Humor and Laughter

JYOTSNA VAID
Texas A & M University

I. Defining Humor seriousness, or even danger, in a situation by deflecting

II. Humor and Laughter its adverse effects and by offering an alternative to
III. Ontogeny of Laughter despair. Humor can provide a way for individuals to
IV. Phylogeny of Laughter
conceal or to reveal their vulnerabilities; it can also
enable them to transcend a predicament. A person
V. Preconditions for the Emergence of Humor
with a good sense of humor is viewed more favorably
VI. What Makes Something Humorous? than one who is humorless. Humor is said to confer
VII. Functional Theories of Laughter and Humor mental flexibility, openness to experience, playfulness,
VIII. Normal Laughter and maturity and, as such, is highly valued whether in a
IX. Disorders of Laughter coworker, a leader, a friend, or a prospective mate.
X. Conclusion
Indeed, the ability to be playful and humorous appears
to be universally valued.
Although scholarly interest in the nature and
functions of humor has a long history, empirical
inquiry into the forms, uses, and biological bases of
GLOSSARY humor and laughter is fairly recent. This article
Duchenne display A characteristic facial display accompanying addresses both well-researched and relatively unex-
laughter in humans involving the joint contraction of lip and eye plored questions pertaining to humor and laughter.
muscles. The following are some of the questions that are
gelastic seizure Typically unprovoked, stereotyped, and inap- considered here: What is humor? What is the role of
propriate laughter accompanying an epileptic seizure. laughter in humor? What are the ontogenetic and
phylogenetic origins of laughter? How does humor
work, cognitively? What is the functional significance
The ability to perceive and produce humor, highly valued of humor? What are the varieties of laughter and how
across human societies, has been the subject of much are they perceived? What kinds of disorders of laughter
philosophical interest but little sustained, empirical have been documented and what do they suggest about
study. This article summarizes the variety of theore- the neural circuitry underlying laughter and humor?
tical perspectives and available research relevant to the
study of humor, including its nature, its origins in
ontogeny and phylogeny, its behavioral manifestation
in laughter, its psychological and evolutionary signifi- I. DEFINING HUMOR
cance, and its hypothesized neural mediation. Al-
though responding with humor may appear to be a Humor normally requires at least two individuals (the
way of evading a serious problem, it might also turn humor initiator and the receiver) who cooperate in
out to be a rather effective way of counteracting the setting up what Gregory Bateson termed a play frame,

Encyclopedia of the Human Brain Copyright 2002, Elsevier Science (USA).

Volume 2 505 All rights reserved.
506 HUMOR AND LAUGHTER

in which the parties involved tacitly agree that what is whereas crying emerges at birth and smiling at
inside the frame is not to be taken seriously. A play approximately 2 or 3 weeks of age, the characteristic
frame may also be set up by a single individual in expiratory movement of laughter does not appear until
response to a perceived incongruity in an event or approximately 4–6 months. However, cases of so-
situation. Verbal humor is a particular form of skilled called gelastic (or laughing) seizures in neonates
language use in which at least two disparate meanings indicate that neural and physiological structures
are interwoven into a text by making use of ambiguity, subserving laughter are in place at birth. Laughter’s
polysemy, intertextuality, or inconsistency in such a innateness is further suggested by the fact that it is
way that the listener is led to expect one meaning but observed in deaf–blind children, even those who could
actually experiences the other. The pleasure of humor not have learned about it by touching people’s faces.
is thought to arise upon the sudden recognition of the Although laughter initially occurs involuntarily, whe-
mismatch between the expected and the experienced ther in response to tickling or peek-a-boo games or as a
meaning. reaction to a sudden change in the sensory environ-
ment, such as an unexpected noise, over the course of
development laughter becomes more regulated, under
II. HUMOR AND LAUGHTER voluntary control, and elicited in response to cognitive
and social stimuli rather than physical stimuli per se.
In his 1872 monograph, The Expression of the
Emotions in Man and Animals, Charles Darwin noted
that ‘‘Joy, when intense, leads to various purposeless
movementsFdancing about, clapping the hands, IV. PHYLOGENY OF LAUGHTER
stamping, etc., and to loud laughter.’’ Laughter is
indeed commonly seen as an expression of joy, Laughter is estimated to be 7 million years old. Like
happiness, or amusement. It typically occurs in other vocalizations, such as moaning, sighing, and
informal social situations, usually in the presence of crying, laughing is thought to have preceded speech
a close friend, sibling, caregiver, or intimate. and, like these vocalizations, it may also have a
Although laughter is normally taken to be an communicative function.
indicator that the laugher is in a happy emotional Although laughter is often claimed to be unique to
state, it does not exclusively or necessarily signal such a humans, behaviors similar to human laughter have
state. One can clearly find something amusing without been observed in other primates and something akin to
it actually making one laugh out loud. Similarly, one laughter has even been argued to be present in rodents.
can feign laughter even when one is not actually In 1997, Signe Preuschoft and Jan van Hooff examined
amused. Other emotional states that give rise to variations in the contexts and social functions of
laughter include scorn, embarassment, and nervous- bonding displays in various primate species, including
ness. It remains to be determined whether the laughter Old World primates, macaques, baboons, great apes,
in these different states is morphologically different. and humans. They noted a silent bared-teeth display
Under certain clinical conditions, laughter can be (also called a grin, grimace, or smile) associated with
triggered in adults without their ability to control it. In inhibited locomotion, evasive and protective body
such cases it occurs for no apparent reason and usually movements and postures, and grooming and em-
without accompanying positive affect. The occurrence braces. This display was observed in all macaque and
of such dissociations between the motoric act of baboon species and humans. A relaxed, open-mouth
laughter and associated affective or cognitive states display (‘‘play face’’), marked by a widely opened
raises the possibility that laughter is under the control mouth but without pronounced baring of the teeth,
of a variety of different neural structures and systems was observed in all macaque and baboon species, in
at different levels and that these may be selectively each of the great apes, including the chimpanzee,
disrupted in pathology. bonobo, orangutan, and gorilla, in humans, and in
more distant species such as vervets and squirrel
monkeys.
III. ONTOGENY OF LAUGHTER One variant of the play face noted in some species
was an open mouth, bared-teeth display with associ-
Laughter and crying both appear to be innate mechan- ated staccato breathing and bursts of vocalization.
isms in humans, although laughter’s onset occurs later: This latter display, which Preuschoft and van Hoof
 HUMOR AND LAUGHTER 507

termed the ‘‘laugh face,’’ appeared strikingly similar to VI. WHAT MAKES SOMETHING HUMOROUS?
human laughter. It was accompanied by boisterous
body postures, brusque movements, mock biting, Speculating about the causes of mirthful laughter in
playful chasing, and evasive, repetitive glancing move- human adults, Darwin noted that ‘‘[s]omething incon-
ments rather than a tense gaze. The laugh face was gruous or unaccountable, exciting surprise and some
found only in certain macaques, mandrills, geladas, the sense of superiority in the laugher, who must be in a
orangutan, the bonobo, and, of course, humans, and it happy frame of mind, seems to be the commonest
occurred primarily in social play contexts, although in cause.’’ In comparing laughter arising from actual
some cases it was observed to accompany solitary play, tickling and that arising from the tickling of the
but in these cases it occurred only when conspecifics imagination, he noted:
were near. The presence of an audience thus appears to
From the fact that a child can hardly tickle itself,
be at least facilitative, if not necessary, for releasing the
or in a much less degree than when tickled by
laugh face display.
another person, it seems that the precise point to be
In 1987, Preuschoft and van Hooff proposed a
touched must not be known; so with the mind,
functional significance of play that is relevant to the
something unexpectedFa novel or incongruous
current analysis. They noted that in bonding behaviors
idea which breaks through an habitual train of
(i.e., those involving care, sex, and affiliation), it is
thoughtFappears to be a strong element in the
important to deemphasize competition and focus on
ludicrous.
shared interests. Play is one form of bonding behavior.
An important element of play is the partner’s un- Darwin’s insights anticipate many subsequent the-
expected performance of an expected behavioral act. oretical accounts of humor. With respect to its
Preuschoft and van Hooff suggested that the ‘‘essence cognitive basis, an early formulation of the mechan-
of play seems to be to actively bring about incongruous isms underlying humor may be found in the work of a
and unexpected behaviors and to interpret them as Gestalt psychologist, Norman Maier. Maier noted
nonthreatening.’’ This, they argued, allows for affec- that a key element of humor is a sudden and
tive and cognitive mastery of incongruous situations. unexpected restructuring of the elements of a config-
Whereas laughter in nonhuman primate species is uration, not unlike that experienced during a flash of
found principally in play contexts and primarily occurs insight. A humorous narrative manipulates our ex-
in the young of the species, laughter in humans occurs pectations by leading us down a garden path only to
in a variety of social contexts throughout the life span. present us with an altogether different conclusion than
Moreover, there is no evidence that nonhuman the one we were led to expect. Inasmuch as the
primates have anything comparable to the human conclusion disrupts the way in which we have been
ability to produce, comprehend, and enjoy humor. thinking about the events in the narrative, we are
totally unprepared for it. After a momentary confu-
sion of thought, we experience the newly restructured
configuration with clarity; the amusement arises when
V. PRECONDITIONS FOR THE EMERGENCE we realize how we were misled. In 1992, Wyer and
OF HUMOR Collins proposed that diminishment is a key element in
humor elicitation: For humor to be elicited, the new
Various factors have been proposed as preconditions perception of the situation must in some sense be
for the emergence of humor perception in a species, diminished in importance in comparison to the
including sociality, an exploratory drive, a system of apparent reality that was first assumed.
communication (to allow for the members of the Subsequent theorists have elaborated on these
species to express and communicate beliefs and accounts. For example, cognitive approaches charac-
feelings), an ability to mentally represent and order terize humor perception in information processing
the physical world, and an ability to construct possible terms as involving at least two and possibly three
imaginary worlds. Additional criteria proposed in- stages: a setup stage, a stage in which the incongruity is
clude a capacity to use logic and reasoning indepen- recognized, and a stage of incongruity resolution.
dently of affective or real-world constraints, a capacity Although there is debate regarding whether humor
to deceive or misrepresent the truth, a capacity for appreciation requires that the incongruity be satisfac-
collaboration, reciprocity, compassion, and a theory torily resolved (with some arguing that incongruity
of mind. recognition per se is sufficient to experience humor),
508 HUMOR AND LAUGHTER

theorists generally agree on the importance of the of humor with aggressive or sexual impulses has
juxtaposition of two or more mental representations characterized subsequent accounts as well, although
for humor to be perceived, following Arthur Koestler, there is no current consensus as to whether hostility is
who, in The Act of Creation, described humor as an inherent in humor.
example of bisociative thinking, which juxtaposes and Proponents of the so-called superiority theory (e.g.,
brings together two disparate matrices of thought. Aristotle, Plato, Hobbes, and Bergson) view laughter
Recent theoretical formulations in cognitive science as reflecting a moral stance on the part of the laugher.
have used concepts such as frame shifting and For Bergson, laughter asserts the human values of
conceptual integration of mental spaces to formalize spontaneity and freedom and therefore erupts when-
the processes underlying humor perception. To date, ever a person behaves rigidly, like an automaton:
there has been little cognitively oriented experimental ‘‘Humor consists in perceiving something mechanical
research on humor comprehension and even less on the encrusted on something living.’’ Hobbes described the
processes underlying humor generation. passion of laughter as ‘‘nothing else but sudden glory
It has also been acknowledged that it is not enough arising from a sudden conception of some eminency in
to postulate incongruity as a prerequisite of humor ourselves by comparison with the infirmity of others,
because, in certain situations, incongruity may give rise or with our own formerly.’’ Superiority is asserted not
to other emotional reactions, such as fear or appre- just by laughing at others’ defects but also by showing
hension, rather than humor. A humorous rather than that one can laugh at (and rise above) one’s own
apprehensive reaction is more likely when the receiver imperfections.
does not have too much invested in the content of the In contrast to negative uses of humor, as in put-
humorFthat is, when the humor does not threaten the down or derisive humor, recent clinically based
receiver’s beliefs or feelings in any profound or research has directed considerable attention to positive
disturbing way. Second, a humorous reaction is more aspects of humor. Humor is increasingly viewed as a
likely when the receiver was led to expect something useful mechanism for coping with stress and regulating
dangerous that turns out not to be. Indeed, Rama- affect. In 1999, Galloway and Cropley proposed that
chandran suggests that laughter may have evolved as a laughter may reduce some existing mental health
false alarm signalFthat is, as an immediate signal to problems and a sense of humor may moderate the
conspecifics that some potentially threatening event perceived intensity of negative life events.
has trivial rather than terrifying implications. This
notion is similar to the diminishment view of humor
developed by Wyer and Collins.
B. Social/Anthropological

Social and ethological theorists (e.g., Martineau) have

VII. FUNCTIONAL THEORIES OF LAUGHTER
viewed laughter as a marker of group membership and
AND HUMOR solidarity (for those who share a joke) or exclusion (for
those who are the ‘‘butt’’ of the joke) and as a means of
Theories about the uses of laughter and humor range
maintaining social control and group cohesiveness.
from psychological and social/anthropological ap-
Anthropologists have described ‘‘joking relation-
proaches to metaphysical and evolutionary ones.
ships’’ in various traditional societies; these refer to
ritualized teasing between specific kin in which the
receiver is required to take no offense. In an incisive
A. Psychological analysis, anthropologist Mary Douglas suggested that
joking is a subversive act in that it levels hierarchy and
Among psychological theories, that of Sigmund represents a triumph of intimacy over formality and of
Freud, as described in Jokes and Their Relation to the unofficial values over official ones. Not surprisingly,
Unconscious (1963), is one of the earliest. For Freud, this subversive aspect of humor is particularly evident
jokes allow a temporary expression of socially unde- among members of minority groups who may use
sirable impulses from the subconscious. Freud notes humor to portray reality in a way that exposes social
that tendentious humor makes possible ‘‘the satisfac- inequities. Much feminist humor and political humor,
tion of an instinct (whether lustful or hostile) in the face particularly that found in Central and Eastern Eur-
of an obstacle that stands in its way.’’ The association opean countries while under authoritarian rule, is of
 HUMOR AND LAUGHTER 509

this type. So-called ‘‘gallows’’ humor also has elements counterproductive; that is, things that our usual form
of this subversive aspect of humor. of reasoning might lead us into but that, in a larger
sense, would be undesirable. This view fits with a
related notion of humor as a device for pointing out
C. Metaphysical and sharing counterexamples, or providing disconfir-
matory evidence, which in turn is useful in reasoning
Although laughter appears to be associated with and problem solving. Chafe further suggests that the
happiness and joy, some have taken the position that fact that laughter provides an audible signal to others
laughter arises precisely from the experience of suffer- may be significant because laughter signals to the
ing. The philosopher Nietzsche, for example, argued receiver that the laugher is in a humor state and thus
that humor was invented precisely because of the cannot be taken seriously; moreover, the laughter may
extent to which humans suffer. This view is echoed in in turn infect others present, rendering them tempora-
existentialist philosophy. For the most part, though, rily disabled as well.
humor has not been taken seriously in classical The notion that humor is a disabling device that is
philosophical thought in the Western tradition, which ultimately adaptive is an intriguing hypothesis and one
has tended to regard it as irrational, irresponsible, and that readily lends itself to empirical test. For example,
frivolous. In contrast, Eastern traditions, such as one could ask whether the effect attributed to humor
Hindu, Buddhist, and Sufi worldviews, regard humor depends on the type of humor, the type of problem
as an appropriate reaction in the face of the vicissitudes from which the laugher has been disrupted, the
of death, disease, and aging; humor here is seen as an relevance of the humor content to the problem
insight that all human aspirations are ultimately content, the presence of others, etc.
comical and that wisdom ensues from that insight.
2. Humor as Social Learning
According to G. Weisfeld, humor provides exposure to
D. Evolutionary fitness-relevant scenarios in a nonserious, playful
context, motivating the practice of social skills that
Laughter is clearly an important aspect of human will be useful in serious contexts. In this view, humor is
nature. All societies appear to value it. It has distinct basically a form of social–intellectual stimulation that
facial and vocal manifestations. It emerges sponta- allows for the seeking and practicing of social skills
neously in early childhood and persists through the life that are fitness enhancing at some later point. Laugh-
span and it is an intensely pleasurable social activity. ter, according to Weisfeld, conveys appreciation and
All these aspects suggest that laughter may be a gratitude to the humorist for having provided such
reasonable candidate for a psychologically adaptive stimulation.
characteristic. Indeed, it has been accorded a special
place in several recent theoretical accounts of human
evolution. Aside from the false alarm theory men- 3. Humor as Status Manipulation
tioned earlier, at least five different evolutionary The notion that mirthful laughter serves to create and
accounts of laughter and humor may be distinguished. solidify group boundaries forms the centerpiece of an
evolutionary account of humor proposed by Richard
Alexander in which humor is seen as a way of
1. Humor as a Temporary Disabling Mechanism
favorably manipulating one’s status in a group to
When we are truly amused by something, more often improve one’s access to resources for reproductive
than not it takes us by surprise and distracts us from success. Specifically, it is proposed that humor devel-
whatever it was that we were in the middle of doing or oped as a type of ostracism, a means of manipulating
thinking. If we are heartily laughing, we quite literally one’s social status in a group (and thereby one’s access
cannot think of, let alone do, anything else. Wallace to critical resources) by facilitating bonds with certain
Chafe suggests that this disruptive effect of humor members and ostracizing others, through explicit or
may be evolutionarily significant (i.e., that humor’s implicit exclusion.
basic adaptive function is a disabling one). Chafe Alexander’s theory views humor primarily as a
proposes that the humor state may have arisen precisely device for establishing dominance and thereby for
in order to keep us from doing things that might be being more competitive for critical resources. He
510 HUMOR AND LAUGHTER

suggests that humor, as a form of social scenario our attraction to novelty. Creativity not only allows
building, gives one an edge in learning how to one to find unpredictable solutions to problems but
negotiate in fitness-relevant domains. Humorous put- also provides inherent pleasure because of its protean,
downs may be more successful than direct criticism or unpredictable nature. Humor encapsulates these ele-
insults as a way of establishing dominance since they ments of creativity and it is therefore not surprising
are indirect and therefore more face-saving ways of that a sense of humor is highly desired in a prospective
manipulating status than direct displays of hostility or mate. In this view, humor evolved because of its
strength. In Alexander’s model, all humor, whether or importance in courtship and mate choice.
not it has an ostensible butt or target, essentially The five evolutionary hypotheses emphasize differ-
developed in the service of status manipulation. ent elements of humor. Humor is hypothesized to
disrupt our routine for the better, to teach and reward
us in our social interactions in fitness-relevant do-
4. Humor as Vocal Grooming
mains, to unify a group but also create group divisions,
An emphasis on vocal grooming characterizes an to create and strengthen affective bonds, and to attract
evolutionary view of language proposed by Robin and sustain a partner. Although there is evidence
Dunbar that has implications for humor. Dunbar consistent with each of these views, research designed
speculates that human language evolved as a vocal to examine the assumptions and claims of the different
extension of physical grooming, allowing, as it were, accounts is clearly warranted.
the simultaneous grooming across space of more than In the remaining sections we review theory and
one partner and thereby the facilitating of social bonds research on the expression and neural mediation of
with larger groups of animals. Language fosters social normal and abnormal manifestations of laughter and
bonding by permitting the exchange of gossip, i.e., humor.
socially relevant information (specifically, who is
doing what with whom), as well as the sharing of
one’s own experiences, actual or desired. The fact that VIII. NORMAL LAUGHTER
laughter typically accompanies speech suggests that it
facilitates and cements the social bonding process. A. Visuomotoric and Physiological Aspects
In this view, the pleasure in shared laughter is
analogous to the pleasure of actual physical touch. Laughter is a motoric and vocal activity requiring the
Humor has in fact been characterized as a form of coordinated action of 15 different facial muscles and
vicarious touch. The importance of touch for primate associated clonic contractions of the thoracic cage and
emotional development is well documented. Laughter the abdominal wall. Laughter produces spasmodic
may have served to attract like-minded others in the skeletal muscle contractions, tachycardia, changes in
bonding process; as Konrad Lorenz noted, finding the breathing pattern, and increases in catecholamine
same thing funny is not only a prerequisite to a real production. Hearty laughter increases heart rate,
friendship but also very often the first step to its blood pressure and respiratory rate, and muscular
formation. Thus, just as language facilitates social activity.
bonding, laughter accompanying speech, and humor- Physiologically, laughter is the opposite of crying.
ous discourse with or without laughter, could have Although the upper half of the laughing face appears
evolved as a way of solidifying affectional bonds. indistinguishable from the crying face, the lower half
and the respiratory pattern in laughter are the reverse
of those of crying. In laughter a characteristic facial
5. ‘‘The Wit to Woo’’: Humor as Mate Attraction
display known as the Duchenne display is invoked.
The most recent evolutionary proposal regarding This display involves the joint contraction of the
humor emphasizes its importance in courtship and zygomatic major muscles (i.e., pulling the lip corners
suggests that the creativity and unpredictability that is back and upwards) and the orbicularis oculi muscles
inherent in humor is seen as attractive to a prospective (i.e., raising the cheeks and causing the eyes to
mate. In this account, proposed by Geoffrey Miller in wrinkle). Indeed, the movement of the eye muscles
The Mating Mind, creativity is seen as not just a may serve as a marker for distinguishing emotional
random by-product of chaotic neural activity but also from voluntary (forced) laughter since the orbicularis
as something that evolved as an indicator of intelli- oculi muscles do not contract during voluntary laugh-
gence and youthfulness and as a way of playing into ter. Laughter is also accompanied by the flaring of
 HUMOR AND LAUGHTER 511

nostrils, mandible retraction, and brightening and pulses in a laugh cycle, although the number can range
sparkling of the eyes. from 9 to 12 depending on lung volume. There are
In terms of respiration, laughter typically begins usually about 5 pulses per second. At the level of
with an abrupt exhalation followed by rhythmic laryngeal movements, a laugh pulse can be further
expiration/inspiration cycles, which may or may not subdivided into the number and duration of vibratory
be phonated as ‘‘ha ha ha.’’ No inspiration preceding cycles of the vocal cords. In one analysis, the duration
the laugh is needed since laughter is produced at a low of laughter pulses varied from 30 to 100 msec.
lung volume. The rhythmic pattern of laughter re- Phonation in laughter pulses involves a series of
spiration is produced by contractions of muscles that stereotypic laryngeal adjustments that include four
are typically passive during normal expiration, i.e., the stages: an interpulse pause (a moment of quiet
diaphragm, the abdominal (rectus abdominus), and aspiration in the periods between voicing), adduction
the rib cage muscles (triangularis sterni). These (closing) of the arytenoid cartilages, vibration of vocal
muscles work together with the larynx. In crying, the chords, and abduction (opening) of the arytenoid
saccadic contractions occur mainly with inspiration, cartilages.
whereas in laughing they occur with expiration; in both Several features in the acoustic signal may serve to
cases the contractions are accompanied by short, cue the degree of positive affect experienced by the
broken sounds. Autonomic correlates of laughter laugher. Widening versus narrowing of the pharynx is
include arousal and muscle tension followed by known to affect voice quality and may signal friendly
relaxation of muscle tone, dilatation of cutaneous versus scornful laughter, respectively. Other cues may
vessels of the face, neck, and hands, lacrimation, and be provided by harmonics, melodic contour, and
fatigue. duration. The melodic contour, including the intona-
In 2000, Niemitz, Loi, and Landerer examined tion and pitch contour, may be particularly informa-
specific visuomotoric aspects of the human laughing tive of emotional state and meaning. The vocal
face and their affective interpretation by human raters. ligaments are more likely to be tensed under conditions
The expressive facial movements of 45 videotaped of arousal or anticipation, giving rise to laughter that
laughing adults and 13 children were shown to more has a rising melodic contour. The lips and the cheeks
than 100 adult raters. The results suggested that visual are typically elevated in joyful smiling or laughter;
aspects of laughing faces that are judged as cheerful, contraction of the muscles involved in the Duchenne
winning, or generally positive were those in which (i) display changes the form of the mouth opening,
the laughs were fairly long, ranging from 3 to 6 sec; (ii) constraining the vowel sounds that can thus be
there were rapid eye and mouth movements during the produced. Indeed, one study showed that listeners
first second of the expression of laughter; (iii) there was can reliably infer a smiling from an unsmiling version
repetition of the mouth and eye movements; and (iv) of the same spoken message purely on the basis of the
there was almost complete closing of the eyelids, voice.
sometimes repeatedly, for 0.1–1.5 sec. There is a need Fewer than a dozen studies have been conducted on
for more such research quantifying how the various the acoustic characteristics of laughter. Almost all
laughing expressions are decoded and interpreted, not have used adult participants. One study examined the
just in the visual channel but also acoustically. acoustic characteristics of a group of 30 male and
female adults from whom one laugh was elicited under
each of four conditions: social, humor, tension release,
B. Acoustic Aspects and tickle. Significant differences in duration, inten-
sity, and mean frequencies of three vowel formants
Willibald Ruch and Paul Ekman proposed a useful above the fundamental frequency were found between
framework for describing the structure of laughter. the humor and social laughs and the tension release
They define a laughter bout as all the respiratory, and tickle laughs. Moreover, listeners could reliably
vocal, facial, and skeletomuscular elements involved in classify the laughs into the different types, suggesting a
a particular laughter event. A bout of laughter, in turn, communicative aspect to the laughs. The mean funda-
consists of an onset (the prevocal facial component), mental frequency of laughter for males (175 Hz) was
an apex (involving vocalization or forced exhalation), higher than that for females (160 Hz) and highest for
and an offset (the postvocalization part, typically a the tickling condition. In another study, a group of 11
smile that fades out). The apex contains laugh cycles, male college students were recorded while they viewed
(i.e., repetitive laugh pulses); there are typically 4 a videotape of comedic storyteller Bill Cosby. Analyses
512 HUMOR AND LAUGHTER

of 55 bouts of laughter showed that the mean peak Martin, the benefits of humor may also extend to the
fundamental frequency while laughing was more than physiological level. For example, laughter is known to
twice as high as that when the subjects were counting. increase respiratory rate and clear mucus. It is entirely
Moreover, an extended range of frequencies was conceivable that laughter may thus be beneficial to
exhibited during laughter, with a difference of 344 Hz patients with chronic respiratory conditions such as
between the lowest and the highest range observed. In emphysema. The increased heart rate and blood
general, there was considerable variability both within pressure accompanying laughter can exercise the
and between subjects on all the measures studied. myocardium and improve arterial and venous circula-
A limitation of existing studies with adults is the lack tion, allowing a greater flow of oxygen and nutrients to
of naturalistic social interaction contexts. Studies with tissues. This in turn may facilitate the movement of
children fare better in this regard. One study recorded immune elements useful in fighting infections. Muscle
laughter among four 3-year-olds during three sessions relaxation following hearty laughter may break the
of spontaneous free play between mother and child. spasm–pain cycle in patients with neuralgias or
An acoustic analysis revealed that the duration of rheumatism. A reduction in laryngeal muscle tension
laughter syllables in children was about the same as accompanying laughter may help patients with vocal
that found in adult laughter (i.e., about 200–220 msec). fold pathology to produce a more relaxed voice, and it
The total duration of the average laugh was also similar may facilitate the recovery of phonation in patients
to that of adults. The main difference between adult with psychogenic dysphonia, an inability to phonate
and child laughter was in fundamental frequency, with during speech despite intact articulation and the
most children’s laughter having a frequency in the absence of any identifiable pathology of the larynx.
upper range of adult female laughter (400–500 Hz) or, Finally, the increased catecholamine levels associated
in the case of one type of laughter (squeals), even higher with laughter may be responsible for the beneficial
(nearly 2000 Hz). Four distinct laughter types were effects humor is thought to exert on mental functions,
noted in the child sample: (i) comment laughter (i.e., such as alertness and creativity. Many of these claims
laughter occurring in conversational contexts that of the healthful benefits of laughter and humor have
lasted about one-fifth of a second), which was sub- yet to be subjected to systematic study.
divided into dull comment and exclamatory comment
laughter; (ii) chuckle laughter, which lasted half a
second and tended to occur in situations provoking D. Neuroanatomical Hypotheses
more excitement than comment laughter; (iii) rhyth-
mical laughter, which lasted 1–15 sec; and (iv) squeal There has been very little direct empirical investigation
laughter, which lasted half a second, had a very high of neuroanatomical correlates of normal laughter. A
fundamental frequency, and usually occurred without hypothetical neural circuit was first theorized in 1924
a break. The latter two kinds were more prevalent in the by Kinnier Wilson. According to Wilson’s model,
context of physical stimulation, anticipation, and fear. laughter is produced by a medullary effector center
It appears, therefore, that different types of sponta- that links the seventh nerve nucleus in the pons with the
neous laughter have different acoustic characteristics. 10th motor nucleus in the medulla and with phrenic
Moreover, laughter differs in different social contexts. nuclei in the upper cervical cord. This center is
Additional studies are needed with different kinds of modulated by the cerebral cortex and limbic structures
communication dyads and with different age groups to by means of an integrative center in the mesial
examine in more detail the relationship between thalamus, hypothalamus, and subthalamus.
acoustic distinctions in laughter production and their Activity of the laughter center is thought to be
affective interpretation by listeners. The specific ways determined by a voluntary pathway (corticobulbar
in which spontaneous versus contrived laughter may fibers) and fibers extending from the orbital surface of
differ, both structurally and neurobehaviorally, also the frontal lobes through the bulbar nuclei. Input to
warrant study. these fibers comes from an involuntary pathway (the
basal ganglia), which appears to be inhibited by the
voluntary one. No single cortical area supplies the
C. Health Aspects origins for the voluntary and involuntary fibers; the
input derives from diffuse cortical regions including
An implicit belief underlying most accounts of humor the frontal, premotor, motor, parietal, temporal, and
is that it confers psychological benefits. As reviewed by hippocampal regions.
 HUMOR AND LAUGHTER 513

The notion of hypothalamic integration of cortical Alzheimer’s disease. These patients are overcome by
and limbic control on the brain stem laughter center uncontrollable laughter and crying that is usually
remains to be substantiated, although it is consistent appropriate to the situation and accompanied by
with clinical observations. mood alteration.
Very few patients actually meet the criteria for PLC.
However, researchers do not concur on the ideal
IX. DISORDERS OF LAUGHTER classification system and whether PLC must involve a
dissociation between the expressed affect and the
Disorders of laughter and crying are very rare. mood. It is acknowledged that the degree of volitional
Although strong emotional reactions are not uncom- control of PLC varies from complete absence to some
mon following brain injury, the most common reaction control.
described is fear. Medical research on laughing and The most common conditions associated with PLC
crying disorders consists largely of case reports, with are summarized in the following sections, subdivided
few systematic, large-scale investigations. Abnormal into disregulatory and excitatory conditions.
laughter, defined as laughter that is involuntary and
inappropriate to the situational context, is most often
seen in generalized affective or cognitive disturbances A. Disregulatory Conditions
such as psychoses (e.g., schizophrenia). Individual
cases of hysterical laughter spells, marked by silly, One disregulatory condition giving rise to abnormal
unrestrained, unmotivated, and unprovoked laughter, laughter is amyotrophic lateral sclerosis (ALS). In
have been reported, as has epidemic hysterical 25% of ALS patients there is bulbar involvement; of
laughter. these, 30–50% develop PLC. Patients without bulbar
In many cases of abnormal laughter, the motor act involvement do not develop PLC.
of laughing may be dissociated from its emotional Multiple sclerosis is another condition that in 7–
aspect. Such a condition has been termed pathological 10% of patients leads to PLC, sometimes with
laughter. Kinnier Wilson defined pathological laugh- euphoria. Characteristics of so-called pseudobulbar
ter and crying (PLC) as ‘‘a sequel to and consequence palsy associated with multiple sclerosis (as with
of a recognizable cerebral lesion or lesions in which bilateral strokes) include dysarthria, dysphagia, bifa-
attacks of involuntary, irresistible laughing or crying, cial weakness, and weak tongue movements but
or both, have come into the foreground of the clinical preserved coughing, yawning, laughing, and crying.
picture.’’ This definition draws attention to the im- In one large-scale study, PLC was observed in 15%
paired control and the episodic aspect of the abnormal of stroke patients 1 month after the stroke, in 21% at 6
emotional expression. months after the stroke, and in 11% a year following
In 1994, Shaibani, Sabbagh, and Doody proposed the stroke. Abnormal emotional reactions were parti-
four criteria to distinguish pathological laughter and cularly associated with lesions in the left frontal and
crying from normal laughter and crying. First, PLC is temporal regions.
inappropriate to the situation since it occurs sponta- A rare syndrome of PLC is associated with acute
neously or in response to nonspecific stimuli or infarctionFthe so-called ‘‘fou rire prodromique,’’ first
inappropriate, arbitrary stimuli (e.g., one patient with described in 1903 and in only a few cases since. In these
a left cerellar hematoma showed pathological laughter cases, laughter lasts from between 15 minutes to 24
following left hand tremor). Second, PLC is unmoti- hours and is almost always followed by death. Lesions
vated; that is, there is no relation between the affect in this syndrome typically involve the left internal
and observed expression, nor is there relief or mood capsule-thalamus, left basal ganglia, or ventral pons.
change afterwards. Third, PLC is involuntary; that is, This syndrome contrasts with pseudobulbar palsy
it has its own pattern and occurs against the patient’s since it is not recurrent and, unlike epileptic laughter, it
will. Neither the duration nor the content of PLC can is not associated with electroencephalograph (EEG)
be controlled, and patients do not gradually change changes or confusion.
from smiling to laughing but have a sudden, brief A host of extrapyramidal disorders, including
outburst without any warning. Fourth, PLC differs Parkinson’s and Wilson’s disease, are also associated
from emotional lability, which refers to an exaggerated with PLC. The syndrome of Angelman, a genetic
emotional response to a normal stimulus. The latter disorder characterized by mental retardation and stiff
characterizes patients with multiple sclerosis and puppet-like movements in children between the ages of
514 HUMOR AND LAUGHTER

2 and 6 years, is also associated with frequent bursts of cingulate gyrus whose laughter appeared to be an
laughter. A variety of toxins, such as nitrous oxide and irrepressible motor behavior; resection of the lesion
insecticides, have also been related to pathological eliminated the laughing behavior. They suggest that
laughter. Finally, various malignant brain stem tu- the premotor mesial system acts as an interface
mors, such as clival chordoma and pontine glioma, between the limbic loop, which includes the anterior
have been implicated in pathological laughter. cingulate gyrus, and the motor loop, which includes
the supplementary motor area. Biraben et al. propose
that cases of gelastic seizure originating in the anterior
B. Excitatory Conditions cingulum involve a critical functional disconnection
between the motor loop and the limbic loop within the
The most common excitatory condition that is asso- mesial premotor system. The laughter arising in these
ciated with pathological laughter occurs in epileptics. cases reflects a behavioral output from a motor
Laughter as part of an epileptic seizure was documen- program separated from all motivation.
ted by the neurologist Trousseau in 1873 and was
Laughter in one other epileptic patient reviewed, a
described in Dostoevsky’s novel The Idiot. In 1957, the
nongelastic case, occurred after cortical electrical
term ‘‘gelastic epilepsy’’ was coined to refer to epileptic
stimulation of the lateral border of the rostral part of
fits in which laughter is the only or the most common
the supplementary motor area. The laughter was
symptom. More than 160 cases of gelastic epilepsy
natural, rather than forced, and was accompanied by
have been reported in the literature. Gelastic epilepsy
a general feeling of amusement toward the environ-
often occurs in patients with hypothalamic hamarto-
ment at large. Biraben et al. suggest that such laughter
mas and precocious puberty, in patients with complex
may reflect an imbalance between the mesial and
partial seizures and temporal lobe origin seizures, and
lateral premotor systems.
in children with infantile spasms. The seizures usually
begin in infancy or childhood and are associated with
cognitive decline in later years. In patients with these
seizures, involuntary, mechanical giggling typically C. Neuroanatomical Hypotheses
occurs as the initial ictal behavior before an alteration
in consciousness. The duration of the laughter in Evidence for the neuroanatomy of pathological laugh-
patients with hamartomas is usually less than 30 ter and crying comes from the following sources: a
seconds and the seizures occur several times a day. limited number of autopsy reports; studies of congeni-
In a recent review of published reports of seizures tally malformed infants; case reports of patients with
involving laughter, Biraben and colleagues proposed pathological laughter following neurological disease;
the following neural hypotheses for the genesis of EEG activity and electrical stimulation of the cortex;
laughter: injection of a barbiturate to the right or left cortex,
typically done in patients with epilepsy to determine
1. Laughter arising as a reactional behavior (i.e., in the language-dominant hemisphere; and studies of
response to a pleasant feeling or mirth: Only a few such humor comprehension in patients with left versus right
cases have been observed, all involving seizures with a hemisphere damage.
temporal focus. In one such case, involving stimulation The autopsy data are of limited value in localizing
of the left temporobasal region (fusiform and para- laughter since the patients had suffered diffuse brain
hippocampal gyri), the individual experienced a pathology. A review of such cases revealed no single
change in the semantic connotation of stimuli (things cortical lesion as causing PLC. Together with clinical
became funny); in another case with stimulation in the case reports, the autopsy data indicate that both
same region, the modification experienced was percep- unilateral and bilateral lesions that affect the descend-
tual (things changed in a funny way). Biraben et al. ing tracts to the bulbar nuclei can cause PLC, as can
suggest that the laughter in these cases might be a lesions on either side in the anterior limb and genu of
physiological response to a modified cognitive process. the internal capsule adjacent to the basal ganglia,
2. Laughter arising as a forced action or an thalamus, hypothalamus, and pons. Even localized
automatism: Seizures with a frontal focus have brain stem lesions can cause pathological laughter.
characteristically produced laughter described as Case studies do not explain why some patients develop
forced and unmotivated. Arroyo and colleagues pathological laughter while others develop pathologi-
describe one patient with a cavernoma of the anterior cal crying or both. Studies of congenitally malformed
 HUMOR AND LAUGHTER 515

newborns with severe anencephaly but with preserved review, Jaak Panksepp observed that incisive research
pons and medulla indicate that newborns do not smile in this area has only just begun and that substantive
but can cry; those with intact midbrains can both cry research remains meagre. Many basic questions re-
and smile. main, such as the occurrence and significance of
The predominant neuroanatomical account of pa- laughter in play and other social contexts; the role of
thological laughter and crying, first proposed by laughter and humor in courtship, mate choice, and
Wilson, regards it as arising from a loss of direct relationship maintenance; what different kinds of
motor cortical inhibition of a laughter and crying laughter communicate and to whom; and the neural
center located in the upper brain stem. However, this circuitry responsible for normal laughter and feelings
explanation does not fully account for the range of of mirth and joy.
phenomena observed in PLC. A recent alternative
account, developed by Parvizi and colleagues, in light
See Also the Following Articles
of new neuropathological findings, suggests that PLC
is caused by dysfunction in circuits that involve the CREATIVITY d DEPRESSION d EMOTION
cerebellum (as well as the cortex) and influence brain
stem nuclei.
There is evidence that the cerebral hemispheres may Suggested Reading
play different roles in the control of emotional states.
Biraben, A., Sartori, E., Taussing, D., Bernard, A., and Scarabin, J.
In normal subjects the right hemisphere appears to be
(1999). Gelastic seizures: Video-EEG and scintigraphic analysis
specialized for the perception and expression of of a case with a frontal focus; Review of the literature and
emotion, particularly negative emotion. Lesions in pathophysiological hypotheses. Epileptic Disorders 1(4), 221–
the right hemisphere have been found to impair 228.
prosodic and lexical expression of emotion. Moreover, Brownell, H., and Stringfellow, A. (2000). Cognitive perspectives on
humor comprehension after brain injury. In Neurobehavior of
in epileptic subjects, injection of the right hemisphere
Language and Cognition: Studies of Normal Aging and Brain
with intracarotid sodium amytal has elicited unpro- Damage (L. Connor and L. K. Obler, Eds.), Kluwer Academic,
voked laughter, whereas left-sided injection has tended Boston.
to produce bouts of crying. Patients with crying Galloway, G., and Cropley, A. (1999). Benefits of humor for mental
seizures tend to show right-sided foci, whereas those health: Empirical findings and directions for further research. Int.
J. Humor Res. 12(3), 301–314.
with gelastic seizures show mainly left-sided foci.
Goel, V., and Dolan, R. (2001). The functional anatomy of humor:
Patients with right hemisphere damage show a Segregating cognitive and affective components. Nature Neu-
preserved sensitivity to the surprise element of humor rosci. 4(3), 237–238.
but a diminished ability to establish narrative coher- Hull, R., and Vaid, J. (2001, July). Cognitive basis of incongruity in
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Barbara, CA.
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have been reported in patients with frontal lobe Methodological issues and research findings. Psychological Bull.
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Neurosci. 11(2), 253–258.
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X. CONCLUSION (2001). Pathological laughter and crying: A link to cerebellum.
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