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Laughter is often considered to be the product of humour. However, laughter is a social emotion, occurring most often in interactions, where it is associated with bonding, agreement, affection, and emotional regulation. Laughter is underpinned by complex neural systems, allowing it to be used flexibly. In humans and chimpanzees, social (voluntary) laughter is distinctly different from evoked (involuntary) laughter, a distinction which is also seen in brain imaging studies of laughter.
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The social life of laughter
Sophie Scott, Nadine Lavan, Sinead Chen, and Carolyn McGettigan
Abstract
Laughter is often considered to be the product of humour. However laughter is a social emotion,
occurring most often in interactions, where it is associated with bonding, agreement, affection and
emotional regulation. Laughter is underpinned by complex neural systems, allowing it to be used
flexibly: in humans and chimpanzees, social (voluntary) laughter is distinctly different from
evoked (involuntary) laughter, a distinction which is also seen in brain imaging studies of
laughter.
Keywords
Emotion; voice; conversation; laughter; motor control
When do we laugh, and why?
Human beings are immersed in laughter: it is a pervasive non-verbal expression of emotion
(1,2), which is universally recognised (3), and is more like a different way of breathing than
a way of speaking (4) (see Box One). Laughter is mediated strongly by social context: we
are 30 times more likely to laugh if we are with someone else than if we are on our own (1,
2), and we laugh most of we can see and hear someone (even if this is via a computer)
compared to voice or text interactions (5). Laughter is also highly behaviourally contagious,
and can occur when primed solely by another’s laughter (2). Observational studies have
shown that laughter is commonly found in conversations, where it occurs at the rate of
around 5 laughs per 10 minutes of conversation (6). Strikingly, this is a much higher rate
than the amount of laughter that people self-report (6). Furthermore, if we are asked about
when we laugh, we report that we laugh at jokes and humour (1, 2): however, observational
studies show that not only does most laughter occur in conversations, within those
conversations most laughter is associated with statements and comments, rather than jokes
(1, 2, 6). Furthermore, in conversation, the person who laughs most frequently is the person
who has just spoken, indicating that laughter is frequently not a reaction to someone else’s
utterance (6). Much conversational laughter, therefore, is an intentional, communicative act.
For example, conversational laughter in both spoken and signed conversations is often timed
to occur at the end of utterances (1): this commonality across modalities underlines the
voluntary aspect of much of laughter production, since (at least in theory) a person could
simultaneously sign and laugh if she wished to.
Consistent with this emphasis on the social roles of laughter, researchers have suggested that
there are two different kinds of laughter, and that these are differentiated by how they are
elicited: the laughter can be either driven by outside events (reactive, involuntary laughter)
or be associated with a more intentional communicative act - i.e. more controlled, deliberate
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Trends Cogn Sci. 2014 December ; 18(12): 618–620. doi:10.1016/j.tics.2014.09.002.
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laughter (7). This has been overtly compared to a distinction between spontaneous and
controlled smiling (7). Work with chimpanzees has revealed a similar distinction between
laughter which is generated in reaction to being tickled, versus laughter which is produced
during play (with the aim of making play last longer (8)).
Neural systems involved in the perception and production of laughter
The neural control of vocalization in humans is considered to involve two cortical systems
acting on midbrain and brainstem motor structures, where lateral premotor and motor sites
control the production of learned vocalizations such as speech and song, and a midline
system involving the anterior cingulate and supplementary motor area is associated with the
production of involuntary vocalizations, such as expressions of emotion (Jurgens, 2002).
Within this framework of voluntary and involuntary vocalizations, Wild et al. (2003) (10)
propose a model of laughter production, largely based on studies of pathological laughter, in
which they identify a coordinating centre for laughter in the brainstem comprising the
periaqueductal grey (PAG) and the upper reticular formation in the control of changes in
facial expressions, respiration, and vocalization. These structures are proposed to receive
excitatory inputs from cortical sites in the basal temporal and frontal lobes, as well as from
structures in the limbic system including the hypothalamus and basal ganglia. Here, the
lateral premotor cortices are implicated not in the basic production of laughter, but rather in
the suppression and regulation of spontaneous laughter vocalizations. In line with this view,
a recent neuroimaging study (11) found that the voluntary suppression of laughter during
tickling involved greater BOLD responses in lateral sensorimotor cortex compared with
when the participants were free to laugh when tickled (and when they voluntarily produced
laughter without tactile stimulation). Another key difference between voluntary and
(relatively) involuntary laughter in this study was a greater engagement of the hypothalamus
during tickling laughter (compared with voluntary or suppressed laughter), and an additional
correlation between activation in the PAG with the frequency of laughter episodes during
tickling. Both of these findings suggest a central involvement of these structures in the
production of basic, more automatic, expressions of positive experience.
Studies of the neural correlates of laughter perception have identified sensitivity to the social
significance of laughter signals. In a study of passive listening to emotional vocalizations
(12), sites on the lateral premotor and primary motor cortices showed a correlation with
valence, being greater to positive sounds (laughter and cheers of triumph) and lower for
negative sounds (screams of fear and expressions of disgust). This could reflect the
contagiousness of laughter and the greater tendency for positive vocalizations and associated
facial movements to occur in the context of social groups. However, more recent work (13)
showed that direct comparisons of the neural responses to voluntary and involuntary
laughter recordings gave no difference in motor cortex activation. Rather, the degree of
engagement of sensorimotor cortices when listening to both types of laughter was related to
individual differences in participants’ accuracy at classifying voluntary and involuntary
laughs and “posed” and “real”, respectively. This could be evidence that the listener engages
in some level of sensorimotor simulation as a mechanism for evaluating the social meaning
of heard vocalizations, rather than exhibiting a basic sound-to-action response. In the same
study, we observed preferential responses to involuntary laughter in bilateral auditory
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cortex, while responses to voluntary laughs were greater in ventromedial prefrontal cortex.
This profile of auditory cortex engagement suggests that, on the one hand, there are acoustic
hallmarks that automatically register the presence of intense, “real” laughter in the brain of
the listener. In contrast, the absence or reduced prominence of such cues in voluntary laughs
engages mentalizing strategies in medial prefrontal cortex, to support the interpretation of
why the laugh has been produced, and what it means. Figure 1 summarizes the key sites
identified in studies of the production and perception of laughter.
Taking laughter seriously
Laughter is more than a positive emotional expression: its social use may extend to the
management of affective states within interactions. Laughter is one of the positive emotional
expressions which is expressly linked to a physiological reduction in the stressful reactions
to negative emotions (e.g. fear, anger, disgust), in a way which may be more effective than
other ways of managing negative emotions (e.g. suppression) (14, 15). Positive emotional
expressions have been associated with the down-regulation of negative emotions in
conversations between couples (14, 15): couples who reported the highest levels of marital
satisfaction also showed the most ‘skillful’ use of positive affect (e.g. laughter) to regulate
negative emotions during a difficult conversation (15). Laughter is therefore not simply a
common emotional vocalization, which we use to establish and maintain social bonds (1,2):
laughter may also simultaneously function as an essential behaviour for helping to ‘de-
escalate’ negative emotional experience, with a positive role in both the short term affective
state of the interaction, and the longer term state of relationships. Understanding the
behavioural and neurobiological bases of laughter will mean more than thinking about jokes:
it could provide a vital link between human language, relationships and emotional states.
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Box One
Breathing, speaking and laughing
Upper panel shows oscillograms for speech and laughter, middle panel shows
spectrograms for laughter and speech, lower panel shows chest expansion dynamics for
metabolic breathing, laughing and speaking. The time scale on the x axis is the same for
each (breathing, laughing and speaking). Both speaking and laughing are distinctly
different from metabolic breathing, in terms of chest wall movements. Laughter is
characterized by very rapid contractions of the intercostal muscles, resulting in large
exhalations followed by individual bursts of laughter: the spectral modulation of laughter
by supra-larangeal structures is minimal. Speech shows a fine pattern of intercostal
muscle movements, which are used to maintain constant sub-glottal pressure at the larynx
and to provide pitch and rhythm to the speech. Unlike laughter, speech also shows
considerable spectral complexity reflecting movements of the supra-laryngeal
articulators.
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Figure 1. Voluntary and involuntary laughter in the brain
The coordination of human laughter involves periaqueductal gray and the reticular formation
with inputs from cortex, the basal ganglia and the hypothalamus [10]. The hypothalamus is
more active during reactive laughter than voluntary laughter [11]. Motor and premotor
cortices are involved in the inhibition of the brainstem laughter centres, and are more active
when suppressing laughter than when producing it [11]. Laughter perception involves
premotor cortex and SMA [12], while auditory and mentalizing regions showed differential
engagement by involuntary and voluntary laughter [13].
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... As such, laughter may involve innate mechanisms rather than learned vocal behaviour [5]. Laughter-like vocalizations are evoked not only in an emotional situation or by tickling, but also, and more frequently, in a social context in which laughter serves as a means of communication, as in humans [6,7] the great apes and rats [4,8]. While the production of speech requires the voluntary control of articulators, such as the tongue, the jaw and the soft palate, that of laughter depends solely on changes in breath control, subglottal pressure and laryngeal tension, which lead to a stereotypical pattern of respiration that is associated with rhythmic 'ha, ha'-like sounds. ...
... They are of particular relevance in distinguishing between the motor control of a postulated voluntary and involuntary pathway of vocalization. While speech can be produced only when the voluntary pathway is engaged, laughter depends greatly on the involuntary one, especially when it is associated with an emotional signal stimulating mirth and is not strategically employed for communication [7,23,24]. Studies involving monkeys reveal these pathways to be largely independent of each other [25,26]. The voluntary pathway controls neurons in the brainstem, which, via the motor cortex, innervate motor effectors either directly or indirectly through the reticular formation [27,28]. ...
... On the basis of available data in humans, a separation of these two pathways is less apparent. It has been postulated that during laughter the two pathways interact at the level of either the ( pre)-supplementary motor area [7,22], the anterior cingulate gyrus [30,31] or the primary motor cortex [15,31], possibly via inhibitory or modulatory processes [7,15]. However, subcortical regions have not been discussed concerning their interaction. ...
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Who has not experienced that sensation of losing the power of speech owing to an involuntary bout of laughter? An investigation of this phenomenon affords an insight into the neuronal processes that underlie laughter. In our functional magnetic resonance imaging study, participants were made to laugh by tickling in a first condition; in a second one they were requested to produce vocal utterances under the provocation of laughter by tickling. This investigation reveals increased neuronal activity in the sensorimotor cortex, the anterior cingulate gyrus, the insula, the nucleus accumbens, the hypothalamus and the periaqueductal grey for both conditions, thereby replicating the results of previous studies on ticklish laughter. However, further analysis indicates the activity in the emotion-associated regions to be lower when tickling is accompanied by voluntary vocalization. Here, a typical pattern of activation is identified, including the primary sensory cortex, a ventral area of the anterior insula and the ventral tegmental field, to which belongs to the nucleus ambiguus, namely, the common effector organ for voluntary and involuntary vocalizations. During the conflictual voluntary-vocalization versus laughter experience, the laughter-triggering network appears to rely heavily on a sensory and a deep interoceptive analysis, as well as on motor effectors in the brainstem. This article is part of the theme issue ‘Cracking the laugh code: laughter through the lens of biology, psychology and neuroscience’.
... Entre las emociones positivas, la risa puede verse como la única expresión omnipresente de alegría, observable en todas las culturas, especies y generaciones que surge en los seres humanos alrededor del cuarto mes de vida (Ruch y Ekman, 2001). Además, la risa promueve fuertemente la vinculación y facilita las interacciones sociales (Martin y Lefcourt, 1983, 2004Scott et al., 2014). En los seres humanos, la risa también se puede utilizar como un mecanismo de afrontamiento (coping mechanism), para aliviar el estrés y anular momentáneamente las emociones negativas. ...
... La percepción de la risa por sí sola no activa una red neuronal tan vasta y, lo que es más importante, no recluta al sistema dopaminérgico responsable de la recompensa y la motivación (Lavan et al., 2016;McGettigan et al., 2015;Wild et al., 2015;Wild et al., 2003), del cual se sabe que mejora el aprendizaje y la memoria (Gruber et al., 2014(Gruber et al., , 2016Kang et al., 2009;Patil et al., 2017). La risa se sustenta en sistemas neuronales complejos, lo que permite la existencia de dos formas distintas de risa dependiendo de la manera en que se provoca: (1) involuntariamente, en respuesta a eventos externos, lo que se conoce como risa espontánea o real, y (2) voluntariamente, en situaciones sociales, con el objetivo deliberado de crear lazos afectivos, lo que ha sido denominado risa social (Gervais y Wilson, 2005;Scott et al., 2014). En los primeros estudios científicos de la risa, los neurólogos se centraron en sus variantes patológicas (Brissaud, 1895;Nothnagel, 1889;Poeck, 1969). ...
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Las emociones positivas son valoradas y buscadas en nuestra vida diaria, pero aún no se comprende bien el rol funcional que juegan en la cognición humana. A pesar de ciertas similitudes, las emociones positivas y negativas parecen depender de vías neuronales distintas y, por tanto, podrían influir de manera diferente en funciones cognitivas como la memoria. En este artículo de revisión, se presentan los efectos cognitivos y los fundamentos neuronales de las emociones positivas, centrándose en las especificidades de las emociones inducidas por el humor. Posteriormente, se describe la influencia beneficiosa del humor sobre la memoria y se analizan los posibles mecanismos neuronales a través de los cuales el humor puede mejorar la memoria en el cerebro humano.
... First, it will be highlighted how the two simulationist systems depend on two distinct motor pathways, i.e., an emotional and a volitional one. Indeed, according to a wellaccepted "dual pathway" account of the neural control of the human voice (Jürgens, 2009), the production of emotional and nonemotional laughter is controlled by two independent networks, respectively, housed in emotional and volitional motor regions (Lauterbach et al., 2013;Scott et al., 2014;Gerbella et al., 2021;Zauli et al., 2022). Second, it will be shown that both emotional and volitional motor systems can be activated during the processing of others' emotional expressions, albeit their recruitment is triggered by different tasks and for different reasons. ...
... According to the "dual pathway" account of the neural control of the human voice, laughter production is also controlled by a voluntary network, originating from the ventral part of the motor system (housing the motor representations of mouth actions) and controlling the volitional production and modulation of laughter (Lauterbach et al., 2013;Scott et al., 2014;Gerbella et al., 2021). Imaging studies showed that this region is active during the observation (Jabbi et al., 2008) and the voluntary imitation of smiles (Leslie et al., 2004;Hennenlotter et al., 2005;van der Gaag et al., 2007) and during the production of voluntary, involuntary, and inhibited laughter (Wattendorf et al., 2013). ...
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Here it will be advanced the hypothesis that the ability to interact with the emotions displayed by co-specifics depends on two distinct simulationist systems, i.e., “motor simulation” and “emotional mirroring.” Although these systems are typically conceived as alternative and mutually exclusive models, the first aim of this chapter is to present empirical evidence demonstrating that they are coexisting systems that differ in terms of input, output, neural circuitry, and cognitive function. The former exploits motor knowledge to assist the visual system during the identification of potentially ambiguous emotional stimuli, while the latter conveys emotional contagion and social bonding. To clarify the functional difference between the two systems, it will be made reference to the notion of automatic abduction, as it has been originally described in the Peircean semiotic tradition. It will be argued that only “motor simulation” – but not “emotional mirroring” – can be thought of as part of an automatic sensorimotor abduction, that is, a sensorimotor inferential process which allows the exploitation of motor knowledge to assist the visual system during the identification of potentially ambiguous emotional stimuli. Instances of simulations triggered by “motor simulation” and “emotional mirroring” will be considered, respectively, in terms of motor and affective habits. The former will be characterized as ignorance-based kind of habits, whereas the latter will be defined as knowledge-based habits. Finally, the mechanism of “motor simulation” will be discussed in the light of the predictive processing framework.
... Humans and non-human primates use numerous vocalizations for maintaining social bonds and proximity with their conspecifics. Laughter is a universally recognized positive social expression occurring frequently in human social interactions [1,2] and it is used for promoting social bonding [2,3]. Numerous other primates [4,5] and rodents [6] also use laughter-like vocalizations for conveying prosocial motivation. ...
... Humans and non-human primates use numerous vocalizations for maintaining social bonds and proximity with their conspecifics. Laughter is a universally recognized positive social expression occurring frequently in human social interactions [1,2] and it is used for promoting social bonding [2,3]. Numerous other primates [4,5] and rodents [6] also use laughter-like vocalizations for conveying prosocial motivation. ...
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Media naturalness theory and social information processing theory make competing predictions regarding the effectiveness of different modes of communication at creating and maintaining emotionally intense social relationships. We explored how the duration of interaction and the form of laughter influenced happiness in communication modes with different levels of media naturalness. Forty-one participants completed a 14-day contact diary, recording interactions across face-to-face, Skype, telephone, instant messaging, texting, and e-mail/social network sites. Increases in duration of interaction positively predicted happiness only for face-to-face interactions, offering partial support for the media naturalness hypothesis. Laughter positively predicted happiness in all but one of the communication modes, with real and symbolic laughter having similar effects, a result consistent with the social information processing theory. © 2012 Wiley Periodicals, Inc.
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Laughter is an instinctive, contagious, stereotyped, unconsciously controlled, social play vocalization that is unusual in solitary settings. Laughter punctuates speech and is not typically humor related, speakers often laugh more often than their audience, and mate speakers are the best laugh getters. Laughter evolved from the labored breathing of physical play, with the characteristic "pant-pant" laugh, of chimpanzees and derivative "ha-ha" of humans signaling ("ritualizing") its rowdy origin. Laughter reveals that breath control is why humans cart speak and chimpanzees cannot. The evolution of bipedality in human ancestors freed the thorax of its support role in quadrupedal locomotion, a critical step in uncoupling breathing from running, providing humans with the flexible breath control necessary for speech and our characteristic laugh. Tickle, an ancient laughter stimulus, is a means of communication between preverbal infants and mothers, and between friends, family, and lovers. Because you cannot tickle yourself, tickle involves a neurological self/nonself discrimination, providing the mostprimitive social scenario.
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Although human laughter mainly occurs in social contexts, most studies have dealt with laughter evoked by media. In our study, we investigated conversational laughter. Our results show that laughter is much more frequent than has been described previously by self-report studies. Contrary to the common view that laughter is elicited by external stimuli, participants frequently laughed after their own verbal utterances. We thus suggest that laughter in conversation may primarily serve to regulate the flow of interaction and to mitigate the meaning of the preceding utterance. Conversational laughter bouts consisted of a smaller number of laughter elements and had longer interval durations than laughter bouts elicited by media. These parameters also varied with conversational context. The high intraindividual variability in the acoustic parameters of laughter, which greatly exceeded the parameter variability between subjects, may thus be a result of the laughter context.
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The burst of laughter that is evoked by tickling is a primitive form of vocalization. It evolves during an early phase of postnatal life and appears to be independent of higher cortical circuits. Clinicopathological observations have led to suspicions that the hypothalamus is directly involved in the production of laughter. In this functional magnetic resonance imaging investigation, healthy participants were 1) tickled on the sole of the right foot with permission to laugh, 2) tickled but asked to stifle laughter, and 3) requested to laugh voluntarily. Tickling that was accompanied by involuntary laughter activated regions in the lateral hypothalamus, parietal operculum, amygdala, and right cerebellum to a consistently greater degree than did the 2 other conditions. Activation of the periaqueductal gray matter was observed during voluntary and involuntary laughter but not when laughter was inhibited. The present findings indicate that hypothalamic activity plays a crucial role in evoking ticklish laughter in healthy individuals. The hypothalamus promotes innate behavioral reactions to stimuli and sends projections to the periaqueductal gray matter, which is itself an important integrative center for the control of vocalization. A comparison of our findings with published data relating to humorous laughter revealed the involvement of a common set of subcortical centers.
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Laughing is examined auditorily and acoustico-graphically, on the basis of exemplary speech data from spontaneous German dialogues, as pulmonic air stream modulation for communicative functions, paying attention to fine phonetic detail in interactional context. These phonetic case descriptions of laughing phenomena in speaker interaction in a small corpus have as their goal to create an awareness of the phonetic and functional parameters that need to be considered in the future acquisition, acoustic analysis and statistical evaluation of large spontaneous databases.
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Previous studies have demonstrated that 1 function of positive emotion is the undoing of physiological arousal produced by negative emotions. These studies have used single-subject paradigms, in which emotions were induced by films in college-age individuals. In the present study, we examined the relationship between physiological down-regulation and positive emotion in a sample of 149 middle-aged and older married couples engaged in a 15-min discussion of an area of marital conflict. During the conversation, autonomic and somatic physiological activity was measured, and emotional behaviors were recorded and subsequently coded. We found that during 20-s periods of down-regulation (where physiology transitioned from high arousal to low arousal), couples showed an increase in positive emotional behavior compared with periods without down-regulation. The finding was quite robust, suggesting that the undoing effect of positive emotion generalizes across age, sex, and marital satisfaction. The advantages of using positive emotion as an emotion regulation strategy are discussed.
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Emotional signals are crucial for sharing important information, with conspecifics, for example, to warn humans of danger. Humans use a range of different cues to communicate to others how they feel, including facial, vocal, and gestural signals. We examined the recognition of nonverbal emotional vocalizations, such as screams and laughs, across two dramatically different cultural groups. Western participants were compared to individuals from remote, culturally isolated Namibian villages. Vocalizations communicating the so-called "basic emotions" (anger, disgust, fear, joy, sadness, and surprise) were bidirectionally recognized. In contrast, a set of additional emotions was only recognized within, but not across, cultural boundaries. Our findings indicate that a number of primarily negative emotions have vocalizations that can be recognized across cultures, while most positive emotions are communicated with culture-specific signals.