to form words in terms of phonemes different from those employed by
humans, one mustplace the advent of protolanguage ability in terms of development
of intelligence. Given the impressive capacities of chimpanzees in manipulating
signs, assigning an earliest possible date becomes difficult. So for the emerging
moral community this chronological hypothesis must remain vague, but potentially
very early. Given the available evidence for technological problem solving, Homo
erectus seems a likely guess for definitive emergence of moral communities and,
presumably, of proto-legal "rules."
LOCUS OF SELECTION
In its broadest outline, the argument underlying these eight hypotheses is so
generally accepted that it approaches being a truism to which deference is paid in
every anthropological textbook. A particular ape developed increasing lability over a
long period of time, and eventually became technologically very inventive, verbally
symbolic and moral in that order. However, there is actually a choice between two
major theories explaining how selection came to favor such lability. The strongly
predominant one (e.g. Leakey, 1961) holds that this was in the technological field
(invention of tools, etc.) while the other (e.g. Kummer, 1971a) favors the field of
social behavior as the locus for development of increasing lability and intelligence.
In extant non-human primates, it is obviously difficult to weigh the degree of lability
manifested in the social versus the technological sphere. But intuitively it seems that
lability is expressed far more strongly and with more variety in the social field than
in the technological field. On this basis, I join the minority to suggest that early in
the evolution of cultural capacity, selection favoring greater social lability could
easily have been the crucial factor. Kummer's ( 1971) arguments concerning benefits
of adaptive modifiability in social structure are highly suggestive here.
THE FULL SCENARIO

In thinking about various spheres of primate social behavior as to which are the most
labile, conflict interference is a well-qualified candidate, since this behavior involves
deliberate use of a number of different behavioral strategies to achieve an
impressively complicated end
(Boehm, 1981d). By tying the evolution of social lability to this particular behavior
in its relation to social dominance, it is possible to construct a more specific
evolutionary scenario than is usually generated in the course of such speculations.
The immediate purpose is to explain the genesis of morality. But since "morality"
has been defined to consist of deliberate problem solving, the argument also explains
something about the earlier development of inventive cultural capacity, as well as
setting the stage for the emergency of law.
For lack of a better concept, one might treat the "aesthetic sense," broadly
conceived, as the locus of human problem solving ability (Boehm 1981b), since this
involves not only a bias in favor of pattern consistency, but a rather considerable
intuitive capacity for what academicians call "systems analysis." Conflict
interference in non- human primates provides a good example of this aesthetically
oriented problem-solving capacity in action: elsewhere (Boehm, 1981c), I have
argued that restoration of social harmony is the object, and that interferers make
rather complicated calculations with respect to social dynamics.
Conflict interference of higher primates not only involves a strong manifestation of
lability, but is proto-moral. For both reasons, I have chosen this behavior as the
logical area in which morality itself developed. This hypothesis is relatively specific,
yet well supported by the slender evidence available. It is difficult to identify
definitely by homology any other form of proto-morality in extant non-human
primates, as they behave in their natural habitats without human tutelage.
The scenario based on the eight hypotheses is actually rather simple. A behaviorally
labile and largely arboreal ape, exhibiting both social dominance hierarchy and some
conflict interference, moved into terrestrial habitats. There, due to increased predator
pressures, group size and social cohesiveness became crucial criteria for selection.
Whether or not genetic submission mechanisms were relatively weakly developed
up in the trees, selection favored labile conflict interference over genetic submission
mechanisms, even though both sources of variation surely were available. The
assumption is that interference was more effective than submission, given the new
terrestrial selection pressures. It is also possible that lability was being positively
selected on other bases, so that interference gained a selective edge over submission
mechanisms with some outside help, as it were. Technological invention remains a
logical candidate, here.
When chimpanzees interfere in conflicts, which is very infrequently
in comparison with macaques and baboons, it appears that the interferer's cues are
more subtle, and that responses are more sensitive. Since these apes manifest a good
deal of dominance behavior in general, this marked difference cannot be explained
simply by saying " that there is less dominant aggressiveness floating around in
these species. Nor does it appear that individual submission is relatively more
developed in the apes to a degree that submission controls dominance so effectively
that interference is seldom necessary. Rather, it appears that interference is carried
out {and responded to) much more frequently by nuance.
I have suggested that in terrestrial prehominids, asocial sensitivity greater than that
of apes was selected mainly because this made control of dominant aggression more
efficient. Selection in this direction not only enhanced the capacity of subordinates
to respond efficiently, but also enhanced dominance interactions in general.
Small sacrifices of time and energy and fewer risks of bodily injury were demanded
by these interactions, as they became based less on physical attacks or serious
threats, and more on subtle indications which were expertly "read" by subordinates.
And this, in turn, led to Ian increased capacity for communication by non-verbal
signals. Of " course, leaders and followers alike became prepared genetically to be
more socially sensitive. As a result, it also became easier for followers to sanction
leaders when leaders acted agonistically not as peace- makers but as aggressors.
At this point, the protohominid we are discussing had developed a stronger degree of
proto-morality than any extant non-human primate, expressed through an
increasingly subtle tradition of conflict interference. It may be assumed also that this
highly social and still rather feisty creature was more similar to chimpanzees than to
any other extant primate. At the same time, its capacity for self-conceptualization
was being extended to include a relatively advanced, if still rudimentary, sense of
group membership. Concomitantly, the genetic basis for its problem-solving
"quotient" was growing, possibly with some additional help from technological
invention.
Verbal symbolic communication was probably necessary to formation of actual
moral communities, since as defined these collectivities developed shared evaluation
understandings about preferred and non- preferred behavior, and practiced
sanctioning accordingly. When such understandings are generally known and are
sanctioned, it is possible to speak not only of a sense of "right" and "wrong," but of
"rules." It is difficult to imagine the development of such under-
standings without the advantage of a proto-language. But then, it is not easy to
imagine a proto-language in the first place. With respect to the development of
morality and of spoken language, it seems likely the two developed concomitantly,
along a gradual continuum. This means that language ability was not selected just
because it permitted better communication useful to the subsistence quest. It was
also selected because of its contribution to more effective social control, as a key
mechanism which permitted this socially aggressive primate to reap the benefits of
cooperation in predator defense, in the subsistence quest, and possibly in winning
encounters with other primates.
As I have indicated in Hypothesis 8, it is difficult to place the transition from proto-
morality to full morality in time. However, given the general intelligence and
aesthetic capacity of Homo erectus, as inferred from stone artifact production and
from the fact that large game was hunted, it seems possible that the moral
community arrived that early. The only other thing needed is development of a
proto- linguistic capacity to label behaviors and to assign them values publicly.
This scenario proposes a straight line development of conflict interference into
conflict management, as the original basis for morality. This constitutes a narrow
view of morality, compared with morality as we know it today. But potentially the
moral regulation of sexuality, property and homicide, as well as moral rules which
apply to veracity, are all intimately related to conflict resolution. All are likely
causes of conflict, and as conflict management became more sophisticated, it is
logical that its causes would be anticipated and dealt with in advance. With this
assumption, the incest taboo falls by the wayside for a candidate as the catalyst
which produced moral regulation of social life. I suggest instead that it was more in
the political sphere that morality developed, and that at some point after the advent
of the moral community, the egalitarian political ethos arose as aside effect.
IMPLICATIONS OF THE ARGUMENT

Moral communities of our precursors, whenever these arrived, signaled a new
development in the evolution of social organization. Social sanctioning was no
longer fully automatic, as in dominance hierarchies of chickens. Nor was there only
a small element of deliberate inventive problem solving, as in the individual conflict
interference of certain monkeys and apes. Social control became not only much
more deliberate and insightful, but also collective. And this
brought about the genesis of morality and the basis of law. The scenario is based on
the assumption that our precursors may have started with relatively strong genetic
preparations for dominance, but with relatively weaker genetic preparations for
submission. In any event, instead of developing strong genetic submission
mechanisms needed to inhibit this dominance in their new terrestrial niche, they
developed more effective inhibitory mechanisms in the area of conflict interference
as a more labile behavior. As a result of increasing lability, genetic preparations for
dominance itself became weaker and less specific, but still remained stronger
relative to submission, compared with other terrestrial primates.
For all the attention the issue has received in the past seven years, the relation of
genes to labile behavior remains a realm of mystery. But still-prevalent
environmental explanatory fashions must not be allowed to obscure the fact that
humans are programmed to learn in rather specific ways (Pulliam & Dunford, 1980),
and in terms of rather predictable basic emotional reactions. It is at least interesting
to ask at this point whether contemporary humans may be somewhat better prepared
genetically to inhibit such behavior, both as submitters and as dominators who
remain responsive to submissive signals (obviously, in humans none of these
behavioral preparations could be very specific).
This speculative hypothesis fits with the evolutionary scenario. If plausible, it helps
to explain a phylogenetic anomaly in the human species. Our species is distinctive
for several reasons, one being that we develop moral communities, another being
that we practice genocidal warfare. A relative preponderance of genetic dominance
over genetic submission has already been hypothesized to be germane to the
development of morality as an inhibitory mechanism which controls dominance. It
may also be highly germane to the explanation of how warfare developed in our
species. Humans appear to be admirably equipped with a potential for homicidal
attack, which cultural development sometimes fosters. Potentially we also have a
relatively low level of responsiveness to submission, as far as genetic preparations
are concerned. It is not difficult to see why serious warfare might develop, as long as
it does not threaten species extinction, since so often conspecific killing is
effectively inhibited only within the primary group, or within a set of allied primary
groups.
To explain the existence of warfare in terms of morality, one must call on another
manifestation of the contemporary moral community, namely ethnocentrism. The
dual moral standard inherent in ethno-
centrism (Le Vine & Campbell, 1972) allows outgroupers to be treated like animals
to which normal rules regulating homicide do not apply. Xenophobia exhibited by
extant terrestrial primates (Southwick, 1974) suggests that sources of variation in
this direction were available in our precursors, and this facilitates arguments that
ethnocentric moral double standards may be quite ancient-as ancient as the moral
community itself.
The result is a species which has been able to practice warfare for at least ten
millenia without destroying itself. One reason this has been possible may be that
while warfare involves a special moral license to commit homicide, it seldom
remains entirely ungoverned by rules. In this sense, 'international law' may be nearly
as ancient as the law which prevails within moral communities.
CONCLUSIONS
In conclusion, I suggest that the origin of the moral community, as a precondition
for the emergence of law, was the product of selective pressures attendant upon the
change from an arboreal to a terrestrial adaptation. Conflict interference provided an
increasingly efficient inhibitory mechanism to control destructive side effects of
social dominance and of culturally reinforced human propensities to be self-
assertive in general, where these resulted in socially troublesome behavior. This was
a highly potent instrument of adaptation, by which humans usefully contributed to
their own adaptive success on a deliberate, perceptive basis. As cultural selection
became more salient, genetic preparations for dominance were weakened, being
augmented by selection favoring individual social sensitivity in leaders and
followers alike. Group self-conceptualization and verbal symbolic communication,
which probably arrived in tandem, made possible the development of moral
communities. In these communities, group sanction emerged as the most powerful
instrument for regulation of individually assertive behaviors, particularly those
which very obviously disrupted cooperation or disturbed social equilibrium needed
for group stability.
I have not spoken at all of duty, obligation, or other favorite abstract entities
favored by moral philosophers, although elsewhere I have suggested (Boehm,
1981c) that extant non-human primates exhibit something like a rudimentary sense
of social responsibility. But my argument is highly consistent with the philosophical
notion that moral behavior is intentional (Stent, 1980), since I have emphasized
that moral behavior involves finding sophisticated solutions to perceived social
problems. It is on this basis, that I have treated the kind of conflict interference
exhibited by extant non-human primates as the prototype for moral behavior in the
human line. My interpretation is that this labile behavior involves some rudimentary
social sophistication, is highly intentional, and is oriented to an aesthetic
appreciation of social harmony, as well as to protective concerns for individuals and
to needs to assert dominance.
This interpretation may seem dangerously "psychological" or "mentalistic" to
ethologists who remain loyal to astrict behaviorist tradition. But arguing as I have
from an interface point joining natural science and behavioral science, I have taken
it as my perogative to be liberal, yet reasonably careful, in the interpretation of data
from primate ethology. In their specificity, the hypotheses I have developed now
await the scientific test of relative plausibility, in competition with alternative
accounts of the origin of morality and law.
Some bases of aggression and

their relationship to law
PAUL BOHANNAN
Dean, Social Sciences and Communication, University of
Southern California, Los Angeles, CA 90007, USA
Aggression has been studied from several vantage points: the behavioral, the
somatic, the social, the cultural and the experiential. Each of these vantage points
dictates a different method of interfering with or "doing something about"
aggression. Each can be "corrected"; by behavior control, by surgery or drugs, by
altering the social system, by changing cultural values and by reconstructing the
interpretation of experience. Law can be seen as a cultural device for turning a social
dyad into a triad in order to "solve" a conflict at the same time that it
reinstitutionalizes some cultural norms of basic social institutions in a legal context.
All of the various vantage points must be recognized as law is reinstitutionalized
from the primary institutions of society.
.
Both scientists and judges must consider the biological basis of human behavior if
they are to deal adequately with it. What can they learn from each other? I am here
concerned with one topic in which both have a stake: aggression. No one questions
that aggression springs from biological roots, that it is affected by social norms,
altered by cultural values, tempered by experience, that it can become abnormal
during somatic or mental disease, and often leads to confrontations with the law.
Other such many-faceted subjects include alcoholism (Madsen, 1974), bastardy
(Laslett, Oversteen & Smith, 1980, has a good bibliography, centered on
demography but covering many other dimensions of the subject), death itself.
Aggression, as a topic of scholarly concern, has had immense difficulty shaking
loose from religious attitudes and from centuries of moral philosophy, some of
which is relevant in the discussion of the relationship of biology and law (Masters:
this volume), but some of which may be used as blinders rather than as a beacon.
There is still a tendency on the part of most Westerners to assume that aggression
and destructiveness are the same thing-they are not.
A period in the study of aggression seems to have come to an end in the early 1970s.
During that period, the major question was the biological basis of aggression-
reaction to the "discovery" by ethologists that there is a biological dimension, and
the claim of the more bold that aggression is therefore "innate." The big argument
was whether aggression was innate or acquired-it is, of course, both. The definition
of aggression sometimes swings from one criterion to another in the middle of a
thought, thereby making this very complex word a muddle. The best summarizers
from that period are Storr ( 1968) and Fromm (1974).
Since those days, there has been, in some behavioral sciences, a movement away
from mere aggression and toward the topic of conflict management (an avatar of
"conflict resolution").
Studies of human aggression are to an unfortunate degree beset by another difficulty,
one they share with criminology in general: investigators are urged, either by their
sponsors or their consciences, to find out what we do about some problem before
they have formed any very clear opinion of what the problem is , and certainly
before we understand the action chains that underlie it. The policy question gets
stuck into the mix too early.
Although most students of aggression insist that the subject is complicated and
requires input from many sciences, few of them have been willing to go beyond lip
service to get an overview of their topic.
We can well understand, of course, that no one is "expert" in all those sciences. But
not being an expert in something is no license to disregard it, especially if we
recognize our "limits of naivete" as Adamson Hoebel warned us above.
Research on aggression can be easily split into five spheres or categories, depending
on the definition the researcher employs. There are those scientists who focus on:
1) aggressive behavior,
2) the bodily, somatic infrastructure (either in its entirety or in specialized
areas),
3) the social groups that are involved in aggression,
4) bellicose or peaceful culture, and how culture can be manipulated and
forced into non-aggressive molds,
5) the fall-out of the experience of aggression.
It is usual that the researcher, having taken up one of these positions, uses it as a
causal lever to indict one of the other spheres. The literature is full of comments
abo!,1t how aggressive behavior is "caused" by our hormones or our brains, by the
inept or faulty structure of society which is therefore inadequate to contain or
provide alternatives to aggression, by our culture with its bellicose values, by our
unfortunate or craven individual experiences. Far less attention is given to the
premises behind the complicated ideas of cause and effect-they are usually naive or
unexamined-or to interaction among the various spheres.
In this paper, I shall review some of the literature on aggression, paying special
attention to how these points of view have led to different attempts to alter
aggressive behavior; I shall then turn to the law.
CORRECTING THE BODY

There is no doubt that there is a biological infrastructure underlying aggressive
behavior. The scholars who have looked at it can be divided into two sorts: those
interested primarily in the brain and the central nervous system who want to know
how aggressive behavior correlates with brain function, and those concerned with
chemistry who want to know how hormones, enzymes, peptides and other chemicals
affect behavior. Some writers, of course, concentrate on the correlation of the two.
They have found, to summarize quickly, that the brain
centers unequivocally associated with aggression are the amygdala and the
hypothalamus, but that different types of aggression seem to erupt from different
locations in those and other brain parts, and often involve other regions of the brain-
and apparently no brain function is associated simply with one specific brain part. In
human beings, the matter is even more complex because the cerebral cortex is
involved extensively.
Chemicals such as hormones and peptides also affect aggressive behavior: adrenalin
and the male sex hormones are the traditionally studied agents. Bartley Hoebel's
paper in this volume deals with peptides, which have been identified comparatively
recently as chemical agents involved in the complex brain mechanism guiding or
rewarding allbehavior, including aggressive behavior. Again, the true state of affairs
is immensely complicated, but we are beginning to accumulate considerable
information about the biological elements that underlie aggressive behavior .
Consonant with these two approaches-using discoveries of physiology and
biochemistry as starting points for applying a policy of reducing aggression-are at
least two ways in which attempts have been made to change aggressive behavior by
altering the body: surgery and drugs.
To take only one example of chemical control of aggression, both stimulants and
sedatives have been administered to school children in order to get the behavioral
response (including reduction of aggressive behavior) that the doctors and/or
teachers desire. Stimulants such as Ritalin have been found to calm the behavior of
hyperactive children and to increase their attention span. The drugs, on impeccable
evidence, are neither narcotic nor physically addictive, but tolerances increase so
that dosages have to be increased. Psychological dependencies may develop if the
child associates acceptance at home or in the classroom with the drug. The drugs are
withdrawn slowly when the child's behavior pattern is that desired by the authorities-
with the hope that the behavior pattern can persist on less and less of the drug.
According to rules of the Federal Drug Administration of 1970, a child must be
diagnosed as a case of "minimal brain dysfunction" before the drugs can be
prescribed. Since all hyperactive children do not by any means show brain
dysfunction, this practice seems to be an attempt to vitiate any unscientific judgment
of "abnormal" behavior and hence to prohibit use of drugs merely to control
behavior. It is difficult to assess the degree to which attempts to control such
programs have been successful (and, as far as that goes, just what
minimal brain dysfunction means-it seems to be a term made up to categorize those

for whom drugs are effective).
The scope for treatment by such drugs is apparently large-the manufacturers of
Ritalin estimated in 1970 that 2,000,000 American children would "benefit" from
the drug. But consumption of the drug for controlling aggression and "hyperactivity"
in children did not reach that proportion, and apparently treatment with the drug is
less frequent now.
There is a considerable literature about surgery to control behavior, admirably
summarized and evaluated by Valenstein (1973). Such methods are very old.
Trephining the skull was one in many parts of the world, from early times. However,
it was only after the later eighteenth and early nineteenth century, with the
development of neurophysiology as an experimental science, that scientific rather
than religious or magical rationales began to be applied to brain manipulation. The
earliest published account of psychosurgery dates from 1891 (Valenstein, 1973:
266). Lobotomy was first carried out by a Portuguese surgeon in the middle 1930s
(Valenstein, 1973: 53-54) and I had become infamous by the middle 1950s. By the
late 1930s the temporal lobe of the brain-and specifically the amygdala and the
hypothalamus-had become implicated in aggressive and sexual behavior. Surgery on
that portion of the brain was found to reduce aggressive behavior in monkeys, cats,
dogs, rats and other experimental adults; it had been used also to control the effects
of Parkinson- ism and epilepsy in human beings. Temporal lobectomies on human
beings were first carried out in the early 1950s. An operation called cingulotomy has
been performed on a number of sufferers from psychomotor epilepsy to reduce
aggression. Operations on both the amygdala and the hypothalamus have had results
that the surgeons called good-they relieved the symptoms of aggressive rage in an
impressive proportion of cases, although Valenstein has harsh words to say about
their evaluations: "there is no convincing evidence that stimulation of any brain
region specifically activates or inhibits one and only one motivational system"
(1973: 198), and even more damaging: "One recurrent difficulty in evaluating all
psychosurgical procedures is that it is usually not possible to make an independent
judgment about the results. Clinical reports generally are written in a very subjective
style, leaning heavily on impressions of the ward staff. It is rare indeed that any data
obtained from objective tests are included in the reports. The surgeons are often too
busy to become involved in time-consuming behavioral testing, and they seldom
have
adequate training in this field anyway" (Valenstein., 1973: 219-220). What the law
did or did not do to encourage or prohibit these types of treatment is unfortunately
beyond the scope of this paper .
CORRECTING THE CULTURE

In 1971 Leon Eisenberg published this statement in Science.
Learning may not account completely for human aggression, but the social
forces in contemporary society that encourage its development are so
evident that preoccupations with hypothesized biological factors is almost
quixotic.
There is little doubt that culture effects the nature and amount of aggression.
Changing culture is the most popular way of trying to alter aggressive behavior. The
documented results are also least convincing. We shall examine only three of the
many points: television violence, myths about the nature of masculinity and
femininity and face-saving devices.
(1) Research has indicated over the years that aggressive acts on television are
numerous-up to a dozen violent acts an hour, but the number varies not only with the
year the count was taken, but with the definition of a violent act and the
predilections of the observers. It is also true that high incidence of violence in a
program is correlated with its rating-high violence goes with high ratings. We have
to ask why. Is it a characteristic of television as a medium that a lot of action filling
the small screen is more riveting than other kinds of material? Is it more basic-part
of the soma or spirit of the human creature? Does violence represent a demand of the
culture or of members of the society? We only know for sure that, whether it is
"damaging" or not, television is a good way to spread the culture of violence-a lot of
violent criminals have apparently learned at least some of their techniques and
maybe some of their values from television.
(2) The place of agonistic behavior and aggression in creating the cultural view of
masculinity is well understood. There is a great deal written, most of it in snippets,
about machismo in Latin cultures, about warrior culture, and about the various
modes by which young men- and sometimes men who are not so young-think they
must maintain their masculine images through aggressive behavior. There is much
less written, but enough that it could profitably be brought together, on the
masculine roles that different societies associate with lack of
violence—priests, shamans, judges and perhaps many others, who may be masculine
without being aggressive. Except for Ashley Montagu ( 1976 ), anthropologists have
been backward in studying this problem. And Montagu's material deals almost
entirely with small, peripheral groups that would seem to have been on the verge of
extinction-many of them may have been exhausted rather than non-violent. Some
anthropologists have certainly noted such matters about '" individual societies they
themselves have studied, but surveys of this literature are hard to find.
There was a profound change in the cultural definition of masculine behavior in the
United States in the late 1960s. It went along with dress-alike styles: the message
was that people did not have to take on agonistic roles in order to be masculine.
Many of the older generation -and no small proportion of the younger generation-
were disturbed by these changes. It seems just as apparent that, as this paper is
written, we are in the middle of a back-swing.
(3) On an impressionistic level, it seems that in many societies with different
cultural traditions a great deal of aggression occurs because there are few or no
cultural norms providing honorable alternatives to aggressive acts. How much
fighting do people-especially young boys-have to do because there is no honorable
way to get out of it? I know of no study of face-saving devices, or how to improve or
extend them. The material in the literature is sparse, but such a study, if if could be
made, would be invaluable.
The degree of bellicosity of a culture may thus be directly associated with the fact
that there is no other recognized way out of a situation in which violence is
traditionally employed. The available data are few because few ethnographers have
ever looked at the problem in this way. But it is obvious that bellicose values in
culture have an immense impact on aggressive behavior.
CORRECTING THE SOCIAL STRUCTURE

We shall here mention only two devices that might be built into the social order for
the purpose of reducing aggressive interaction: one of them involves particular
methods for avoiding polarization (most often within large, particularly intercultural
or international social situations); the other deals with law and order .
Complex groups must reach a state of polarization before war can break out. I would
like to see a study made of a number of wars, showing the steps by which such
polarization occurred, with particular
.
attention paid to the choice points that led ultimately to the polarization and the
resultant conflict. I think that struggles like the War Between the States and the
Hundred Years War would provide a good beginning point because the polarization
in those wars was never total. Many wars could then be examined in a comparative
study to get a broader view.
A common reaction to smaller scale, non-international situations in which our
cultural values tell us there is "too much aggression" is likely to result in a demand
for "law and order." An historical example is to be found in the reorganization of the
police in the United States in the 1840s and 1850s after the phenomenal rise of the
crime rate after the War of 1812. It is possible, today, to see that in those years the
crime rate rose in response to the destruction of the traditional community and the
problems attendant on the creation of decent living conditions in the new industrial
city. The fact is that the crime rate rose precipitously. The police were not sensibly
organized -in fact, they were scarcely organized at all. The tasks of maintaining
order were piecemeal and poorly coordinated among a whole series of officials-
constables, magistrates, night-watch, port guard, militia and others. When authorities
attempted to create city-wide or state-wide police systems, there was violent
opposition-not just from those with vested interests, but also from a civilian
population who feared a police state even more than they feared the rise in the crime
rate. "Liveried1' policemen, they felt certain, would expropriate the civil rights of all
citizens.
The police system in any country works as long as it takes into consideration a
respect for social structure and culture, a certain degree of tolerance about human
behavior, and the strictures of the evolved human organism. But human behavior
can be constrained only so far. Choices that powerful political systems or leaders
leave open to ordinary people may be narrowed and the price for certain of those
options made very high. But people ultimately make their own choices, even as they
feel constrained, even as they buckle under, hating themselves almost as much as
they hate their oppressors. Thus, the limits of "law and order" are built into the
somatic mechanisms of behavior, and burnished by experience and cultural value
systems.
Social change, therefore, will always follow "too much violence" because such
change is the result of a feedback mechanism, with the reference level for "too
much" set by a working combination of biology and culture. If the resultant swing is
too far in the direction of law and order-that is, if crime is reduced and anti-crime
activities are
felt to be oppressive-the same kind of public and individual reaction occurs as
happened with the rise of the crime rate: when political power becomes too
oppressive, resistance will mount, requiring ever more oppression.
In short, “law and order” —whether it be “too much” or “too little" —will not
work if it is out of proportion to the cultural experience and expectations. Yet, police
systems have to be improved and their organization changed as society becomes
more complicated and culture more developed. The imbalance is probably a constant
feature.
CORRECTING THE EXPERIENCE
Every society has ideas about which particular violent behavior is understandable
and forgivable. It would seem that this point puts the scientists in the same boat as
the victims and the violators. Both experience the biological bases of violence and
the cultural and social context of violent behavior. For the scientist, all violent
behavior in which he can empathize with both victim and perpetrator is "under-
standable." If he can empathize with only the victim, then it is not. The standards of
empathy may be-but may not be-quite different for the violent criminal or the
victim.
As we all know, there are only two or three ways of "correcting"' experience. One is
by therapy, whether it be insight-based psycho- therapy, or social learning therapy,
or operant conditioning. A second is by a sort of conversion experience-it need not
be a religious one- that casts the past and one's part in it into a completely new light,
causing the individual to think about his past experience in different terms, and
hence changing both expectations and behavior. The third would seem to be
extensive travel and immersion in a foreign culture, which provides a background
and a basis for a new view of the self .
There is some doubt in my mind about a fourth alternative: the power of education
in altering experience. Freud said some place that he had trouble remembering what
he had learned and just as much trouble forgetting what he had experienced. Yet I
am convinced that without some way of purposefully altering experience-that is,
getting people to evaluate their personal history and their current lives by new and
different criteria-we cannot fully succeed in altering behavior in such away as to
arrive at our social and cultural goals.
CORRECTING BEHAVIOR
Behavior seems not to be an independent variable in the way the foregoing topics
are. Violent, aggressive behavior is, apparently, to be controlled by manipulating the
other variables. Most often, attempts to change behavior have built on the axiom that
if you change the soma, or the culture, or the society, or the way we comprehend our
experience, then behavior will "automatically" change.
I have found it convenient to use a simple tetrahedral model to examine these
matters. The advantage is that a tetrahedron is a solid figure with four sides, each of
which touches the others. The sides of the tetrahedron can be labeled: soma, society,
culture and experience. Anyone who wants to try to control aggressive behavior is
very likely to land firstly on one of the sides. Too often they forget that there are
other sides, and that all the sides are connected. It may be that you cannot change
aggressive behavior by changing only one side of the tetrahedron: every dimension
may have to be altered. Each of these sides has been said to "cause" violent
behavior, and attempts have been seriously made to try to alter them often in
complete disregard for the other dimensions of the problem. Very few attempts have
been made to change two dimensions, let alone the entire figure: almost nobody
works on all the faces at once.
It is my contention that one of the problems underlying this conference is how to
consider all the surfaces and angles of that tetrahedron (on which the biophysical
dimension of man touches on every other dimension), as we discuss and evaluate
human legal behavior.
In the rest of this paper, I shall review briefly how law fits into this matter of dealing
with or altering aggressive behavior.
AGGRESSIVE BEHAVIOR AND THE LAW

At this conference we are involved in examining the relationship between biological
explanations of behavior and a particular set of cultural norms called "laws." Among
the different kinds of somatic- ally based behavior that require the attention of the
legal profession, aggression ranks among the most important. In studying aggression
in its multidimensional aspects, a major goal is to find ways to control and utilize the
aggressive capacities of the human being for pro-social ends. We have to be sure
that we involve every surface of the tetrahedron. A sub-question is, then: what is the
relationship between
.
Gruter and Bohannan [I 57]
the biological dimensions of aggression { or of every other kind of behavior) and the
functions of law?
Law can be seen as a cultural device that evolved to do two things at once: {1)
socialy it turns a conflicted or "adversary" dyadic relationship into a triadic group in
which a third party interferes in order to "solve" the conflict, thereby getting two
surfaces {culture and society) into the picture, then {2) it reinstitutionalizes some of
the norms of basic institutions like the family, the business firm or the political
group into the realm of a more complicated, secondary institution, the jural
institution. {This process of reinstitutionalization may, of course, work backwards-in
complex societies such as our own, many innovations begin in the jural and
legislative institutions and are then required to be institutionalized in the primary
institutions.)
Thus, the law is a cultural means of controlling social relationships in such away as
to reduce physical aggression or solve the unacceptable results of aggression.
Indeed, the law, as it is perceived in the Western world, is itself a very interesting
tool to contain our biologically based aggression. It is founded, in Western societies,
on what we call the adversary process {with aggressive behavior by competing
parties), and it is triadic in nature: two-party conflicts are settled by the interference
of a third party in the person of policemen and judges. Obviously, we know from
legal anthropology that there are a lot of jural mechanisms for controlling
aggression, even in our own society, that are not in fact triadic. But triadic forms are
nevertheless central to all highly developed legal systems. Thus, if the principals do
not bring their case before a judge, thus turning their dispute into a triad, the state or
some other organization will do so. The law in complex societies supercedes
disputes, but one of its basic concerns is with dyadic disputes-and it deals with them
in a triadic way.
Aggression in the two-group can lead to death, to flight or to relationships of
dominance and submission. The aggression mayor may not be expressed as
violence-indeed even violent acts of domination may be followed by reassurances or
atonement. But when a third party is brought in, whether as judge or as peacemaker,
the three- group is a solution to hostility in two-groups.
What, then, is aggression? Aggression among human beings is a drive originating in
the brain; it may be triggered by hormonal and chemical processes; it may be
inhibited or expressed under the influence of cultural values, social structures and
past experience {the latter of which may include cost-benefit calculations
concerning the social and cultural sanctions of a specific piece of behavior).
Adversary
relationships are social relationships in which two aggressive persons claim

conflicting rights. Bellicosity is the cultural norm of using aggression more or less
freely to achieve given ends. Conflict can occur in the drives, in the experience, in
the society or in the culture.
We are therefore talking about at least five different, if inter- connected, things: (1)
the drive of aggression, as it is or is not (2) expressed in hostile behavior, in (3)
adversary social relationships more or less in accordance with ( 4) bellicose or
peaceful cultural values, and (5) the experience of the organism. When the law deals
with aggression, it must necessarily deal with all of them, including the biological
processes that are part of the experience.
In this view, the legal system is a cultural device for reducing hostile behavior to
predictable social dyads in accordance with a set of cultural values that can be more
or less bellicose. It sometimes takes experience into account, as it does when it
defines extenuating circumstances. However, so far scholars and practitioners of law
have paid little attention to the biological dimensions vf aggression. Yet, effective
legal sanctions-"good law"-should surely take all dimensions into account along
with the other aspects. Legal behavior has foundations in biology just as it has in
society and culture and psychology and history.
Law already deals with aggressive behavior on some levels. That it does not yet deal
adequately with the somatic or, biological dimension seems to concern all of us who
are here. Law is traditionally concerned with behavior, with the principles and the
content of social relationships, and with cultural values about peace, stability and
conflict. So far little attention has been paid in the legal context to analysis of the
biological foundation of aggression. But the facts are that this biological trait can be
used for pro-social ends, that law is one of the fundamental human means for
insuring a livable society, and that we need all the help from the profession of the
law we can get.
[159]
Legal and primary-group social controls

DONALD T. CAMPBELL
Department of Social Relations,
Lehigh University, Bethlehem, PA I8oI5, USA
The ultrasociality of human beings, unlike that of the social insects, is beset
by competition among social cooperators. Four social mechanisms to deal
with this problem are discussed: mutual monitoring, internalized restraint,
legal control and market mechanisms. Each is investigated in sociobiological
perspective.
This essay is a preliminary sketch of how sociobiology may be united with the
traditional sociological issues of social control and bureaucracy. Human beings, like
all fertile animals, are in direct competition with their kin (except their identical
twins) and neighbors for the food and shelter available in their environment. This
ubiquitous condition creates an obvious interest in controlling how other individuals
behave, best elaborated in David Wilson's (1980) deterrence theory. It can be
analyzed as a conflict of interests between the group-as-a-whole and each individual
as to how that individual should behave, thus getting into the problem of social
control. However, I believe it is more appropriate to reserve that mode of analysis
for the conflict of interests that emerges with social and ultrasocial forms of life.
In a wide range of ecologies, cooperation such as big animal hunting and irrigated
grain fields increases food resources and shelter from predators. But individuals are,
once again, in competition for the size of their shares of these increased resources.
At this level, however, a novel form of collective interest emerges: individual
competition for maximum share of the resources jeopardizes the benefits of
cooperation and the cooperative organization itself. Greedy quarrelling for
maximum share reduces the pool of resources to be shared. In ecologies where
cooperation can double or quadruple the per capita resources available, there is a
payoff for effective social control that protects the efficacy of cooperation from
individual greed. That such mechanisms are rare and fragile is the conclusion of
analyses coming both from the mathematical models of evolutionary biology
(Haldane, 1932; Williams, 1966; Wilson, 1975; Chapter 5; Wilson, 1980; Boorman
& Levitt, 1980), and the social sciences (Von Neumann &
.
Morgenstern, 1944; Hardin, 1968; Olson, 1968, Schelling, 1971). Masters {this
volume) presents the issues in terms of "prisoner's dilemma" examples. My own
point of view {Campbell, 1972. 1975, 1979, 1982) can be summarized thus.
Ultrasociality refers to the most social of animal organizations, with full time
division of labor, specialists who gather no food but are fed by others, effective
sharing of information about sources of food and danger, self-sacrificial effort in
collective defense. This level has been achieved by ants, termites and humans in
several scattered archaic city-states. Inclusive fitness, kin-selection, and structured
deme theory {Wilson, 1980) adequately explain moderately social forms such as the
semi social wasps and baboons. In the social insects, the further route to
ultrasociality has been achieved by caste sterility, almost entirely removing genetic
competition among the cooperators; this route has not been available for human
urban societies. Instead, cultural evolution {including norms inhibiting human
selfishness, deceitfulness and cowardice) has been required.
The human route to ultrasociality is not fully understood, but a detailed
examination of present understanding of the route of the termites {extreme
inbreeding alternating occasionally with outbreeding which produces a generation of
siblings more closely related to leach other than to own offspring, making possible
caste sterility) and ; the route of the ants {haplodiploidy, which makes sisters more
closely ; related to each other than to own offspring, which when combined with sex
ratios favoring females, stabilizes brood care for parental ; offspring at expense of
own fertility) makes clear that the human route must have been different-it did not
involve the sterility of the cooperating specialities. Among the conceptual tool
available for reconstructing the human route are reciprocal altruism {clique
selfishness), moralistic aggression to punish defectors from reciprocal-altruist pacts,
the in-group as a socially inherited reciprocal-altruist pact, socially evolved beliefs
promising transcendant purposes, posthumous rewards for altruistic contribution to
group welfare at own expense, and transcendent sanctions against self-serving
behavior that jeopardizes group welfare. In general, biologically evolved supports
for preferences for altruistic behavior on the part of other group members do not
have the costs to inclusive fitness that tendencies to own self- sacrificial altruism
have. Moral norms, socially evolved with or
.
without biological support, thus tend to be focused on other persons' behavior, not
one’s own..
Much hypothesized cultural evolution must achieve a kind of "group selection"
precluded among vertebrates at the purely biological level and achieved by
invertebrates only through caste sterility. " The models of cultural evolution of Boyd
& Richerson (1980) help here. Non-linear, multiple-social-parent transmission, with
a majority amplifying effect, pushes face-to-face groups to internal unanimity in the
absence of selection. This provides the raw material of ingroup homogeneity and
group-to-group heterogeneity prerequisite for group selection. Such selection would
come through differential group success, differential growth, conquest with cultural
imposition, voluntary attraction of converts, imitation etc. As a byproduct, striking
group-to-group differences also occur in functionally neutral beliefs and customs.
These acquire a secondary function as indicators of ingroup membership,
designating fellow reciprocal-altruists of the same clique.
This perspective on human social evolution is more loyal to the details of biological
evolution than are current over-emphases on kin selection theory which by omission
imply that human ultrasociality can be explained by the same evolutionary
mechanisms that explain the social insects. This more complex understanding of
human evolution produces a sociobiology much more readily reconciled with the
traditional understandings of the social sciences and the humanities. Light is thrown
on human ambivalence, deceit, cowardice, disloyalty; on the specific content of lists
of sins, commandments and taboos; on human intuitions of justice, equality and
equity; on the dynamics of ethnocentrism, nationalism and war; and on self-seeking
and nepotistic distortions of collectively-rational bureaucratic roles.
Emphasis on group selection is central to my argument. It is a minority point of
view. The issue was the focus of an important half- day subgroup discussion at
Monterey Dunes involving Alexander, Durham, Masters and myself, focused on
Durham's paper for illustrative purposes. The discussion has led me to back off
somewhat from an implication of my flamboyant title of 1975, "On the Conflicts
between Biological and Social Evolution and between Psychology and Moral
Tradition." For most sociobiologists and evolutionary theorists (Alexander, 1979),
sociocultural evolution cannot persistently produce behavior tendencies that result in
a net loss of biological inclusive fitness. Cultural evolution is dependent on
biologically evolved capacities; cultural items that reduced inclusive fitness would
never have
evolved. Cultural evolution thus cannot conflict with biological evolution or reduce
biological inclusive fitness. I should have emphasized the conflict between
behavioral tendencies produced by biological evolution and those produced by
social evolution as mediated through moral indoctrination, beliefs about the
supernatural, and social organization. Some degree of group selection is always
going on. Haldane's and subsequent analyses point out that behavior traits that
benefit the group but involve sacrifice of individual inclusive fitness will be
undermined by individual competition within the group. The benefits to average
individual inclusive fitness that could come from self- sacrificially altruistic social
cooperation are thus precluded by the collectively self-defeating "genetic
competition among the cooperators" (Haldane, 1932; Williams, 1966). Any
mechanism that can overcome this self-defeating tendency will result in great gains
in biological inclusive fitness for the average gene, the average individual, the
breeding pool and the organized group. Therefore, cultural evolution of behavior
tendencies that furthers group effective self- sacrificial altruism would be strongly
selected for. On the other hand, as Hamilton ( 1964) has emphasized, the stronger
these tendencies, and the more collective prosperity they produce, the greater the
inclusive fitness payoff to being a successful "free rider" (Olson, 1968), parasitical
on the cooperative efforts of others without risking the costs to own inclusive
fitness.
Let us examine the issue in terms of a skeletonized model of the cultural evolution
of a general moral preaching. Assume some degree of "direct" effect of teaching-that
people tend, however slightly, to believe what they are told, and act according to
their beliefs. According to the mathematical models of Boyd & Richerson (1980), a
cultural evolution of moral preachments from parents to offspring under an analog
of individual selection social transmission would produce moral preachings of the
nature "Get others to cooperate but be a free-rider yourself." "Don't be a sucker."
"Better a procreating coward than a dead hero." When this model is expanded to
include multiple, across- lineage, social-transmission of "parenting," the same type
of moral teaching is favored in their linear model with individual selection. Only in
their non-linear group-selection of teaching and preaching do we get conditions that
would select for public moralizing that favors group advantage at individual cost.
No doubt self-serving preachings such as the above often go on in the privacy of
homes, directly or indirectly, consciously or unconsciously. We can envisage on an
individual selection basis a double
standard of preaching. an altruistic morality for exhortation to others, a self-serving

one for own offspring. I anticipate that in the long run such a system would not work
to produce complex social coordination, even though it would end up with the
altruistic preachings heard by the offspring generation being many times more
numerous than the selfish ones. I am more hopeful for another analysis: if one's own
private preachings of selfish opportunism to one's offspring undermined the
tendency of others to behave cooperatively, and if the payoffs for cooperation were
high, situations could occur in which it would be of net inclusive fitness advantage
to preach the altruistic message to one's own offspring even if they behaved
accordingly. Does it require something like group selection to create a situation in
which such a choice is presented to an individual? Or can it be achieved purely on
an individual selection basis?
At Monterey Dunes we did not discuss the group selection issue for the termites and
ants, but I have elsewhere been challenged by Alexander for upholding the old-
fashioned thesis that group selection is required for their ultrasociality too. With due
attention to a considerable portion of the relevant literature I have come (Campbell,
1982) to the conclusion that where proto-social insects find themselves in an
ecology in which multiple brood care is required, it becomes in the inclusive fitness
interests of an auxiliary brood carer to distribute a fertility inhibiting pheromone to
fellow auxiliary brood-carers even at the expense of having own fertility inhibited.
At this initial stage the insects are only semi-social, the mother or sister "queen" still
gathers food for herself and for the immature individuals just as do the sterile
"workers." There is no structural differentiation or behavioral division of labor
except for fertility. Preserving this sterile worker caste effectively removes genetic
competition among the cooperators, and makes possible the subsequent evolution of
ultrasociality, with the adult queen and adult soldiers being fed by the adult workers.
It is the preservation of this sterility that makes the unit of selection the cooperating
social nest, including the fertile queen. Selection for socially effective self-sacrificial
altruism is not undermined be genetic competition among the cooperators, since the
cooperators are all sterile. Now in the formal mathematical models of "group
selection" in population genetics, only groups of fertile organisms are considered;
since there is no group selection for groups of queens and nests (indeed
they compete most ruthlessly), the social insects are rejected as instances of group
selection. I must be more careful about my terminology. What I and Wynne-
Edwards (1962) and William Morton Wheeler (1928) were talking about is in
today's usage better called "selection by social organization and not by participating
individual" (but the word "group" used to be a near synonym for social
organization).
SOCIAL CONTROL MECHANISMS
Mechanisms of social control serve two functions: coordination and restraint.
Coordination is the function of routing, routinizing and information sharing which
reduces chaos and maximizes coordination in ways that do not oppose human nature
or individual preference. As a pure type, coordination is supported by individual-
dispositional preferences, assisting the individual to choose from a set of neutral
behavioral alternatives the one that optimizes group function, the one that is for this
reason preferred by the individual once designated, for it optimizes a group function
from which the individual profits, and at no individual cost relative to other
alternatives.
Social controls involving restraint also focus on optimizing efficacious collective
action. But in contrast to coordination, they appear in situations in which individual
dispositions favor some action other than the collectively optimal one. As we have
seen, given the vertebrate ecology of genetic competition among cooperators,
evolutionary biology makes some degree of such bias inevitable, and directly
predicts the direction of the individual-dispositional bias against which the restraint
is directed. Thus food sharing is essential for division-of-labor social organizations:
genetic competition predicts a bias in the direction of under-generosity, not over-
generosity. Truth telling is essential for the information sharing which produces the
great economy of cognition (Campbell, 1965b) which makes social life so much
more productive than individual-the direction of bias is for self-serving dishonesty,
not for self-sacrificial dishonesty. Audits on tax returns are based on anticipated bias
in the direction of under-paying, not overpaying. A bureaucrat's personnel choices
will over-favor own children and other kin, rather than be biased against them. Self-
saving cowardice in battle is more common than foolhardy bravery. All of these
obvious and well-known biases have not been able to eliminate genetic competition
among the cooperators. All create problems of social control of the type I have
designated as restrain
For purposes of this paper, four mechanisms of social control can be delineated.
(1) Mutual monitoring
This includes face-to-face approval and disapproval, ostracism, conformity pressure,
shame and pride. All group members share in both detecting violations and
enforcing sanctions.
(2) Internalized restraint
This category includes processes of conscience or superego, the pain of guilt feelings,
and the fear of supernatural sanctions. There is self- monitoring of norm violation
and self-punishment.
(3) Legal control
Overt rules about offenses and punishments are in this mode. The detection of
violations and the enforcement of penalties are delegated to specialists such as
police, militia, tax collectors and judiciary. Sanctions include job loss,
imprisonment, fines, exile and death. Rational bureaucratic systems are included in
this category, as is government by administrative regulation.
(4) Market mechanisms
In such processes the intelligently selfish choices of all individuals curb the greed of
individuals by making it unprofitable, as in the "invisible hand" of laissez-faire
economics and libertarian political theory.
The last two, legal control and market mechanism, are currently the focal alternative
poles being advocated in political discussion. Western democracies are based on a
compromised mixture of the two. It seems to me that sociobiology has an important
critique to make of each of them.
The first two, mutual monitoring and internalized restraint, seem unduly neglected
in modern considerations of legal control and market mechanisms. Including them
will help us understand both why the last two have worked when they have, and why
they so often fail.
The first three, mutual monitoring, internalized restraints and legal control,
represent to some extent an evolutionary sequence, partly one of biological
evolution, but mainly a product of cultural evolution.
Mutual monitoring covers most of the "primary group" or "face-to- face" group
social controls described in the older sociology. The
anthropological concept of "shame" culture overlaps. Primate sociality as reviewed

by Boehm, Goodall, Gruter and Itani {this volume) exists, at this level. Trivers' {
1971) concepts of an innate predisposition to ; form "reciprocal altruist" pacts, and
to show "moralistic aggression" when such compacts are violated, and Axelrod and
Hamilton's {1981) "tit-for-tat" contingent cooperation and retaliation belong here.
These are important concepts urgently needing elaboration beyond the dyad to
multi-person groups. The ethological study of human facial expression, the
autonomic nervous system and hormonal reactions associated with face-to-face
disapproval, opinion-minority status and lying all fit in here.
In my judgment, it will turn out that many of the mechanisms that make mutual
monitoring effective are stubbornly innate in human beings, and that they can be
counted on to be at work creating a kind of ingroup solidarity, homogeneity of
belief, and discipline in even arbitrarily assembled aggregates of persons who
repeatedly interact in small groups. Indeed, research on social processes in
experimental and social psychology laboratories show that such group formation
processes begin to occur in as little as two hours. This fact is of extreme importance
in understanding the dynamics of bureaucracy and large- scale organizations, but is
still overlooked in human organizational theory.
In practice, reciprocal altruism can be translated as clique selfishness. While it
may be true that a complete theory of games analysis would show cooperation to be
mutually beneficial even if everyone were in the cooperating group, in practice
reciprocal-altruist pacts and the inhibition of parasitical free-riding by mutual
monitoring are the most common and most feasible for small groups, as the
economist Olson {1968) has shown. In addition, both the lower primates and
humans achieve their group solidarity in a context of competition with, or threat
from, other groups of conspecifics. The formation of an ingroup solidarity is always
accompanied by an outgroup hostility, as ubiquitously noted in studies of
ethnocentrism {LeVine & Campbell, 1972). One of the most ubiquitous principles
of that literature is that ingroup social control is enhanced under conditions of
outgroup threat.
It is the common circumstance of modern social organization, in public or private
bureaucracies, that they are made up of many separate face-to-face groups,
connected by messages and messengers. It follows from our mutual monitoring
principle that each face-to-face assemblage tends to become an ingroup whose
solidarity tends to be motivated by treating other units as outgroups. In this process,
orders
from higher administration can be reacted to as though they were enemy

impositions, and cooperative efforts to subvert them can be supported by mutual
monitoring. The engineering department and the sales department can play ingroup
and outgroup. roles to each other. Customers and clients can become outgroups
disrupting cozy ingroup patterns. Given human nature, both as observed and
predicted from evolutionary tendencies, such occurrences are not occasional,
isolated instances, but are unavoidable, universal tendencies. If they fail to produce
major problems for large organizations it is because other factors keep them in
check, not because they are absent.
Thus mutual monitoring as a means of social control is effective at the small group
level, but can easily become organized around purposes that are contrary to the
larger group's collective interests. Keeping these interests co-aligned is one of the
major unsolved problems of organizational design. Max Weber gave us a theory of
bureaucratic rationality that failed to take these processes into account. Some such
optimistic assumption that ''as it is planned so shall it be carried out seems endemic
among legislators and administrative designers as ever larger bureaucratic structures
get created. Yet actual studies of bureaucracies (e.g. Blau, 1963) support the popular
concept of inflexible, lethargic, self-serving bureaucracy. Sociobiology thus
provides two grounds for predicting distortions of bureaucratic rationality. The first
is the individual's selfish and nepotistic biases, particularly biasing when
exemplified by those in high administrative rank. The second is the clique
selfishness, the tendency for face-to-face ingroup formation with clique solidarity
interests athwart those of the larger collective.
Bureaucracies, large organizations, states (Masters, this volume) would work with
bureaucratic rationality if the only control requirements were those of coordination,
none of restraint, if individuals had no interests divergent from those of the
collective, if they were self- guiding and self-monitoring solely in terms of collective
interests at the highest level of the social organization in which they participate.
While it is turn-of-the-century-old-fashioned to use the social insects in such
discussions (Campbell, 1975), I do find in them an enlightening comparison as to
what human beings are not but conceivably could have been-a role that science
fiction can also play. The ants, termites and bees are such self-monitoring automata
in their social systems, executing their micro-purposes with extemporaneous
intelligence in the face of local obstacles, micro-purposes which fit into generally
competent collective action (in the absence of such novel
.
ecological features as ant poison). The 12 or so pheromones with which they

exchange signals {Wilson, 1971) have the function of coordination, not restraint.
None of their division of labor goes into monitoring each other’s behavior for
compliance. A major reason for this, as I see it, is that the elimination of genetic
competition among the cooperators has made the inclusive fitness interests of each
worker and soldier exactly that of collective nest or hive survival.
Internalized restraint, traditionally achieved through religion and the awe- and fear-
inspired morality accompanying it, has been the central focus of my major
contribution to sociobiology { Campbell, 1975, 1965a, 1965b, 1972, 1979, 1982). A
dozen or so archaic city- states and nations {each probably independently) achieved
ultrasociality, with fulltime priests, governors and soldiers who gathered no food,
being fed by the workers, with granaries and large civic buildings and often with
apartment-house concentrations. All were theocracies. All invested human effort
heavily in temples, funeral monuments and graves for their rulers. All believed in
supernatural gods and god- I stories far more incredible {from a modern scientific
point of view) J than those of the simpler human societies that have remained for
anthropologists to study-and those are incredible enough. The economist and
evolutionist must ask what inclusive fitness function led to the selection of this
apparently lavish waste of human energy, tools and the sacrifice of useful
domesticated animals and servants. These belief and action systems were obviously
products of a cultural evolutionary process.
While the Boyd arid Richerson {1980) model predicts the occurrence of bizarre
beliefs that are neither functional nor dysfunctional, it would not predict such a
widespread uniformity of such apparent violations of economic and biological
efficiency. We are thus required to seek a function. I see that function in achieving
social control of the kind of restraint that is effective {to some degree) even when
human supervisors and policemen and worldly rewards and punishments are not
present to shape individual behavior in the collectively optimal form. The beliefs
about Valhallas that reward brave soldiers killed in battle, or the hells that punish
cowards, thieves and liars, were legitimated and made more credible by the royal
funeral waste testifying to the leaders' belief in an afterlife. The fit between the
biases in human nature {that follow from the fact of genetic competition among
competitors) and lists of religious sin and temptation explicit or implied in the
commandments is also convincing. The emphasis on carnal, biological, human
nature in the Christian tradition
(at least) further fits. The genes say "Thou shalt covet"; ultra-social human culture
says (or used to say) "Thou shalt not covet." While the detailed picture is more
complex and confusing than this, I know of no better evolutionary explanation for
religious moralizing and temple building or for the current energy investment in
religion, even though other functions are no doubt also served.
Even though self-monitoring, moral, dutiful persons (whose effective altruism goes
far beyond situational opportunism or long range hedonism) come out of non-
religious families, the general culture from which they come has both religious
ancestors and neighbors. But whether or not a religious justification is required for
such a culturally- induced self-monitoring restraint, internalized moral norms
actually acted upon are of obvious use in a social organization. The seeds of world
tragedy, of great losses in current human inclusive fitness such as Alexander (this
volume) refers to, lie in the fact that self-monitoring internalized social control
systems are necessary for effectiveness in national warfare, and are enhanced (just as
is mutual monitoring) by ingroup-outgroup polarization between religions or
nationalistic versions of religion. Tragically, nation-worship seems less incredible,
less supernatural, to the modern secularized mind than do the reifications of
collective interests found in the great religions of purportedly universal scope.
Legal control is a universal feature of all modern nations, going hand in hand with
administrative bureaucracy. It has worked reasonably well in many nations for many
historical periods. It is not conspicuously successful in most of the new third world
nations, and its failure is being announced in some well developed ones, including
such one- time paragons of national success as England and the United States. Given
our sociobiological model of human nature, it is remarkable that it ever worked. As
a pure form, without mutual monitoring or internalized restraint, with all detection
of non-compliance and delivery of sanctions delegated to specialists, the required
size of this specialist corps becomes unaffordably large, even if the self-interest and
nepotistic biases of these legal-control specialists is disregarded.
The judicial system is more heavily used to further self-and-nepotistic interests than
to curb them. The concept of "loophole," the literal legalistic interpretations of the
law that subvert announced legislative intent, illustrate one aspect of the problem.
Language is an inherently imperfect tool, which is used effectively only when shared
contexts fill in the meaning, as the concept of "indexicality" points out (Putnam,
1975; Barnes & Law, 1976). But law in its effort to
achieve universality and fairness must pretend that words have situation-free
constant meanings, denying their indexicality. Written and interpreted "literally," the
wordings allow loopholes and unintended uses never envisaged in their construction.
Perhaps more than half of our lawyers and legal research efforts are now employed
to serve collectively biasing individual interests.
One must conclude that legal social control can only have worked well when
supported by mutual monitoring and internalized restraint. This internalized
restraint would have to cover not only restraint from violating the letter of the law,
but also restraint from violating the public-interest announced intent of the law. It
would also have to involve restraint in use of the law against fellow citizens in the
service of opportunistic greed.
If the internalized restraint that makes legal social control work has to a
considerable degree been based on consciences coming from social indoctrination
employing transcendent religious belief, then secularization and also religious
pluralism in which ethical norms apply only to co-religionists, both may lead to self
defeating political efforts to increase the burden of social control delegated to legal
and bureaucratic processes.
Market mechanisms in which the intelligent selfishness of
all curbs the selfish greed of others, are the methods of social control recommended
by laisse-faire economics, in which the only governmental interference with human
nature allowed is the protection of private property and inherited wealth. Mancur
Olson (1968) employs the tools of mainstream economics which are characterized
by the hope that an "invisible hand" emerging from competition will provide the
restraints needed for collective action. But he shows that this hope is wrong, that
basic economic assumptions show that if each person is intelligently selfish in
choices, "collective goods," including the benefits of cooperation, will be lost. He
concludes that to solve the "free-rider" problem, compulsion is needed. For small
groups whose members cannot readily join and quit, mechanisms such as I have
included under mutual monitoring suffice. For larger groups, he judges legal
compulsion is required. Thus he recommends compulsory taxation and compulsory
union membership. I find his analysis compelling, and in keeping with sociobiology,
except for the unexamined assumption that legal compulsion can be made to work.
In agreement with Olson, I am inclined to reject the efficacy of market mechanisms,
even though I admire their efficacy for social system- tuning in labor supply
allocation. (The mixture of government
controlled intervention in income and services redistribution with j market

mechanisms, which Friedman ( 1962) has introduced in the "negative income tax"
and the "voucher system" for education and housing subsidies seem to me worth
thorough consideration.) Market mechanisms, like legal control, seem to have
worked well in some settings in the past. My conclusion is again that they only did
so when supported by the internalized moral restraints and pride which the culture of
traditional beliefs and mutual monitoring provided.
The task of thinking through the problems in social control that are created by
genetic competition among the cooperators has just begun.

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Law, Biology and Culture

The Evolution of Law

1 The foundations in law and morality

2 MARGARET GRUTER—Biologically based behavioral research and the

15 RICHARD D. SCHWARTZ—On the prospects of using sociobiology in the

7 E. ADAMSON HOEBEL—Anthropology, law and genetic inheritance

34 MANFRED REHBINDER—Questions of the legal scholar concerning the

47 The search for the missing pieces: in biology

50 JANE GOODALL—Order without law

62 JUNICHIRO ITANI—Intraspecific killing among non- human primates

74 PAUL D. MacLEAN—A triangular brief on the evolution of brain and law

90 HUBERT MARKL—Constraints on human behavior and the biological

101 RICHARD D. ALEXANDER—Biology and the moral paradoxes

111 BARTLEY G. HOEBEL—The neural and chemical basis of reward: new

129 The search for the missing pieces: in social science

134 CHRISTOPHER BOEHM—The evolutionary development of morality as

147 PAUL BOHANNAN—Some bases of aggression and their relationship to

159 DONALD T. CAMPBELL—Legal and primary-group social controls

171 ROGER D. MASTERS—Evolutionary biology, political theory and the

Richard D. Alexander- Museum of Zoology, Insect Division, University of

John H, Beckstrom -School of Law, Northwestern University; .357 East

Christopher Boehm –Dept. of Anthropology & Sociology, Northern

Paul Bohannan -Dean, Social Sciences and Communication, University of

Donald T. Campbell -Maxwell School of Citizenship and Public Affairs,

William Durham -Department of Anthropology, Stanford University;

Lawrence M. Friedman -School of Law, Stanford University; Stanford,

Jane Goodall- Post Office Box 727, Dar es Salaam, Tanzania

Joachim H. Gruter -158 Goya Road, Portola Valley, California 94025/USA

Margaret Gruter -158 Goya Road, Portola Valley, California 94025/USA

Bartley Hoebel -Department o/Psychology, Princeton University; Princeton,

E. Adamson Hoebel- 2273 Folwell Street, St. Paul, Minnesota 55108/USA

Helmut Hofer -Department of Zoology and Comparative Anatomy,

Jay G. Hook -Department of Psychology, University of Houston; Houston,

Junichiro ltani -Faculty of Science, Laboratory of Physical Anthropology,

Paul D. MacLean -Laboratory of Brain Evolution and Behavior; National

Hubert Markl -Department of Biology, University of Konstanz; Postfach

Roger D. Masters -Department of Government, Dartmouth College; Hanover,

Manfred Rehbinder -School of Law, University of Zurich; CH-8032 Zurich,

Ernst Schuermann -Goethe Institute, San Francisco, California 94100/USA

Richard D. Schwartz -College of Law, Syracuse University; Syracuse, New

Harvey Wheeler -Institute of Higher Studies, Carpinteria, California

MARGARET GRUTER and PAUL BOHANNAN

This first Monterey Dunes conference is an integral part of what is

Although we talked about it very little during the conference, many

The Foundations in Law and Morality

Biologically based behavioral research

Can we find precursors of human cultural characteristics among primates? Eugen

Ehrlich (1913) formulated his "Fundamental Principles of the Sociology of Law"

Can we discern functions of organization and protection, a kind of “inner order,” at

Biologists postulate that evolutionary changes {including the continuous

An essential part of this mechanism is the "internal reward system" {Routtenberg,

THE FACTS OF LAW

The oldest fact of law, according to Ehrlich's observations, is usage. "Usus" or

Order can be maintained in human societies by various means among them

Examples from non-human societies demonstrate the emergence of new behavioral

In Ehrlich's opinion, usage is determined by economic pressures and the results of

Ethological observations document changes in group behavior brought about by

non-human. The concept of dominance is related to Ehrlich's second fact of law.

Domination as a ìfact of Lawî

Another mechanism in animal societies that serves to protect weaker or "dominated"

Even in non-human primates, this behavior is not dictated absolutely by biological

It is possible that a sense of well-being similar to that originally

generated by hormonal changes during the childbearing period can be experienced

As new research in neuropharmacology elucidates the sites and causes of these