THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3,265-290

Printed in the United States ol America

Human suicide:
a biological perspective
Denys deCatanzaro
Department of Psychology, McMaster University, Hamilton, Ontario,
Canada L8S 4K1

Abstract: Human suicide presents a fundamental problem for the scientific analysis of behavior. This problem has been neither appreciated nor
confronted by research and theory. Almost all other behavior exhibited by humans and nonhumans can be viewed as supporting the behaving
organism's biological fitness and advancing the welfare of its genes. Yet suicide acts against these ends, and does so more directly and
unequivocally than any other form of rnaladaptivc behavior. Four heuristic models are presented here to account for suicide in an evolutionary
and sociobiological framework. The first model attributes suicide to the extraordinary development of learning and cultural evolution in the
human species. Learning may make human behavior so independent of biological constraints that it can occasionally assume a form entirely
contrary to the principles of biological evolution. The second model attributes suicide to a breakdown of adaptive mechanisms in extremely
stressful novel environments. The third model involves kin and group selection, arguing that in limited circumstances suicide may occur
because of beneficial effects it has on other, surviving individuals who share the suicidal individual's genes. The last model suggests that suicide
should be tolerated by evolution when it has no effect on the gene pool. This model holds particular promise in accounting for aspects of suicide
not attributable to culture. The evidence indicates that suicide is most common in individuals who are unlikely to reproduce and unable to
engage in productive activity; such individuals are least capable of promoting their genes. A complete explanation of suicide may derive only
from an analysis of its biological significance.

Keywords: suicide; self-destruction; sociobiology; evolution; altruism; life-threatening behavior

Human suicide, unlike almost all other behavior, acts directly
against the survival of the behaving organism. It consequently
presents a special challenge for general theories of behavior
and a particular one for current sociobiological theory. It is
generally held that evolution selects for behavior that
promotes the existence of the organism and its genes, yet
suicide, which acts directly against these ends, is not uncom-
mon. The purpose of the present discussion is to examine the
clinical and sociological literature on suicide from a biological
and evolutionary perspective. The anomaly that suicide pres-
ents will be outlined, and some heuristic models that might
explain this anomaly will be presented. The feasibility of
conducting research to test the validity of evolutionary
models will then be discussed, and some specific research
strategies will be suggested.
Evolution and behavior
Much attention has recently been given to the relation
between behavior and evolution (see reviews by Barash 1977;
Dawkins 1976; Maynard Smith 1978; Parker 1978; Wilson
1975). It has been explicitly or implicitly held that, in general,
an organism's behavior is oriented toward promoting its
genes. It is reasoned that behavioral predispositions conducive
to the welfare of the organism's genes will survive in natural
selection, while less adaptive predispositions will be elimi-
nated from the gene pool. This is not a teleological position,
attributing "purpose" to behavior, but rather suggests which
types of behavioral predispositions survive in natural selec-
tion. Nor does it entail that an individual is "conscious" of the
effect of his behavior on his genes, but rather that he will
probably inherit factors leading his behavior to advance his
genes.
Any behavior supporting the individual's survival and
reproduction will probably promote his genes. If an individ-
ual survives through reproductive age, producing and foster-
ing children, any genes contributing to his health and repro-
duction will survive into future generations. However, an
individual's behavior may contribute to the welfare of his
genes despite the fact that he may not reproduce. This can
occur because other related individuals share some of his
genes, and his behavior may promote the survival and repro-
duction of those individuals (see Hamilton 1964). According-
ly, the concept of "inclusive fitness" has been developed. The
inclusive fitness of an organism encompasses not only his
individual biological fitness but also the fitness of his genes as
they exist in other organisms. An individual may also behave
for the benefit of unrelated individuals not sharing his genes;
this has been explained through the concept of "reciprocal
altruism," wherein some tacit or explicit contract ensures
reciprocity (Trivers 1971).
An overview of human and other animal behavioral
patterns suggests that almost all such patterns readily fit this
framework. Almost universally, organisms actively avoid
circumstances that threaten their existence or health, unless
they have not learned or are incapable of learning the
negative contingencies of such circumstances. Conversely,
organisms generally seek situations in which food, water,
shelter, physical comfort, and the like are available to
improve their welfare; they also invest a considerable amount
of time and energy in reproductive activity. It is not difficult
to explain such behavior. Through natural selection, orga-
nisms that behave in ways that are conducive to survival and
reproduction will pass on their genes, while the genes of
organisms that do not do so will be eliminated from the gene
pool. Thus any genes predisposing toward behavior that
promotes survival and reproduction will have a selective
advantage over other genes.
In adopting these notions one does not assume that all

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 deCatanzaro: Biology of suicide
behavior is under direct genetic control. Behavioral pheno-
types may be strongly influenced by conditioning processes,
which select for adaptive behavior within each individual
without requiring genetic change. The learning processes
themselves, rather than specific response topographies, may
have been selected by evolution; indeed, the uniformity of
learning processes across individuals and species (cf. Hinde
and Stevenson-Hinde 1973; Nevin and Reynolds 1973) and
indications of nonassociative physiological mediation (see
Gallistel 1973; Mogenson and Phillips 1976; Stein 1978)
suggest that such processes have an inherited, biological basis.
Also, many motivational variables that interact with learning
may be influenced by innate factors. This follows from
evidence of nonassociative physiological mediation of aspects
of feeding, drinking, sexual behavior, aggression, pain avoid-
ance, and emotion (see Barchas, Akil, Elliott, Holman, and
Watson 1978; Carlson 1977; Conner 1972; Gorzalka and
Mogenson 1977; Schildkraut and Kety 1967; Stein 1978). It is
thus likely that behavioral development involves complex
interactions between learning and innate variables; the innate
component should reflect countless generations of selection
for adaptiveness, and we should thus be able to account for
most behavior in terms of evolutionary contingencies that
affect it.
The problem of suicide
Suicide is anomalous within this evolutionary perspective. In
contrast to almost all other behavior, suicidal behavior acts to
decrease rather than increase the biological fitness of the
behaving organism. Indeed, suicide is behavior operating
directly against the behaving organism's survival. Since
behavior oriented toward propagating the organism's genes
should generally promote the organism's existence, suicide
requires a special explanation in behavioral theory.
A cursory analysis of the problem might suggest that
suicide should be very rare. Individuals committing suicide
remove their genes, at least those carried specifically by the
individuals themselves, from the gene pool. Consequently,
any genes that they carry that permit or support suicidal
behavior should be selected against. Conversely, individuals
possessing genes acting to reduce the likelihood of suicidal
behavior should have a strong selective advantage over those
lacking such genes. Given such selection pressures, suicide
should be at best an infrequent event. Yet suicide is a fairly
common cause of death in humans, while a number of other,
"subsuicidal" behavioral patterns in humans, also acting
against the behaving individual's survival, may be even more
common.
Suicidal behavior occurs in diverse forms. While there have
been a number of attempts to develop nosological systems
(e.g. Durkheim 1951; Pokorny 1974; Shneidman 1968; 1969),
there is no universally accepted framework for classifying
suicidal acts. Many investigators (e.g. Kreitman 1977; Stengel
1973) feel tha,t the factors underlying completed suicide
differ from those underlying attempted but unsuccessful
suicide, in that the latter involves methods with a low proba-
bility of success and may relate to reinforcement accruing to
the surviving individual. Others (e.g. Menninger 1938;
Roberts 1975) have suggested that a broad range of life-
threatening behavioral patterns, such as excessive risk taking,
alcoholism, and heavy consumption of other drugs, must be
included in a definition of suicide. Menninger (1938) has.
called these "chronic suicide," since the self-destructive
behavior is dispersed over time. Shneidman (1969) has spoken
of "sub-intentioned death," which involves the individual's
playing some partial, covert, or unconscious role in facilitat-
ing his own death; he has suggested that this may be charac-
teristic of a large number of human deaths. There is also
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stereotyped, repetitive self-injurious behavior, which occurs
in schizophrenic, retarded, and other psychiatric populations
and which may or may not bear some resemblance to suicidal
behavior (deCatanzaro 1978; 1979; deCatanzaro and Baldwin
1978). All of these behavioral patterns have as their major
apparent consequence a self-produced decrease in biological
fitness.
In the present discussion, unless otherwise specified, the
term "suicide " refers only to completed suicide. Completed
suicide is perhaps the most unambiguous form of self-destruc-
tive behavior, in that it is difficult to conceive of any
secondary gain that could accrue to an individual not surviv-
ing his suicidal act.
Suicide has been evident throughout recorded human
history (Farberow 1972). Records from early Western civili-
zation suggest that it occurred frequently among the Greeks,
Romans, and early Christians, and perhaps to a lesser extent
among the Jews. In the Orient, it was reported in early
Brahman and Buddhist writings and condemmed with sever-
ity in Muhammadan teachings. Suicide occurs in a wide
variety of human cultures today, with varying frequency,
including such diverse groups as North American Indians,
Europeans, Chinese, Japanese, and East Indians (see Farbe-
row 1975).
Suicide currently accounts for a substantial portion of all
human deaths. Overall, it ranked as the ninth most frequent
cause of death in the United States in 1975, occurring at a rate
of 12.7 per 100,000 population (Demographic Yearbook
1976-1977). Rates in many other countries are considerably
higher, and generally are greatest in the developed countries.
In 1975, the highest rates were in Hungary, with 38.4, and
Finland, with 25.1 per 100,000 population. Generally, men
commit suicide more frequently than women, while the
frequency of successful suicide increases as a function of age,
being most common in the elderly (Diggory 1976; Linden
and Breed 1976). However, suicide accounts for a larger
portion of all deaths in young adults, where in many countries
it is the third or fourth most frequent cause of death for that
age group.
These statistics reflect only the deaths that are formally
registered as suicide. Many investigators believe that this
represents a considerable underestimate (e.g. Stengel 1973).
Many suicides may be registered as accidental deaths to spare
relatives embarrassment, or because of uncertainty about the
cause. Moreover, these statistics reflect only completed
suicides attributable to discrete acts. If death resulting from
chronic subsuicidal behavioral patterns were included, the
rates for suicide would surely be much higher.
Models of suicide
On the basis of evolutionary theory and existing data on
suicide, some hypotheses that might explain the anomaly of
suicide can be suggested. These models outline potential
relationships among suicidal acts, pressures of natural selec-
tion, and documented proximate determinants of suicide.
They are developed here only for their heuristic value in
suggesting research strategies. The models explain suicide as
resulting from independence of learned behavior from natu-
ral selection, pathological conditions, actual selection favoring
suicide, or evolutionary tolerance of this behavior.
These may also be viewed not so much as models, but as
factors that could each contribute to the occurrence of
suicide. There is no reason to assume that the factors are
mutually exclusive; rather, it is likely that some combination
is involved.
I: Independence from biological evolution. The first
model of suicide attributes this behavior to the human species'

extraordinary development of the capacity to learn. Such
learning ability may make behavior sufficiently flexible and
independent of biological evolution to allow the occasional
maladaptive behavior, such as suicide, to arise.
In many respects, human suicide may be unique in the
animal kingdom. There are only limited examples of suicidal
behavior in other species, and these may be qualitatively
different from human suicide. Among insects, there are many
behavioral patterns in which the behaving insect dies in
protecting the social group to which it belongs. For example,
a honeybee worker will usually die when it stings. It is
believed that such self-sacrificing behavior occurs because it
improves the fitness of reproducing individuals sharing the
genes of the suicidal individual (Wilson 1971). Among
nonhuman vertebrates, however, clear examples of suicidal
behavior are lacking. The reputed mass-suicidal behavior of
the Norwegian lemming may actually be a population
dispersion phenomenon in which death is incidental to the
presence of major geographic barriers (see Clough 1965).
Some mammals, particularly primates, will, under limited
circumstances, engage in chronic self-injurious behavior, but
it is not established that this is suicidal (see deCatanzaro,
1978; 1979). Another exception may have occurred in a case
recently reported by Jane Goodall (1979), in which one
eight-year-old chimpanzee became lethargic, ceased eating,
and died following the death of his mother, although this is
subject to other interpretations. There may also be cases in
many species in which organisms sacrifice their lives for
offspring (see Hamilton 1964; 1970; 1971; Wilson 1975),
although human suicide also occurs in many other situations. It
remains possible that suicidal acts occur in other vertebrates
but are undocumented because of a lack of thorough
ethological and laboratory research. Current evidence,
however, suggests that behavior oriented directly toward the
behaving individual's death is almost uniquely human.
Human behavior in general can be distinguished from that
of other animals in a number of relevant respects, notably
communicative ability, complexity of the social structure, and
flexibility of the behavioral repertoire (Washburn 1978;
Wilson 1975). These differences have made possible the
development of cultural evolution, which permits changes in
behavior, social patterns, and technology without the direct
influence of biological evolution (see Alexander 1979;
Durham 1979; Irons 1979). Human behavior probably relies
on learning to an extent not paralleled in other species.
Learning allows changes in behavioral phenotypes within
each individual's lifetime, and thus engenders fine
adjustment of behavior to the environment. Many behavioral
patterns are thus neither directly controlled by genetics nor
subject to natural selection. Consequently, occasional
behavior that might not be adaptive in a biological sense, and
that could not arise through biological evolution, might occur
because of extraordinary learning experiences.
Accordingly, suicide may be an entirely learned behavior,
and its especially high incidence in humans may be
attributable to an extraordinary development of learning
capacity in this species. Skinner (1969) argued briefly that
suicide must result from learning. He reasoned that any
genetic mutation favoring suicide would quickly eliminate
itself, and that therefore suicide must result from ontogenetic
rather than phylogenetic contingencies. Much of the available
evidence concerning suicide can be construed as support for a
model based on learning and cultural evolution.
At the very least, many of the methods employed to
commit suicide must be learned. Most common methods
today, such as those involving guns, drugs, and the
automobile (see McCulloch and Philip 1972; Stengel 1973)
depend on human technological innovations. These methods
and their effects would have to be learned. However, other
methods, such as stabbing, burning, or hanging involve older
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deCatanzaro: Biology of suicide
technologies, while methods such as drowning, jumping from
high places, and exposure to the elements must have always
been available to man.
Furthermore, complex and unusual cognitive events
frequently underlie suicide (see Levenson 1974; Neuringer
1976; Shneidman 1957; Tripodes 1976); these events
presumably result largely from an individual's learning
experiences, and may be especially related to his learning
about death. For example, some individuals may believe that
following suicide they will enter an afterlife that is better
than their present life or that they will rejoin a lost relative.
Other individuals may see their successful suicide as
producing desired effects on their environment, but not
perceive clearly the relation of this act to their personal
fitness. Such suicides may be motivated by the desire to have
an impact on the environment, such as improving the welfare
of others, taking revenge on others, or increasing the respect
or attention received from others. Suicide may also be seen as
a way of removing oneself from pain or other aversive events.
Normal, non-life-threatening behavior directed toward these
ends might increase the fitness of the behaving individual;
but individuals committing suicide to achieve such ends do
not, of course, survive to experience increased fitness. It is
conceivable that, because of such aberrant cognitions or
logical errors, suicidal individuals falsely perceive that their
behavior increases fitness when it actually has the opposite
effect.
Another indication of a mediation or modulation of suicide
by learning is the involvement of modeling, or learning
through the observation of others' behavior (Bandura 1971).
Suicides are often clustered in location and time. They may
increase with reports of suicide in the news media or be
brought on by suicide among friends (Hankoff 1961; Phillips
1974). The involvement of one type of modeling process,
contagion, is illustrated by the recent mass suicide in
Jonestown, Guyana. The simultaneous suicide of several
hundred individuals must indicate a major contribution of
learning and suggestion to the occurrence of suicide.
However, this does not necessarily indicate that suicide is
exclusively determined by learning. Modeling may modify
the probability of suicide and determine its temporal and
physical location. It may not, however, by itself provide
sufficient motivation for suicide. In the Jonestown case, for
example, many of those attracted to the cult may have been
inclined toward suicide before joining. Stress and desperation
may thus also have been major determinants of their suicides;
these factors will be discussed in more detail below.
On a larger scale, the prescribed patterns of behavior in a
culture may determine when, where, and in whom suicide
will occur. Previous suicides in specific circumstances may
act as models, teaching members of a culture that suicide is
appropriate if similar circumstances are encountered.
Although suicide occurs in some form in most cultures, there
are differences in the situational determinants and forms
across cultures. For example, the custom of self-immolation of
a woman on the funeral pyre of her husband has been
particularly common in India (Venkoba Rao 1975). In Israel,
suicide is almost equally frequent in females and males,
contrary to trends elsewhere (Headley 1975). In general,
suicide is relatively infrequent in predominantly Catholic
countries, which may reflect either that church's strictures
against suicide or a tendency to certify suicide under other
modes in those countries (Farberow 1972). There are
numerous other differences among cultures and subcultures
with respect to suicide (see deVos 1968; Farberow 1972).
Although differences in the recording of suicide statistics
among cultures cannot readily be factored out, cross-cultural
variation does strongly suggest that learning and culture are
among the major determinants of suicide.
Some evidence also suggests a role of learning in
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 267
 deCatanzaro: Biology of suicide
subsuicidal forms of behavior. In some cases, completed
suicide may be the unintentional result of such subsuicidal
behavior, thus making the suicide a result of learning
processes. This might occur where attempted suicide,
attributable to reinforcement expected after survival of the
act (see Kreitman 1977; Stengel 1973), inadvertently
produces suicide. Also, more rarely, self-injurious behavior,
which may be partly attributable to reinforcement
contingencies (see Bachman 1972; Carr 1977; deCatanzaro,
1978; 1979), can sometimes lead to death.
The first model, then, reconciles suicide with functional
analyses of behavior by maintaining that it results from
learning and culture. Since humans have a highly developed
capacity to learn, their behavior may be sufficiently
independent from the constraints of natural selection that it
can occasionally act contrary to this selection. Selective
pressures favoring a high degree of behavioral flexibility may
outweigh negative pressures accompanying occasional
incidental maladaptive behavior. Indeed, there are strong
indications that learning is involved in the etiology of suicide.
It can determine the method employed and the time and
place of suicide. It is less clear that the underlying
motivational events are entirely due to learning, although
learning probably modulates such motivation.
II: Suicide as pathology. The second model of suicide
would account for this behavior in terms of a poor fit between
the organism's inherited capacities and the environment. In
general, the genes that are passed on in evolution may have
survived because they facilitate adaptation, but in unusual
environments they may fail to do so. Furthermore, genes
frequently mutate or recombine, often producing
maladaptive phenotypes. These processes may engender
pathological conditions that facilitate the occurrence of
suicide.
An individual may inherit genes that, in interacting with a
normal range of ecological conditions, would make him
well-adapted. These genes would thus usually be favored in
natural selection. If the individual is exposed to environments
outside of those normally experienced by members of the
species, his genotype may fail to ensure that his behavior is
adaptive. The possession of a normally adaptive genotype
does not prevent an organism from being damaged by
physical injury, disease, or congenital malformation. Nor does
it prevent psychological malfunctioning because of an
impoverished early environment, severe social isolation, or
extremely stressful experience. Such conditions may be so
extreme and unusual that the organism does not have an
inherited capacity allowing him to cope. Indeed, this is true
by definition of any extreme stress (Selye 1973). Under
extreme stress of any sort, then, disorganized and
maladaptive behavior may develop despite the fact that the
individual's genes would predispose toward adaptive
behavior under more normal circumstances. Such
disorganization of behavior may facilitate suicide.
Concordant with this model is evidence that suicide bears a
fairly consistent relation to stressful experience. Prolonged
stressful experience is almost ubiquitous in the backgrounds
of suicidal individuals, while some exceptionally stressful
experience is frequently an immediate antecedent of the act
of suicide (see Ferrence, Jarvis, Johnson, and Whitehead
1975; Kiev 1977; McCulloch and Philip 1972; Stengel 1973).
Divorce, loss of a loved one, unemployment, poor health,
terminal illness, financial problems, alcoholism, and
disordered personal relationships frequently constitute such
stressful experience. Indeed, suicide is probably very rare in
the absence of some sort of stress or severe difficulty in
coping. While there are reports of suicide among individuals
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who appear to be "gifted," some stressful experience is
usually found in these cases (Shneidman 1976b).
Also in agreement with this model is some evidence (see
Lester 1972; Stengel 1973; Temoche, Pugh and MacMahon
1964) that suicide is correlated with other psychological
disturbances. Its frequency in psychiatric populations is
generally higher than in the population at large, and it
appears to be especially associated with affective disorders.
However, it is questionable whether suicide is universally
related to some other conventional category of psychological
disorder. Stengel (1973) estimates that on the average only
one-third of the people who commit suicide have been
suffering from any neurosis, psychosis, or severe personality
disorder; the majority of people who commit suicide have not
been under any psychiatric treatment. It may thus be
difficult to form any generalization about the relation
between suicidal acts and mental disorder without some
expansion of the latter concept.
Another notion consistent with the present model is that
current high rates of suicide reflect the fact that modern
society makes excessive demands on many individuals. Since
these demands may be historically novel, many individuals
may not be equipped to cope and may consequently display
maladaptive behavior such as suicide. There is some evidence
of an increase in suicide in recent times (see Diggory 1976;
Shneidman 1976a), although the rise may not be dramatic
and may also reflect improvements in the accuracy of
demographic data.
The greatest difficulty with the pathology model is that it
does not fully account for the transition between stress-
induced disorganization of behavior and the occurrence of
behavior oriented against survival. One possibility is that
cognition might become sufficiently disorganized under
severe stress to enable learning variables to produce suicide.
Another possibility is that mechanisms mediating
physiological pain may not adequately inhibit self-injury in
times of severe stress; evidence indicates that pain perception
is markedly reduced during stress by endogenous chemicals
known as endorphins, which act on receptors in the brain (see
review by Barchas et al. 1978). In such ways stress may
facilitate suicidal behavior; the actual occurrence of suicide,
however, may require the interaction of these factors with
other variables.
There are some other, limited conditions under which
genes may predispose an organism to behave maladaptively;
these may for convenience be included in this model.
Evolution involves genetic mutation and recombination as
well as natural selection. Such processes produce
heterogeneity and allow genetic change, but may also
produce maladapted organisms. The involvement of
mutation in suicide would seem quite unlikely, given the
frequency of suicide, although it might be involved in
exceptional cases or associated with some forms of self-
injurious behavior (Lesch and Nyhan 1964). Also, because the
adaptive value of any gene depends on its genetic milieu,
maladaptive gene combinations might occur despite the
adaptiveness of component genes in other circumstances
(Caspari 1967; Herskowitz 1973). This could conceivably
influence the occurrence of suicide in that, for example,
polygenic control of affective conditions (see Gershon,
Bunney, Leckman, Van Eerdewegh, and DeBauche 1976)
might occasionally produce individuals prone to severe
depressive disorders, which in turn may be conducive to
suicide.
Although this model may not account for all aspects of
suicide, it does appear to explain very well the chronic
self-injurious behavior that occurs in primates raised in social
isolation and in autistic, schizophrenic, and retarded humans.
As I have discussed elsewhere (deCatanzaro 1979), such

self-injurious behavior is almost universally associated with
physiological, genetic, and developmental abnormalities that
may directly produce the behavior.
The present model, then, proposes that suicide may result
from a disorganization of behavior accompanying the
occurrence of normally adaptive genes in unusual
environments or the occurrence of novel genes or gene
combinations in more normal environments.
Ill: Suicide as altruism. In the third model, I would like to
consider whether some of the selection processes favoring
"altruistic" behavior, discussed by the sociobiologists, might
be involved in suicide. This model suggests that self-
destructive behavior may occur because it has beneficial
effects for individuals other than the suicidal individual. This
would imply that an individual's inherited motivational
constitution is such that he may occasionally sacrifice his own
fitness for others, which might suggest that genetic factors
could indirectly favor suicide.
Prosuicidal genetic factors that were expressed in all
individuals possessing them would strongly select against
themselves. However, such factors might survive and even
prosper if expressed in conditions where they confer a benefit
on surviving individuals sharing the genes of the suicidal
individual. If evolution selects only for behavioral dispositions
favoring the individual, suicide may not have genetic
underpinnings. But in considering the fact that an
individual's genes are shared by others, a genetic basis for
suicide becomes conceivable. The model does not require that
such prosuicidal genes directly control the topography of
suicide, but rather that the inherited motivational constitution
of individuals be such that they could behave for the benefit
of others at their own expense.
Hamilton (1964; 1970; 1971) proposed that behavior
sacrificing the fitness of the behaving organism may develop
through its benefits for related individuals. This process,
which has been termed "kin selection" (Maynard Smith
1964), occurs when the loss of one individual supports the
survival of close relatives. Improved fitness of relatives is
sufficiently great that the loss of one individual's genes is
outweighed by enhanced fitness of identical genes in
relatives. Kin selection might be adapted to explain suicide by
attributing it to prosuicidal genetic factors expressed when
the individual is or perceives that he is a burden on relatives.
Relatives would share genes directly or indirectly favoring
suicide, but because their circumstances were better, these
genes would not be expressed in them.
Kin selection probably could not explain the majority of
cases of suicide. Frequently, the suicide of an individual is the
source of considerable grief to surviving kin (McCulloch and
Philip 1973), which is difficult to reconcile with the notion
that this death is for their benefit. Furthermore, suicide often
occurs when an individual has little contact with his family
(Stengel 1973); in fact, protracted isolation from family and
other members of society is among the best predictors of
suicide (see Batchelor and Napier 1953; Breed 1972; Ganzler
1967; Leese 1969; McCulloch and Philip 1972; Sainsbury
1955; Schneider 1953; Stengel 1973; Worden 1976).
Nevertheless, given that individuals may be motivated to
act for the benefit of kin, and given the variance in individual
motives underlying suicide, kin selection may play a role in a
minority of cases. This is suggested by references in some
suicide notes to benefits accruing to surviving kin (see, for
example, Shneidman 1976c, p. 270). Also, there are some
indications that certain family patterns, such as broken
homes, economic difficulties, and the consistent receipt of
blame for family problems, commonly occur in the
backgrounds of suicidal individuals (Jacobs 1971; Richman
1971); it is conceivable that kin selection might operate in
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deCatanzaro: Biology of suicide
such conditions.
Other evolutionary mechanisms can be invoked to explain
suicide. Wynne-Edwards (1962; 1971) has argued that
behavioral traits may be affected by selection among groups
or subpopulations within a species, because reproductively
isolated subpopulations compete and have different gene
frequencies. Stanley (1975) has argued that selection above
the species level may induce more genotypic change than
does selection acting on individuals. Through selection at
these levels, genetic traits that, when expressed, decrease an
individual's chance of survival, might increase in frequency.
This may occur in that some competitive advantage accrues
to that individual's population, which shares these traits,
despite or because of his loss. For example, in a human tribal
grouping, the loss of members experiencing certain extreme
stresses and thereby taxing common resources might improve
the general fitness of the tribe. Tribes in which burdensome
members were lost through suicide might fare better than
those in which suicide did not occur.
Some notions of suicide might be subsumed by a group-
selection hypothesis of human suicide. Durkheim (1951), in
his pioneering work on suicide, states that one form of this
behavior could be classified as altruistic. Altruistic suicide
occurred in individuals who felt very much part of a group
and sacrificed themselves for the good of that group or of
society in general. Freud (1924), on the basis of clinical
observations, postulated the existence of a "death instinct" in
humans which only surfaced under limited circumstances.
Furthermore, there are indications that feelings of guilt and
other negative self-attitudes may be correlated with suicide
(Kaplan and Pokorny 1976). Negative self-concepts may be
created by the behavior of other members of society toward
these individuals; these self-concepts may help to elicit
suicide.
Farberow (1972) has described "institutional" or "social"
forms of suicide, involving self-destruction that a society
demands of a member. Institutional suicide has occurred in
many forms throughout recorded history. Farberow discusses
an example from a primitive tribe, in which an individual
accused of transgressing the tribe's taboos would climb to the
top of a palm tree and plunge headfirst to his death. Another
example is found in other cultures, where in times of food
shortage, the old and sick were expected to sacrifice
themselves for the good of the group. Also, throughout
history, in times of war men have jeopardized and sacrificed
their personal existence for the benefit of the social group to
which they belong. These forms of suicide are compatible
with a group-selection hypothesis, but they might equally be
attributable to cultural evolution and may thus not involve an
elicitation of special suicidal tendencies.
There is considerable controversy about the importance of
group selection in evolution (cf. Dawkins 1976; Wiens 1971;
Wilson 1975; Wynne-Edwards 1962; 1971). Currently, many
people appear to support the view that the role of group
selection has been relatively minor. For example, Dawkins
(1976) argues very strongly that the dominant force in
evolution is individual selection, and that most of the animal
behavioral patterns attributed by Wynne-Edwards to group
selection can be explained in other ways. He has also argued
that selection due to the differential reproductive success of
individuals proceeds much more rapidly than selection due to
the differential success of whole breeding populations.
Therefore, traits causing individuals to increase their genetic
representation in future generations will win out over traits
favoring the population or species. Any individual within a
group who behaves in a self-serving manner when altruistic
behavior is called for will be likely to pass on genes supporting
self-serving behavior; through such processes individual
selection may be more powerful than group selection.
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 269
 deCatanzaro: Biology of suicide
It may nonetheless be premature to dismiss entirely the role
of group benefit in human behavior. Most theorizing about
group selection has dealt with nonhuman animals. During
most of human history there appears to have been an
extraordinary degree of intraspecific aggression organized at
a group level. While complete annihilation of one group by
another may have been rare, some differential reproductive
success would probably accrue from success or failure in such
competition. Furthermore, the human situation may be
complicated by the coevolution of culture and biology (see
Durham 1979). Cultural factors have interacted with
biological evolution in a complex manner that makes an
analysis of biological changes exceedingly difficult.
Also, much of the current theory of selfishness and altruism
has been based upon concepts of gene sharing among
individuals. The coefficients of relationship most commonly
used (Hamilton 1964; Wright 1922) give a brother or parent
one-half of an individual's genes, a cousin one-eighth, and so
on. According to this model, one begins to approach a zero
relationship rapidly in considering individuals progressively
less related in a familial sense. Yet this takes into account only
gene sharing by proximate descent and not gene sharing by
kind. Also, it considers only a few generations rather than the
totality of the species' history, which might produce much
more complex interrelationships among us all. Surely there is
some sense in which individuals within a species are more
related than individuals across species. King and Wilson
(1975a) have reported that the overlap between human and
chimpanzee DNA and amino acid sequences is 99% and that
the variation is even less within the human population. While
this may reflect differences in variability of structural and
regulatory genes (King and Wilson 1975b), it does suggest
that the situation may be more complex than we have
assumed. Although an offspring physically inherits one-half of
each parent's genes, many genes do not differ between his
parents and may also not vary substantially within a
population. Thus more than one-half of the individual's genes
may be identical to his parents' when one is considering genes
in kind (see also Washburn 1978). Even employing
conventional formulas and considering the most proximate
generations, gene sharing among naturally aggregated
individuals may be quite complex; this is illustrated by the
careful calculations of genealogical relationships within
Yanomano villages by Chagnon (1979). I do not wish to imply
that the totality of gene sharing among individuals should be
indicative of commonality of interest, nor that more closely
related individuals should not share especial interests. Rather,
I am suggesting that we have not yet defined adequately the
point of zero relationship and thus of no commonality of
interest. The situation may be exceedingly complicated if we
fully consider the parameters of population genetics (see also
Wilson 1975, pp. 119-20).
Assessment of the role of altruism in human suicide thus
must await further developments in the theory of gene
sharing and altruism, as well as further data on the impact of
suicide on others. Tentatively, given the indications that
suicide may have little or negative impact on kin and the
current weight of theory against group selection, it may be
best to turn our attention to other variables.
IV: Evolutionary tolerance of suicide. The final model
presents a promising alternative to the second and third
models in accounting for aspects of suicide not attributable to
culture. This model would explain suicide on the basis of
special ecological and social characteristics of those
committing the act. These characteristics may make suicide
immune to the pressures of natural selection.
Suicide is an act of desperation. It occurs in extreme
circumstances (Lingens 1972), particularly when an
individual feels incapable of coping with his present
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circumstances and expects little improvement in the future
(Bogard 1971). Farber (1968) has argued that suicide occurs
when people envision not only the present but also the future
as negative. He asserts that suicide is unlikely when people
anticipate acceptable life conditions and have a sense of
competence, but likely when acceptable life conditions are
seriously and permanently threatened. Indeed,
"hopelessness" is an excellent predictor of true suicidal intent,
being much better than "depression, " which may be merely a
transitory state (Beck, Kovacs, and Weissman 1975).
Human suicide occurs, then, in individuals experiencing
extreme difficulties in coping and who expect little improve-
ment in the future. Given our assumption that coping behav-
ior promotes the individual's genes, difficulty in coping
would seem to entail difficulty in advancing the welfare of
one's genes. Conversely, suicide is relatively rare in individu-
als who are well adapted; well adapted individuals may, by
definition, be those most capable of reproducing or otherwise
promoting their genes.
The difficulties in coping associated with suicide may be of
a diverse nature, but are generally severe enough to impede
behavior that effectively promotes the individual's genes. For
example, suicide is frequently related to economic hardship,
commonly occurring in the unemployed or those who have
suffered some severe economic loss (McCulloch and Philip
1972). Where economic hardship occurs with little
probability of improvement it is unlikely that an individual
will be able to provide adequately for himself and for others
sharing his genes. Suicide is also common in the terminally ill,
the elderly, and those with severe physical disabilities
(Dorpat, Anderson, and Ripley 1968; McCulloch and Philip
1972; Weiss 1968). Such individuals may show low
probability of engaging in future reproduction and would be
relatively unable to engage in productive activity promoting
the welfare of others sharing their genes. Furthermore,
adaptive behavior such as proper eating, sleeping, sexual
behavior, grooming, and working habits may break down
under chronic stress or depression (Beck 1967; deCatanzaro
and Gorzalka 1979); such conditions may illustrate how
self-promoting behavior breaks down in states that may
precede suicide.
Suicide appears to bear a remarkable relation to
reproductive success. Suicide is much more common among
single and divorced individuals and those with unstable
marriages than among those with stable marriages (Dublin
1963; Durkheim 1951). Those with stable marriages would
probably be relatively likely to reproduce and able to
promote the welfare of offspring. Indeed, individuals with
children are among those least likely to commit suicide
(Breed 1966; Dublin and Bunzel 1933). Also, recent divorce
or difficulty with members of the opposite sex is often a
proximate antecedent of suicide (McCulloch and Philip
1972).
Furthermore, as discussed above, social isolation is a good
predictor of suicide. While this may be due in part to
voluntary social withdrawal during presuicidal depression,
there is considerable support for the notion that social
isolation can precipitate suicide (see Stengel 1973). Socially
isolated individuals may be less likely than other members of
the population to form successful social relationships enabling
them to advance their genes. Not only would they be unable
to bring their genes into future generations without stable
relationships to members of the opposite sex; without contact
with kin they would be incapable of behaving to improve
their inclusive fitness, advancing their genes as they exist in
relatives. Consistent with this is evidence that suicide is most
common in the developed countries, where the family unit
has become relatively less stable and smaller, and thus where
individuals have less opportunity to promote the welfare of
relatives sharing their genes.

Another excellent predictor of suicide is age; suicide
frequency increases as a direct function of age, particularly in
men (Diggory 1976; Linden and Breed 1976). Although some
cases have been reported, suicide is extremely rare among
children, almost never occurring in children under ten and
infrequently between ten and fourteen (Diggory 1976; Otto
1972; Shaffer 1974; Toolman 1968). Productive and
reproductive capacity may not be established by these ages,
and many of the difficulties in adjustment experienced by
children may be transitory. Suicide becomes more frequent in
adolescence and early adulthood (Jacobs 1971; Hendin 1976),
which could relate to increasing differential productive and
reproductive success. With increasing age, individuals may be
less likely to engage in reproductive activity, and
consequently less able to spread their genes. Furthermore,
with increasing age, some individuals may be less capable of
engaging in productive activity that promotes the welfare of
related individuals such as existing offspring. It is interesting
that there are sex differences in both the frequency of suicide
and its relation to age (Linden and Breed 1976). This sex
difference could conceivably reflect differences between the
sexes in reproductive strategies and nurturance of offspring
(see Daly and Wilson 1978).
A low capacity to promote one's genes thus appears to
account well for all of the best predictors of suicide. I would
now like to discuss why this relationship might occur.
If an individual's present and future behavior is unlikely to
change the status of his genes, there may be no ecological
pressures preventing his suicide. If an individual is unlikely to
reproduce, unable to support himself and his family
adequately, and unable to contribute to the welfare of other,
reproducing individuals sharing his genes, his death may not
affect the frequency of genes he carries. Suicide would
consequently not eliminate any genes from the gene pool that
were not already eliminated. Thus, under the limited
ecological conditions in which suicide appears to occur, there
may be no selective pressures to prevent it. Conversely,
suicide may be rare in relatively well-adapted individuals
because their death would have an impact on the gene pool.
In its weakest form, this factor would mean that suicide
that occurred for other reasons would not be influenced by
selective pressures. Thus, for example, a suicide that occurred
because of cultural influences or behavioral disorganization
consequent to stress might not cause any impact upon the
gene pool. In a stronger form, this factor might mean that
selective pressures have favored the development of a
variable motivational constitution in members of the species.
Such a constitution might favor behavioral efforts toward
adaptation and self-advancement when such advancement
was possible, but not when such advancement was no longer
possible. It is also conceivable that this factor would allow
cultural pressure or species benefit (perhaps mediated by
culture more than genes) to influence behavior so that it
operated against one's survival when one consumed resources
without being productive. Where there is "no reason to live, "
any otherwise trivial reason not to live might influence
behavior.
This model may require that selective pressures act
differentially on the same individual's behavior under
different circumstances. When an individual is well-adapted,
selective pressures act against suicidal behavior. If the same
individual subsequently loses fitness and the capacity to
further his genes, as might be the case, for example, in old age
or severe illness, these selective pressures would no longer
operate. Because of this, what might be favored in evolution is
the capacity to vary in the tendency to protect one's existence.
It is not unusual for natural selection to favor behavioral
predispositions that act differentially in different
environments. Sexual, courtship, aggressive, hibernation,
migration, and other behavioral patterns of many species all
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deCatanzaro: Biology of suicide
occur under limited environmental conditions. Genes whose
expression depends on environmental factors are by no means
rare (Caspari 1967; Herskowitz 1973).
If an individual's propensity to promote his survival varies
in parallel with selective pressures, as described above, this
variation must have some inherited, biological basis. Such a
basis could conceivably consist of physiologically mediated
emotional or motivational variables. There is growing
evidence that much of what we commonly call "emotional"
behavior is in part mediated by organized biochemical
systems acting upon and within the brain (Barchas et al. 1978;
Levine 1976). These systems involve neurochemical,
hormonal, and other neuroregulating substances which
respond in characteristic manners to stressful, novel, or other
stimuli. Such evidence comes from pharmacological
manipulations that affect specific neuroregulatory systems
and biochemical assay data that correlate with behavior. For
example, injection of central catecholamine stimulants may
induce subjective feelings of well-being or remove protracted
states of depression, while catecholamine antagonists may
induce or exacerbate states of depression. However, suicide
may relate more to serotonin than catecholamines; a number
of postmortem studies of brain samples of human suicides
indicate that 5-HIAA, a metabolite of serotonin, is
substantially reduced in suicides as compared to controls (see
review by Murphy, Campbell, Costa 1978). Indeed,
reserpine, which depletes catecholamines and serotonin, has
been reported to induce suicide (see Lester 1972; Weil-
Maherbe 1972). Physiological systems of this sort, responding
to stress and environmental stimuli, and controlling
behavioral predispositions, may provide the inherited
variable substrate required by this model.
One other set of data may have bearing upon this model
and also perhaps on the third (altruism) model. Several
investigators (see Farber 1977) have noted that suicide
appears to cluster in families. While this could be due to
common learning experiences in such families, it may also
indicate that genetic factors underlie suicide. Kallman (1949)
compared suicide rates in twins, finding higher concordance
for monozygotic than dizygotic pairs. While Kallman did not
believe that he had established a relationship between
genetics and suicide, Lester (1968), reanalyzing these data
with more sophisticated statistical methods, concluded that
they did indicate such a relationship. Other evidence
(Gershon et al. 1976) suggests that affective disorders, which
are often precursors of suicide, are probably influenced by
genetics. These data may suggest genetically produced
variability among individuals in the propensity to commit
suicide. However, the model does not require a large amount
of variability among individuals; rather, it suggests that a
tendency for individuals to vary in self-preserving and self-
promoting behavior is widespread in the population.
The final model suggests, then, that suicide is most
common when an individual is incapable of promoting his
genes. This situation may mean that suicide is immune to
selective pressures in that it would not affect the gene pool.
Conclusions and research suggestions
These models are not necessarily mutually incompatible, nor
do they necessarily exhaust all possibilities. Each model
explains some, but not all, aspects of suicide, which perhaps
indicates that some combination of factors is responsible.
Suicidal acts are complex and diverse, and there is variability
in the documented proximate etiological factors. Different
factors may contribute to different cases, and it is likely that
an interaction of several factors is necessary in any case. The
validity of these models remains to be established by future
research, which will have to determine the relative contribu-
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 Commentary/deCatanzaro: Biology of suicide
tion of the factors they suggest. The study of human suicide
also presents many problems that are not encountered in
other research endeavors; good discussions of these problems
can be found elsewhere (e.g. Neuringer 1962).
There is a substantial need for work reconciling the find-
ings of human psychology and sociology with evolutionary
theory. Any science of human behavior must be integrated
with other developing areas of science. This is especially true
with respect to biological science, since behavioral processes
are a function of biological organisms. Behavior is a means by
which organisms adapt to their environments, much as physi-
ological processes act toward this same end. If we accept that
organisms are subject to natural selection, we must accept that
their behavior is also a function of such selection.
It is common practice to analyze animal behavior in terms
of its potential biological adaptive value. However, this kind
of analysis has not been practiced in psychological and
sociological approaches to human behavior, perhaps because
of artificial academic disciplinary boundaries. There is no
reason why human behavior cannot be subjected to the
scrutiny of evolutionary reasoning just as animal behavior and
physiology are. Human behavior is as much a product of
biological variables as is the behavior of other organisms.
Although learning and cultural variables produce much of the
topography of human behavior, we must still consider the
ways in which culture and learning interact with biological
constraints.
Many may be concerned with the testability of hypotheses
about evolutionary effects on behavior. Hypotheses about
evolutionary history or the relations between selective pres-
sures and behavior in numerous previous generations may
indeed be difficult to test. Presumably, very little trace is left
of suicidal events that occurred one thousand or one million
years ago. Nevertheless, information about current suicidal
acts and their proximate determinants is readily available.
Through systematic examination of this information we may
gain insight into the ultimate determinants of suicide.
For example, it may be difficult to obtain data directly
testing the proposition that "suicide has been tolerated by
evolution only when it has occurred in individuals who have
had little capacity to promote their genes." This statement
may or may not be true, but it is difficult to test its implica-
tions with respect to selection processes occurring over several
human generations. Many of its corollaries, however, are
readily tested. One corollary is that "suicide is currently most
likely in those with little capacity to promote their genes."
This statement is testable through the careful collection of
case histories, including data on the social situations of those
committing suicide. The corollary, "suicide is very unlikely in
nonreproducing individuals who can support the welfare of
reproducing relatives," is similarly quite testable. By gather-
ing together such information we can piece together a picture
that may begin to explain suicide. However, without more
general hypotheses, we may be unable to generate more
readily testable corollary hypotheses, and our power to
explain may be considerably reduced.
The situation with the other hypotheses presented is simi-
lar. We can gain a substantial amount of information about
the involvement of learning in suicide by collecting informa-
tion on the backgrounds of those exhibiting this behavior.
Cross-cultural studies may be especially valuable. The
involvement of genetics is also not impossible to factor out.
Studies probing the frequency of suicidal acts in related
individuals raised in separate environments would be particu-
larly fruitful; the application of these and other behavior-
genetic techniques has provided important information about
genetic involvement in other kinds of behavior (e.g. Gershon
et al. 1976). Further study of depression and suicide elicited
by pharmacological agents may shed light on possible physio-
logical and genetic determinants, as would additional autopsy
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data on physiological correlates of suicide. Information about
the involvement of group and kin selection may come from
analyses of the social situations of suicidal individuals.
Hypotheses regarding evolutionary processes can be exam-
ined through logical analysis and computer modeling tech-
niques. One can analyze the plausibility of hypothetical
evolutionary processes by simulating them mathematically.
Furthermore, hypotheses about ongoing evolutionary pro-
cesses are not untestable in any absolute sense: we may have
difficulty devising means of testing them directly because of
limitations on the length of time any one researcher can
sample, but, over generations, a sufficiently large sample of
information may be gathered to test the validity of such
hypotheses.
Further advances in many areas of research, then, could
contribute to the validation of the heuristic models presented
here. Perhaps the major thrust of future work should involve
careful investigation of the ontogeny of suicide and the classes
of individuals displaying this behavior. Such work would not
be entirely new, of course. However, in classifying etiological
factors and classes of people we musb have some conceptual
framework that orders the information we collect. That
conceptual framework may be best developed by formulating
theories about the biological roots and ultimate causes of
suicide. There is a substantial need for general theories that
account for the many disparate findings and that help to
generate research hypotheses. Any general theory of suicide
must account for the biological anomaly that this behavior
presents.
Open Peer Commentary
Commentaries submitted by the qualified professional readership of
this journal will be considered for publication in a later issue as
Continuing Commentary on this article.
by Hymie Anisman
Department of Psychology, Carleton University. Ottawa, Ont., Canada K1S5B6
Depression and suicide:
stress as a precipitating factor
As indicated by deCatanzaro, the mechanisms underlying suicidal
behavior cannot readily be subsumed within a single theoretical
framework. Among other things, the antecedent conditions associated
with suicide vary considerably (depression, psychosis, chronic drug
abuse, terminal illness, and maintaining honor, as well as "(or king and
country"). Not unexpectedly, attitudes toward suicide vary among
cultures and may be dependent on the conditions precipitating the
suicide (witness the condemnation of the misanthrope and the worship
of the martyr).
The proposition that suicide is related to the individual's capacity to
promote his genes (reproductive success) is not one that can be
readily assessed through experimental methods, despite the provi-
sional suggestions for research advanced by deCatanzaro. Moreover,
evaluation of the evidence reviewed by deCatanzaro suggests that
attempts to relate gene propagation to suicide are gratuitous. In
particular, a series of social conditions associated with suicide is
described (e.g., separation, divorce, isolation), and it is suggested that
each of these represents an instance where the probability of gene
propagation is reduced. However, each of these examples also
represents an instance where the individual is confronted by uncontrol-
lable stress or an instance where the individual does not have access
to social factors that might help in buffering aversive insults. Thus, the
source of suicidal intent can be derived from more conventional
models than that offered by deCatanzaro. Put simply, upon exposure
to aversive stimuli, behavioral and physiological changes occur in
order to meet environmental demands. If the stress cannot be dealt

with effectively through behavioral means, then the burden of coping
rests principally on physiological mechanisms. Under such conditions,
use of central neurotransmitters, such as norepinephrine and seroto-
nin, will increase and may exceed the rates at which these amines are
synthesized. As a result, a net decline of these neurotransmitters may
be expected (see reviews in Anisman 1978; Stone 1975; Welch and
Welch 1970). Interestingly, if the stress is sufficiently protracted,
physiological accommodation may occur in the form of increased
activity of synthetic enzymes, thereby resulting in amine levels
approaching control values (Kvetnansky, Mitro, Palkovits, Brownstein,
Torda, Vigas, and Mikulaj 1976; Weiss, Glazer, and Pohorecky 1976).
The initial stress-induced depletion of brain amines, and failure of
amine levels to increase with protracted stress, may result in clinical
depression, which in turn may precipitate suicide.
The available data largely support a stress model such as that
described here. Stress has been considered as a precipitator of
depression, although it cannot be denied that stress may simply
increase the incidence of hospitalization of already depressed individu-
als (see discussions in Akiskal and McKinney 1973, 1975). As
indicated earlier, stress will reliably induce depletion of brain mono-
amines, provided that the stress is one over which control is not
possible (Anisman, Pizzino, and Sklar 1980; Weiss et al. 1976). Of
particular relevance to the present discussion, the amine depletions
are also dependent on the age of the animal (Ritter and Pelzer 1978),
as well as on social housing conditions (Modigh 1976; Welch and
Welch 1970). It will be recalled that such factors are also associated
with the incidence of depression and suicide (see deCatanzaro).
Finally, considerable data are available indicating that the monoamines
most affected by stress are those that underlie clinical depression
(Murphy, Campbell, and Costa 1978; Schildkraut 1978). To be sure,
inconsistent and contradictory evidence has been reported concerning
the role of any one neurotransmitter in depression. Nevertheless, it
does appear that both norepinephrine and serotonin, and possibly also
dopamine and acetylcholine, are intricately related to the depressive
symptomatology.
In considering a stress-depression model it should be remembered
that the response to stress may vary widely between individuals.
Accordingly, apparently equivalent stresses may lead to pathology in
one individual but have little effect on a second individual. These
individual differences may be due to any number of experiential,
environmental, and organismic variations, and a role for genetic
factors is certainly not being dismissed in the analysis of depression. It
has been reported that concordance rates for depression are higher in
monozygotic than in dizygotic twins, even if the twins were reared
apart. At the same time the concordance rates do not approach values
that would preclude an environmental contribution to the affective
illness (see review in Winokur 1978). It is likely that genetic factors
predispose individuals to depression provided that certain environmen-
tal conditions are present. In a recent review of the literature Gershon
(1978) indicated that catechol-O-methyl transferase, an enzyme
involved in catecholamine degradation, may serve as a genetic marker
for depression. These data support the hypothesis that catechol-
amines (or associated enzymes) are linked to depression and may be
genetically transmitted. However, this cannot be taken to support the
contention that depression or suicide is related to differential reproduc-
tive success. At this stage in the analysis of factors that promote
suicide it is not advantageous to rely on a sociobiological model such
as that presented by deCatanzaro.
by D. Caroline Blanchard
Bekesy Laboratory of Neurobiology. University of Hawaii. Honolulu, Hawaii 96822
Biological variation and suicide
Complex phenomena such as cancer or aggression - or suicide - may
involve a number of diverse processes with different antecedents and
different evolutionary outcomes. The functional or adaptive signifi-
cance of the component processes treated under one rubric may vary
tremendously, and progress in understanding them depends largely on
an adequate analysis of the different events involved.
OeCatanzaro's "suicide" is not defined, but examples given suggest
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Commentary/deCatanzaro: Biology of suicide
a range of events from participation in war through direct self-
destruction "for cause" such as terminal disease, to acts associated
with depression or other mental illness. It seems obvious that separate
consideration of these types of events is likely to be more fruitful than
lumping them together.
Depression- and mental-illness-related suicide, for example, proba-
bly represent malfunction of a set of brain systems of which the normal
functioning is extremely adaptive. This view is quite different from
deCatanzaro's "pathology" category, which involves the failure of a
system to function adaptively because of poor fit with unusual environ-
mental circumstances. The point is that significant numbers of suicides
involve endogenous depression not primarily related to environmental
stress. There is increasing evidence of neurochemical pathology for
such individuals as well as familial incidence rates suggesting that
genetic factors (Pare and Mack 1971) are in large part responsible for
the specific brain-chemistry changes associated with endogenous
depression. The problem, then, is not fit with a changed environment,
but malfunction with deleterious effects even in very favorable environ-
ments. The fact that such cases represent only about 30% of suicides
does not indicate that malfunction is a poor explanation of suicide: it is
a very good explanation for one specific type of suicide.
The remaining types are more complex. For many of them deCatan-
zaro's "pathology" model is probably the most relevant: human
capacities for memory and reasoning are normally adaptive, as is the
ability to feel and respond to pain, and a tendency to predict and
anticipate future events. When all of these traits combine to present an
individual with a very dim view of his future prospects, the result may be
suicide. In such cases (e.g. terminal illness, profound social disruption,
or ostracism) the future reproductive potential of the individual may be
limited in any event, and his inclusive fitness only minimally affected by
suicide. However, even if every such suicide did result in a drop in
inclusive fitness, the contributory processes (e.g. memory) are still so
adaptive that they might be expected to be relatively little influenced by
their combined consequence in certain situations.
There may, however, be one area in which human cognitive abilities
influence suicide (or phenomena involving hopelessness in general).
People do not recall or anticipate pain with anything like the vividness
of their impressions of past or anticipated joy. Might this not represent
the result of negative selection against those individuals who have
been able accurately to represent to themselves their own future
prospects for pain and suffering? Like all evolutionary questions, this
one is practically untestable at present, but it would be interesting to
investigate some possible mechanisms involved in memory distortions
or painful and pleasant events.
by Edward G. Carr
Department of Psychology. State University of New York at Stony Brook. Stony
Brook. N.Y. 11794
Suicide: comments on deCatanzaro's
diathesis-stress model
The conceptualization of suicide that deCatanzaro argues most
strongly in favor of can be characterized as a diathesis-stress model.
The diathesis is polygenic in nature; the stress consists of any of a
number of common but serious life problems.
The first question is how to test this model. DeCatanzaro suggests
that one major corollary, namely that "suicide is currently most likely in
those with little capacity to promote their genes," is easily tested by
examining case histories. However, he forgets that this "corollary" is
really nothing more than a summary statement based on a series of
studies conducted by other investigators who examined case histories
involving the terminally ill, the elderly, alcoholics, those involved in
disrupted social relationships, and the like. Thus, there is an inherent
tautology here. One cannot test a model by relying upon the same set
of data used to construct the model in the first place. In contrast, the
genetic aspect of the model can be tested in a noncircular fashion.
Behavior-genetic techniques have proven useful in understanding
schizophrenia and certain adult affective disorders. Perhaps the same
methods of analysis could be employed to study suicide, as deCatan-
zaro himself suggests. In this regard, one would think that studies of
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 273
 Commentary / deCatanzaro: Biology of suicide
monozygotic and dizygotic twins reared together and apart could
provide the most critical tests.
The second question concerns what clinical advantages accrue
from adopting a model with a strong biological component. Presum-
ably, an important reason for developing models of suicide is to help
do something about the problem. In this respect, the primary clinical
advantage of knowing whether there are individuals who possess
genes that facilitate suicidal behavior would be in terms of prevention.
That is, such individuals, once identified, could be taught how to cope
with environmental stressors that would normally interact with their
genetic endowment to increase the probability of suicide. Once they
acquired such coping skills, they should be less likely to respond to
stress in a suicidal manner.
In the absence of a systematic body of data demonstrating an
important role for genetic factors in suicidal behavior, it would be a
serious mistake to underestimate the impact of social learning vari-
ables in the etiology and maintenance of suicidal and subsuicidal
behaviors. At present, a sociobiological approach to the problem,
while intriguing, appears at best to offer the possibility of understanding
the subtle modulating effects that result from constitutional factors. At
worst, such an approach may lead researchers to overemphasize
biological variables at the price of ignoring coping skill deficits and
powerful modeling and social reinforcement factors whose role in the
origin and control of suicidal behavior has been clearly documented.
by Richard Dawkins
Department of Zoology. University of Oxford. Oxford OX 1 3PS, England
Domesticity, senescence, and suicide
DeCatanzaro's introductory section "Evolution and behavior" is sensi-
ble and correctly forestalls some of the misconceptions to which social
scientist readers might be prone. I do, however, have some critical
remarks to make about his treatment of suicide itself.
I am not qualified to judge his review of the sociological literature on
suicide, so I will confine myself to two miscellaneous points. The first is
a minor statistical worry. What can it possibly mean to say that suicide
"ranked as the ninth most frequent cause of death in the United States
in 1975," and that, in young adults in many countries "it is the third or
fourth most frequent cause of death"? Obviously everything depends
on how the other causes of death are classified. Is "disease" one
cause of death, or are "cancer" and "heart failure" classified sepa-
rately? Depending wholly on how finely subdivided the other causes of
death are, we may be very impressed, or very unimpressed, with the
importance of suicide as a cause of death. My second remark about
the human evidence is more constructive. It is that it would have been
very interesting to have been told something about the incidence of
suicide in bushmen, or other tribes who might be supposed to be living
in an environment similar to the one in which they have been subjected
to natural selection. This brings me to the part of the article that I am
qualified to criticize.
Of deCatanzaro's four models, the first two do not seem to me to be
clearly distinguished. I would lump them together as the "domestic
animal hypothesis." A domestic animal lives in an environment other
than that in which its genes were naturally selected. Domestic pigs are
obviously hopelessly ill-suited to avoid predators, and their behaviour
in the presence of a wolf might well look like suicide to us. It would not
surprise us, however, because we suspect that the wild ancestors of
modern pigs would have behaved very differently. As R. D. Alexander
has observed, Darwinians do not worry unduly about moths' habit of
immolating themselves in candle flames. Candles, indeed noncelestial
light sources generally, are recent innovations in the world of moths,
and we are prepared to accept that natural selection due to candles
has not yet had time to work on their gene pools. The very sensible
orienting technique of maintaining a fixed compass bearing towards
light rays coming from infinity can become suicidal if the moth tries to
do the same thing to light rays radiating out from a candle: it describes
a neat logarithmic spiral into the flame. My first impulse whenever
somebody bring up some odd aspect of modern human behaviour and
defies me to "explain that with your selfish genes if you can," is to tell
274 THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3
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them the moth story. That is my inclination in the case of suicide.
There is another, and rather less obvious, way in which domestica-
tion can enter into the argument. Suicide, even in a wild population,
requires a Darwinian explanation only if there is some genetic contribu-
tion to variation in the tendency to commit suicide. This is so
fundamental, by the way, that it is rather surprising that deCatanzaro
waited until near the end of his article before mentioning it. Let us
accept, for the sake of argument, that there is such genetic variation in
human populations. Then, "gene for suicide" means a gene that,
compared with its alternatives (alleles) at the same chromosomal
locus, makes individuals more likely to kill themselves given a specified
environment. It follows that a gene for X in environment A may turn out
to be a gene for Y in environment B. Ridley and I have already made
this point with respect to homosexuality (Ridley and Dawkins in press).
A gene for suicide in the environment of modern western society may
have had some completely different effect in Pleistocene Africa. That
effect could have been almost literally anything, for development is
such a complex process. It does not have to bear any resemblance to
suicide; a modern gene "for" suicide might, in the Pleistocene, have
been a gene "for" resistance to some disease. Nearly all mutations
have many effects anyway: the outwardly visible behavioural effect that
interests us may be the tip of an internal biochemical iceberg. If a
biochemically advantageous gene has disadvantageous behavioural
side-effects, selection may work on other "modifier" genes to change
the side-effects. But if the side-effects only manifest themselves in
modern "civilized" environments, we cannot expect the necessary
modifiers to have been selected.
Turning to deCatanzaro's hypothesis that suicide might be favoured
by kin selection, my principal feeling is that the domestic animal
hypothesis is much more likely. But I would like to correct one rather
important error in his treatment of the theory of kin selection. "The
coefficients of relationship most commonly used . . . give a brother or
parent one-half of an individual's genes, a cousin one-eighth, and so
on. . . . Surely there is some sense in which individuals within a species
are more related than individuals across species . . . the overlap
between human and chimpanzee DNA and amino acid sequences is
99% . . . and the sequences vary even less within the human popula-
tion. . .. Thus more than one-half of the individual's genes may be
identical to his parents' when one is considering genes in kind."
DeCatanzaro here falls for what I have elsewhere criticised as "Wash-
burn's Fallacy" -misunderstanding number 5 of my "Twelve Misun-
derstandings of Kin Selection" (Dawkins 1979). Let me briefly explain.
It is, of course, perfectly true that all of us share more than 90% of our
genes. The calculations of coefficient of relationship take all that for
granted and refer to the extra relatedness of close relatives over and
above the commonly shared genes. DeCatanzaro is aware of this, but
he doesn't understand why. His intuitive feeling is that altruism towards
nonkin may be favoured by selection, because even nonkin share the
vast majority of their genes. But let us express this intuitive feeling in
terms of a simple genetic model and see where it leads. Imagine a
gene, U, for universal altruism. Possessors of U are likely to commit
suicide for the benefit of any members of their own species. U is shared
by 99% of the population, so at first sight it seems that U is, in effect,
caring for copies of itself, in just the same way as genes for kin altruism
are supposed to. But now look at the 1% unaccounted for. Suppose
that this minority possesses a gene K for kin altruism. K individuals
sacrifice themselves only for close kin. It is now easy to see that the K
gene is bound to spread through the population at the expense of the
U gene. This is because the U gene, by its indiscriminate altruism,
benefits its rival, the K gene, every bit as much as it benefits copies of
itself. The K gene, on the other hand, since it causes individuals to
discriminate in favour of close kin, statistically benefits copies of itself
at the expense of its U rival. I hope this very crude model will give at
least some intuitive understanding of the nature of deCatanzaro's
error.
Finally, deCatanzaro's fourth model does not, it seems to me,
deserve the preference that he gives it. He suggests that suicide is
most likely to occur among people who are relatively unlikely to
reproduce anyway, for example the old, the sick, and the divorced. No
doubt many critics will object that these categories of people have

more obvious reasons for a statistical tendency to suicide than
concern for their genes. It is not clear to me that such common-sense
proximal explanations are necessarily more parsimonious than deCa-
tanzaro's type of functional explanation, but in any case I want to raise
a different kind of problem. "If an individual's present and future
behavior is unlikely to change the status of his genes, there may be no
ecological pressures preventing his suicide. . . . Suicide would conse-
quently not eliminate any genes from the gene pool that were not
already eliminated." DeCatanzaro might be right if it were literally true
that suicide was confined to individuals who have not the slightest hope
of ever reproducing. Clearly, however, he is only talking about a
statistical tendency. The genetic impact of death in an elderly bachelor,
say, is less than the genetic impact of death in a newly married young
man, and selection would penalize a tendency to suicide in the young
more than in the old. But consider the options open to an elderly
bachelor. It may be true that he is quantitatively unlikely to reproduce
anyway, but he is still even less likely to reproduce if he kills himself
than if he soldiers on. DeCatanzaro's argument is incomplete. His
suggestion of a quantitative weakening of selection against suicide in
certain classes of people needs to be combined with some positive
reason for suicide. He does, indeed, suggest that his fourth model
might be combined with one of the other three.
Let me suggest another, and I think rather fruitful, way in which he
might develop his fourth hypothesis. It is that he should combine it with
the Medawar-Williams theory of senescence (Williams 1957). Any new
mutation, as we have seen, is likely to have many different effects
("pleiotropy"). These effects will not necessarily make themselves felt
at the same time in life. Some will affect the individual in early
embryology, others in childhood, others in young adulthood, others in
middle age, others in old age. It is rare for any one of a mutation's
effects to be advantageous, and the chance of two of its effects being
advantageous is negligible. If a mutation has a beneficial effect at one
particular time in the individual's life history, its effects at other times of
life are likely to be disadvantageous for purely statistical reasons. A
mutation that has a bad effect in youth will tend not to persist in the
gene pool, however beneficial it may be in old age. A gene that has a
beneficial effect in youth, on the other hand, will tend to be passed on
to future generations, however lethal its effects in old age. This is, very
briefly, the Medawar-Williams theory: senile decay is due to the
accumulation in the gene pool of lethal or sublethal genes that have not
been removed by natural selection because of beneficial early effects,
and because their deleterious effects do not make themselves felt until
after the individual has had time to reproduce and pass them on.
If there is a genetic tendency to suicide (a big "if," but there has to
be if deCatanzaro's paper has any point at all), the genes concerned
are properly described as lethal, albeit they work via complex behav-
ioural routes. Their presence in the gene pool could be explained in the
same way that the Medawar-Williams theory explains the presence of
other lethal genes. I find this a more complete explanation of deCatan-
zaro's observations on statistical biases in suicide statistics than his
fourth hypothesis on its own. But my main reaction to the paper is the
one I gave earlier, that the whole problem he raises may be a
nonproblem. Darwinians need worry about suicide only if it is under
genetic control, and then only if it is widespread in wild animals
(humans), observed in the environment in which their genes were
naturally selected.
by Jack D. Douglas
Department ot Sociology. University ol California, San Diego. La Jolla, Calif. 92093
Baechler's theory of suicide
Camus said that the only serious philosophical question is suicide.
That is wrong even in the strict sense intended. The biologist, who is
concerned with questions of physiology and evolutionary history,
realizes that self-knowledge is constrained and shaped by the
emotional control centers in the hypothalamus and limbic system of
the brain. These centers flood our consciousness with all the
emotions - hate, love, guilt, fear, and others - that are consulted by
ethical philosophers who wish to intuit the standards of good and
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Commentary/'deCatanzaro: Biology of suicide
evil. . . . Self-existence, or the suicide that terminates it, is not the
central question of philosophy. The hypothalamic-limbic complex
automatically denies such logical reduction by countering it with
feelings of guilt and altruism (Wilson 1975, p. 3.)
Thus begins Edward Wilson's famous work Sociobiology. Wilson
appears to deny the seriousness of the question of suicide, and,
perhaps, even the possibility of suicide, by arguing that the emotions
flooding consciousness will determine what the individual does and
that the genes ultimately determine those emotional inputs. Of course,
Wilson later greatly modified this assertion of biological determinism,
especially in On Human Nature (1978). But, while he did attempt to
explain one seemingly unadaptive form of behavior, homosexuality, he
did not return to the question of suicide. This is most unfortunate, since
suicide not only poses a great challenge to the theory of evolution, as
deCatanzaro makes so clear, but also, being far more restricted to
human beings than homosexuality, will ultimately reveal far more to us
about human nature. Suicide has special bearing on the question of
biological and cultural determinism versus individual freedom.
DeCatanzaro has certainly done a masterful job of bringing a mass
of interdisciplinary research and theory to bear on suicide and relating
this to basic questions about the theory of evolution. He has shown
beautiful restraint and interdisciplinary balance in evaluating the many
complex factors that seem to be involved in the many complex forms
of suicide. But I believe there are two ways in which he could
strengthen his argument.
First, there is the vital question of the validity and reliability of the
data. It is all too clear that there are very broad statistical biases in the
official statistics on suicide (Douglas 1967). There are now at least
three comparative studies of medical examiner and coroner proce-
dures for certifying the causes of death that reveal how these biases
come about. (See, for example, Douglas 1971, pp. 79-132.) Some of
the works deCatanzaro draws on rely heavily on official information.
Fortunately, whether by happenstance or by design, some of the major
biases in the official statistics undercount cases in the direction
supportive of his major arguments. Any experience in the medical
world quickly reveals that the intentional killing of oneself (suicide), and
intentional cooperation in "pulling the plug" (ending medical support),
are far more common among terminally ill patients than is indicated by
official statistics. Doctors, nurses, and others cooperate tacitly in not
noticing, hence hiding, these suicides. Since most of these patients are
older and are all, according to medical prognosis, doomed to die soon,
their deaths fit the model of "evolutionary tolerance." Some of these
deaths also involve altruism because they remove a burden from the
families.
DeCatanzaro could also have strengthened his argument greatly by
carefully considering the arguments of Jean Baechler in Suicides
(English translation 1979; originally published 1975). Starting with the
conclusion that official information on suicide is often invalid, generally
highly unreliable, and always very shallow, Baechler carried out the
most extensive investigation of case studies yet accomplished, both
within Western cultures and cross-culturally. His study is interdisciplin-
ary in the same creative spirit as that of deCatanzaro. But Baechler
was able to propose a systematic theory of suicide that goes beyond
specifying the possible contributing factors, the level of theorizing that
deCatanzaro feels constrained to remain at because of the more
limited data with which he was dealing.
Though the details are far too complex to present here, Baechler
has summarized his theory very clearly. He argues, as do most
theorists of human nature today (e.g., Wilson 1978), that any adequate
theory of behavior must take into consideration the initial genetic inputs
(human nature); the lifetime of experience and learning that build upon
the genetic inputs but are by genetic programming partially autono-
mous; the relatively stable personality tendencies to react in specific
ways to specific situations that result from the interactions of the
genetic inputs with experience; and the immediate (concrete) situa-
tions that people encounter in their lives. (One part of the experience is
the so-called culture variable, but it is vital to see that "culture" is
important only insofar as it directly affects the experience of individu-
als. In all civilizations cultural experience is quite pluralistic, so that
different individuals may experience different parts of the culture.)
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 275
 Commentary/deCatanzaro: Biology of suicide
Baechler, unlike most social scientists, but very like geneticists and
behavioral biologists, recognizes that "life is a struggle" (or full of
conflict). Each individual, presumably as a result of genetic program-
ming, tries to protect himself in the struggle, and allow himself to
continue actively pursuing the struggle, by building personal "security"
for the self. This is done primarily by using two basic strategies, which
are both social strategies because of the fundamental importance of
social relations in human life. These are the strategies of power over
others and dependency upon others. Most people use a combination
of these two because this is the optimal strategy for building personal
security, but those thwarted in one may pursue the other almost
exclusively, thus making themselves more vulnerable to emotional
insecurity from defeats in the struggles of life. Pursuit of either or both
strategies may in itself demand that one risk death by resorting to
suicidal actions in certain situations. (This would, for example, be true
when a suicidal gesture used to get revenge, which enhances one's
power, leads to death.) But the use of these strategies may also fail.
(For example, one may be rejected in love-dependency.) When this
happens in extreme ways, thus producing extreme feelings of rejection
or failure, then individuals respond with varying degrees of depression
and hopelessness, presumably because of genetic and personality
differences (which may be amplified or damped by cultural experi-
ence). It is only in extreme situations that "one can be driven to the
point where one can no longer act and seize on the possibility of
leaving the fight" (Baechler 1979, p. 455).
We must note that the individual may "seize" the possibility. We
must also note that the individual may or may not choose to risk his life
by pursuing his strategies to extreme positions. For, ultimately, after all
of the powerful predisposing factors are considered, human beings do
have some freedom of individual choice. "The conclusion is unambigu-
ously imposed: suicides, in all their different meanings, are neither
perversions nor scandals. They are coeval with the human condition,
for human life is an endless struggle where each of us by his own free
will risks his life each moment" (Baechler 1979, p. 451).
There will always remain some element of individual mystery to
suicides, as there will to all human action, because man is "con-
demned" by genetic programming to make his own choices in the full
light of the genetically determined emotional inputs from the
hypothalamic-limbic system, the situations he faces, and his aspira-
tions. Man is predetermined biologically to be both constrained and
free. Suicides are the outcome of profound personal struggles and the
ultimate choices that are made to deal with those struggles.
by Marshall P. Duke
Department of Psychology. Emory University. Atlanta. Ga. 30322
Feasting on the sociobiology of suicide: somehow I
still feel hungry . . .
While I applaud deCatanzaro's efforts to delineate a biological
perspective on suicide, I came away from the article with a mixture of
puzzlement and disappointment. I almost feel as if I had been prepared
for a sumptuous feast and the waiter stopped serving the food just after
the appetizer. I was intrigued by the author's goal of establishing the
adaptational-biological-evolutionary impact upon suicide and looked
forward to the presentation of the models that were discussed. I am
assuming that other important psychological and sociological models
were deleted in the interests of exposition, so after simply registering
my concern that several viable and empirically supported models were
not discussed, I will proceed with some reactions to the materials
actually included in the target article.
A problem I had initially was accepting deCatanzaro's assumption
that "the innate component [of biological development] should reflect
countless generations of selection for adaptiveness, and we should
thus be able to account for most [!] behavior in terms of evolutionary
contingencies that affect it." Such a statement cries out for an
empirical foundation upon which to rest. If I, as a reader, cannot accept
this assumption (and I'm not sure I can) the worth of the remaining
effort put forth by the author is in question.
However, given a scholarly acceptance of deCatanzaro's assump-
276 THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3
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tion that suicide could in fact be evolutionary based, I shall try to react
to the models described. With regard to the first model, that of suicide
as a learned behavior, it seems that the evidence presented actually
argues against the hereditary component. The presence of suicidal
behavior in humans and not other animals and the fact that "much of
the available evidence concerning suicide can be construed as support
for a model based on learning and cultural evolution" seem to preclude
a strong genetic component. Further, such findings as that suicide is
relatively infrequent in Catholic countries, for example, certainly
suggest that cultural and learning factors are most important - unless,
of course, Catholics have in some way selected genetically for
nonsuicidal genes.
The learning theory model is not the only one with which I had
trouble. I also had some difficulty with the pathology model and the
altruism approach. My main question with regard to the former deals
with whether or not the capacity to cope must, as the author seems to
imply, be inherited; cannot coping methods also be learned? As for the
altruism model, it strikes me that the author is confounding what is with
regard to isolation with what an individual feels with regard to isolation.
The net result is ascription of motive based upon outcome rather than
upon actual experience.
While a longer commentary would afford me the opportunity to react
further to the three models I have very briefly mentioned, in the
remainder of my comments I would prefer to focus on the model
dealing with evolutionary tolerance. Here deCatanzaro states that
suicide is likely to be selected for in individuals who experience
difficulties, become nonproductive, and see little hope for improvement
in the future. It is noted that persons incapable of reproducing or
otherwise promoting their genes are those in whom suicide would be
evolutionary "tolerated." As support for this contention the author
cites data indicating that suicide is more common among single and
divorced persons and those with unstable marriages. Also cited as
"predictors" of suicide are social isolation and increasing age. All of
these factors are seen as "signs" of decreased chances of promoting
one's genes. For example, it is stated that divorce and suicide are
related, and it is thus concluded that suicide is tolerated in the person
who, once divorced, has suffered a reduction in probability of repro-
duction. A more parsimonious and probably more easily supported
explanation, however, might be that suicide and divorce are both the
result of faulty interpersonal abilities and maladaptive learning. Like-
wise, the other "signs" of decreased reproductivity, such as unstable
marriage, nonmarriage and isolation, may also be related to suicide
only by virtue of the fact that all of them may stem from basic
psychological dysfunctions. DeCatanzaro's conclusions concerning
the relations between suicide and signs of reduced ability to promote
the genes may be the result of a common logical error: b and c occur
concurrently; what is their relationship? Conclusion I: b causes c
(divorce causes suicide); or conclusion II: a causes b (a representing
basic interpersonal dysfunction); a causes c; thus b and c occur
together (faulty relating causes divorce; faulty relating causes isolation
and depression).
In addition to making this logical mistake, the author seems to leap
from prediction of suicide to precipitation of suicide. On p. 270 for
example, it is first stated that "social isolation is a good predictor of
suicide." One sentence later the author writes, "social isolation can
precipitate suicide." Again, I am left with the question of whether social
isolation causes suicide or whether suicide and social isolation are
both caused by something else again.
Another problematic implication of the evolutionary tolerance
perspective runs counter to data cited by the author. It is correctly
noted that suicide occurs more often in males than in females,
regardless of age, yet males typically continue to be able to reproduce
far beyond the average age for female menopause. If being unfit to
reproduce removes the negative pressure against suicide, one would
expect postmenopausal females to be more suicidal than men ot
equivalent age - statistically this is simply not the case.
While my comments to this point have been theoretical and logical,
my final reactions to the article rest on some scholarly concerns. The
extensive use of secondary sources seems to have laid the author
open to some criticism. For example, I was struck by the statement

that "indeed, reserpine, which depletes catecholamines and serotonin,
has been reported to induce suicide." Having never felt that such a
relationship had been clearly supported, I went to one of deCatanza-
ro's sources (Lester 1972) and found the following statement:
It is claimed from time to time that particular drugs produce depres-
sion and suicidal tendencies in patients. Berger (1967) reviewed the
evidence that rauwolfia serpentia (reserpine) induces suicidal behav-
ior. It is difficult to evaluate the research since the patients given the
drug are often not matched for symptoms and degree of distur-
bance with patients given the placebo. Secondly, those who react
suicidally to the drug may have pre-existing tendencies to depres-
sion. Berger concluded that the bulk of evidence available indicated
that reserpine was more likely than any other central nervous system
depressant to produce depression and lead to suicide. But this
conclusion must be regarded as a tentative one. (Lester 1972, p.
309)
Indeed, Lester |q.v.| presents a very different picture of the reserpine-
suicide relationship.
In what may be seen as a further scholarly lapse, in discussing the
possible relation between genetics and suicide deCatanzaro stated
that "Lester (1968), reanalyzing these (Kallman'sl data with more
sophisticated statistical methods, concluded that they did indicate
such a relationship." In fact, Lester (1968) actually stated, "It must be
concluded, therefore, that it is not possible to dismiss inherited factors
in the determination of suicidal behavior on the basis of present
evidence" (p. 320). There is quite a difference between concluding
that a relationship is indicated and that one cannot yet be ruled out.
I began by mentioning that the target article left me feeling unsatis-
fied. Much of this feeling stemmed from the dearth of clear research
direction at the end of the article. I still don't know where we ought to
go; or whether the genetic approach is worth pursuing aggressively
and enthusiastically. While the evolutionary argument is interesting, it
seems that most of the data support a minimal genetic component
in the act of suicide itself. On the basis of his review of suicide Lester
(1972) has stated: "It is unlikely that the tendency to suicide per se is
inherited, but it is possible that genetic factors play a role in the
formation of personality types susceptible to suicide or in the incidence
of particular mental disorders which facilitiate the appearance of
self-destructive behavior" (p. 25).
In general, I was disappointed by many things in deCatanzaro's
laudable attempt to establish an evolutionary perspective on suicidal
behavior. Had I not been distracted by a number of what seemed to be
forced arguments, illogical conclusions, and scholarly lapses, I might
have come away with a more favorable view of the effort.
by Maurice L. Farber
Department ot Psychology, University of Connecticut, Storrs, Conn. 06268
Suicide as natural selection
Clearly, as deCatanzaro's target article asserts, suicide presents a
theoretical paradox in that it appears to run counter to the process of
evolution. His model 1 considers the possibility that suicide could be
conceived as outside this process. But, if it can be demonstrated that
suicide involves natural selection, it must then be viewed as being
within the evolutionary process and contrary to the model. This
commentary will attempt to place suicide within the evolutionary
framework.
Although it is true that suicide operates against the survival of the
organism, it does so selectively. DeCatanzaro's model IV makes use of
this fact, but, as will be shown, supports its conclusions with somewhat
flawed analysis.
Suicidal behavior is determined not only by situational stresses, but
also by characterological vulnerability to such stress (Farber 1968).
This vulnerability would appear to involve both acquired and innate
components. Of special interest to us here are individuals with these
innate vulnerabilities. In suicide, precisely such people are eliminated.
Just as the slowest-running deer are eliminated by the pursuing wolves,
so too are humans with vulnerable character structures eliminated by
environmental stress. Although it is done by their own hand, the effect
https://doi.org/10.1017/S0140525X00004799 Published online by Cambridge University Press
Commentary/deCatanzaro: Biology of suicide
is the same. Thus suicide can be placed squarely in the process of
natural selection and evolution.
Model IV, by suggesting that sociologically defined groups with
relatively high suicide rates produce, on average, fewer offspring and
provide less favorable conditions for their genes, adds support to the
notion that the elimination of this group from the gene pool is
beneficial.
One must, however, interpose a caveat. Although it is true that these
groups have relatively high suicide rates, many suicides are not in
these categories. Moreover, the old were once young and may earlier
have produced large families. The divorced were once married and
reproductive. The isolated may well at some point have been parents.
There exist many offspring of forebears who have committed suicide.
The contrast between the suicidal and the nonsuicidal with regard to
reproduction, at least, would appear to be somewhat overdrawn.
Nevertheless, there remains some force in the argument. Those
eliminated tend to provide neither good quality nor quantity of genetic
material.
But there is another effect: the loss of valuable genes. Along with
genes vulnerable to stress there may exist desirable gene combina-
tions in an individual, even some for genius. If the suicide is young, he
may well not yet have reproduced. If older, his loss may impair the
welfare of the genes of others. On balance, however, one might
speculate that the net effect of the two countervailing forces is
weighted toward elimination of the undesirable. The infrequent loss of
high-quality genes would not seem to outweigh the probably more
frequent elimination of stress-susceptible ones. Not only can suicide be
"tolerated" by evolution, as deCatanzaro suggests, but, on balance,
from a purely evolutionary point of view and ignoring the personal grief
and social loss, it may well entail a modest gain.
Overall, one can congratulate the author on having addressed
himself incisively to a problem rarely considered by psychologists.
by Gary Frieden
ASt Market Research. Inc.. Los Angeles. Calit. 90046
Self-destructive behavior: suicide, shocks, and worms
DeCatanzaro's approach considering suicide in terms of four models,
is an interesting one. I would like to expand on his points pertaining to
learning, social, and cognitive factors. The purpose of this commentary
is to elevate the importance of these variables in any model of suicide.
DeCatanzaro stated that self-destructive behavior may occur when
organisms have not learned the negative contingencies surrounding
such acts. In a review of self-punitive behavior (Renner and Tinsley
1976), it has been shown that such maladaptive behavior extinguishes
when organisms become aware of the negative contingencies asso-
ciated with behaving in a self-punitive fashion. Similarly, learning
experiences can actually produce self-punitive behavior as well. For
example, one study (Stretch, Orloff, and Dalrymple 1968) has demon-
strated that an aversive stimulus, such as an electric shock, can be
reinforcing because it simultaneously serves as a discriminative stimu-
lus for the absence of punishment: organisms first received avoidance
training in that their responses could indefinitely delay an electric
shock. In a second phase, shocks were presented on a fixed-interval
schedule; nevertheless, the organisms' responses increased! Interest-
ingly, the only way the animal could be cued as to the absence of
shock in a subsequent interval was to cause the shock actually to
occur. Thus, the punishment was really a discriminative stimulus for the
absence of punishment; as a result, it was reinforcing. A most intriguing
anomaly indeed - a punisher and a reinforcer at the same time.
This aspect of learning relates to a point made by deCatanzaro. He
stated that attempted and completed suicide differed in that there
could be secondary gains from attempted suicide (perhaps sympathy
or better treatment from other people), but not from completed suicide.
Yet he later added that completed suicide could result in a removal of
pain or other aversive events - that is, negative reinforcement. It
seems, then, that there are gains to be derived from both forms of
suicide, depending on the individual's needs at the time of the act.
There is another factor that can contribute to self-destructive
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 277
 Commentary/deCatanzaro: Biology of suicide
behavior; deCatanzaro referred to this as "complex and unusual
cognitive events." I would like to expand on this category by briefly
explaining a model of "role confusion" that I am currently developing.
Suppose people were to suddenly wonder who they really were. They
could become confused about whether a previous sample of their
behavior represented merely role playing or their "real" self. At such
times, they may wish to "test" themselves to find out more about
themselves (and to answer the question, "Was that really me?"). This
behavioral test would consist of behaviors similar to the acts that
produced the confusion itself. Lo and behold, people depressed about
the possibility that they are worthless, incompetent, masochistic,
sadistic, and so on, would continue responding in that fashion in an
effort to resolve the role confusion.
The above model was tested (Frieden 1978) by having respondents
role-play as if they enjoyed suffering - that is, as if they were masoch-
ists. Some people were then led to experience role confusion by
suggesting to them that their previous behavior may actually have
reflected their true desires. Those who believed they had acted on their
own accord later chose to be bound, blindfolded, and verbally abused,
and to receive painful electric shocks for twenty minutes. Not only that,
but when later asked if they would be willing to eat a dead worm, 75%
agreed! (No one ever experienced any of the above - once they made
their decision, the study ended, and they were extensively debriefed.)
What does this tell us about suicide? It should make us aware that
the "complex and unusual cognitive events" to which deCatanzaro
refers can be quite powerful indeed. Not wishing to play down the
biological role, I am instead citing the importance of cognitive factors.
When people attribute causality to themselves for unfortunate events
that they did not in reality produce, the self-questioning process can be
maladaptive. In another study (Frieden and Fraser 1978), an inverse
relation was found between one's masochistic tendencies and one's
sense of self-worth - the more subjects enjoyed suffering, the worse
they felt about themselves. Furthermore, those whose self-esteem was
experimentally lowered in the laboratory subsequently chose to suffer
voluntarily. It is not surprising, as deCatanzaro pointed out, that guilt
feelings and negative self-concepts correlate with suicide (Kaplan and
Pokorny 1976).
The learning, social, and cognitive factors, then, cannot and should
not be viewed as having only a minor influence on suicidal behavior.
Whether the gene pool is enhanced or unaffected by one's committing
suicide, the tragic act itself has multiple causes, all of which deserve
further study.
Acknowledgment
I wish to thank Scott C. Fraser, at the University of Southern California, for
providing valuable assistance with the understanding and implementation of this
research.
by William J. Hamilton III
Division of Environmental Studies. University of California, Davis, Calif. 95616
Do nonhuman animals commit suicide?
DeCatanzaro's evaluation of the biological basis of suicide follows
logically from the current attempt to evaluate all aspects of human
behavior in the context of comparative animal behavior and compara-
tive analysis of diverse human societies. This is at least a strong
alternative to the classical psychiatrists' motivational theories.
This approach is effective and needs no elaboration by me. No
doubt some of those concerned with suicide prevention will fault any
attempt to consider suicide as an adaptive process. But it is clear that
in the case of some human hunter-gatherer societies the kin selection
hypothesis for suicide as an adaptive process has both strong logical
and empirical support.
Human kin selection hypotheses can be placed in direct contrast to
Skinner's (1969) contention that suicide must be learned because it is
always maladaptive. I single out this evolution-culture dichotomy
because Skinner's alternative automatically throws support to explana-
tions lying on the cultural side, an alternative that critics of genetic
determinism (hard-core sociobiology) should be quick to seize upon.
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Of course Skinner's (1969) argument was presented before kin
selection had much of a chance to be developed theoretically or with
significant examples (as is now the case) for bird and mammal
species.
DeCatanzaro's discussion of the extent of suicide by animals can be
extended. He notes that suicide has not been commonly identified by
ethologists in field situations and suggests the possibility that this may
result from inadequate observations. In my opinion, however, we are
today not in a situation where this is a very probable explanation.
Rather, the main message I derive from comparative animal behavior is
that, in spite of the intensive study of thousands of animal species in
nature, suicidelike behavior is rarely found, and where it occurs it
probably has its basis in parental behavior or kin selection. It is not
expressed by individuals who have direct residual reproductive poten-
tial, in striking contrast to most human suicide.
Because, as deCatanzaro points out, suicide has been a little-
treated subject in this context, any analysis will be incomplete. My own
consideration identifies four patterns of self-destructive behavior:
Emigration. Animals may make one-way movements to areas
where they do not survive. Many animals move to and from alternative
ranges. Such migrations, although they are hazardous and may involve
massive mortality, are clearly adaptive, involving movement to and
from seasonally available resources and breeding grounds. To the
extent that emigrators or their descendants do not eventually return to
the site of origin, they contribute nothing to the population of origin and
can therefore not influence the adaptive characteristics of the source
population. But to be adaptive the return need not be immediate and
can involve the progeny of the emigrators. This is so, for example, in
the case of the painted lady butterfly, Vanessa cardui, which resides in
Mexico. Northward emigration from there in the summer involves a
wave of movement including three generations extending to Canada.
The fall return mirrors the exodus, and the annual return to Mexico is in
essence a multigeneration migration and therefore adaptive (Arthur
Shapiro, personal communication).
The sometimes spectacular and apparently one-way movement of
tropical butterflies and moths may not involve a return to the area of
origin. If not, the adaptive explanation must resort to parental behavior
or kin selection arguments involving relief of population pressure upon
progeny or other relatives living under submarginal conditions. The
pattern in the case of a classical emigrator, the Norwegian lemming
{Lemus lemmus), appears to involve movement up and down slopes to
areas of seasonally preferred habitat (Kalela 1962). For some individu-
als this may involve movement to places where they cannot survive
(Clough 1965), but this is little different from the imperfection of
long-distance migration by birds, whales, and insects, which may
sometimes include massive mortality. In the face of completely biohos-
tile conditions, Caribbean lizards may leap into the sea and swim and
float away with a high probability of death (Schoener, personal
communication). But the consequence of remaining in an environment
without adequate food is certain death, and movements of terrestrial
lizards into the open sea can, in fact, be best identified as an optimal,
albeit high risk, strategy. Additional supportive evidence for this
interpretation is that lizard species living in coastal environments and
on small islands are, in fact, better able to stay afloat for prolonged
intervals than interior mainland species (Schoener, personal communi-
cation). Thus, positive selection for swimming ability by a terrestrial
animal is established.
Completed suicide by social insects. The significance of self-
destructive behavior by "individual" social insects is unresolved. There
are at least two theoretical alternatives to the treatment of nonrepro-
ductive worker members of social insect societies. One is to treat them
as biological individuals like any other animal population and to include
them in comparative discussions. This tactic was freely used by E. O.
Wilson (1975) in his now classical book, Sociobiology. An alternative is
to consider such "individuals" as in every sense involuntary evolution-
ary slaves of the controlling reproductive caste. This alternative
suggests that these individuals have no control over their biological
destiny other than that imposed by their genetic and facultative control
of the reproductive members of the colony. I strongly favor this
alternative and would treat a worker honeybee, for example, as no

more in control of its destiny than is the stinging cell of a hydra or an
anemone. The genetic reproductive potential of asexual insect workers
acts through the survival of the parent and since the parent controls
the progeny and its actions, it seems incorrect to include the behavioral
actions of these asexual workers as comparatively analogous to social
phenomena in general and suicide in particular. In other words, a
worker honeybee is not committing suicide when it stings a bear on the
nose, pulls away its stinger, and loses its life, any more than a lizard is
doing so when its tail breaks away in the extreme emergency of an
attack. Energy is gone, but reproductive potential remains if the
evasive tactic is successful. Suicide of a biological individual is not
involved.
Extreme parental investment. Parental investment in progeny is
made in proportion to the optimum fitness of progeny. If the optimum
strategy is to maximize reproduction once and for all, as is the case for
those salmon species that spawn and die, then this is a route
life-history strategies can take. A spawning salmon can be said to be
committing suicide. But it is an adaptive strategy and an evolutionary
choice of alternatives. Some salmon species have evolved, as an
alternative, a less complete allocation of energy resources to spawn-
ing, a return to the sea, and the possibility of returning in subsequent
years to spawn again. All of this is potentially explicable in terms of
classical natural selection, and a learning process is not implicated. All
other well studied examples of parental investment (Trivers 1972)
seem to tit well into an adaptive evolutionary framework and thus lie
outside the current discussion.
Antipredation behavior. Animals sometimes take unnecessary
risks in the presence of predators. All mobbing behavior and alarm
calling falls into this category. An extensive literature supports adaptive
explanations of these phenomena involving primates, birds, and
mammals.
A different logic has been advanced to explain the relationship of the
risk-taking behavior of certain butterfly species to their vertebrate
predators. Adults or larvae may offer themselves to predators when
senescent (postreproductive) (Blest 1963) or when doomed by surely
lethal internal parasites (Shapiro 1975). One adaptive benefit of such
behavior may be the indoctrination of predators in the case of
distasteful and colorful moths (Blest 1963). Shapiro's clever explana-
tions of the apparently suicidal behavior of parasitized nymphalid
butterfly larvae seem plausible, but the phenomenon is largely un-
studied, and a more secure understanding of it requires further study.
DeCatanzaro does not make a great distinction between self-
mutilation in captivity and in nature. The paraphernalia (artifactual
basis) for suicide is more available in captive situations. In addition, it
seems likely that many of the self-destructive propensities of captive
organisms are the result of the expression of behavior patterns that
would be adaptive in nature. For example, wolves and other canines
range widely daily, regardless of their hunting success (except when
denning). This insures that they do not overhunt local areas, that they
encounter territorial transgressors, and that they develop an accurate
learned inventory of the prey within their space. The same behavior,
expressed in captivity, may result in a pacing behavior resulting in
physical damage and an early death. Other forms of subsuicidal
behavior by captive primates are also associated with an environment
that does not include the social or physical conditions that would
reorient this self-destructive activity in nature.
Most contemporary human populations are to some extent out of
evolutionary context and live in self-imposed captivity. I therefore feel
that the only viable comparison with animals is to caged creatures.
DeCatanzaro notes that there are age-specific expressions of
behavior. Few evolutionary oriented biologists would deny that this is
true and that for many animal species, and probably for humans, this
is, in part, a heritable set of traits. Can this lead to the expression of
maladaptive behavior at any age? By definition, maladaptive charac-
ters cannot be selected for and, given even marginal reproductive
prospects, there is going to be selective pressure against suicide. To
evaluate the costs and benefits of suicide it will be necessary to
evaluate individuals who have made unsuccessful attempts at suicide
(e.g. by leaping from tall buildings and surviving) and to compare this
with the positive and negative benefits to relatives of completed
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Commentary/deCatanzaro: Biology of suicide
suicides. If it can be shown that there is any significant increase in
reproductive success for surviving individuals, even if it is massively
less than that for individuals who do not express such behavior,
selection against the maladaptive (suicidal) behavior will persist. The
more difficult part of the equation to analyze is: what would have been
the benefit to relatives if the suicide had been successful? The difficulty
of accurately making such a measurement leaves this analysis in the
area of speculation, probably forever.
In conclusion, I must answer the question of animal suicide with a
probable no. Few if any animals in nature commit completed suicide.
The closest analogues appear to be emigrations of insects and
self-sacrifice of doomed insects to predators; both are poorly studied
phenomena. In captivity, the expression of subsuicidal behavior is, by
contrast, commonplace. Perhaps the greatest insight an animal-human
comparative analysis offers is that humans have, in their transition from
nature to a form of self-imposed captivity, voluntarily created out-
of-evolutionary-context environments where zoolike self-destructive
behavior is commonplace.
Humans were probably a highly kin selected species. Thus, a part of
their behavioral repertoire may have been shaped by earlier evolution-
ary conditions where self-sacrifice via suicide was an adaptive
process. But study of contemporary suicide by humans may be no
more meaningful to evolution than is the study of analogous behavior
by animals in zoos and laboratories.
by L. D. Hankoff and William J. Turner
Department of Psychiatry and School of Medicine. State University ot New York at
Stony Brook. Stony Brook, N.Y. It 794
A nontheory of suicide
The heterogeneity of suicidal behavior is a commanding issue in any
effort to understand the many forms that suicidal behavior takes. The
classification of suicidal behavior must take into consideration the
broad range of individuals, acts, and situations involved. Considering
the basic parameters of age, sex, and fatal versus nonfatal forms, a
matrix of eight cells based simply on a dichotomization of each of
these three variables is possible. Obviously, a young woman who
carries out an act of attempted suicide poses a picture and a challenge
to our understanding radically different from that of an old man who
commits suicide.
The age distribution of suicide varies with sex in most western
countries. Among men suicide risk increases with age, whereas among
women the age rate curve is a flatter one which falls off in the fifties.
The risk of suicide is invariably higher among men than among women.
In the case of attempted suicide, both of these relationships are
changed. Attempted suicides occur more frequently among women
than men, and the risk of an attempt decreases with age for both
sexes, by and large. The peak for suicide attempts is around 20 years
of age for both men and women, in contrast to the older age peaks for
fatal suicides in both sexes.
Impressive efforts at a total view of suicidal behavior date back
many decades. Durkheim's (1951) volume, which was first published in
French in 1897, presents a four-way classification of suicidal fatalities
that has stood the test of time and remains valuable in stimulating
continuing research. Durkheim divided suicide into egoistic, anomic,
altruistic, and fatalistic types, relating each to a variety of real situations
that he observed in his own data from France in the nineteenth century
and a wide variety of other sources dating back to the eighteenth
century and covering all of Europe.
Durkheim, as the father of modern sociology, emphasized and
clarified social influences in the genesis of suicide. His understanding
remains a valuable vantage point for our current view of suicide. While
the emphasis in most modern approaches has been on psychopathol-
ogy, and the refinements in diagnostic classifications in psychiatry
have helped establish firm links with suicidal behavior, the influence of
social factors remains quite evident in many cases.
The most dramatic example of suicide in the past decade was the
Jonestown, Guyana, incident which involved over 900 individuals.
Clearly, this large group of men and women of all ages must have
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 279
 Commentary/deCatanzaro: Biology of suicide
included many who were entirely free of psychopathology and who
were dedicated to suicide on the basis of social and peer-group
influences. Durkheim allowed for combinations of his four suicidal
categories, and it is very likely that the Jonestown mass suicide would
have been classified by Durkheim as an example of altruistic-fatalistic
suicide. Durkheim used the term "altruistic" in a specific sociological
sense, roughly equivalent to our understanding of socially prescribed
and formally defined behavior, not as altruism in the sense of behavior
devoted to the welfare of an individual or a group.
Durkheim was aware of suicide in traditional and primitive societies
although his data were obviously not nearly as exact as those relating
to the industrialized countries of Europe. The data on suicide in
underdeveloped, traditional, and primitive societies indicate that het-
erogeneity considerations hold for these societies as well. It is often
assumed that primitive society, free of urban and industrial pressures,
is relatively free of suicide. This is not the case, and relatively high
suicide rates have been associated with specific groups. For example,
B. Malinowsky spoke of the frequent suicides among the Trobriand
Islanders he studied and Peter Freuchen made the same observation
of the Eskimos with whom he lived for many years.
Suicide and biological factors. The clear-cut findings relating
suicide to biological factors are few in number. No genetic pattern in
terms of inheritance has emerged in relation to suicidal behavior. On
the other hand, the relation of suicidal risk to affective disorder is well
established. The affective-disorder patient has a risk at least fifteen
times that of the general population of dying by suicide. The few
biochemical studies of suicidal individuals suggest that there are
biogenic amine changes in specific diencephalic and brainstem areas
that may distinguish the suicidal from the nonsuicidal brain. However,
such findings are few in number and may not be distinguishing a
suicidal affective-disorder patient from the nonsuicidal affective-
disorder patient.
Any correlation between suicide and survival of the human species
does not appear strong in view of the fact that most suicidal
fatalities occur among individuals who are beyond the child-bearing
period. It seems unlikely, therefore, that suicide affects mankind's
survival one way or the other, in evolutionary terms. While genetic and
biological factors may be at work, the likelihood of determining such a
connection depends upon an understanding of the heterogeneity of
suicidal behavior. Does our biological theory relate suicide to biological
correlates in the old or the young, in men or women, or in fatal suicides
or attempted suicides? Genetic factors may be significant in terms of
some suicides, but it is necessary to isolate that subgroup of suicidal
behavior in order to demonstrate significant correlations and arrive at
any proof of the theory. Because of the wide variety of suicidal
behaviors and suicidal individuals, any theoretical framework is
doomed to some success since there is something in suicidal behavior
for every point of view. The totality of theoretical efforts in terms of
suicide emphasizes again the heterogeneity of this form of behavior
and the need to seek specific theories for specific and operationally
defined subgroups within the broad group of suicidal behaviors.
by Morton G. Harmatz
Department ol Psychology, University ol Massachusetts. Amherst. Maes. 01003
The biological perspective on suicide: to be or not to
be - is that sociobiology?
The examination of suicide from an evolutionary and sociobiological
perspective has led deCatanzaro to some interesting speculations
about suicide and the special challenge it poses for sociobiological
theory. The models for suicide that are presented and compared are
meant to provide a springboard to further research and analysis of the
problem. The discussion is stimulating, but there are problems at the
very core of the analysis that raise serious questions about the value of
a sociobiological analysis of suicide as presented here.
One problem emerges at the very beginning of the discussion with
deCatanzaro's use of the term suicide. He approaches it simply as a
behavior. Suicide is not a behavior in the formal sense of that term, but
rather a descriptive label for the outcome of a large variety of other
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behaviors. This is more than hair splitting over the fact that the word
suicide is a noun and not a verb. Thinking about suicide as a single
behavior creates an illusion of a single behavioral act, which may be
convenient for discussing the sociobiological transmission of the
activity but is a serious distortion of reality. In a more sophisticated and
appropriate analysis suicide would be approached as the outcome of
any of a large variety of motivations, in response to varied antecedent
conditions, with totally different behavioral responses used to produce
that outcome. To use so vague and overinclusive a term in an attempt
at scientific analysis is bound to lead to great confusion - and it does.
The oversimplification of the concept of suicide provides a shaky
foundation to the models deCatanzaro puts forward for discussion.
Our understanding of suicide is subject to further distortion by his view
that it does not have environmental outcomes for the victim. DeCatan-
zaro states that "completed suicide is perhaps the most unambiguous
form of self-destructive behavior, in that it is difficult to conceive of any
secondary gain that could accrue to any individual not surviving his
suicidal act." This statement ignores the human ability to experience
"gains" from symbolic acts, fantasized events, future projects for the
self, and so on. DeCatanzaro's view of gain is much too narrow to
encompass people's cognitive ability to go well beyond concrete
reinforcers. Such a statement also ignores the literature on suicide.
Much of the research on suicide postmortems has demonstrated the
imagined impact the suicidal act is meant to have on the world left
behind. Biographical and novelistic presentations of the suicidal experi-
ence are even more persuasive in their support of just how very
important the "gains" are to the person contemplating or committing
suicide. Clinicians who have worked with potential suicides and suicide
attempters are quickly made aware of just how important is the
ideation about the aftereffects of the act. It is the rare individual who
does not have a desirable outcome in mind - that is, a strong
reinforcement for the event. Not infrequently there is a cognitive
distortion of the fact that they themselves will no longer exist after the
suicidal act to enjoy the outcome. The clinician's therapeutic activity
must often be directed toward correcting this distortion. Put in this light,
suicide is clearly more consistent with learning principles than the
unexplainable, simple behavioral act without gain as proposed by
deCatanzaro.
Even if we could accept deCatanzaro's view of suicide as behavior
with no gain for the person, we would have to question strongly his
defense of a genetic transmission of that behavior. The weight of the
empirical evidence is much more supportive of a sociopsychological
determination of suicide. Differences in suicide rates in different
countries, religious groups, and times of the year (seasons, holidays)
and in response to social upheaval or stress (war, famine) all support
environmental determination. The sociopsychological argument finds
strongest support in the changes in the suicide rates in cultures
undergoing dramatic social change such as industrialization, urbaniza-
tion, and political upheaval. In these examples the genetic pool is the
same yet the rate reflects environmental determinants at both social
and individual levels.
DeCatanzaro proposes four models for suicide, each representing a
different perspective on the act. Each perspective is well presented,
with strengths and weaknesses clearly stated. While he argues in his
conclusion that these models are not mutually exclusive. deCatanzaro
does favor the fourth one as the best support for a sociobiological
position. This model indicates that suicide becomes possible when
individuals are less capable of reproducing their genes. This model
attempts to integrate evolutionary and environmental factors by
proposing a release from constraints against suicide when the gene
pool will not be affected. It is not clear however, how this change is
mediated in the individual, nor how such a releasing mechanism would
be communicated within the species.
As a central argument for the fourth model, deCatanzaro states that
suicide occurs in those individuals who are expecting little improvement
in their future. It could well be argued that suicidal ideation is actually an
attempt to create a desirable future from a highly undesirable present.
This is suicide as a "rite de passage" to a better existence, unrealistic
as it may be. A clinical sign of concern is the sudden change to a joyful
mood in the potentially suicidal person, which is frequently the time of

greatest danger. Suicide is seen as the solution, not the problem.
Again, a learning interpretation seems to fit the data better.
We have argued that much of the confusion in the discussion results
from the way deCatanzaro uses the concept of suicide. These
criticisms do not necessarily mean there is no sociobiological compo-
nent to self-injurious behavior. An alternative approach is to move
away from the confusion created by the concept of suicide and look to
a more general state of the organism as the mediator of these potential
responses. Depression may well prove to be the more appropriate
mechanism for understanding suicide as a sociobiological event.
Evidence is mounting that humans and other animals appear to have a
neurophysiological capability for experiencing depression. The depres-
sion appears to be activated by any of a large number of environmen-
tal and physical events. This could be the biological mechanism for the
removal of weak or damaged organisms from the group and thus have
selective value for the species. The depression could lead to any of a
number of behavioral outcomes based on social and individual learning
- self-injurious behavior being but one possibility. Other organisms may
simply "waste away," while still others may succumb to natural
predators. The work on learned helplessness could easily be incorpo-
rated into this model. The phenomenon of depression provides a much
better fit to both the arguments raised by deCatanzaro and may well
prove to be the better construct on which to expend the research effort
called for by deCatanzaro in his conclusion.
by David Lester
Richard Stockton State College. Faculty of Social and Behavioral Sciences.
Pomona. N.J. 08240
The categorization of suicide
Suicide is far from easy to categorize. What kind of behavior is it? Does
it stand by itself in a class of behaviors, or is it rather a manifestation of
a broader class of behaviors? On the whole, deCatanzaro discusses
suicide as a behavior standing alone, although at times he mentions
the relation of suicide to attempts at suicide and the relation of suicide
to depression. The categorization of suicide has important implications
for the biological perspective that deCatanzaro proposes.
For example, let us assume that suicide is an aggressive behavior,
and therefore appropriately compared with other aggressive behaviors
such as homicide and assault, as Henry and Short (1954) have
proposed. We would then discuss the relevance of aggression for the
survival of the individual organism's genes, and deCatanzaro's Model II
("Suicide as pathology") would be concerned with environments that
make externally directed aggression impossible, thereby facilitating the
appearance of internally directed aggression (Henry and Short 1954).
Or, alternatively, let us categorize suicide as an extreme manifesta-
tion of depression, as deCatanzaro does in his section "Conclusions
and research suggestions." We would then be concerned with the
relevance of depression (and its underlying physiological bases) for
the survival of the organism's genes. Again, suicide may be seen as a
rare pathological outcome of such a process.
Many investigators consider suicide to be a symptom of psychiatric
illness. Estimates of the number of suicides who are psychiatrically ill
vary widely. DeCatanzaro cites a figure of one-third, yet other investi-
gators set this estimate as high as 94 percent (Lester 1972). To
consider suicide as merely a symptom of psychiatric illness would shift
our approach to the models proposed.
Broadening the concept of suicide would also have certain implica-
tions. Viewing completed suicide as similar to attempted suicide and
other minor manifestations of suicidal tendencies would make it
possible for us to place suicide in a context in which it could be seen as
beneficial to the organism. Commonly, suicidal episodes are seen as
crises (Tabachnick 1970), and crises can be seen as periods for
potential psychological growth. Too frequently, psychologists view
periods with negative affect as bad for people, and they try to eliminate
them. The viewpoint of existentialists and some humanistic psycholo-
gists, in contrast, is that such periods must not be avoided, but rather
explored in depth, so that the individual can grow from this experience
(Laing 1967; Lester 1977). In this case, suicide may be seen as a
crisis and growth period, in which helpful environmental supports were
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Commentary/deCatanzaro: Biology of suicide
absent, resulting in a diminution rather than an enhancement of the
individual's life.
Suicide In modern and ancient times. A second comment
worth making is that we may be distorting our analysis of suicide if we
focus solely on suicide in modern times. In deCatanzaro's discussion
of Model IV ("Evolutionary tolerance of Suicide"), he notes that suicide
is much more common in the elderly. In primitive times, the life
expectancy of humans was only twenty years or so. Few people
survived into what we now call adulthood. Thus, suicide was probably
much rarer in ancient times (let's say five or ten thousand years ago)
than it is today.
The advanced technology of modern times, leading to the survival of
people into what we now call "old age" may have led to the
appearance of pathological behaviors because of the "abnormality"
of such long-term survival. It may be that evolutionary processes will
eventually lead to the selection of genes and individuals more suited to
modern environments. Perhaps the existence of behaviors such as
suicide simply indicates that technology advances much more quickly
than the natural selection of genes appropriate to the environment.
Similarly, suicide may have been useful as an altruistic behavior in
the sense of deCatanzaro's Model III ("Suicide as altruism") in ancient
times. In primitive times, food supplies were most likely often sparse,
leading to such behaviors as suicide among the elderly and infirm, the
murder of neonates born soon after the birth of another child, and
other behaviors that served to limit the population.
The advances in technology that have increased food supplies for
most nations have served to make these population-limiting behaviors
obsolete and unnecessary. However, changes in the genes may lag
behind the rapid advances of technology. This speculation about the
living conditions in ancient times has been applied by others to the
existence of an incest taboo in modern times (Slater 1959) and, in a
more analogous situation, to the benefits of having genes that predis-
pose an individual to schizophrenia (Jonas and Jonas 1975; Osmond
and Hoffer, n.d.).
by F.V. Wenz
Department ot Sociology. University of South Carolina. Spananburg. S.C. 29303
Heredity, environment, and culture in suicide
I wish to compliment Dr. deCatanzaro for an excellent review of the
literature on the present state of biogenetically oriented research and
theorizing about various manifestations of self-aggression. Since my
own work has not brought me into direct contact with this research, I
should like to bring out some of the weaknesses of this orientation with
respect to suicide. My remarks are addressed against the strict genetic
determinism of human behavior in general, and suicide in particular.
The writer states that "suicidal behavior acts to decrease rather
than increase the biological fitness of the behaving organism." Well,
physicians, other university graduates, and the gifted (i.e. intellectually
superior) are regarded by many investigators as high suicide risk
groups. On the other hand, prisoners, hospitalized mental patients, the
military, and the never married, who comprise about five percent of the
population 15 years of age and older, are also high suicide risk groups.
Does that mean that these diverse groups are biologically unfit? Is
there a ubiquitous genetic precursor to suicide?
The oft-quoted statement by Linus Pauling and Ralph Gerard that
"behind each twisted thought lies a twisted gene" gives succinct
expression to the convictions of sociobiologists that suicide will
ultimately be understood, at least in part, as reflections of basic
genetic evolutionary processes. DeCatanzaro's hypothesis that suici-
dal individuals are those least capable of promoting their genes may be
correct. But while our ignorance still exceeds our knowledge by a wide
margin, there can be no doubt that continuing biological scrutiny of
self-injurious behavior may yield some increments in theoretical under-
standing.
If we ask, "Why do newborn ladybugs eat their brothers?" or "Why
does the female praying mantis sometimes eat her mate?", we are
usually satisfied with the answer, "Well, that's the way they are." When
we ask ourselves, "Why do human beings kill each other or themselves
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 281
 Commentary / deCatanzaro: Biology of suicide
at so great a rate?" such a dispositional explanation does not satisfy
us. We do not wish to hear that human beings kill themselves because
"that's the way we are." So we search for clues that seem to trigger
the suicide in the genes or the environment.
The possibility that animals commit suicide is one of those intriguing
suggestions that has been proposed regularly by scientific observers.
Research has demonstrated that animals do die under the impact of
environmental and psychological stress. Furthermore, animals may
hasten their own deaths through specific self-injurious acts. It appears
very likely that suicide may occur in animals in the absence of a
knowledge or concept of self.
My general social science background has led to the tentative
conclusion that no unitary explanation of the nature of suicide will
contribute significantly to the advancement of research on suicide. A
biological view of suicide oversimplifies both social Darwinism and the
human condition by reducing suicide, however defined, to the level of
genes. What one must not lose sight of is the important fact that, while
genetic data may provide a basis for the understanding of the human
species as a species, it cannot go very far in explaining attitudinal and
behavioral differences among human populations. Humans are
animals, but they need not necessarily behave like other animals.
Human evolution is unique insofar as it is regulated by an additional
unique variable, namely culture, rather than just the usual two variables,
heredity and environment. Any explanation that attributes suicide to
heredity only, to social factors only, or to environmental factors will be
unproductive. Explanations of the occurrence of suicide in individuals,
and of their distribution in populations should take into account
knowledge of how heredity, culture, and environment interact to
produce the observed distributions. DeCatanzaro shows he agrees
with this notion by stating that "it is thus likely that behavioral
development involves complex interactions between learning and
innate variables." However, the author overstates the case in the
statement that "the innate component should reflect countless genera-
tions of selection for adaptiveness, and we should thus be able to
account for most behavior in terms of evolutionary contingencies that
affect it."
I believe that a biological framework is necessary for an understand-
ing of human behavior, but it is important to investigate the role of
culture more adequately in population and genetic studies of human
behavior. For instance, operational definitions of culture are needed so
that cultural variables may be quantified, and persons and groups can
be classified accordingly. The major problems we now face in research
on suicide result from the fact that we do not have adequate informa-
tion about the necessary and sufficient conditions for its occurrence.
Research that would uncover a necessary condition or single factor
that must be present for a specific type of suicide to occur, regardless
of whatever other factors are involved, would be a major contribution
to any of the four models discussed by deCatanzaro. Suicide occurs in
people who have little in common - except their behavior. In fact, the
cause(s) of suicide are so different that we suspect that there may be
many kinds of behaviors masquerading under the single label of
suicide.
Genes are responsible for our capacity to acquire culture. Genes
also form the basis for other capacities, the wide potential range of
behaviors. Different human populations may have somewhat different
potentials, but it must be borne in mind that while genes may provide
different probabilities for behavior they do not produce the actual
behavior itself. Different environmental conditions can produce the
same behavioral results with different genetic backgrounds. Individuals
with very similar genetic backgrounds can be very different.
Furthermore, humans can overcome certain genetic disabilities
through culture. If there is a genetic predisposition toward suicide, this
predisposition can be modified by culture. If more and more humans
come to wear glasses as the result of "bad eye genes," the success
of the human species will not be threatened. Eyeglasses are a
legitimate substitute for better eyes.
Whatever may be the final verdict on the heredity-culture-environ-
ment-suicide controversy, there remains no doubt that genetic studies
are as important as psychological and sociological studies of suicide.
Any inquiry into the nature and significance of genetic variables in any
282 THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3
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disease or human behavior is always legitimate. For example, our
present knowledge of the association between blood groups and
disease indicates that few, if any, genes are entirely neutral from the
standpoint of evolution. They may have positive or negative selective
value for survival. In the individual case of suicide we may be able to
afford to ignore genetics, but to do so in terms of populations might be
a mistake.
by Richard D. Wetzel
Department of Psychiatry, Washington University School of Medicine and the
Jewish Hospital of St. Louis, St. Louis, Mo. 83tW
Suicide: the need for a cognitive perspective
Research shows that suicide is usually a premeditated act (Stengel et
al. 1973; Sainsbury 1955). Since it appears to be the result of a
decision, some attention has been directed to how suicidal people
think. Neuringer and his associates (Neuringer 1968, 1973, 1974,
1976; Neuringer and Lettieri 1971; Levenson 1974) have suggested
that suicidal individuals have a lifelong tendency to think dichotomously
(in either/or terms) about all concepts and that this tendency leads to
their suicidal decision. Wetzel (1975a, 1975b, 1976b) was able to
replicate Neuringer's findings but also showed that the tendency to
think dichotomously applied only to concepts relevant to the crisis (e.g.
life, self). He further showed that evaluations of life and self changed
over time. These changes were highly correlated with changes in
suicidal intent, depression, and hopelessness.
Beck and his associates (Minkoff et al. 1974; Beck et al. 1975) have
suggested that suicide is a function of a sense of hopelessness. They
have shown that suicidal intent is more closely correlated with hope-
lessness (as they have operationalized it) than with depression. Their
results have been replicated using different measures of depression
and different populations (Wetzel 1976a, 1980). Beck (1972) has
described the cognitive triad in depression as a negative view of
oneself, a negative view of one's current life situation, and a negative
view of the future (hopelessness). While it is clear that cognitive
aspects of depression correlate more with suicidal intent than do other
components of the depressive syndrome, no research to date shows
that any one member of the cognitive triad correlates more with
suicidal intent than the others. The available data show that all three
are highly correlated with suicidal intent and load on the same factor
(Wetzel 1976b).
Beck's theory suggests that depression (and hopelessness as an
aspect of it) may be a function of values (schemata) learned early in life
and maintained through cognitive distortions of experience (Beck
1976; Beck et al. 1979). No data are available on this point. No family
studies on cognitive distortions have been done. There is an obvious
need for family studies looking at the relation among "irrational"
beliefs, cognitive distortions, and depression/hopelessness/suicide.
Since it is clear that depression runs in families, one would expect
these cognitive factors (if causally related) to be familial as well. The
appropriate research design and data analysis might allow one to
distinguish multifactorial genetic transmission from nongenetic familial
transmission. Currently available data exclude single-gene models for
the transmission of unipolar affective disorder.
Cohort analysis of Canadian and American suicide rates shows that
there are marked cohort and secular trend effects (Hellon and
Solomon, in press; Murphy and Wetzel, in press; Wetzel and Murphy, in
preparation). The data indicate clearly that cohorts show consistently
high or low rates predictable from the cohort's rates at ages 15-19.
While hardly confirming Beck's theories about the time of origin of
cognitive structures and processes leading to depression/hopeless-
ness/suicide, these sociological findings are obviously compatible. It
would be intriguing to have the data to assess the correlation between
cohort and other sociological predictions of suicide rates and the
cognitive variables implicated in suicide.
An attempt, like that by deCatanzaro, to choose between models,
while fascinating, is clearly premature. The social, familial, and genetic
bases for the cognitive content and processes of suicidal patients

need to be worked out in detail before conclusions about suicide's
evolutionary significance are attempted.
by David Sloan Wilson
Division of Environmental Studies, University ot California, Davis, Calif. 95616
Suicide, beanbag genetics, and pleiotropy
The idea that a given trait or behavior is coded by a single gene was
labeled "beanbag genetics" by Mayr (1963). Although long ago
abandoned by geneticists, the beanbag approach is still used by many
ecologists, ethologists, and sociobiologists, mostly for simplicity and
for lack of a general alternative model. Suicide and other self-sacrificial
behaviors pose a problem for the beanbag approach because they are
so obviously selected against, unless they benefit others who share the
same gene. However, many of these problems disappear when more
sophisticated genetic models are employed.
Fundamentally, we wish to explain how evolution can produce
maladaptive individuals, or maladaptive traits in otherwise well adapted
individuals. The three most obvious candidates are novel environ-
ments, genetic load, and pleiotropy. That individuals may behave
against their own interest in environments to which they are not
adapted is straightforward and needs no more discussion. That sexual
reproduction can produce bad genetic combinations also needs little
elaboration. The concept of pleiotropy recognizes that a gene can
code for not one, but several behaviors. It could conceivably explain
suicidal tendencies in "normal" individuals when they are placed in
certain unusual circumstances, and is therefore more interesting.
Simply put, many adaptations involve both costs and benefits, and
natural selection is expected to maximize the net gain. Maladaptive
behavior may be selected if it is a necessary part of a larger behavioral
complex whose overall effect on fitness is positive. As one pertinent
example, it is possible that an unavoidable physiological trade-off
exists between vigor and longevity. Under some conditions a short-
lived, vigorous animal will leave more offspring than a long-lived,
sluggish animal, in which case early senescence is the price paid for an
exuberant youth (e.g. Stearns 1976).
A parallel argument for suicide would be as follows. Consider a
species whose individuals find themselves placed permanently in one
of two "circumstances" with probabilities p, and p2 (= 1 — pt). The
maximum fitness that can be obtained within each circumstance is F,
and F2. The individuals possess a single behavior pattern that allows
them to realize a fraction q, of the fitness in circumstance 1, and q2 of
the fitness in circumstance 2. The expected fitness of the individual is
therefore
W - q,p,F, + q2p2F2.
As an example, let q, - 1 and q2 = 0. In other words, individuals are
maximally fit in circumstance 1 and suicidal in circumstance 2. Several
interesting conclusions emerge from this simple model. First, suicidal
behavior will have only minor effects on fitness if either p2 or F2 is very
small; that is, if either the probability of encountering, or the fitness
associated with the second circumstance is minute. Of course as a
"beanbag" trait, q2 should increase no matter how small its effect on
fitness, but as a pleiotropic trait q2 will not increase if q, decreases
along with it. (In ecological jargon, this model is identical to the
question of whether a species should specialize in a coarse-grained
environment; Levins 1968).
Second, redefine p, and p2 to be the proportion of time spent by all
individuals in the two circumstances. Now q2 - 0 is highly disadvanta-
geous, no matter how small the value of p2 and F2, because suicide
prevents a return to the more desirable circumstance. To summarize,
this simple model predicts that suicide will be most prevalent in
circumstances that are infrequent and contain few prospects for
reproductive success and little opportunity for change. This of course
matches the observations, indicating that suicide does not pose a
problem for evolutionary models of human behavior, although nonevo-
lutionary models could also be constructed to explain the same
patterns.
With respect to kin and group selection, deCatanzaro's statement
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Response/deCatanzaro: Biology of suicide
about overall proportion of shared genes is a common technical error.
The overall frequency of the gene is irrelevant. It is merely necessary
that the gene frequency among the recipients of the altruistic behavior
be greater than among the population at large (Dawkins 1980; Wilson
1980). However, kin and group selection may still be important under a
variety of circumstances. If the prospects for personal fitness are
extremely low, even a small benefit to recipients may be justified. A
society can make suicide adaptive if it threatens the disgrace of the
whole family as the only alternative, or glorifies the family as a result
(Ghiselin 1974). Suicide may be justified if it carries large-scale
implications for societal change.
Many of the above points are also made in deCatanzaro's interest-
ing target article. This note is intended to emphasize that most of his
arguments can be easily incorporated into a purely biological model,
when a realistic genetic structure is considered. Suicide may be
advantageous in itself or a necessary cost of behavior that is advanta-
geous when averaged over all circumstances. In both cases, the
improbability of change emerges as a critical variable, either by
predisposing the individual toward the benefit of others, or by causing
suicidal situations to have very little "weight" in the calculation of
average fitness. I suggest that suicide can best be understood, not as
an isolated behavior, but as one manifestation of humans' complex
adaptive machinery.
Author's Response
by Denys deCatanzaro
Department of Psychology, McMaster University, Hamilton, Ontario, LBS 4K1,
Canada
Human suicide: toward a diathesis-stress hypothesis
At the outset, I would like to restate my original intentions in
outlining alternative biological interpretations of human
suicide. My analysis was not intended to provide definitive
answers with respect to the biological anomaly of suicide -
that would surely be premature. Rather, my purpose was to
stimulate examination of this phenomenon at a level of
analysis radically different from that of the existing literature.
I am pleased that my target article has succeeded in eliciting
some insightful elaborations and criticisms, but I must also
draw attention to some unfortunate misinterpretations.
The level of analysis that I undertook has been notably
absent from the sociological and psychological literature on
suicide. There is a need for a framework that integrates the
otherwise apparently disparate findings in these areas; such a
framework would surely require a global perspective rooted
in, or at the very least concordant with, biological science.
The sociological literature indicates remarkable consistencies
in the social ecology of suicides, yet there has been no
complete attempt to account for the genesis of this social
ecology. In psychology, behavior has long been viewed in
terms of its orientation toward reinforcers and away from
punishers, which implicitly, but not explicitly, addresses the
relationship of behavior to biological fitness. Both disciplines
appear to be in need of infusions of some of the fertile notions
that are being developed in the areas of sociobiology and
behavioral ecology.
For the sociobiologist, human suicide may present a special
challenge. No other behavior appears to be so contrary to the
principles of natural selection, at least when analyzed at a
superficial level. Such problematic phenomena may provide
tests of the limits of sociobiological reasoning. Indeed, it is
undoubtedly wrong to attribute adaptiveness to all the fine
details of behavior; there is no exact correspondence between
selective pressures and behavior. Our task is to find out
through research and analysis how precisely human behavior
is determined by natural selection.
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 283
 Response/deCatanzaro: Biology of suicide
My outline here follows that of the target article, and I
address each commentator's suggestions as they arise in this
sequence. At times this sequence may break down as I discuss
relationships among the models. At the end, I present a
tentative synthesis that integrates the diverse notions
presented by myself and the commentators.
Evolution and behavior. My suggestion that innate
influences on behavior should reflect generations of selection
for adaptiveness and that we should thus be able to account
for most behavior in terms of evolutionary contingencies is
questioned by Wenz and Duke. I was, however, allowing for
the influence of learning here, as should be clear from the
context of this statement. Also, I was simplifying to set up the
problem that suicide presents biologically. The subsequent
discussions, especially those in the pathology model, define
limits on this statement. But, where the numerous patho-
genic events do not occur, this statement should hold in full. It
is supported by a growing literature on the behavior of
organisms living in the conditions in which - they have
evolved. I would thus point out to Duke that there is empiri-
cal support for this contention in the large literature on
behavioral ecology.
Harmatz has questioned my treatment of suicide as behav-
ior. There is no doubt that what I refer to is a class of
behavioral patterns, with the constituents being linked by a
common consequence and some sort of common antecedent
motivational condition (the expressed "desire to die"). In this
sense any deliberate self-injurious act appears to be just the
same as any other goal-directed behavior. Also, I did not
contend that the behavioral topography itself or the mode of
suicide was anything other than learned behavior; what is
interesting is the origin of the antecedent motivational condi-
tions and why such learning can occur.
Although his title is quite clever, Harmatz has missed the
point in his assertion that I have ignored a human ability to
experience gains from symbolic acts. Such symbolic gains
cannot be directly translated into gains in biological fitness.
My concern was with biological fitness as defined by success
in gene propagation. There is no doubt in my mind that
humans are influenced by symbols; such symbols, where they
bear on suicide, are subsumed by factors in models I and II.
But they are not directly representative of biological fitness,
although they may relate in many interesting ways to biologi-
cally meaningful gains and losses.
Frieden also briefly fell into this trap in asserting that relief
from pain in dying, as negative reinforcement, constitutes a
gain for the individual. What matters in biological terms is
not the individual's conception of gains or losses (although this
is of great interest on other grounds) but rather the true
impact of his actions upon gene frequency. A cognition that
relief from pain justifies suicide may relate to any of the
models (perhaps especially model I), depending on the nature
of the pain.
The incidence of suicide. My assertion that suicide is the
ninth most common cause of death was quite correctly
questioned by Dawkins, who suggested that this depends on
how finely such causes are classified. This does not diminish
the problem the frequency of suicide presents, however.
Douglas raises the important point that statistical biases are
present in official statistics on suicide. This does render
tentative a number of the arguments based on statistical
tendencies in suicide, although he notes that removal of such
biases in many cases supports the conclusions I have drawn.
Hankoff & Turner have contended that suicide is hetero-
geneous in nature. I have no quarrel with this notion and
would suggest that this heterogeneity is largely responsible for
the fact that the biological causes of suicide are not obvious.
But the idea that a simple nosology would neatly isolate
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diagnostic categories is tenuous. Such nosological systems
have almost always failed to bring order to the diversity of
syndromes of abnormal behavior. In considering schizophre-
nia, depression, aphasia, child psychosis, suicide, or almost
any of the major forms of behavior disorder, we must not
forget that each individual is different. Different etiologies
may yield convergent symptomatologies; more often, disor-
ders result from a synergism of diverse factors. I doubt that
Durkheim's subclassifications of suicide, being founded on a
small and old data base, retain much utility or construct
validity. The diversity of suicidal behavior is probably best
accounted for by differential concentrations of many dispa-
rate determinants acting convergently or synergistically in
different individuals. We must accordingly consider a broad
range of etiological factors and their interactions, a unique
combination for each individual.
Model I. Some novel expansions of the notion that complex
and unusual cognitive events may underlie self-damaging
behavior are presented by Frieden. There can be no doubt
that suicide derives in part from human cognitive and behav-
ioral flexibility. The involvement of cognitive variables is not
inconsistent with any of the models, and it is not my intention
to play down the roles of cognition and learning as proximate
determinants of suicide. I deliberately did not provide an
extensive discussion of these issues because so many other
such discussions are already available, limiting myself instead
to a brief overview and an interpretation of the biological
meaning of such factors.
Wetzel has argued for a cognitive perspective on suicide.
Again, cognition is clearly involved at a level of analysis that
differs from my own; but even when we have sorted out the
role of cognition in suicide we are left with a fundamental
problem from a biological standpoint; this is the anomaly
outlined at the beginning of the target article. Developments
in the study of cognitive aspects of suicide will surely clarify
some of the biological issues. Otherwise, however, Wetzel s
comments do not really address the issues I have raised.
Hamilton's discussion of suicide by nonhuman animals is a
very helpful examination of a seldom treated subject. His
conclusions lend support to the notion that attributes specific
to contemporary human culture are responsible for suicides.
This topic has implications not only for model I (where it is
placed only for convenience) but for each of the other models
as well. As will be elaborated below in reference to other
commentaries, contemporary human culture supports the
existence of genotypes in ecologically novel situations. This
results in a gene expression different from the one that was
responsible for the gene's diffusion. This converges with
Hamilton's suggestion that self-destructiveness is most
common in nonhuman animals when they are in captivity, as
well as with the notions advanced by Dawkins, Lester, and
Wilson (and possibly Anisman and Blanchard). This brings
us to a discussion of the pathology model (II).
Model II. The involvement of stress-induced central amine
dysfunctioning in suicide is suggested by Anisman. These
elaborations are informative and readily accommodated or
subsumed by the pathology model. It is very reasonable that
stress-induced abnormalities in neuroregulators and neuro-
modulators facilitate or induce maladaptive behavior.
However, in considering this we must also consider interac-
tions with cognitive variables, and how these variables in turn
are influenced by selective pressures and genetics. The "de-
pression " construct covers a rather amorphous and complex
set of phenomena, only some of which may relate to suicide.
As Beck, Kovacs, and Weissman (1975) pointed out, "depres-
sion " is not so highly correlated with suicide as is "hopeless-
ness."
Blanchard also offers some helpful elaborations of model

II, along much the same line as Anisman. The idea that a
significant number of suicides derive from endogenous
depression not primarily related to environmental stress is an
important consideration. It suggests a convergence of stress-
induced neurochemical pathology and familial neurochemi-
cal pathology, which may adequately characterize some of
the variance in human depression. The question it raises is
how pathological conditions conducive to suicide might arise
in the absence of stress. As Blanchard mentioned, genetic
variance seems to be involved, although stresses from con-
genital factors or environmental toxicology remain possibili-
ties in some cases. The genetic component would be accom-
modated by my statements about genetic recombination.
Lester also offers some strong arguments in favor of model
II. The suggestion that an increase in human longevity may
contribute to suicide is quite useful. Indeed, this strengthens
the notion that gene expression in historically novel environ-
ments is responsible. The well-documented increase in suicide
rates as a function of age is concordant with this idea. As age
increases, the probability that a gene is expressed in novel
circumstances increases; consequently, the probability that it
will predispose toward a maladaptive phenotype increases.
This converges nicely with some of the ideas of Dawkins,
Hamilton, and Wilson, which I treat below.
Wenz's concern about suicide in the gifted can be handled
by the factors in model II. Some constitutional factors condu-
cive to suicide may be concentrated differentially in different
individuals because of gene recombination; this could occur
irrespective of the viability of other genes the individuals
carry. The same stress or aberrant environment might thus
affect individuals differentially. It could also be argued that
in our culture gifted individuals are often streamed into
demanding and competitive environments.
I would like to add an illustration of how gene expression in
novel environments may facilitate suicide. As I mentioned in
the target article, technology has produced many new meth-
ods for committing suicide. These methods may increase the
likelihood of suicide because they are readily available for
impulsive use. Also, many methods may overcome natural
mechanisms that might deter suicide; for example, barbitu-
rate overdose may overcome pain mechanisms that might
help inhibit other methods. The availability of such methods
is historically novel and thus not something for which we
would be biologically adapted.
Models I and II considered jointly. That models I and II
need not be distinguished from one another is argued by
Dawkins. Human flexibility and some of our current novel
environmental circumstances are of course inextricably
linked, and the differentiation was in part for convenience of
treatment. However, model I involves the notion that human
flexibility, while highly adaptive, has as a by-product occa-
sional maladaptive, learned behavior. The major notion
underlying model II is that individuals now frequently
encounter conditions for which they are not biologically
prepared. Quite probably, maladaptive by-products of flexi-
bility are largely responsible for unbiological environments,
but neither is necessary or sufficient for the other, and either
could occur independently. Model II also includes some
features that could have been treated separately, for example,
that genetic load or gene recombination might be involved in
suicide.
Wilson presents some interesting elaborations of points
made in several of the models; these may be especially
relevant here. I might first state clearly that I did not suggest
that a single gene could underlie suicide. It is more likely that
suicide would be influenced by multiple genes, albeit in
indirect and subtle manners. Such genes could be pleiotropic,
each affecting a range of phenotypes. Pleiotropy could mean
that one highly advantageous trait might outweigh another
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Response / deCatanzaro: Biology of suicide
one that was very occasionally maladaptive, allowing a gene
supporting both to flourish. However, such a gene might
ultimately fail in competition with one allowing the advan-
tage without the cost. For pleiotropy to be involved, as Wilson
states, "maladaptive behavior [must be some] necessary part
of a larger behavioral complex whose overall effect is posi-
tive." Thus, we would have to find linked to suicide some
larger trait whose overall effect is positive. This subsumes my
point in model I, that suicide may derive from a behavioral
flexibility that is generally adaptive but that occasionally
engenders maladaptiveness such as suicide. In another sense,
pleiotropy belongs with genetic load and gene recombination
in model II, as means whereby genes predisposing toward
maladaptive traits can continue to be present in the gene
pool.
An example may illustrate some relationships between
pathological conditions and associative processes. Modeling
(Bandura 1971) is a process whereby individuals learn vicar-
iously by viewing the behavior of others and its consequences.
In normal conditions, such a process is highly adaptive
because it allows the learning of complex behavioral
sequences without an extended trial and error process. First,
modeling may sometimes facilitate suicide, thus permitting
an occasional negative by-product because the overall process
is so adaptive. Thus, due to the rareness of such a by-product,
selective pressures may not have induced a sufficiently fine
degree of adaptation to eliminate it. However, processes such
as modeling may currently produce more of a negative
by-product than they did during our evolutionary history.
Since modern culture may expose more individuals to ecolog-
ically novel environments, there may be more opportunity for
maladaptive behavior such as suicide to arise from such
processes.
Model III. According to Hamilton kin selection is respon-
sible for most self-destructive behavior in nonhuman animals
living in their natural ecological circumstances (i.e. where
their genes evolved). This might suggest a similar involve-
ment of altruism in suicides of humans living in more natural
environments. This accords with Lester's attribution to
altruism of suicide in ancient and less developed cultures. As I
mentioned in the target article, in modern societies altruism
appears (on the surface) to be involved in only a minority of
cases.
This potential influence of factors favoring altruism can be
extended by considering potential interactions with factors
developed in model II. Some capacity toward altruism
presumably exists in the human population because of factors
such as kin selection, group selection, and reciprocal altruism.
Given that we now so frequently find ourselves in situations
for which we are not biologically prepared, tendencies
toward altruism might occasionally be expressed where they
do not truly serve inclusive fitness. Hence, traits attributable
to factors of model III may be expressed in abnormal condi-
tions because of factors in model II. (Could this relate to the
fact that individuals committing suicide have so frequently
been isolated from kin?) This also seems consistent with the
last two paragraphs of Lester's commentary.
Both Dawkins and Wilson criticized my point about
coefficients of relationship. I realize that kin selection should
work approximately as Hamilton (1964) explained when it is
applied to the diffusion of a novel gene for altruism. I was also
not trying to repeat Washburn's (1978) point, nor did I argue
that altruism should depend on the totality of gene sharing.
Undoubtedly, an individual shares more genes with close kin
than with individuals from the general population. But the
Hamilton-Wright coefficients provide a ratio scale when only
an ordinal scale is justified. We cannot define a zero point. I
doubt, consequently, that we can insert the Hamilton-Wright
coefficients, given their ordinal scale properties, into some
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 285
 Response/deCatanzaro: Biology of suicide
simple calculus to compute the tendency toward altruism.
Considering genes shared by kind as well as by common
descent (a distinction that becomes hazy in considering the
totality of generations), there should be a progression away
from gene sharing between individuals who are far removed
from one another spatially, temporally, ecologically, racially,
phylogenetically, and so on. There are discontinuities in this
progression, one of which may lie between kin and other
subpopulation members, with the extent depending on the
composition and size of the subpopulation; other discontinui-
ties may lie between subpopulations, still others between
species, and so on. Another consideration is that if an individ-
ual or even a family is removed from a subpopulation, all of
its genes, with the exception of any specific mutations, will
probably remain in the total subpopulation, albeit in smaller
proportion. The necessary and sufficient conditions for
altruism toward another individual or toward the totality of
some group would depend upon numerous parameters; this
issue must not be closed prematurely.
Let us ignore for a moment the cost of altruism to the
behaving individual. I agree with Wilson's comment that for
kin and group selection to operate, it is merely necessary that
gene commonality with recipients of altruism be greater than
with some population at large. But even allowing for the
discontinuities mentioned above, I am not convinced by
Dawkins (1976) that kin and group selection are not just two
different points along a continuum. The potential complexi-
ties of gene sharing within different levels of aggregation of
human individuals, especially within smaller aggregations
that characterized earlier periods of our evolutionary history,
place estimates of "r " based on one or two generations in
some perspective. Such estimates assume random panmictic
breeding in a population of infinite size; such assumptions do
not entirely hold for realistic human populations.
Dawkins appears to have misinterpreted my point here;
the two views are not as far apart as he assumes. I am simply
less inclined to dismiss group selection. There is good reason
to assert that individual selection is generally more powerful
than kin selection, which in turn is more powerful than
selection among larger groups. The strongest arguments
against the importance of group selection rest upon mathe-
matical models pitting group selection against individual
selection in the determination of some trait. But again the
assumptions may be too limited. We must consider what will
occur if individual selection is rendered impotent by poor
prospects for personal fitness. This brings me to the next
model (perhaps "factor" would have been a better term),
which elaborates my point.
Model IV. The last model presents the notion that as
productive and reproductive capacity declines, there is
progressively less evolutionary reason why the individual
should behave in a self-preserving manner. This is really just
an extension of model II, because it describes an additional
condition under which an individual's genes need not be
expressed in adaptive or self-serving ways. This condition
would affect many of the same individuals as those affected
by the factors of model II, providing further reasons why such
individuals might not be expected to behave in a self-
preserving manner.
This unification of models II and IV is also supported by a
number of the commentaries as I discuss below; but first I
would like to consider the implications of this dimension for
model III, to demonstrate further how all four models could
be unified.
The importance of kin and group selection depends on the
extent to which these have to compete with individual selec-
tion. Self-serving behavior would generally propagate the
individual's genes more readily than would kin-serving
behavior, which might in turn propagate his genes more than
286 THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3
https://doi.org/10.1017/S0140525X00004799 Published online by Cambridge University Press
subpopulation-serving behavior, and so on. But if the pros-
pects for personal fitness became very low, would not even a
small benefit to kin outweigh a personal loss? Similarly, if the
prospects for both personal fitness and behavior that helped
kin became low, might not group selective pressures begin to
have some influence over the fate of the individual's genes?
This is a major dimension that is missing in discussions
belittling group selection.
As Wade (1978) has cogently argued, group selection may
be quite powerful when it operates in the same direction
as - or is simply not opposed to - pressures of individual
selection. I note that Wilson's comments also support this
point when he says that "kin and group selection may still be
important under a variety of circumstances. If the prospects
for . . . fitness are extremely low, even a small benefit to
recipients may be justified." These notions again raise the
question of how precisely selective pressures operate in deter-
mining human behavior. A fruitful empirical enquiry might
focus on whether altruism is increased in times of personal
desperation.
Another important consideration is the role of human
culture in this process. Societies may frequently compel
altruism from some members by placing various behavioral
constraints on the probabilities for successful gene propaga-
tion. This would be true in wartime, for example, if the only
alternative to military service were severe personal disgrace.
It may also characterize some of the forms of institutional
suicide discussed by Farberow (1972).
Dawkins's suggestion that model IV be integrated with the
Medawar-Williams hypothesis (Medawar 1957; Williams
1957) is most helpful. I agree wholeheartedly that this logic
enriches model IV; I also think it can be viewed as an
extension of model II. Suppose that through some course of
events, a gene-environment combination arose that allowed
the occurrence of suicide (indeed, this much has surely
happened). If a gene in this combination expressed itself
when an individual retained a capacity to promote the
continued existence of his genotype, the gene would be
deleterious and would remain infrequent in the population. It
would be rapidly eliminated if expressed when the individual
was otherwise likely to reproduce and foster children.
However, if it were expressed only after reproductive age or
in younger individuals who were under stress and unlikely to
reproduce, and only when circumstances also prevented
activity that indirectly promoted the individual's genotype, it
would remain in the population. It is thus conceivable that
genes permitting suicide, expressed only in conditions where
the prospects for personal fitness are poor, could have accu-
mulated in the population. Like the factors in model II, this
describes conditions under which the individual's genes need
not predispose toward adaptiveness.
A caveat is necessary here. There is no claim here that
genes directly predispose toward suicide. Such a contention
would be implausible given the importance of conditioning in
human development and the consequent complexity of rela-
tionships between genotypes and behavioral phenotypes in
humans. Rather, it suggests that multiple genes could subtly
contribute to the motivational aspects of suicide. In all likeli-
hood, such genes would not be specific to suicidal behavior. A
weak genetic or innate basis for suicidal motivation is
conceivable because of the nature of the social-ecological
situation in which suicide appears to occur.
Harmatz argues that evidence that environmental factors
are associated with suicide is contrary to a genetic involve-
ment in suicide. Again, he has missed the point. Gene
expression varies with the environmental context, as I and a
number of the commentators have discussed. Thus, there is by
no means a conflict between the weak sort of genetic involve-
ment I speak of and the evidence he cites.
Anisman's argument that the high-risk populations of

model IV are stressed individuals points out the linkages with
model II. The conditions necessary for the factors in model II
appear to converge at many points with those necessary for
the factors in model IV. Of course, the "stress" construct
requires a concrete definition when used in this context.
Model IV really describes further conditions under which
gene expression need not be adaptive; thus, depending on our
definition of "stress" and "pathology," we may ascribe such
attributes to individuals without the capacity to propagate
their genes. Also, a loss in the capacity to propagate one's
genes is probably associated with a substantial demand on the
organism's coping apparatus.
Duke's assertions about the logic of this section address a
similar concern that the data reviewed also support a pathol-
ogy model. He suggests that we can interpret the correlations
of stress, social isolation, and poor productivity and reproduc-
tive capacity with suicide in terms of a number of different
causal relationships. What he does not realize is that, for
factors discussed in model IV to hold, no particular proxi-
mate causal relationship among these variables is necessary.
All that would be required is that suicide consistently be
associated with behavioral dysfunctions preventing gene
propagation. That association, no matter what the complexi-
ties of proximate causation, would mean that suicide would
have a minimal impact upon the gene pool. I really made no
strong assertions about the order of causation here, nor would
I need to. For example, I did not suggest that divorce causes
suicide, and his quote "social isolation can precipitate
suicide" is taken out of context. He has missed the point here
and has overlooked a number of qualifiers.
Duke's suggestion that sex differences in suicide are incon-
sistent with model IV overlooks some relevant aspects of
male-female differences in gene-propagation strategy. As is
true in many other species, human females would seem to
have been more prone than males to engage in kin-favoring
and nurturant behavior over human evolutionary history.
Consequently, a postmenopausal female (although we might
wonder why menopause can be correlated with severe
depression) may retain a capacity to promote her genes as
long as she continues to act in some kin-favoring manner.
Duke's worries about referencing here do not undermine
my statements. First, I did not imply that there is a decisively
demonstrated relation between reserpine and suicide; I only
said that such a relation has been reported. Nor did I make
any conclusive statements about Kallman's (1949) data.
Again, I feel that Duke has not read carefully enough to see
the qualifiers. I do not see the degree of difference that he
implies between my statements on these two issues, taken in
context, and the sources he cites. Also, I believe that some
license for secondary referencing (i.e. citing thorough reviews
that elaborate, support, or qualify points) must be allowed in
any endeavor operating at this interdisciplinary and global
level.
Farber's discussion seems to lend support to the notions
developed in model IV. He provides a second potential
answer to Wenz's query about suicide in the gifted when he
says that "the infrequent loss of high-quality genes would not
seem to outweigh the probably more frequent elimination" of
low-quality genes. He interprets suicide as an agent of natural
selection rather than as an anomalous behavior that is
contrary to natural selection. This may reduce to questions
about the locus of determination of suicidal behavior and the
locus of action of selective pressures. To the extent that some
residual reproductive capacity is lost when suicide occurs, his
interpretation may be correct. In line with the notion in
model II that modern suicide may be increased by the
exposure of genes to novel environments, suicide may indeed
be one force serving to return us to genetic equilibrium with
our environment.
Regarding Farber's further comments, there is no problem
https://doi.org/10.1017/S0140525X00004799 Published online by Cambridge University Press
Response/deCatanzaro: Biology of suicide
in model IV with the fact that offspring of forebears who
have committed suicide exist, at least, provided that at the
time of suicide the forebears were not capable of promoting
their genes through behavior increasing the reproductive
capacity of their offspring. The model hinges upon whether
or not suicide occurs when the individual has no residual
reproductive and productive capacity.
Testing the hypotheses. According to Carr a tautology is
inherent in my suggestion that we examine the social ecology
of suicide to determine further the relation of the act to gene
propagation. This is not really accurate. It is commonplace in
science to develop a hypothesis on the basis of preliminary
data, then to probe the same phenomena further to determine
the validity of the hypothesis. Implicit in my statements is a
suggestion that new data be collected and analyzed in light of
the hypotheses.
One further important point can be made regarding behav-
ioral-genetic studies. Such studies only address sources of
variability among individuals and do not really test whether
innate factors contribute to suicide. For example, if genes
indirectly permitting suicide in some environments were
widely and fairly uniformly distributed in the population,
there could be no observed difference between monozygotic
and dizygotic twins. Variance among individuals is not
entirely reflective of innate (if opposed only to learned)
influences on behavior, since environmental influence on
gene expression may be as important as associative (learning)
processes. As a further example, we call some fixed action
patterns in nonhuman species innate because they are not
influenced substantially by learning, yet they may vary little
across different members of the species.
A synthesis. My discussion is quite accurately character-
ized by Carr as a diathesis-stress model. The majority of the
commentators' remarks add support to such a model, as do
the integrations of the diverse factors I have discussed in this
response. I think that without prejudging the relative contri-
butions of the various factors, we are now in a position to
present a relatively unitary framework that nonetheless
allows for differential causation in different individuals.
Let us first consider the diathesis. At one level, genes have
to be involved even in suicide; all behavior is a function of
gene-environment interactions. Inevitably, there will be some
who will reduce my arguments to some gross oversimplifica-
tion, seeing absurdity in a notion that genes influence suicide.
Surely, any involvement of innate factors in behavior so
complexly determined as suicide is weak, subtle, and indirect.
No one would contend that there is a gene directly predispos-
ing toward suicide. However, many genes undoubtedly
predispose rather directly against suicide, as must those
controlling physiological pain, reinforcement, and punish-
ment mechanisms. The largest part of the question is why
factors such as these occasionally fail in this respect.
A more fruitful controversy concerns the extent to which
suicide derives from associative processes (learning) as
opposed to nonassociative processes more closely related to
environmentally controlled gene expression. However, this
distinction may also be unclear, since associative processes
interact in complex ways with nonassociative motivational
variables. This brings us to the relations between the factors in
models I and II. The most likely relationship is that human
flexibility and culture have, over our evolutionary history,
given rise to our current situation, wherein human genotypes
are frequently exposed to novel environments (although some
other relationships have been discussed above).
I have outlined several ways in which indirect innate
influences on suicide might occur; these have been expanded
upon by some of the commentators. A good argument can be
made that some tendency toward altruistic self-sacrificing
THE BEHAVIORAL AND BRAIN SCIENCES (1980), 3 287
 References / deCatanzaro: Biology of suicide
behavior may occur in the human species. Constitutional
factors facilitating maladaptive self-destructive behavior may
also be carried in the human population through genetic load
or pleiotropy. Also, when an individual is exposed to chronic,
intractable stress, there is no basis for adaptive genetic expres-
sion. In this sense, stress might be equated with exposure to
historically or ecologically novel circumstances, conditions
other than those in which the organism's genes evolved. There
seems to be a consensus among the more biologically oriented
commentators that the weighting of this factor should be
increased, and I agree. The factors in model IV simply extend
this notion by defining additional, often similar, circum-
stances in which gene expression need not be adaptive. When
an individual's capacity to propagate his genes declines, there
is no ecological reason why his genotype should predispose
him toward self-preserving behavior. This last factor may in
turn facilitate altruism.
There can be no doubt that modern human culture
frequently places stresses on many individuals by placing
them in situations for which they are not genetically
prepared. Biological evolution has not caught up to cultural
evolution. This is suggested by Dawkins's moth-flame analo-
gy. Hamilton's review of self-destructiveness in nonhuman
animals underlines how high rates of suicide may be specific
to modern human culture or to animals in captivity. This
notion is also suggested by the high rates of suicide in
developed as opposed to nondeveloped countries and by the
involvement of modern technologies that facilitate impulsive
suicidal acts. Indeed, modern society has increased longevity,
as pointed out by Lester, which again exposes genes to
conditions differing markedly from those in which they
evolved. Such stress-induced maladaptive behavior may be
mediated in part by the physiological variables emphasized
by Anisman and Blanchard, and by aberrant cognitions and
learning experience as emphasized by some of the other
commentators.
A number of commentaries have alluded to variance
among individuals in terms of the characterological condi-
tions conducive to suicide. This variance may derive from
differential concentrations of predisposing factors and of
aberrant or extreme environments engendering self-destruc-
tive behavior. Undoubtedly, every individual case involves a
different combination of some subset of the diverse factors we
have considered.
There may be some residual occurrences of suicide that
may not be "pathological, ' or that may not fit a stress-
diathesis model, but that may nonetheless derive from some
of the factors discussed above. These might include suicides
that are altruistic in nature and that truly serve the individu-
al's inclusive fitness. They may also include suicides in socie-
ties in equilibrium with their environment, although here
some "pathology " may also occur.
None of this is meant to supersede careful empirical
research, and at best any conclusions at this point must be
tentative. However, it would seem important that we attempt
periodically to integrate at a global level relevant findings
from a broad range of disciplines.
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