The difference of being human: Morality

Francisco J. Ayala1
Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697

In The Descent of Man, and Selection in Relation to Sex, published they may be the most obvious. Other notable anatomical dif-
in 1871, Charles Darwin wrote: “I fully . . . subscribe to the judg- ferences include the reduction of the size of the jaws and teeth
ment of those writers who maintain that of all the differences and the remodeling of the face; reduction of body hair and
between man and the lower animals the moral sense or conscience changes in the skin and skin glands; modification of the vocal
is by far the most important.” I raise the question of whether tract and larynx, with important implications for spoken lan-
morality is biologically or culturally determined. The question of guage; opposing thumbs that allow precise manipulation of
whether the moral sense is biologically determined may refer ei- objects; and cryptic ovulation, which may have been associated
ther to the capacity for ethics (i.e., the proclivity to judge human with the evolution of the nuclear family, consisting of one mother
actions as either right or wrong), or to the moral norms accepted and one father with their children.
by human beings for guiding their actions. I propose that the Humans are notably different from the apes and all other
capacity for ethics is a necessary attribute of human nature, animals in anatomy, but also and no less importantly in their
whereas moral codes are products of cultural evolution. Humans functional capacities and behavior, both as individuals and so-
have a moral sense because their biological makeup determines cially. Most fundamental are the advanced intellectual faculties,
the presence of three necessary conditions for ethical behavior: (i) which allow humans to categorize (see individual objects as
the ability to anticipate the consequences of one’s own actions; (ii) members of general classes), think in the abstract and form
the ability to make value judgments; and (iii) the ability to choose images of realities that are not present (and, thus, anticipate
between alternative courses of action. Ethical behavior came future events and planning future actions), and reason. Other
about in evolution not because it is adaptive in itself but as a nec- distinctive functional features are self-awareness and death
essary consequence of man’s eminent intellectual abilities, which awareness; symbolic (creative) language; tool making and tech-
are an attribute directly promoted by natural selection. That is, nology; complex and extremely variable forms of cooperation
morality evolved as an exaptation, not as an adaptation. Moral and social organization; legal codes and political institutions;
codes, however, are outcomes of cultural evolution, which science, literature, and art; and ethics and religion (3).
accounts for the diversity of cultural norms among populations Humans live in groups that are socially organized, and so do
and for their evolution through time. other primates. But primate societies do not approach the
complexity of human social organization. A distinctive human
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biological evolution cultural evolution | human uniqueness | moral social trait is culture, which may be understood here as the set of
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norms moral sense non–strictly biological human activities and creations. Culture in
this sense includes social and political institutions, ways of doing

H umans are animals and have evolved from ancestors that were
not human. But our “bodily frame,” as well as the capacities
that stem from it, show also that we are a unique kind of animal,
things, religious and ethical traditions, language, common sense
and scientific knowledge, art and literature, technology, and in
general all of the creations of the human mind. Culture “is a pool
a unique kind of ape, with distinctive features, of which the moral of technological and social innovations that people accumulate
sense is one and, if we are to agree with Darwin, the most im- to help them live their lives” (ref. 4, p. 65). The advent of culture
portant one (ref. 1, p. 67). As Steven Pinker has written, “Morality has brought with it cultural evolution, a superorganic mode of
is not just any old topic in psychology but close to our conception evolution superimposed on the organic mode, which has, in the
of the meaning of life. Moral goodness is what gives each of us the last few millennia, become the dominant mode of human evo-
sense that we are worthy human beings” (ref. 2, p. 34). In this lution. Cultural evolution has come about because of cultural
essay, I will examine morality as a consequential attribute among change and inheritance, a distinctively human mode of achieving
those that determine “the difference of being human.” At issue, of adaptation to the environment and transmitting it through the
course, stands the evolutionary origin of morality. generations (3, 5–9).

Human Uniqueness Moral Behavior

Two conspicuous human anatomical traits are erect posture and I will define moral behavior for the present purposes as the
large brain. We are the only vertebrate species with a bipedal gait actions of a person who takes into account in a sympathetic way
and erect posture; birds are bipedal, but their backbone stands the impact the actions have on others. A similar definition is
horizontal rather than vertical (penguins are a trivial exception) advanced, for example, by David Copp in The Oxford Handbook
and the bipedalism of kangaroos lacks erect posture and is of Ethical Theory (ref. 10, p. 4): “[W]e can take a person’s moral
drastically different from our own. Erect posture and bipedal gait beliefs to be the beliefs she has about how to live her life when
entail other morphological changes in the backbone, hipbone,
and feet and others.
Brain size in mammals is generally proportional to body size. This paper results from the Arthur M. Sackler Colloquium of the National Academy of
Sciences, “In the Light of Evolution IV: The Human Condition,” held December 10–12,
Relative to body mass, humans have the largest brain. The 2009, at the Arnold and Mabel Beckman Center of the National Academies of Sciences
chimpanzee brain has an approximate volume of 300 cm3; and Engineering in Irvine, CA. The complete program and audio files of most presentations
a gorilla’s is slightly larger. The human adult brain is more than are available on the NAS Web site at www.nasonline.org/SACKLER_Human_Condition.
three times larger, typically between 1,300 cm3 and 1,400 cm3. In this essay, the author draws extensively from his work, “What the Biological Sciences
The brain is not only larger in humans than in apes but also much
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Can and Cannot Contribute to Ethics,” chap. 18, pp. 316–336, in Ayala FJ and Arp R, eds.
Contemporary Debates in Philosophy of Biology (Wiley-Blackwell, Malden, MA, 2010).
more complex. The cerebral cortex, where the higher cognitive
functions are processed, is in humans proportionally much Author contributions: F.J.A. wrote the paper.

greater than the rest of the brain when compared with apes. The author declares no conflict of interest.
Erect posture and large brain are not the only anatomical This article is a PNAS Direct Submission.
features that distinguish us from nonhuman primates, even if 1
E-mail: [email protected].

www.pnas.org/cgi/doi/10.1073/pnas.0914616107 PNAS | May 11, 2010 | vol. 107 | suppl. 2 | 9015–9022

 she takes into account in a sympathetic way the impact of her life Aristotle and other philosophers of classical Greece and
and decisions on others.” Altruism may be defined in a similar Rome, as well as many other philosophers throughout the cen-
way as, for example, “unselfish regard for or devotion to the turies, held that humans hold moral values by nature. A human is
welfare of others” (11). Altruism, however, is usually taken to not only Homo sapiens, but also Homo moralis. For the last 20
imply some cost to the altruist for the benefit of others, and this centuries, the foundations of morality were an important subject
is the sense in which I will use “altruism” here. Moreover, “al- for Christian theologians, as in the case of Thomas Aquinas, but
truism” is often predicated on the behavior of social insects and also for philosophers, such as, in the 18th and 19th centuries,
other animals, in which no intentionality is involved but rather Hume, Kant, and others familiar to Darwin, including notably
comes about as a result of genetically determined behaviors. This William Paley (The Principles of Moral and Political Philosophy,
is biological altruism, or altruismb, in contrast to moral altruism, 1785; ref. 13) (Fig. 1) and Harriet Martineau (Illustrations of
or altruismm (12). Political Economy, 1832–1834; ref. 14).
I will use the term “ethical behavior” as a synonym of “moral The theory of evolution brought about the need to reconsider
behavior,” and “morality” and “ethics” as synonyms of each the foundations of morality. We do not attribute ethical behavior
other, except when explicitly noted or contextually obvious that to animals (surely, not to all animals and not to the same extent
they are used with a somewhat different meaning. Some authors as to humans, in any case). Therefore, evolution raises distinctive
use “morality” or “virtue ethics” in a broader sense that would questions about the origins and tenets of moral behavior. Is the
include good feelings in regard to others and exclude inappropriate moral sense determined by biological evolution? If so, when did
thoughts or desires, such as entertaining sexual desires for ethical behavior come about in human evolution? Did modern
somebody else’s wife or wishes that something harmful would humans have an ethical sense from the beginning? Did Neandertals
happen to others. So long as these thoughts or desires are not hold moral values? What about Homo erectus and Homo habilis?
transformed into actions, they will not be included in my use of And how did the moral sense evolve? Was it directly promoted by
“morality.” Actions that may be thought to be evil or sinful in natural selection? Or did it come about as a by-product of some
some moral systems, such as masturbation or eating pork, will other attribute (such as rationality, for example) that was the direct
not be included either in my use of “morality,” so long as the target of selection? Alternatively, is the moral sense an outcome of
actions have no consequences for others. cultural evolution rather than of biological evolution?
Theories of Morality Darwin and the Moral Sense
People have moral values; that is, they accept standards ac- Darwin’s most sustained discussion of morality is in chapter III
cording to which their conduct is judged as either right or wrong, of The Descent of Man (ref. 1, pp. 67–102). The keystone sig-
good or evil. The particular norms by which moral actions are nificance of morality in human distinctness is clearly asserted by
judged vary to some extent from individual to individual and Darwin in the first sentence, already quoted, of chapter III: “I
from culture to culture (although some norms, such as not to kill, fully subscribe to the judgment of those writers who maintain
not to steal, and to honor one’s parents, are widespread and that of all the differences between man and the lower animals
perhaps universal), but value judgments concerning human be- the moral sense or conscience is by far the most important” (ref.
havior are passed in all cultures. This universality raises two 1, p. 67). Darwin (Fig. 2) had started gathering the contemporary
related questions: whether the moral sense is part of human literature on human moral behavior much before the publication
nature, one more dimension of our biological make-up; and
whether ethical values may be products of biological evolution
rather than being given by religious and other cultural traditions.
When philosophers consider theories of morality they distin-
guish between metaethics, normative ethics, and practical ethics
(10). Theories of metaethics seek to justify why we ought to do
what we ought to do. They are the primary concern of philoso-
phers, who favor different theories, such as “divine command”
(God’s commanding is what makes a particular kind of action
moral); “moral realism” (there are moral facts; our moral
judgments are made valid or not by the moral facts); “utilitari-
anism” (the moral value of an action is determined by the
expected benefit to the largest number of people); “positivism”
(there are no objective rational foundations for morality, but
rather moral norms are determined by social agreement or, in
the individual, by emotional decisions); “libertarianism” (moral
values are measured by the extent to which they maximize per-
sonal freedom and limit the role of the state to the protection of
individual freedoms); and several others.
Normative ethics refers to the rules or laws that determine
what we ought to do. Practical ethics considers the application
of moral norms to particular situations, which often involve
conflicting values: will abortion be justified to save the life of
the mother?
In practice, humans justify the set of moral norms they follow
on several, not only one, metaethical doctrines. Thomas Aquinas,
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the 13th century Christian theologian whose authority is highly Fig. 1. William Paley (1743–1805). English theologian who taught at the
respected up to the present, says that some moral laws come University of Cambridge, United Kingdom, and author of The Principles of
from divine authority (worship only one God), others from nat- Moral and Political Philosophy (1785). His best known work is Natural The-
ural law (do not kill, do not commit adultery), and still others ology, or Evidences of the Existence and Attributes of the Deity (1802). Im-
from civil authority (respect private property, pay taxes). age source: www.nndb.com/people/526/000096238/.

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 Fig. 3. Cover page of Darwin’s The Descent of Man and Selection in Re-
lation to Sex, first American edition, published by Appleton and Company,
Fig. 2. Charles Robert Darwin (1809–1882). Photograph by Oscar Gustave New York, in 1871, the same year in which his first English edition was
Rejlander, ca. 1871, the year Darwin published The Descent of Man. Image published by John Murray, London.
source: http://commons.wikimedia.org/wiki/File:Charles_Darwin_photograph_
by_Oscar_Rejlander,_circa_1871.jpg.
foundation, endowing humans with an ethical capacity, was for
Darwin compatible with different cultures manifesting different
of The Descent of Man in 1871 (Fig. 3); indeed, we know from his
notebooks that Darwin was reading the contemporary philo- stages of moral evolution and with different sets of moral norms.
sophical literature about moral behavior in 1837, only a few years
after returning from his trip on the HMS Beagle (1826–1831).
Treatises that he read early on include the aforementioned
Moral and Political Philosophy by Paley (13), which he had al-
ready encountered while a student at Cambridge University, and
the multivolume Illustrations of Political Economy by Harriet
Martineau, published more recently, in 1832–1834 (14). These
two authors, like other philosophers of the time, maintained that
morality was a conventional attribute of humankind, rather than
a naturally determined human attribute, on the grounds of an
argument often advanced nowadays by philosophers and
anthropologists: the diversity of moral codes.
The proliferation of ethnographic voyages had brought to light
the great variety of moral customs and rules. This diversity is
something Darwin had noticed when comparing the prevailing
English and European norms of morality with those of South
American Indians and other native populations elsewhere. But
Darwin would eventually develop a more complex and subtle
theory of the moral sense than his contemporaneous authors;
a theory that, implicitly at least, recognized moral behavior as
a biologically determined human universal but with culturally
evolved differences. For Darwin, the ethnographic diversity of
moral customs and rules came about as an adaptive response to
the environmental and historical conditions, unique in every
different place, without necessarily implying that morality was an
acquired, rather than natural, human trait.
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A variable adaptive response could very well derive from some

fundamental attribute, a common substrate, unique for the whole
human race but capable of becoming expressed in diverse direc- Fig. 4. Theodosius Dobzhansky (1900–1975), a principal author of the
tions. Darwin did not attribute the universality of morality to modern theory of evolution. The In the Light of Evolution (ILE) Sackler col-
supernatural origin but rather saw it as a product of evolution by loquium series is named after Dobzhansky’s well-known statement, “Noth-
natural selection. The presence of a universal and common ing in biology makes sense except in the light of evolution.”

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 Darwin’s two most significant points concerning the evolution I propose that the moral evaluation of actions emerges from
of morality are stated early in chapter III of The Descent of Man human rationality or, in Darwin’s terms, from our highly de-
The two points are (i) that moral behavior is a necessary attri- veloped intellectual powers. Our high intelligence allows us to
bute of advanced intelligence as it occurs in humans, and thus anticipate the consequences of our actions with respect to other
that moral behavior is biologically determined; and (ii) that the people and, thus, to judge them as good or evil in terms of their
norms of morality are not biologically determined but rather consequences for others. But I will argue that the norms according
a result of human collective experience, or human culture as we to which we decide which actions are good and which actions are
would now call it. evil are largely culturally determined, although conditioned
After the two initial paragraphs of chapter III of The Descent by biological predispositions, such as parental care to give an
of Man, which assert that the moral sense is the most important obvious example.
difference “between man and the lower animals” (see quotation
above), Darwin states his view that moral behavior is strictly Moral Behavior as Rational Behavior
associated with advanced intelligence: “The following proposi- The moral sense refers first and foremost to our predisposition
tion seems to me in a high degree probable—namely, that any to evaluate some actions as virtuous, or morally good, and others
animal whatever, endowed with well-marked social instincts, as evil, or morally bad. Morality, thus, consists of the urge or
would inevitably acquire a moral sense or conscience, as soon as predisposition to judge human actions as either right or wrong in
its intellectual powers had become as well developed, or nearly terms of their consequences for other human beings. In this
as well developed, as in man” (ref. 1, pp. 68–69). Darwin is sense, humans are moral beings by nature because their bi-
affirming that the moral sense, or conscience, is a necessary ological constitution determines the presence in them of the
consequence of high intellectual powers, such as exist in modern three necessary conditions for ethical behavior. These conditions
humans. Therefore, if our intelligence is an outcome of natural are (i) the ability to anticipate the consequences of one’s own
selection, the moral sense would be as well an outcome of nat- actions; (ii) the ability to make value judgments; and (iii) the
ural selection. Darwin’s statement further implies that the moral ability to choose between alternative courses of action. These
sense is not by itself directly promoted by natural selection, but abilities exist as a consequence of the eminent intellectual ca-
only indirectly as a necessary consequence of high intellectual pacity of human beings.
powers, which are the attributes that natural selection is di- The ability to anticipate the consequences of one’s own actions
rectly promoting. is the most fundamental of the three conditions required for
In the ensuing paragraph of chapter III, before proceeding to ethical behavior. Only if I can anticipate that pulling the trigger
a discussion of how morality might evolve, Darwin makes an will shoot the bullet, which in turn will strike and kill my enemy,
important distinction: “It may be well first to premise that I do can the action of pulling the trigger be evaluated as nefarious.
not wish to maintain that any strictly social animal, if its in- Pulling a trigger is not in itself a moral action; it becomes so by
tellectual faculties were to become as active and as highly de- virtue of its relevant consequences. My action has an ethical
veloped as in man, would acquire exactly the same moral sense dimension only if I do anticipate these consequences.
as ours . . .. [T]hey might have a sense of right and wrong, though The ability to anticipate the consequences of one’s actions is
led by it to follow widely different lines of conduct” (ref. 1, p. 70). closely related to the ability to establish the connection between
According to Darwin, having a moral sense does not by itself means and ends; that is, of seeing a means precisely as a means,
determine what the moral norms would be: which sorts of actions as something that serves a particular end or purpose. This ability
might by sanctioned and which ones would be condemned. to establish the connection between means and their ends
Darwin’s distinction between the moral sense or conscience on requires the ability to anticipate the future and to form mental
the one hand, and the moral norms that guide the moral sense or images of realities not present or not yet in existence.
conscience on the other, is fundamental. It is a distinction I will The ability to establish the connection between means and
now elaborate. Much of the post-Darwin historical controversy, ends happens to be the fundamental intellectual capacity that
particularly between scientists and philosophers, as to whether has made possible the development of human culture and
the moral sense is or is not biologically determined has arisen technology. An evolutionary scenario, seemingly the best hy-
owing to a failure to make that distinction. Scientists often affirm pothesis available, proposes that the remote evolutionary roots
that morality is a human biological attribute because they are of this capacity to connect means with ends may be found in the
thinking of the predisposition to make moral judgments: that is, evolution of bipedalism, which transformed the anterior limbs of
to judge some actions as good and others as evil. Some philos- our ancestors from organs of locomotion into organs of manip-
ophers argue that morality is not biologically determined but ulation. The hands thereby gradually became organs adept for the
rather comes from cultural traditions or from religious beliefs, construction and use of objects for hunting and other activities
because they are thinking about moral codes, the sets of norms that improved survival and reproduction; that is, which increased
that determine which actions are judged to be good and which the reproductive fitness of their carriers. The construction of tools
are evil. They point out that moral codes vary from culture to depends not only on manual dexterity, but on perceiving them
culture and therefore are not biologically predetermined. precisely as tools, as objects that help to perform certain actions;
that is, as means that serve certain ends or purposes: a knife for
Moral Judgment vs. Moral Norms cutting, an arrow for hunting, an animal skin for protecting the
The question of whether ethical behavior is biologically de- body from the cold. According to this evolutionary scenario, nat-
termined may, indeed, refer to either one of the following two ural selection promoted the intellectual capacity of our bipedal
issues. First, is the capacity for ethics—the proclivity to judge ancestors because increased intelligence facilitated the perception
human actions as either right or wrong—determined by the bi- of tools as tools, and therefore their construction and use, with the
ological nature of human beings? Second, are the systems or ensuing improvement of biological survival and reproduction.
codes of ethical norms accepted by human beings biologically The development of the intellectual abilities of our ancestors
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determined? A similar distinction can be made with respect to took place over several million years, gradually increasing the
language. The question of whether the capacity for symbolic ability to connect means with their ends and, hence, the possi-
creative language is determined by our biological nature is dif- bility of making ever-more complex tools serving more diverse
ferent from the question of whether the particular language and remote purposes. According to the hypothesis, the ability to
we speak—English, Spanish, Chinese, etc.—is biologically de- anticipate the future, essential for ethical behavior, is therefore
termined, which in the case of language obviously it is not. closely associated with the development of the ability to con-

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 struct tools, an ability that has produced the advanced technol- older function or coexist together with it. Feathers seem to have
ogies of modern societies and that is largely responsible for the evolved first for conserving temperature, but were later co-opted
success of humans as a biological species. in birds for flying. The beating of the human heart is an exap-
The second condition for the existence of ethical behavior is tation used by doctors to diagnose the state of health, although
the ability to advance value judgments, to perceive certain this is not why it evolved in our ancestors. The issue at hand is
objects or deeds as more desirable than others. Only if I can see whether moral behavior was directly promoted by natural se-
the death of my enemy as preferable to his survival (or vice lection or rather it is simply a consequence of our exalted in-
versa) can the action leading to his demise be thought of as telligence, which was the target of natural selection (because it
moral. If the consequences of alternative actions are neutral with made possible the construction of better tools). Art, literature,
respect to value, an action cannot be characterized as ethical. religion, and many human cultural activities might also be seen
Values are of many sorts: not only ethical but also aesthetic, as exaptations that came about as consequences of the evolution
economic, gastronomic, political, and so on. But in all cases, the of high intelligence.
ability to make value judgments depends on the capacity for The second issue is whether some animals, apes or other
abstraction; that is, on the capacity to perceive actions or objects nonhuman primates, for example, may have a moral sense, how-
as members of general classes. This makes it possible to compare ever incipient, either as directly promoted by natural selection or
objects or actions with one another and to perceive some as more as a consequence of their own intelligence.
desirable than others. The capacity for abstraction requires an The position that I will argue here is that the human moral
advanced intelligence such as it exists in humans and apparently sense is an exaptation, not an adaptation. The moral sense
in them alone. consists of judging certain actions as either right or wrong, not of
I will note at this point that the model that I am advancing choosing and carrying out some actions rather than others. It
here does not necessarily imply the ethical theory known as seems unlikely that making moral judgments would promote the
utilitarianism (or, more generally, consequentialism). According reproductive fitness of those judging an action as good or evil;
to the so-called “act consequentialism” the rightness of an action acting in one way or another might be of consequence in pro-
is determined by the value of its consequences, so that the moting fitness, but passing judgment by itself would seem unlikely
morally best action in a particular situation is the one, the con- to increase or decrease adaptive fitness. Nor does it seem likely
sequences of which would have the most benefit to others. I am that there might be some form of “incipient” ethical behavior
proposing that the morality of an action depends on our ability that would then be further promoted by natural selection. The
(i) to anticipate the consequences of our actions, and (ii) to three necessary conditions for there being ethical behavior are
make value judgments. But I am not asserting that the morality manifestations of advanced intellectual abilities.
of actions is exclusively measured in terms of how beneficial their It, indeed, rather seems that the target of natural selection was
consequences will be to others. the development, which happened mostly through the Pleisto-
The third condition necessary for ethical behavior is the ability cene, of advanced intellectual capacities. This was favored by
to choose between alternative courses of actions. Pulling the natural selection because the construction and use of tools, made
trigger can be a moral action only if you have the option not to possible by advanced intelligence, improved the strategic posi-
pull it. A necessary action beyond conscious control is not tion of our biped ancestors. In the account I am advancing here,
a moral action: the circulation of the blood and the process of once bipedalism evolved and after tool-using and tool-making
food digestion are not moral actions. Whether there is free will is became practiced, those individuals more effective in these
a question much discussed by philosophers, and the arguments functions had a greater probability of biological success. The
are long and involved (e.g., refs. 15–18). Here, I will advance two biological advantage provided by the design and use of tools
considerations that are common-sense evidence of the existence persisted long enough so that intellectual abilities continued to
of free will. One is personal experience, which indicates that the increase, eventually yielding the eminent development of in-
possibility to choose between alternatives is genuine rather than telligence that is characteristic of H. sapiens.
only apparent. The second consideration is that when we con- A related question is whether morality would benefit a social
front a given situation that requires action on our part, we are group within which it is practiced and, indirectly, would also
able mentally to explore alternative courses of action, thereby benefit individuals who are members of the group. This seems
extending the field within which we can exercise our free will. In likely to be the case, if indeed moral judgment would influence
any case, if there were no free will, there would be no ethical individuals to behave in ways that increase cooperation, or
behavior; morality would only be an illusion. A point to be made, benefit the welfare of the social group in some way; for example,
however, is that free will is dependent on the existence of a well- by reducing crime or protecting private property. That is, the
developed intelligence, which makes it possible to explore al- moral sense that had evolved as an exaptation associated with
ternative courses of action and to choose one or another in view high intelligence could eventually become an adaptation, by fa-
of the anticipated consequences (Fig. 4). voring beneficial behaviors.

Adaptation vs. Exaptation Group Selection in Human Populations

I will now consider explicitly two issues that are largely implicit in I have asserted that patterns of actions beneficial to the tribe or
the previous section. The moral sense, as I have proposed, social group might, in humans, be favored by natural selection.
emerges as a necessary implication of our high intellectual This brings up the issue known as “group selection.” Evolu-
powers, which allow us to anticipate the consequences of our tionists generally contend that group selection based on altruistic
actions, to evaluate such consequences, and to choose accord- behavior is not an evolutionarily stable strategy. Altruistic be-
ingly how to act. But is it the case that the moral sense may have havior within an animal population would benefit the population
been promoted by natural selection in itself and not only in- itself, so that a population consisting of altruists would do better
directly as a necessary consequence of our exalted intelligence? than a population consisting of selfish individuals. This would be
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The question in evolutionary terms is whether the moral sense is group selection: the population as a whole benefits from the
an adaptation or, rather, an exaptation. Evolutionary biologists behavior of its individuals. But this state of affairs is not evolu-
define exaptations as features of organisms that evolved because tionarily stable in an animal population. The reason is that
they served some function but are later co-opted to serve an mutations that favor selfish over altruistic behavior will be fa-
additional or different function, which was not originally the vored by natural selection, because the behavior of an altruistic
target of natural selection. The new function may replace the individual implies a cost. The altruistic individual as well as the

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 rest of the population will benefit from the behavior of the al- Yet, when looking at the world of life as it exists today, it would
truist. A selfish individual also benefits from the behavior of the seem that there is a radical breach between human intelligence
altruist, but the selfish individual does not incur the cost implied and that of other animals. The rudimentary cultures that exist in
by the altruistic behavior. Thus, selfish behavior will be favored chimpanzees (19, 20) do not imply advanced intelligence as it is
within the population. Natural selection will thus eliminate ge- required for moral behavior.
netically determined altruistic behaviors. A different explanation of the evolution of the moral sense has
Of course, it is admitted that it might be the case that pop- been advanced by proponents of the theory of “gene–culture
ulations with a preponderance of altruistic alleles would survive coevolution” (5, 21–24). It is assumed that cultural variation
and spread better than populations consisting of selfish alleles. among tribes in patriotism, fidelity, sympathy, and other moral-
This would be group selection. But typically there are many more izing behaviors may have occurred incipiently in early hominid
individual organisms than there are populations; and individuals populations, starting at least with H. habilis. This cultural vari-
are born, procreate, and die at rates much higher than pop- ation may have, in turn, selected for genes that endowed early
ulations. Thus, the rate of multiplication of selfish individuals humans with primitive moral emotions. Primitive moral emo-
over altruists in a given population is likely to be much higher tions would in turn have facilitated the evolution of more ad-
than the rate at which altruistic populations multiply relative to vanced cultural codes of morality. Repeated rounds of gene–
predominantly selfish populations. cultural coevolution would have gradually increased both the
There is, however, an important difference between animals moral sense itself and the systems of moral norms. That is, the
and humans that is relevant in this respect. Namely, the fitness evolution of morality would have been directly promoted by
advantage of selfish over altruistic behavior does not necessarily natural selection in a process whereby the moral sense and the
apply to humans, because humans can understand the benefits of moral norms would have coevolved.
altruistic behavior (it benefits the group but indirectly it benefits The gene–culture coevolution account of the evolution of
them as well) and thus adopt altruism and protect it, by laws or morality is, of course, radically different from the theory I am
otherwise, against selfish behavior that harms the social group. advancing here, in which moral behavior evolved not because it
As Darwin wrote in The Descent of Man: “It must not be for- increased fitness but as a consequence of advanced intelligence,
gotten that, although a high standard of morality gives but which allowed humans to see the benefits that adherence to
a slight or no advantage to each individual man and his children moral norms bring to society and to its members. The extreme
over the other men of the same tribe, yet that an advancement in variation in moral codes among recent human populations and
the standard of morality and an increase in the number of well- the rapid evolution of moral norms over short time spans would
endowed men will certainly give an immense advantage to one seem to favor the explanation I am proposing. Gene–culture co-
tribe over another” (ref. 1, chap. V, p. 159). evolution would rather lead to a more nearly universal system of
The theory of sociobiology advances a ready answer to the morality, which would have come about gradually as our hominid
second question raised above, whether morality occurs in other ancestors gradually evolved toward becoming H. sapiens.
animals, even if only as a rudiment. The theory of kin selection, Empathy, or the predisposition to mentally assimilate the feel-
they argue, explains altruistic behavior, to the extent that it exists ings of other individuals, has recently been extensively discussed in
in other animals as well as in humans. I will propose, however, the context of altruistic or moral behavior. Incipient forms of
that moral behavior does not exist, even incipiently, in non- empathy seem to be present in other animals. In humans, in-
human animals. The reason is that the three conditions required creasing evidence indicates that we automatically simulate the
for ethical behavior depend on an advanced intelligence—which experiences of other humans (ref. 25, chap. 5, pp. 158–199).
includes the capacities for free will, abstract thought, and an- Empathy is a common human phenomenon, surely associated with
ticipation of the future—such as it exists in H. sapiens and not in our advanced intelligence, which allows us to understand the
any other living species. It is the case that certain animals exhibit harms or benefits that impact other humans, as well as their as-
behaviors analogous with those resulting from ethical actions sociated feelings. Empathic humans may consequently choose to
in humans, such as the loyalty of dogs or the appearance of behave according to how their behavior will impact those for
compunction when they are punished. But such behaviors are ei- whom we feel empathy. That is, human empathy occurs because of
ther genetically determined or elicited by training (conditioned our advanced intelligence. Humans may then choose to behave
responses). Genetic determination and not moral evaluation is altruistically, or not, that is morally, or not, in terms of the an-
also what is involved in the altruistic behavior of social insects and ticipated consequences of their actions to others.
other animals. Biological altruism (altruismb) and moral altruism The question remains, when did morality emerge in the human
(altruismm) have disparate causes: kin selection in altruismb, lineage? Did H. habilis or H. erectus have morality? What about
regard for others in altruismm. the Neandertals, Homo neanderthalensis? When in hominid
evolution morality emerged is difficult to determine. It may very
Mind to Morality well be that the advanced degree of rationality required for
The capacity for ethics is an outcome of gradual evolution, but moral behavior may only have been reached at the time when
it is an attribute that only exists when the underlying attributes creative language came about, and perhaps in dependence with
(i.e., the intellectual capacities) reach an advanced degree. The the development of creative language. When creative language
necessary conditions for ethical behavior only come about after may have come about in human evolution is discussed in ref. 3.
the crossing of an evolutionary threshold. The approach is gra-
dual, but the conditions only appear when a degree of in- Moral Codes
telligence is reached such that the formation of abstract concepts I have distinguished between moral behavior—judging some
and the anticipation of the future are possible, even though we actions as good, others as evil—and moral codes—the precepts
may not be able to determine when the threshold was crossed. or norms according to which actions are judged. Moral behavior,
Thresholds occur in other evolutionary developments—for ex- I have proposed, is a biological attribute of H. sapiens, because it
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ample, in the origins of life, multicellularity, and sexual re- is a necessary consequence of our biological makeup, namely our
production—as well as in the evolution of abstract thinking and high intelligence. But moral codes, I argue, are not products of
self-awareness. Thresholds occur in the physical world as well; biological evolution but rather of cultural evolution.
for example, water heats gradually, but at 100 °C boiling begins It must, first, be stated that moral codes, like any other cultural
and the transition from liquid to gas starts suddenly. Surely, systems, cannot survive for long if they prevailingly run in out-
human intellectual capacities came about by gradual evolution. right conflict with our biology. The norms of morality must be by

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 and large consistent with human biological nature, because One interesting behavior, associated with a sense of justice,
ethics can only exist in human individuals and in human societies. or equal pay for equal work, has been described by Sarah
One might therefore also expect, and it is the case, that accepted Brosnan and Frans de Waal (26, 27) in the brown capuchin
norms of morality will often, or at least occasionally, promote monkey, Cebus paella. Monkeys responded negatively to un-
behaviors that increase the biological fitness of those who behave equal rewards in exchanges with a human experimenter.
according to them, such as child care. But the correlation be- Monkeys refused to participate in an exchange when they wit-
tween moral norms and biological fitness is neither necessary nor nessed that a conspecific had obtained a more attractive reward
indeed always the case: some moral precepts common in human for equal effort. Is the capuchin behavior phylogenetically re-
societies have little or nothing to do with biological fitness, and lated to the human virtue of justice? This seems unlikely, be-
some moral precepts are contrary to fitness interest. cause similar behavioral patterns have not been observed in
How do moral codes come about? The short answer is, as other primates, including apes, phylogenetically closer to
already stated, that moral codes are products of cultural evolu- humans. Cannibalism is practiced by chimps, as well as by
tion, a distinctive human mode of evolution that has surpassed human cultures of the past. Do we have a phylogenetically
the biological mode, because it is a more effective form of ad- acquired predisposition to cannibalism as a morally acceptable
aptation: it is faster than biological evolution and it can be di- behavior? This seems unlikely.
rected. Cultural evolution is based on cultural heredity, which is The interpretation of the capuchin monkeys’ behavior as an
Lamarckian, rather than Mendelian, so that acquired charac- incipient sense of justice (26) has been challenged by other
teristics are transmitted. Most important, cultural heredity does investigators. Silberberg and collaborators (28) have shown that
not depend on biological inheritance, from parents to children, the capuchins rejected a reward whenever a more desirable re-
but is transmitted also horizontally and without biological ward was visible to them, not just whenever the more desirable
bounds. A cultural mutation, an invention (think of the laptop reward was offered to other individuals.
computer, the cell phone, or rock music) can be extended to Schiff and de Waal (29) observed also that chimpanzees rejec-
millions and millions of individuals in less than one generation. ted a reward when they observed another chimpanzee obtaining
In chapter V of The Descent of Man, entitled, “On the De- a more attractive reward for equal exchange with the human ex-
velopment of the Intellectual and Moral Faculties during Pri- perimenter, although the tolerance for inequity increased with
meval and Civilized Times,” Darwin writes: “There can be no the social closeness among the chimpanzees. However, this in-
doubt that a tribe including many members who, from pos- terpretation of inequality rejection has also been challenged in the
sessing in a high degree the spirit of patriotism, fidelity, obe- case of the chimpanzees. The chimpanzees’ rejection may be at-
dience, courage, and sympathy, were always ready to give aid to tributed to a breach in their expectations, rather than to a sense of
each other and to sacrifice themselves for the common good, equality (30, 31).
would be victorious over most other tribes; and this would be Moral codes arise in human societies by cultural evolution.
natural selection. At all times throughout the world tribes have Those moral codes tend to be widespread that lead to successful
supplanted other tribes; and as morality is one element in their societies. Since time immemorial, human societies have experi-
success, the standard of morality and the number of well- mented with moral systems. Some have succeeded and spread
endowed men will thus everywhere tend to rise and increase” widely throughout humankind, like the Ten Commandments,
(ref. 1, pp. 159–160). although other moral systems persist in different human socie-
Darwin is making two important assertions. First, that mo- ties. Many moral systems of the past have surely become extinct
rality may contribute to the success of some tribes over others, because they were replaced or because the societies that held
which is natural selection in the form of group selection. Sec- them became extinct. The moral systems that currently exist in
ond, Darwin is asserting a position of moral optimism, namely humankind are those that have been favored by cultural evolu-
that the standards of morality will tend to improve over human tion. They were propagated within particular societies for rea-
history precisely on grounds of group selection, because the sons that might be difficult to fathom but that surely must have
higher the moral standards of a tribe, the more likely the suc- included the perception by individuals that a particular moral
cess of the tribe. This assertion depends on which standards are system was beneficial for them, at least to the extent that it was
thought to be “higher” than others. If the higher standards beneficial for their society by promoting social stability and
are defined by their contribution to the success of the tribe, success (25, 32). Cultures, of course, do not evolve as completely
then the assertion is circular. But Darwin asserts that there are differentiated units. Rather, cultures often incorporate elements
some particular standards that, in his view, would contribute to from other cultures. “Far from being self-preserving monoliths,
tribal success: patriotism, fidelity, obedience, courage, and cultures are porous and constantly in flux. Language . . . is a clear
sympathy. example” (ref. 4, p. 66).
The norms of morality, as they exist in any particular culture,
Moral Norms and Natural Selection are felt to be universal within that culture. Yet, similarly as other
Parental care is a behavior generally favored by natural selection elements of culture, they are continuously evolving, often within
that may be present in virtually all codes of morality, from primi- a single generation. As Steven Pinker has pointed out, western
tive to more advanced societies. There are other human behaviors societies have recently experienced the moralization and amor-
sanctioned by moral norms that have biological correlates favored alization of diverse behaviors. Thus, “smoking has become
by natural selection. One example is monogamy, which occurs in moralized . . . now treated as immoral . . . At the same time many
some animal species but not in many others. It is also sanctioned behaviors have become amoralized, switched from moral failings
in many human cultures, but surely not in all. Polygamy is sanc- to lifestyle choices. They include divorce, illegitimacy, working
tioned in some current human cultures and was more so in the past. mothers, marijuana use and homosexuality” (ref. 2, p. 34). Ac-
Food sharing outside the mother–offspring unit rarely occurs in ceptance by individuals or groups of particular sets of moral
primates, with the exception of chimpanzees—and, apparently, in norms is often reinforced by civil authority (e.g., those who kill or
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capuchin monkeys (26, 27)—although even in chimpanzees food commit adultery will be punished) and by religious beliefs (God
sharing is highly selective and often associated with reciprocity. is watching, and you’ll go to hell if you misbehave). But it is
A more common form of mutual aid among primates is coalition worth noticing that the legal and political systems that govern
formation; alliances are formed in fighting other conspecifics, al- human societies, as well as the belief systems held by religion, are
though these alliances are labile, with partners readily changing themselves outcomes of cultural evolution, as it has eventuated
partners. over human history, particularly over the last few millennia (33).

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