The Adaptive Value of Humor and

Laughter
Glenn E. Weisfeld
Department of Psychology, Wayne State University, Detroit, Michigan

Most explanations for humor neglect important types of humor, such as tickling and
word play; or raise difftcult evolutionary theoretical problems, such as group selection,
dubious fitness benefits, and excessive complexity of design; or ignore the data on humor
and laughter. The present theory was based on the following observations. Tickling
entails a mock attack at vulnerable body spots, and may provide youngsters with prac-
tice in defending themselves. The child’s laughter is pleasant and encourages the tickler
to persist. Similarly, juvenile primates including children encourage roughhousing by
laughter and other emotional expressions. We also laugh at humorous content that
provides striking counter-examples (incongruities), as in word play, or that informs us
about fitness-relevant topics such as sexual, aggressive, and social poise scenarios. The
present theory is that the pleasure of humor motivates us to seek out poignant, fitness-
enhancing input of this sort. Laughter evolved to allow us to continue to receive amuse-
ment. Laughter is a pleasant social signal that prompts the humorist to persist in provid-
ing this edifying stimulation. In response to true wit, laughter conveys appreciation and
gratitude-an intention to reciprocate for having received a stimulating idea. Thus,
humor benefits both humorist and laughter. This theory and others are evaluated in
the light of evolutionary principles and relevant data.

KEY WORDS: Humor; Laughter; Reciprocal altruism; Emotion.

THE ADAPTIVE VALUE OF HUMOR AND LAUGHTER

T
his is a preliminary attempt to apply evolutionary theory and previ-
ous research results to the question of the adaptive value of humor.
Many inquiries into the function of humor have either neglected
evolutionary principles or omitted reference to much of the re-
search that bears upon the question. Most theories fail to account for some
of the major forms of humor, including laughter during tickling and agonistic
play in primates, and word play. Many theories do not account for the content

Received July 13, 1992; revised March 5, 1993.

Address reprint requests to: Glenn E. Weisfeld, Department of Psychology, Wayne State Uni-
versity, Detroit MI 48202.

Ethology and Sociobiology 14: 141-169 (1993)

0 Elsevier Science Publishing Co., Inc., 1993 0162-3095/93/$6.00
6.55 Avenue of the Americas, New York, NY 10010
142 G. E. Weisfeld

of humor, especially its incongruity element,
meaning of humor and laughter.
This article will begin with a definition of
for humor having an evolved basis and hence
a functional explanation for humor. Next this
the major types of humor. Finally, alternative
critically reviewed.
or for the social context and
humor, proceed to arguments
a function, and then present
explanation will be applied to
explanations of humor will be

DEFINING HUMOR

Humor, or humorous stimuli, are notoriously difficult to define or classify

(McGhee 1979). This is especially true when humor is delined in terms of
the cognitive processing of humorous stimuli (e.g., Apte 1985); it is hard to
specify what makes something funny.
Sometimes humor is defined in terms of its characteristic expression,
or display-laughter. Observed laughter is highly correlated with subjective
ratings of funniness (Chapman 1983). However, other emotional states, nota-
bly triumph and anxiety, are sometimes accompanied by laughter or smiling
(Monro 1951). And we are often amused without laughing or smiling, espe-
cially when alone. Smiling is far less specific as an indicator of humor, since
it can accompany almost any pleasurable state. (See LaFrance [1983] for an
ethological description of smiling and laughter as facial expressions.)
Because of these limitations of defining humor by its cognitive properties
or displays, I favor referring to its characteristic affect, which might be
termed humor appreciation. Laughter is useful as a measure of the amount
of humor appreciation occurring in a research setting, but the affect is para-
mount. Laughter can occur without anything being funny, but these situa-
tions are-not funny. Therefore, for purposes of this essay, humor apprecia-
tion is a distinct, pleasurable affect that often is accompanied by laughter.
Defining humor mainly by its affect agrees with common practice for
other emotions (cf. Zajonc 1980). Emotions theorists usually employ affec-
tive labels such as hunger, thirst, fear, and interest, rather than cognitive or
stimulus terms such as food perception and food, water detection and water,
interestingness and novelty, or danger detection and dangerousness.
But there is more to an emotion than just its characteristic affect and
display. Ekman (1984) and Scherer (1984) viewed the universal human emo-
tions as comprising several adaptive facets. These facets always include a
specific affect, an overt behavioral tendency, neural mediators, and eliciting
stimuli. They may or may not include viscera1 adjustments, displays, and
higher cognitive processing. Information about all of its facets is useful for
identifying a basic human emotion.
I agree with Zajonc (1984), however, that affects are primary. Displays
and visceral adjustments may be absent or ill defined, as in thirst and the
esthetic emotions, and higher cognitive processing may not occur. The trou-
 Adaptive Value of Humor and Laughter 143

ble with defining an emotion by its particular behavioral tendency is that the
latter can take multitudinous forms, especially in primates. But all emotions
possess a single distinctive affect and an adaptive behavioral tendency, and
whatever displays and visceral adjustments occur.

DOES HUMOR HAVE AN EVOLVED BASIS?

Does humor have a species-wide, evolved basis whose function can, in the-
ory, be discerned? “Evolved basis” means that the behavior was selected
for specifically in evolution, rather than being an incidental by-product of
the general behavioral capacities of the species (e.g., bicycle riding by bears).
Several research strategies, pioneered by Darwin (1872), are commonly used
to test for an evolved basis for a given behavior. These strategies include
testing for cross-species prevalence, early (presocialization) onset, specific
neurophysiological mediation, presence in related species, and stereotypy
(Weisfeld 1982). Some of these strategies for identifying an evolved behavior
will now be invoked in testing for an evolved basis for humor.
The first of these strategies is to see if the trait occurs throughout the
species. If it does, then an evolved basis is probable (Barlow 1977; Eibl-
Eibesfeldt 1975).
Humor and laughter seem clearly to be universal in our species, and to
be likely to have an evolved basis (Darwin 1872; Hinde, 1974). Assertions
to the contrary by some anthropologists (e.g., Birdwhistell 1970) seem to be
based upon the observation that norms concerning the overt exhibition of
facial expressions including laughter vary across cultures, and not upon evi-
dence of any culture lacking humor or laughter (Apte 1985). All universal
emotions seem to be expressed (or suppressed) under somewhat different
circumstances in different cultures, but this does not contradict the universal
presence and therefore the likely evolved basis of these emotions (Ekman
1973).
Another research strategy for identifying traits with an evolved basis is
to test for early onset (Tinbergen 1951). Variants of this strategy include
raising an animal in social isolation or otherwise precluding the possibility
of acquiring the behavior through social experience.
Laughter does not appear until about four months of age (McGhee 1979;
also see Sroufe and Waters 1976); one of Darwin’s (1872) children first
laughed at 113 days. Young (1973) argued that this onset was early enough
for laughter to qualify as “innate,” but others may disagree. Plooij (1979)
reported that laughter in response to tickling by the mother emerged at three
months in the chimpanzee. One instance of a human baby laughing when
tickled at five weeks is recorded (F. X. Plooij, personal communication
1992).
Even if the onset of laughter is taken as too late to support its having
an evolved basis, the possibility of an evolved basis still remains. Many
144 G. E. Weisfeld

traits, such as the changes of puberty, are genetically programmed to appear

well after birth.
It is sometimes possible to identify a specific neural structure orpathway
that mediates a particular behavior. If the existence of such a structure can
be demonstrated, this suggests that the behavior itself, like the structure,
has been selected for.
In the case of laughter, some patients with pseudobulbar palsy; bulbar
palsy; or basal ganglia, hypothalamic or temporal lobe pathology sometimes
exhibit “sham laughter” (Bannister 1973; Black 1982; Kalat 1988; Williams
1956). In this condition, laughter appears spontaneously or inappropriately
(see also Haig 1988; Poeck 1969). In pseudobulbar palsy cases, the most
common, the patient usually does not experience pleasure, and laughter is
atypical in form (Black 1982). When ictal laughter arises from a hypothalamic
focus, pleasure is usually absent also, but limbic seizures accompanied by
“fits” of laughter are typically pleasurable. Similarly, laughter can be trig-
gered by nitrous oxide, by electrical stimulation of the anterior globus pal-
lidus (Hassler and Riechert 1961), or by lesions of the right hemisphere
(McGhee 1983b).
Since the hypothalamus and the limbic structures of the temporal lobe
are involved in the expression of numerous other species-wide, evolved emo-
tions, this evidence that laughter too is mediated there suggests that this
expression likewise has an evolved basis. The following neural system seems
to operate generally in emotional expression (Buck 1984), including laughter
(Black 1982): At the cortical level, control is exercised and the expression
can be modified in form (Black 1982); the sham laughter of pseudobulbar
palsy, in which cortical control is interrupted, has been described as infantile
(Ironside 1956). At the effector level, bulbar nuclei evoke the facial, phonic,
and respiratory components of laughter and other expressions. At the hypo-
thalamic and limbic level, integration of these expressional components oc-
curs. Of course, the cerebral cortex is also involved in the appreciation of
humor, especially the right frontal lobe (Brownell and Gardner 1988).
Abnormalities not only in appreciating but also in generating humor can
sometimes occur with neural lesions. Prankish joking and punning (Witzel-
sucht), accompanied by a “fatuous euphoria,” sometimes occur with frontal
lobe lesions (Lishman 1978). Right hemisphere lesions sometimes result in
joking in inappropriate contexts (McGhee 1983b; Brownell and Gardner
1988). Martin (1950) described a remarkable case of a patient who had an
intraventricular cyst removed under local anesthesia. As the surgeon was
swabbing blood from the floor of the third ventricle, the patient burst out
laughing, made jokes, and uttered obscene remarks. This reaction was re-
peated whenever this area of the brain was touched, and only so long as it
was touched. From this case and others, Martin concluded that a “humor
center” exists in the region of the hypothalamus.
The case for an evolved basis for a particular behavior is strengthened
by demonstrating phylogenetic continuity, that is, showing that the trait oc-
 Adaptive Value of Humor and Laughter 145

curs in related species also. Human laughter is thought to be phylogenetically

descended from the relaxed, open-mouth display, or play face expression,
of nonhuman primates (van Hooff 1972). Further, simians laugh in much the
same way as people do, including children at play (Monro 1951; van Hooff
1972; Yerkes 1943). Darwin (1872) compared the sound of human laughter
to the vocalizations of apes being tickled. Goodall (1968) went so far as
to state that “ ‘[llaughing’ (a series of staccato panting grunts) frequently
accompanied bouts of wrestling and tickling” (p. 258) in chimpanzees. Bernt-
son et al. (1989) presented a sound spectograph of the “laughter” of ajuvenile
male chimpanzee at play. Likewise, vocalizations resembling laughter have
been reported to accompany play in various Old World primates: gorillas,
orangutans, entellus langurs, various baboons, and rhesus macaques (Aldis
1975). These comparative observations suggest that humor itself may have
some equivalent in other primate species, and that therefore it is likely to
have an evolved basis in humans (Hinde 1974).
Indeed, humor appreciation-not just laughter accompanying play or
tickling-has been reported anecdotally in captive apes. The following exam-
ples were personally communicated by various trainers to McGhee (1979).
Chimpanzees and gorillas that have learned sign language sometimes give a
series of incorrect linguistic signs, often accompanied by “laughter.” Chim-
panzees have been observed to throw debris at or to urinate on people, and
then to sign “funny”; this sign has also been used to accompany tickling or
chasing games. Apes also sometimes use the sign for “funny” to refer to
human behavior that is silly, unusual, or out of context. And they sometimes
distort a linguistic sign, as when Koko the gorilla placed her thumb in her
ear instead of her mouth for “drink.” From my own experience recounting
these simian witticisms, undergraduates seem to find them highly amusing
(more so than my own attempts at humor), suggesting that humor is indeed
shared by apes and people, at least at the undergraduate level.
A final research strategy is to test for steveolypy of the behavior. Laugh-
ter is clearly stereotypic across cultures, and “not restricted by either the
linguistic or the cultural codes of the laugher” (Apte 1985, p. 252). People
do not use one emotional expression to convey amusement in one culture
and a different expression elsewhere. The expression of laughter is stereo-
typic for our species, and was described anatomically by Darwin (1872).
Acquisition by imitation is unlikely; a lo-year-old congenitally blind and deaf
girl was observed to laugh recognizably (Goodenough 1932).

A UNITARY FUNCTIONAL EXPLANATION OF HUMOR

If humor does have an evolved basis, then its function may be discernible
through comparative analysis. True wit (excluding laughter at play) may be
found only in humans and chimpanzees (and possibly gorillas). What features
distinguish these species from the other primates?
One clue to the function of humor may be the sociality of these species.
146 G. E. Weisfeld

Indeed, humor is an eminently social activity (Bergson 1911). Studies of

children and adults consistently show that laughter occurs in a social context
over 95% of the time (Bainum et al. 1984; Kenderine 1931; Provine and
Fischer 1989; Young 1937). The specific aspect of sociality that is crucial
for the evolution of humor remains to be identified. Perhaps the ability to
impute mental states to others, found in chimpanzees, allows one to joke
about the foibles of others.
Unlike many emotions that often occur in social isolation, such as hun-
ger and fear, humor is eminently social. The social nature of humor is under-
scored by the fact that it possesses a distinct emotional display. In fact, it
has no characteristic overt behavior except for its displays, laughter and
smiling. Most other affects, such as fear and anger, evince a specific overt
behavior, such as flight and aggression, in addition to any display elements.
The prominent role of the display of laughter may offer clues to the
function of humor. Animals tend not to exhibit a display unless they benefit
from influencing another animal thereby. Displays are costly in that they
consume metabolic energy and attract the attention of predators. Thus, theo-
ries that explain laughter as an adaptive overflow of surplus nervous energy
(Freud 1905; Spencer 1860) violate the law of economy (Stanley 1898).
How, then, does laughter affect the receiver? Laughter seems to be
rewarding-in two ways. First, laughter is often described as musical (cf.
amuse), since it consists mainly of voiced vowels (Apte 1985). When we
intentionally amuse someone, we are rewarded by that person’s laughter,
which is pleasant sounding.
Second, laughter is taken as a compliment to our wit. This is not specific
to laughter; other emotional expressions, especially attention and smiling,
are also rewarding. Moreover, expressions of appreciation are given in re-
sponse not just to humor but also to other esthetic forms, such as storytelling
and music. The fact that laughter punctuates conversation even when no
humor occurs (Provine and Fischer 1989) suggests perhaps that we learn that
laughter keeps the other person talking.
How might sending these rewarding signals benefit the laugher? By
laughing, smiling, or otherwise indicating appreciation, one elicits continued
humorous stimulation, which may enhance fitness in a way specified below.
If people did not benefit from receiving this stimulation, they would probably
not have evolved displays that reward those who provide it.
But the humorist too must benefit from eliciting appreciative laughter,
or else the tendency to enjoy it would probably have been selected against.
How do we gain from amusing others? Perhaps humorous stimulation consti-
tutes a service that the laughter appreciates. By laughing, the recipient in
effect acknowledges having received something valuable from the humorist.
Laughter may be an expression of appreciation, or gratitude, for a favor
received.
Trivers (1971) suggested that the capacity for experiencing gratitude
evolved so that we reciprocate for favors received from others, and hence
 Adaptive Value of Humor and Laughter 147

encourage others to offer us additional favors. Humans and chimpanzees

(Teleki 1975) are among the few species known to have evolved a capacity
to reciprocate favors. In order for reciprocal altruism to evolve in a species,
individual members of the group must be able to recognize each other, to
keep track of who has done what for whom, and to pay others back. In
our species, reciprocation and hence mutual benefit are promoted by the
operation of several emotions (Trivers 1971). A nonreciprocator is punished
through the victim’s anger and his own guilt. A reciprocator is rewarded via
the recipient’s gratitude and his own pride (Weisfeld 1980). These emotions
impel us to exchange services equitably.
Consistent with the notion that being amused constitutes a favor, we
appreciate a good joke and acknowledge our appreciation; the humorist is
proud of having amused us, and feels justified in being compensated for his
performance. As expected if the exchange of jokes is a form of reciprocal
altruism, humor is common among friends and coworkers, who socialize
together over long periods (Fine 1983).
Furthermore, as with receiving other forms of favor, we dislike laughing
at our enemies’ jokes; perhaps we wish to avoid being obligated to return a
favor to them. On the other hand, we are quick to express appreciation for
the boss' jokes, so as to show our admiration and liking for her (Kane et al.
1977). Thus, laughter may carry some of the appeasement connotation of
the smile. Laughter is indeed more effusive toward the jokes of a high-status
person (Fry 1977).
But humor seems not to enhance our fitness directly. Therefore the
benefit probably comes later. Delayed payoffs are typical of the Old World
primates, which mature slowly and rely heavily on learning to increase their
behavioral efficiency in adulthood. They learn through solitary play, peer
play, observation, imitation, practice, and sometimes active guidance and
stimulation by the mother. Perhaps humor provides another means of social
learning.
The functional explanation I wish to propose, then, is this: Humor pro-
vides the recipient with information or stimulation that later enhances$t-
ness. Laughter constitutes an evolved, rewarding emotional expression, This
expression has the effect-usually not deliberate-of encouraging the hu-
morist to continue to provide the information or stimulation of humor to
the recipient. Laughter in response to wit often carries a connotation of
appreciation or gratitude, i.e., an implied promise to reciprocate for a favor
received.

RESEMBLANCE OF HUMOR TO PLAY AND INTEREST

This social stimulation explanation of the function of humor is supported by
the resemblance of humor to play (Mannell and McMahon 1982), whose
probable function is to provide practice in essential skills (Boulton and Smith
1992; McDougall 1923; Smith 1982; Suomi and Harlow 1971). One point of
148 G. E. Weisfeld

similarity has already been mentioned: the common phylogenetic origin of

the primate play face and human laughter. In addition, laughter accompanies
play as well as humor. Moreover, the play face and laughter emerge at about
the same time in human (L. J. Stettner, personal communication 1992) and
chimpanzee infants (Plooji 1979). And in both humor and play, some signal,
or metacommunication, often conveys the frivolousness of the interaction
(Fry 1963). For example, laughter and a play face often accompanied pre-
schoolers’ rough-and-tumble play, whereas a frown or stare were associated
with aggression (Blurton Jones 1967, 1972).
Several other parallels can be noted. Like play, laughter tends to occur
when no urgent needs require attention. To appreciate a joke, one needs to
be in a good humor. In a study of undergraduates, anxious individuals exhib-
ited less appreciation of humor (Blank et al. 1983). Similarly, being in a
playful or hedonic mood (cf. Chance 1988) increases receptiveness to humor
(Bariaud 1988; Mannell and McMahon 1982; Martin and Lefcourt 1983).
Martin (1950) found that exposure to sad pictures reduced reported amuse-
ment to subsequently presented humor, whereas exposure to funny pictures
enhanced it. In another study, facial responses and ratings of funniness in-
creased over the first few cartoons of a series (Deckers et al. 1989); cf. the
phenomenon of warming up an audience.
Thus, humor may elevate one’s mood, and may do the same for bystand-
ers. The well-established contagiousness of humor (McGhee 1976) may stem
partly from the relaxing bemusement that results from seeing someone laugh.
If fear is contagious, then lack of fear, as signaled by laughter, may also be
contagious and may put us in a mood to laugh. The contagiousness of humor
has been documented clinically, in the form of veritable epidemics of hysteri-
cal laughter (Izard 1979). One such episode began at a convent school in
East Africa in 1962 and spread to 14 other schools, forcing their closure,
and thence to the girls’ female relatives and finally to neighboring villages
(Bean 1967). Some patients required hospitalization for exhaustion.
A link between humor and play is perhaps supported by the terms used
to refer to humor. These include ludicrous (from ludere, L., to play), funny,
word play, horseplay, slapstick (cf. play fighting), lark (from laken, M.E.,
to play), monkeyshines, trick, and joke (from jocu,, L., a joke or game).
If we regard play as one group of overt behaviors that are prompted by
the emotion of interest, then additional parallels to humor emerge (cf. Ber-
lyne 1972). Interest tends to wane with exposure or familiarity; similarly,
humor ratings by adults declined with repetition of the content (Gelb and
Zinkhan 1985). Subjects who were high on susceptibility to boredom (pre-
sumably the opposite of interest) were found to appreciate humor more (Ruth
1988). Depression can reduce both interest and humorousness, and euphoria
can increase them (Haig 1988). We express appreciation in both cases.
Both interest and humor are intensified if the content is relevant to
fitness, e.g., sex, aggression, or competition (McCauley et al. 1983; McGhee
1979). Boulton and Smith (1992) discussed the possible adaptive value of
 Adaptive Value of Humor and Laughter 149

rough-and-tumble play in providing practice for fighting, predator avoidance,

and hunting. Interest and humor are both maximal when the cognitive con-
tent is moderately challenging to the subject- the cognitive congruency prin-
ciple, or discrepancy hypothesis. Sroufe and Wunsch (1972) found that
younger infants laughed more at cognitively simpler stimuli (tactile and audi-
tory), and older infants at more complex stimuli (visual and social). Similarly,
tickling had to be moderate to elicit laughter in chimpanzees; mild tickling
was ineffective, and strong tickling produced withdrawal (Plooij 1979). Also,
prolonged tickling between juvenile or adolescent chimpanzees usually re-
sulted in screaming.
Consistent with the discrepancy hypothesis, the appreciation of incon-
gruity humor increases with age (Justin 1932), as children’s cognitive capaci-
ties develop. In other research on children, humor appreciation and laughter
have been found to be greatest when the joke content was neither too easy
nor too difficult for the subjects (Bariaud 1988; Whitt and Prentice 1977;
Zigler et al. 1966, 1967), and when the joke depended on the cognitive devel-
opmental stage of the child (Athey 1977) or on a Piagetian operation that the
child had recently mastered, e.g., conservation (McGhee 1976). Similarly,
the appeal of peekaboo to infants in the first year (McGhee 1979) may corre-
spond with the development of object permanence. Senile individuals tend
to lose these Piagetian skills in reverse order of the original acquisition.
Comprehension of conservation concepts declines with age, and appreciation
of jokes on a conservation theme increases concomitantly (Schaier and Cici-
relli 1976).
Presumably, it would be adaptive for people to be drawn to cognitive
content that was appropriately challenging-neither too difficult to be mas-
tered, nor too simple to provide any new understanding. The fact that we
are drawn to appropriately challenging humorous content, e.g., to jokes that
play on our current level of cognitive development, supports the idea that
humor is informative.
Interest and play resemble humor in their ontogeny. All of them seem
to peak in childhood, which is consistent with the idea that they are edifying.
The earlier that one can acquire some bit of information, the longer one can
benefit from it. The lesser appreciation for humor in the elderly (Schaier and
Cicirelli 1976) can perhaps be explained ultimately in terms of a diminished
need for new ideas.
If humor resembles the emotion of interest, why do we need both?
Would not interest suffice to motivate us to focus on stimulation that was
salient to fitness concerns? Perhaps the capacity for humor evolved to en-
courage others, through laughter, to provide especially poignant informa-
tional stimulation. Interest can arise in an asocial context, and occurs in
many asocial species. Humor, however, is eminently social, and seems to
have evolved not only to prompt the recipient to seek out humorous stimula-
tion but also to spur the humorist to continue providing it. Furthermore, the
content of humor often concerns social behavior and especially human foi-
150 G. E. Weisfeld

bles (Bergson 1911); by contrast, inanimate phenomena are usually merely

interesting (Haig 1988).
Another difference between humor and interest seems to lie in the la-
tency of response. Fry (1963) observed that a punchline is essential for
humor, but not for play or games. Jokes that drag on are not funny; jokes
that were rated as more humorous were understood faster (Goldstein 1970).
Professional comedians often emphasize timing (Nathan 1971). Interest, on
the other hand, can be maintained for some time, leading to protracted explo-
ration. Thus humor may be more suited to the rapidity of social interaction.
Although both humor and interest can lack “seriousnes5,” interest is
sometimes elicited by “real” situations requiring an earnest response.
Humor, on the other hand, is usually playful and can be indulged in solely
for its future benefits (cf. Berlyne 1972; Rothbart 1973).
A final difference is that humor may be experienced when we receive
certain characteristic types of stimuli that do far more than arouse interest.
These stimuli may be extraordinarily valuable, and hence a behavioral mech-
anism for rewarding others for presenting them may have evolved. Simi-
larly, other forms of esthetic appreciation, such as olfaction, music, and
visual art, seem to constitute distinct evolved capacities that draw us to
certain stimuli that have generally been adaptive to experience during homi-
nid prehistory (Pugh 1977).
As mentioned above, interest and humor are maximally aroused by stim-
uli that are moderately challenging for the subject. Similarly, esthetic stimuli
that are moderately complex seem to be the most appealing. Presumably
this tendency maximizes learning, since it inclines us to seek out stimuli that
we can profit from discerning but that are not beyond our cognitive powers.
Also adaptively, once we comprehend, or appreciate, the stimulus (problem,
joke, work of art), we experience an increase in pleasure that then dissipates.
This rewards us for our attention and then impels us to move on to other
sources of cognitive stimulation. An old joke is seldom funny, since it con-
veys no new information. Interestingly, little habituation to a series of car-
toons was reported, a result that the authors interpreted as indicating that
cartoons even on the same theme can be highly variable (Deckers et al.
1989).

APPLYING THE THEORY TO VARIOUS TYPES OF

HUMOR

I will now try to apply the theory to some of the different types of humor.

Tickling
If it were especially important for the organism to receive a certain form of
stimulation from others, then laughter might have evolved to reward those
 Adaptive Value of Humor and Laughter 151

who provided this stimulation. One such form of stimulation may be tickling.
Enjoyment of tickling may be a fundamental source of humor and hence a
crucial test for theories of humor (Alexander 1986; Darwin 1872; McDougall
1923). It occurs in chimpanzees as well as humans, and it reliably elicits
laughter in infants as young as four months old (Apte 1985). These facts
argue for an early phylogenetic origin for humor in our species, the biogenetic
law having general, albeit imperfect, validity (Gould 1977).
Tickling may constitute valuable stimulation because it provides prac-
tice in defending vulnerable bodily areas (Alexander 1986). Hayworth (1928)
noted that the tickler plays the role of a friendly attacker, stimulating parts
of the body that are supplied with protective reflexes-the soles, armpits,
ribs, and abdomen. Darwin (1872) characterized these bodily areas as those
that are seldom touched, which is consistent with these areas being vulnera-
ble and hence protected by avoidance and guarding behaviors. Consistent
with this self-protection interpretation, an essential element in tickling seems
to be the surprise of a sudden thrust, or attack; thus, one cannot tickle oneself
(McDougall 1923). If the tickler stops being playful and instead scowls, the
effect is aversive (Leuba 1941; Melzack 1967); an earnest attack needs to
be defended against.
The laughter of a child who is being tickled is itself highly amusing, and
prompts the tickler to persist (Alexander 1986). Darwin (1872) suggested that
a capacity for humor develops in infancy and rewards caretaking by adults,
thus promoting survival. As children and infant chimpanzees get older, they
generally become less ticklish (Plooji 1979); in fact, tickling may become
aversive. This is typical of the developmental pattern for most forms of play.
In most mammals, play drops out by adulthood, presumably because the
mature organism needs to have mastered whatever behaviors it has acquired
through play practice. The fact that tickling likewise declines with maturation
(perhaps because it becomes more aversive) supports the interpretation that
it promotes practice in a vital behavior.
This practice can be provided unilaterally, as when a parent tickles a
child, or it can be mutual, as when two children tickle each other. They do
so in order to hear laughter and to be tickled back. Because tickling requires
no great skill, presumably this laughter lacks the laudatory aspect of laughter
in response to true wit. In this respect tickling is a rather simple, rudimentary
form of humor.

Social Play
Certain other examples of humor also seem to meet the test of providing
valuable stimulation through the playful practice of attack and defense. Brit-
ish and Bushman children exhibited laughter most commonly in association
with chasing and fleeing, hitting and wrestling (Blurton Jones 1972). These
activities are standard games for juvenile primates in many terrestrial spe-
cies, and are accompanied by the play face expression. Practicing these
152 G. E. Weisfeld

agonistic behaviors is of obvious relevance to fitness, especially for male

primates, which tend to play fight more than females (Hutt 1972). Similarly,
performing bodily stunts is a favorite amusement for children (Stone and
Church 1957), as is slapstick.

Joking
Many forms of humor pertain to loss of social, rather than bodily, poise,
These jokes about “ticklish” situations are hard to create and tend to elicit
appreciative laughter. Along these lines, Alexander (1989) stated Lhat humor
can provide practice for social competition. Apte (1985) suggested that jokes
provide instruction on how to avoid embarrassment under difficult condi-
tions.
Various pieces of information suggest that humor often enlightens us
about social foibles. Professional comics often employ a stooge to serve as
the butt of jokes-or, occasionally, to turn the tables on the comic. Many
jokes contain an element of public failure or embarrassment, such as teasing,
ridicule, repartee, pranks, farce, and clowning. Analyzing “trickster tales”
cross-culturally, Apte (1985) concluded that they exercise a normative func-
tion, inasmuch as the trickster’s clumsy, antisocial attempts to satisfy his
biological needs are typically thwarted.
Jokes, because of their inherent playfulness or surrealism, allow people
safely to probe sensitive social issues, and even to expose the foibles of
others (Kane et al. 1977) or themselves. Despite the surreal quality of humor,
humorous stories that are true usually are especially funny (Pugh 1977),
perhaps because the information they convey is ecologically valid.
Not all jokes reveal social foibles, of course. Many jokes have an ele-
ment of incongruity, as discussed below. The social foible element of humor
seems to be absent from simian humor, and to develop relatively late in
humans. Children enjoy incongruity jokes, such as riddles concerned with
causal and logical relations (Athey 1977), but older subjects, in addition to
appreciating incongruity humor, often joke about social issues of special
interest to them. Much of the humor of adolescents pertains to sexuality,
peer norms, and ambivalence toward authorities (Fine 1983; Omwake 1937;
Ransohoff 1975; Sanford and Eder 1984). Jokes told by adults commonly
concern sex, ethnicity, politics, and alcohol (Winick 1976)-all delicate sub-
jects in the U.S.

Word Play
Many examples of wit involve word play. Children learn to speak from age
2 to 5; accordingly, preschoolers frequently laugh about words (Davids
1987)-initially, about the sounds of words (McGhee 1983a). Inasmuch as
we continue to improve our mastery of language throughout life, it makes
sense that word play should be amusing even in adulthood. We treat the
 Adaptive Value of Humor and Laughter 153

“lowly” pun with disdain, perhaps because puns even amuse seven-year-
old children (McGhee 1983a), but we are amused nonetheless.
Is it justifiable to group these various forms of amusement together-do
they all constitute humor? The fact that they all frequently elicit laughter, a
universal emotional expression, suggests a common evolutionary basis.
This is not to say that humor cannot be classified in a biologically mean-
ingful way. To the contrary, it is likely that tickling and social play were the
first forms of humor to evolve, since they occur in nonhuman primates and
develop early. Other forms of humor, such as jokes, word play, and other
witticisms, may be quite different in their neural basis from that of tickling
and play fighting. Further analysis ideally will lead researchers to a generally
accepted typology of humor.

OTHER FUNCTIONAL EXPLANATIONS OF HUMOR

The theory offered here will now be contrasted with other theories of humor.
In some cases, an integration seems indicated. In others, the competing
theory may violate some evolutionary principle.

Incongruity

Many theories of humor entail the notion of abrupt incongruity or paradox

(e.g., Berger 1987; Hayworth 1928; Nilsen, Donelson, Nilsen, and Donelson
1987; also see Keith-Spiegel 1972). Kant (1790 quoted in Haig 1988, p. 16)
described humor as “arising from a strained expectation being suddenly
reduced to nothing.” The word absurd originally meant “out of tune,” and
burlesque derives from burrae, L., nonsense.
Research support for the incongruity interpretation abounds. Incongru-
ity seems to be featured in humor around the world. Shultz (1977) concluded
that incongruity characterized the vast majority ofjokes, riddles, and humor-
ous tales of every culture studied. For example, he himself analyzed 242
Chinese jokes and found that 210 relied on incongruity and its resolution.
Six did not, and the rest were “inscrutable.”
A dose-related effect of incongruity on funniness has even been demon-
strated. Nerhardt (1970) and Deckers and Kizer (1975) found that the greater
the discrepancy between the expected weight of an object to be lifted by the
subject and its actual weight, the more laughter and smiling occurred.
Rothbart (1977) reviewed evidence for incongruity humor through the
life span. Incongruous elements, such as peekaboo and distorted faces, are
appreciated by infants. The bodily distortions and exaggerated movements
of cartoon figures may explain much of their appeal. Langevin and Day
(1972) elicited laughter from children by showing them incongruous pictures,
e.g., of a door with handles and hinges on both sides. McGhee (1979) has
154 G. E. Weisfeld

presented a stage theory of the development of incongruity humor in

children.
Another such theory, the two-stage theory of humor, holds that even
children appreciate simple incongruity, but more sophisticated humor in-
cludes a poignant resolution of the incongruity (Shultz 1972; Suls 1972). In
support of this theory, patients with right hemisphere lesions usually show
surprise at an incongruity (the first stage). However, they may be “unable
to integrate the punch line information with information from the rest of the
joke to provide a coherent package of meanings” (McGhee 1983a).
Pugh (1977) explained the functional value of incongruity humor by not-
ing that incongruity or inconsistency arises from counter examples, which
are especially valuable in Bayesian deduction. For example, the discovery
of one white crow is more instructive than the discovery of yet another black
crow.
Pugh’s interpretation neatly accounts for word play and also agrees with
the present one in emphasizing the social nature of humor. He referred to
humor as “nature’s way of motivating the social sharing of counter exam-
ples” (p. 329). Because Pugh’s approach is also biological, it is not surprising
to find general agreement between the two theories.
However, he maintained that language is necessary for communicating
ideas about counter examples, or incongruity, and that this explains the
absence of humor in nonhuman primates. But as noted above, chimpanzees
can exhibit a sense of humor. Moreover, many examples of humor in our
species ‘do not seem to depend on language, such as sight gags, mime, and
musical jokes. The point is underscored by the fact that some patients with
left hemisphere damage no longer appreciate cartoons with verbal captions
but do respond to cartoons without captions (Kolb and Whishaw 1985). Then
too, much of nonverbal humor may be said to contain elements of incongru-
ity, such as cross-dressing and other “sight gags.” Thus, Pugh’s reasoning
that humor requires language is dubious.
Nevertheless, his explanation of the adaptive utility of receiving “incon-
gruous” counter examples from others fits into the present theory, and in-
deedstrengthens it. As Haig (1988) pointed out, incongruous behavior, such
as acting in a rigid, one-track fashion, is often very funny, perhaps because
it points up a human foible and hence is especially instructive.

Superiority
Another common theory of humor is the “sudden glory” interpretation of
Hobbes (1651), that we regale in the failure or embarrassment of others
(Alexander 1986; Lorenz 1963; Plato 1871). Similarly, Aristotle (1895) held
that defects or deformities are the source of the ludicrous.
Certainly, many jokes can be characterized as put-down humor or prat-
falls. The Greenland Eskimos traditionally resolved disputes by engaging in
public contests of ridiculing each other (Levine 1977). U.S. street gang
 Adaptive Value of Humor and Laughter 155

youths often indulge in competitive banter, in which they attempt to “rank”

the opponent. Chapman (1983) has suggested that a group’s rank order may
be discernible in its pattern of who ridicules whom and who laughs at whom,
just as evaluative comments were observed to flow down the hierarchy dur-
ing a volleyball game (Weisfeld et al. 1984). Supporting Chapman’s interpre-
tation, Coser (196tJ) demonstrated that ridicule traveled down the hierarchy
among hospital workers. Interestingly, preschoolers, in whom the capacity
for dominance hierarchization is nascent, laughed more at an adult victim
than at a child (McGhee 1983a).
Eibl-Eibesfeldt (1989) has pointed out that laughing aloud at someone
is a profoundly aggressive act. He maintained that this vocalization is derived
from the phylogenetically ancient mobbing call; both vocalizations are “sim-
ple, staccato, repetitive” (Provine 1992, p. 3). He noted that, similarly, col-
lective laughter solidifies the group, as discussed in the next section. The
intensity of the negative experience of being ridiculed argues for the impor-
tance of this aspect of humor.
However, many forms of humor lack this competitive element. The
superiority theory fails to account easily for word play, for the laughter that
accompanies children’s play, for the incongruity element in humor, and for
self-deprecating humor. Furthermore, ridicule usually involves three par-
ties-humorist, butt, and bystanders-and so is unlikely to constitute a more
basic form of humor than dyadic examples of humor, such as tickling.
Another problem, pointed out by McDougall (1923), is that another per-
son’s “serious” misfortune, although sometimes gratifying (e.g., Schaden-
fieude [Bateson 1969]), usually is not funny. In fact, research indicates that
the greater the misfortune of the butt of the joke, the less funny the joke
(Zillman et al. 1974). Furthermore, humor is greatest when retaliatory acts
equitably compensate for some provocation, rather than when superiority
of one party results (Zillman and Bryant 1974).
Also, it is difficult to see how the individual’s fitness is enhanced by
laughing at another person’s failure, beyond the advantage entailed by a
rival’s failure per se. In many species, one animal triumphs over another
without the necessity for laughter at the vanquished. As Kane et al. (1977)
put the point, this explanation fails to explain why humor should be used
when clever criticism would serve just as well. Mocking laughter may sum-
mon allies (Alexander 1986), although at the cost of losing the butt’s alle-
giance.
Even if it did not evolve for this purpose, humor may incidentally pro-
vide a means for people to compete in a legitimate, make-believe way, as
Kane et al. (1977) suggested. Perhaps repartee, ridicule, mockery, sarcasm,
and so forth allow people to compete with the understanding that it is “all
in fun”; cf. Radcliffe-Brown (1940) on joking relationships and Freud (1905)
on aggressive outlets.
Humor furnishes a means of establishing one’s superiority not just by
ridiculing others but also by showing that one has got the joke (Alexander
156 G. E. Weisfeld

1986). Even humor at our own expense can demonstrate our superiority by
virtue of the fact that we are self-assured enough to acknowledge our foibles
(Alexander 1989).
But, again, these applications of humor seem to miss the point of explain-
ing why the appreciation of humor evolved in the first place. Why would
simple criticism not suffice? Many behaviors can be performed competi-
tively, such as running, cooking, and even urinating. These incidental appli-
cations do not explain the evolutionary origin of these behaviors.
To the extent that people compete to be witty, i.e., to earn others’
appreciation through joking, humor may be said to be competitive. Also,
jokes at the expense of a butt may be said to be aggressive. But many jokes
lack a butt and hence are not essentially adversarial.
Aggressive individuals do tend to like aggressive humor (Grziwok and
Scodel 1956; Hetherington and Wray 1964; Holmes 1969), and subjects in a
violent country (the U.S.) liked hostile jokes more than did Japanese and
Senegalese subjects (Goldstein 1977). But these results may be due simply
to aggressive subjects often being in an aggressive mood and therefore inter-
ested in aggressive content. Strickland (19.59) angered some subjects and
found that they then preferred aggressive humor; similarly, sexually arousing
other subjects resulted in greater appreciation of sexual humor.
Thus aggressive joking may merely result from heightened interest in
this topic and may not reduce aggressive urges any more than sexual humor
reduces libido. In fact, exposure to aggressive humor can heighten subse-
quent aggression (e.g., Berkowitz 1970), not lower it as seems to occur when
the humorous material distracts the subject from being angry (Baron 1977).
Aggressive humor does not seem to be cathartic, any more than exposure
to aggressive films reduces subsequent aggression. In fact, exposure to ag-
gressive films or models usually heightens aggression. A better interpretation
seems to be that subjects enjoy humor that is salient to matters that interest
them. For example, subjects especially appreciate automobile jokes (or mu-
sical jokes or medical jokes) if they have just been exposed to the correspond-
ing content (Goldstein et al. 1972).
Why does this salience element seem to operate? Individuals who prefer
a particular type of humorous content may seek it out because learning about
salient matters increases one’s fitness. By telling aggressive jokes, one may
explore the topic of aggression and prompt others to tell informative aggres-
sive jokes in turn. Similarly, depressed patients were especially amused by
jokes about depression, suicidal patients by morbid jokes (Haig 1988), and
feminists by anti-male jokes (Chapman and Gadtield 1976).

Social Control

Another theory of humor refers to social control (Bergson 1911; Wallis 1922;
Wilson 1979). According to this interpretation, laughter corrects deviant be-
 Adaptive Value of Humor and Laughter 157

havior. This indeed does occur (Bryant et al. 1983), and criticism applied
with the balm of humor may be easy to accept (Haig 1988).
However, it is difficult to see why tactful criticism or shunning would
not do just as well, or why such a complex behavioral system would have
evolved for an apparently limited purpose (McDougall 1923). Also, this the-
ory makes it difficult to account for word play, children’s laughter at play,
and the incongruity element of humor.
Usually this explanation is cast in terms of the welfare of the group, i.e,
the group is said to benefit from this method of social control (Fine 1983).
However, group selection of genetic traits is probably rare (Williams 1966).
The social control effect of humor may be an incidental by-product of
its original function. The humorist exposes the foibles of the butt of the joke
mainly in order to amuse bystanders and earn their appreciation, and perhaps
to establish her superiority over the butt. This behavior is generally adaptive
to her because she gains credit for providing her listeners with information
about faux pas. Incidental to these effects, group members fear being simi-
larly ridiculed and learn how to avoid similar faux pas; deviant behavior is
thereby reduced. Many forms of humor, including irony, satire, parody,
sarcasm, and caricature, possess this corrective potential.

Group Solidarity
Another explanation holds that sharing jokes enhances in-group solidarity
(Alexander 1986; Martineau 1972) through mutual enjoyment and the
achievement of consensus. Humor can indeed increase self-reported cohe-
sion in group members (Banning and Nelson 1987; Vinton 1989). For exam-
ple, in a Chippewa tribal council, mutual ribbing seemed to enhance solidar-
ity (Miller 1967). The strong contagion effect of laughter and the fact that
the amount of laughter increases with group size both suggest the operation
of “an automatic social synchronization process that predates language”
(Provine 1992, p. 2). Humor does seem to bind groups just as play and other
forms of hedonic activity do.
However, it is difficult to see why sharing a joke should enhance solidar-
ity any more than sharing rituals, dress, values, speech, experiences, hard-
ship, or anything else. The cohesive effect of humor may simply be a special
case of the general tendency of similarity to foster solidarity. Why postulate
the evolution of a special behavioral capacity to accomplish this fitness bene-
fit? Moreover, this explanation does not easily account for word play, tick-
ling of children as young as four months, and the incongruity element.
The observation that humor can increase solidarity is consistent with
the present theory. If humorous exchanges constitute a form of shared expe-
rience and mutual benefit, then solidarity would increase. This might be
especially true for humor directed at outsiders, since the presence of a com-
mon adversary can increase group cohesion. Recall Eibl-Eibesfeldt’s (1989)
comment about the acoustic resemblance of laughter to mobbing calls.
158 G. E. Weisfeld

On the other hand, humor that disparaged the listener or the in-group
would be expected to be divisive. These effects, since they result in both
increases and decreases in solidarity, are unlikely to reflect an evolved func-
tion, but rather are probably side-effects of the phenomenon of humor.

Levity
McDougall (1923) explained humor as relieving us of the unnecessary burden
of feeling sympathy for the fool. But then what communicative purpose does
laughter serve, and why do children commonly laugh at play, and what of
word play?
We do seem to feel less sympathy for someone responsible for his own
ridiculous predicament. However, this tendency seems to be part of the set
of psychological mechanisms for deciding whom to help (Trivers 1971),
rather than having evolved for some independent purpose. Research has
shown, for example, that we feel sympathy toward people in direct propor-
tion to their need for help and our feeling of responsibility, and in inverse
proportion to the cost of helping and to their responsibility for their plight
(Lerner and Simmons 1966).
Humor may indeed be one means of reducing sympathy for those who
are unwise candidates for help, but this would seem to be an incidental effect
of certain forms of humor rather than its main adaptive function. Presumably,
sympathy for the fool could have been reduced without the necessity for the
evolution of humor. Indeed, often we feel little sympathy for someone even
though we do not ridicule him.
Then too, we may need to be somewhat detached (Bergson 1911) or
objective (Eysenck 1942) in order to enjoy humor; perhaps in many instances
this means not being too sympathetic toward the butt of the joke. Thus, lack
of sympathy may facilitate humor, rather than the reverse.

Salubriousness
Somewhat more generally, it has been proposed that humor reduces anxiety
or otherwise provides an escape from life’s unpleasantness (e.g., Berlyne
1960; Clapier-Valladon 1983; Morreaull 1983; Spencer 1860). One would ex-
pect from this theory that anxious individuals would appreciate humor more,
but in fact Blank et al. (1983) found an inverse relationship between these
variables. Anxiety apparently interfered with the appreciation of humor,
rather than being reduced by it.
Another problem with this explanation is that life’s unpleasantness must
be there for an adaptive reason. Nature would not allow psychic pain to be
easily dissipated just so that we felt better. Affects serve our biological needs,
not the other way around.
Other explanations also posit that humor is salubrious (e.g., Chafe 1987;
Freud 1905; Lefcourt and Martin 1986; Rothbart 1973). But all of these health
 Adaptive Value of Humor and Laughter 159

explanations leave unresolved the question of why laughter accompanies

any physiological demobilization that occurs. Why couldn’t we simply be
designed to relax when nothing urgent confronted us, or otherwise to be
healthy without humor and laughter (Alexander 1986)?
A related problem is that sympathetic arousal increases when the
punchline is delivered (Chapman 1976; Godkewitsch 1976) and in proportion
to the subject’s amusement (Langevin and Day 1972; McGhee 1983). One
would expect decreased arousal to accompany the consummatory response,
if the function of humor were relaxation. Similarly, infant chimpanzees that
heard recorded chimpanzee laughter exhibited heart rate acceleration (Bernt-
son et al. 1989).
Moreover, laughter itself consumes calories. Energy expenditure rises
by about 75% during vigorous laughter (McGhee 1983b), and infant chimpan-
zees showed increased movement in response to laughter (Berntson et al.
1989). It is true that muscle tension (Paskind 1932) and heart rate also de-
crease following the punch line-but not in proportion to the funniness of the
joke (Godkewitsch 1976), as expected if the function of humor is relaxation.
Research does suggest that people who enjoy humor tend to be healthier
and better adjusted (Bizi et al. 1988; Dillon and Totten 1989; Martin and
Lefcourt 1983; but see Porterfield 1987). Many famous physicians have en-
dorsed it, including Benjamin Rush, William Osler, and Albert Schweitzer
(Haig 1988). However, any correlation between humor and health can be
explained in other ways. Healthier people may be happier and less anxious
and hence in a better humor. As mentioned above, anxiety has been shown
to detract from the appreciation of humor (Blank et al. 1983; Levine and
Abelson 1959); inadequately mothered infants, who tend to be anxious, may
seldom or never laugh (Bowlby 1965; Mehler 1961; Provence and Ritvo 1961;
Wolff 1969).
On the other hand, in one of the few studies in which humor was intro-
duced as an independent variable, it reduced fear in hospitalized children
(D’Antonio 1988). Similarly, Dworkin and Efran (1967) reported that expo-
sure to humor reduced anxiety and anger.
Humor may indeed reduce anxiety under certain conditions, and may
often be therapeutic. But it is questionable that the capacity for humor origi-
nally evolved for this purpose. Humor may have evolved for some other
reason but then have come to enhance fitness in some additional ways. For
example, humor can enhance the problem solving efficiency of groups (Good-
childs and Smith 1964)-possibly because it increased feelings of cooperation
through its reciprocal altruism element (P. K. Smith, personal communica-
tion 1992).

Communication of Pleasure
Some theorists (e.g., Hayworth 1928; Lorenz 1963) propose that laughter
communicates safety to other group members. But it would seem more ad-
160 G. E. Weisfeld

vantageous and parsimonious to be equipped with warning signals, the ab-

sence of which communicated safety, as is the case in many species.
Haig (1988) has suggested, more generally, that laughter informs others
of our pleasure. However, the adaptive advantage to the individual of so
doing is left unspecified. It is easier to see why the individual might benefit
from communicating distress, as through crying, which summons aid. We
laugh when we are happy, but this constitutes a proximate causal analysis,
not an ultimate one.

Courtship
Grammer and Eibl-Eibesfeldt (in press) have proposed that humor sometimes
functions in courtship. They recorded the spontaneous conversations of Ger-
man young adults who were initially strangers to each other. Men laughed
as often as women in same-sex dyads. Women laughed more than men in
mixed-sex dyads, perhaps because of greater nervousness. Still, women
laughed less often when conversing with a man than with a woman.
Moreover, the more intense the woman’s laughter (vocalized vs nonvo-
calized), the greater her self-reported interest in seeing the man again. Men’s
own laughter did not indicate interest in the woman, however. Men’s interest
was indicated by the intensity of the woman’s laughter; her vocalized laugh-
ter seems to have piqued his interest, whereas nonvocalized laughter sup-
pressed it.
These researchers suggested that a woman’s laughter can function to
communicate interest in a man. They attributed this sex difference to the
submission aspect of “nervous” laughter; that is, the woman signals her
submission to the man.
An alternative explanation is that a man tries to amuse a woman by
joking because if the attempt fails, he can save face by blaming her poor
sense of humor (Walle 1976). Walle observed that men in a bar courted
women by means of risque humor. A woman who was interested in the man
laughed. A woman who was not interested ignored the joke, which could be
rationalized as her having been offended rather than as her having rejected
him.
Grammer and Eibl-Eibesfeldt also found that the more the couple
laughed simultaneously, the greater their interest in each other. This was
explained as a social bonding behavior that serves to solidify the relationship
once mutual interest has been established. This explanation may have some
validity. However, the hearty, vocalized laughter of social bonding would
seem to be incompatible with an explanation in terms of nervous laughter.
With regard to the theory presented here, perhaps the man auditions
for the woman’s interest by trying to amuse her. If he succeeds, she expresses
her appreciation by laughing.
Grammer and Eibl-Eibesfeldt’s research is especially instructive be-
 Adaptive Value of Humor and Laughter 161

cause it involves naturalistic observation of spontaneous humorous behav-

ior. Subjects have difficulty making jokes in laboratory studies, and they
laugh less heartily than in naturalistic studies (Chapman 1983).
Also, social context effects are important and need to be elucidated by
naturalistic research. Grammer and Eibl-Eibesfeldt’s research revealed that
a laughing person avoided eye contact, which the authors interpreted as
ambivalence, presumably about the relationship. Alternatively, perhaps eye
contact, which can connote threat as well as intimacy (Argyle and Dean
1965), means aggressive laughter, i.e., laughing “at” the other person, and
so is avoided in friendly interaction. Laughing men tended to look away
earlier than women, perhaps because men’s gaze may be more threatening
than women’s.
Another sex difference in humor is harder to explain in terms of court-
ship. Boys and men generate more humor than girls and women (Chapman
et al. 1980; Ziv 1984). Goldstein and McGhee (1972) demonstrated this effect
in Belgium, the U.S., and Hong Kong. This sex difference has been found
to emerge at age six, when joking first appears (McGhee 1979)-well before
puberty and the onset of mature sexual motivation.
However, this developmental pattern coincides with the stabilization of
dominance hierarchies (Omark et al. 1975); perhaps this sex difference re-
flects the greater dominance, or competitive, motivation of boys across cul-
tures (Omark et al. 1975). Males may be more highly motivated than females
to gain recognition by joking, and less afraid of the joke misfiring. Another
indication of males’ competitive use of humor is that men tend to prefer put-
down jokes more than women, who favor self-deprecating humor (Zillman
1977).
Boys also appear to compete with each other more than do girls to show
that they have gotten the joke (Chapman et al. 1980), which is also consistent
with greater competitiveness among males. Similarly, repartee, a highly com-
petitive form of humor, is especially common in adolescent boys and men
cross-culturally (Apte 1985). As indicated above and confirmed by Ziv (1984),
women seem to appreciate humor more than do men, but perhaps only in
mixed-sex dyads. Perhaps one gains credit for making someone laugh; being
less competitive, women may be more inclined to concede this credit to men
than the reverse.
In support of this dominance interpretation of joking, dominance among
peers and seeking attention from adults were found to predict “sense of
humor” later in childhood (McGhee 1980). Adult amateur humorists tended
to have been popular, outgoing, and self-confident as children (Ziv 1984).
Professional comics seem to be highly dominant (McGhee 1986). Ridicule
may also be considered a form of dominance, insofar as observational re-
search has shown that people tend to ridicule those of lower status than
themselves (Coser 1960).
162 G. E. Weisfeld

CONCLUSIONS

It seems desirable to identify a single original evolutionary function for

humor. Not only would such an explanation be parsimonious, it also would
be in keeping with most evolutionary explanations of particular emotions
(Darwin 1872; McDougall 1923; Izard 1977; Pugh 1977). According to Haig
(1988), no such unitary explanation has been proposed that applies to all the
major examples of humor, including tickling, humor in apes, agonistic play,
ridicule, and word play.
The tentative explanation proposed here is that humor evolved to induce
the subject to seek out informative social stimulation and to reward others
for providing such stimulation. It may apply to all of these types of humor.
Moreover, it attempts to distinguish derivative effects of humor, such as
competition, group solidarity, courtship, and relaxation, from the fundamen-
tal adaptive value of humor.
The explanation is intended to be consistent with modern evolutionary
theory. It does not rest on assumptions of tension release or pleasure seeking,
avoids group selection reasoning, addresses the fitness benefits of the humor-
ist as well as the laugher, accounts for the fact that laughter is a means of
social influence, and applies to chimpanzees as well as humans. Being an
ethological theory, it recognizes the motivational and affective aspects of
humor, and not just its cognitive properties.
Lastly, the explanation seems reasonably consistent with the available
psychological and anthropological research on humor. For example, it helps
explain why sexual, aggressive, and competitive content is especially funny;
why a playful mood is essential for appreciating humor; why intelligent,
socially competent children (Masten 1986) and adults (Levine and Zigler
1976; Turner 1980) tend to make good humorists; and why incongruity is
humorous. It also accounts for the distribution of anatomical structures tar-
geted in tickling, and the pleasure that humor (except for ridicule) brings to
both humorist and audience.
Other emotions, sometimes referred to as the esthetic emotions, resem-
ble humor in being maximally aroused by stimuli that are salient to fitness
considerations and are moderately complex for the subject. It appears that
in our species, unusual for its great dependence upon learning and cognition,
several distinct emotions evolved to attract us to different forms of stimuli,
of suitable novelty and complexity, from which we can learn. That is to say,
the explanation for humor proposed here is similar to biological explanations
for other esthetic emotions (e.g., Pugh 1977).
One drawback of this explanation may be that it is not very different
from the leading evolutionary explanation for the emotion of interest, namely
that of motivating practice in skills that enhance fitness. As mentioned above,
however, there are some differences between humor and interest. The latter
need not be social, whereas humor almost always is, being accompanied by
a distinctive social display. Humor is usually briefer than interest. Also, we
 Adaptive Value of Humor and Laughter 163

are interested in fitness-salient aversive stimuli as well as attractive ones;

we cannot afford to ignore anything that can kill us. Funny things, on the
other hand, are always fun.
Arriving at a plausible evolutionary explanation for a given emotion is
only one step toward understanding that emotion. As Tinbergen (1951)
stressed, complete understanding of a behavior also requires that we describe
its development and its proximate causation. The development and causation
of humor-what makes something funny-remain quite elusive and merit
further research. Because all of these facets of a behavior are interrelated,
what we learn about one of them should aid our comprehension of the others.
Future work on the function of humor therefore may help our comprehension
of the whole phenomenon.

Jeffrey Meldman contributed over many years to the ideas in this paper. Thanks also to Larry
Stettner, Kevin MacDonald, Peter K. Smith, John Tooby, Robin Wright, Richard Alexander,
Irenaus Eibl-Eibesfeldt, and the anonymous reviewers for their helpful comments.

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