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les articles sur la contagion du
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articles about contagious
yawning
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- Definition : The yawn is a stereotyped and
often repetitive motor act characterized by
gaping of the mouth accompanied by a long
inspiration of breath, a brief acme, and then a
short passive expiration of breath.
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- Synonyms : gap, chasm
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- "...seeing a dog and horse and man yawn,
makes me feel how much all animals are built on
one structure".
- Charles Darwin, 1838 notebook.
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- Introduction
- Yawning is a universal everyday behavior
shared by all vertebrate species. It is a
stereotyped and often repetitive involuntary
motor act characterized by gaping of the mouth
and accompanied by a long inspiration of breath,
a brief acme, and then a short expiration of
breath. Yawning is not merely a simple opening
of the mouth but a complex, coordinated movement
involving a flexion followed by an extension of
the neck and a wide dilatation of the
pharyngo-larynx with strong stretching of the
diaphragm and antigravity muscles (Provine,
1986). Contagious yawning is the urge to yawn
when thinking about, listening to, reading
about, hearing, or viewing yawning. Ethologists
use a better word, i.e., replication, or
behavioral mimicry, whereas neurologists refer
to echokinesis, a term coined by Jean-Martin
Charcot (1825-1893). The widespread familiarity
of contagious yawning is due in part to the fact
that almost everyone has experienced the
phenomenon and it occurs seemingly without
volition. This behavior is well documented in
humans; however, its function and prevalence in
the animal kingdom and the brain mechanisms
underlying it are only beginning to be
understood.
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- Behavioral Framework
- Yawning is morphologically similar in
reptiles, birds, mammals, and fish, and no
environmental input changes the sequence of
movements (Walusinski & Deputte, 2004). This
behavior may be an ancestral vestige maintained
throughout evolution with little variation,
bearing witness to the early phylogenetic
origins of yawning. Like any phylogenetically
old behavior, yawning can be observed early in
ontogeny; that is, at 12 weeks of fetal life in
humans (Walusinski, 2012). Behavioral and
neurophysiological studies provide converging
evidence that yawning occurs preferentially
during rest, periods of drowsiness, and
awakening. Hunger and satiety also trigger
yawning. The frequency of yawning has a
distinctive circadian distribution and occurs
most frequently before and after sleep; that is,
during periods of lower levels of vigilance and
alertness (Walusinski & Deputte, 2004).
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- When animals transition between behaviors,
they are not merely responding in a passive way
to the conditions of the environment, such as
day-night succession. Rather, they are following
internally generated signals produced by
homeostatic processes originating in the
hypothalamus (suprachiasmatic nucleus and
paraventricular nucleus of the hypothalamus,
where yawning is triggered). The resultant
internal rhythm allows for the anticipation of
transitions and triggers behavioral and
physiological changes in accordance with those
transitions. This results in two advantages:
predictability and the possibility to detect the
unexpected. Similar to sleep and hunger, yawning
functions in this way and appears to be
associated with transitions between periods of
high and low activity or arousal. This universal
form of yawning is internally elicited (Gupta,
& Mittal, 2013; Krestel, et al., 2018).
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- But other, externally elicited forms of
yawning exist. Among mammals, morphologically
identical yawns occur in situations relative to
stress, social interactions, and sexuality
(Kubota et al., 2014). Contagious yawning is a
form of social interaction without language,
triggered involuntarily. Nonconscious mimicry
("chameleon effect") refers to an individual's
tendency to imitate a social partner's behaviors
without either party's awareness or intent.
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- Yawning in the animal kingdom
- Yawning conveys information to other
conspecifics about dominance, state of arousal
or tiredness, anxiety, or fear. The combinations
of different types of signals (visual, acoustic,
and olfactory) establish the interaction in
either conspecifics or human counterparts.
Indeed, a cross-species chameleon effect or
contagiousness has now been documented for
yawning between dogs and humans (Buttner &
Strasser, 2014; Harr, Gilbert & Philips,
2009). In nonhuman primates, teeth-bearing
during yawning has led to the suggestion that
this behavior reinforces dominant identity
rather than signaling a threat (Deputte, 1994).
Accordingly, yawning has been compared with
intimidating displays that dominant males
usually show to subordinate males to display
their dominant status. Chimpanzees, like humans,
differ from Old World monkeys in that yawning by
adult males is not a form of ritualized display
expressed in situations of mal-male
confrontation.
- Other ethological studies propose that
yawning might induce relaxation of social
tension and thus be a displacement activity,
considered to indicate changes in behavioral
state for the yawner and simultaneously to
operate as a physiological capacity signal. In
Sprague-Dawley rats, yawning associated with
penile erections suggests that yawning behavior
arises from the ritualization of pre-existing
cues involved in perception of stimulating
chemo-signals in a mating context (Moyaho et
al., 2015).
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- Contagious Yawning
- Contagious yawning has been observed in
nonhuman primates (chimpanzees, bonobos,
stump-tailed macaques, gelada baboons, and
common marmosets) (Demuru, & Palagi, 2012;
Palagi, et al., 2009), in dogs (Harr, et al.,
2009), in wolves (Romero, et al., 2014), in
sheep (Yonezawa, et al., 2017), in budgerigars,
the only non-mammalian specie (Gallup, et al.,
2015), in rats (Moyaho, et al., 2015), and in
elephants (Rossman, et al, 2017), but not in
lemurs (Reddy et al., 2016) and tortoises (Huber
& Wilkinson, 2011). No data are available
for other species. Animal studies have reported
low levels of contagious yawning in dogs,
significantly lower than 33%, the rate observed
in chimpanzees (Harr, et al., 2009). Contagious
yawning seems to be more easily triggered when
models are conspecifics or have a strong social
bond with the observer. The individuals that are
more "reactive" when watching yawning of
conspecifics, are not more reactive when exposed
to other behaviors (Anderson, et al.,
2006).
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- Observational surveys and experimental
analysis of contagious yawning in humans has
revealed occurrence in 65%-75% of the entire
population. There is still no consensus as to
whether women exhibit more contagious yawning
than men (Gallup, & Masse, 2016). At any
rate, individuals who conform spontaneously to
normative social influence, and who are most
receptive to advertising messages, also appear
most susceptible to contagious yawning. People
who scored higher on measures of empathy and
mental state attribution skills were more likely
to show contagious yawning (Palagi, et al.,
2014). These observational data, among others,
are produced for linking contagious yawning and
empathy.
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- How is contagious yawning triggered? Sight
is a powerful stimulant. In spectators viewing a
video showing 30 successive yawns, 55% will yawn
within five minutes. The latency period varies
from a few seconds to five minutes. There is no
need for the face of the yawner to be in a
precise visual plane relative to the subject
receiving the contagion. Face to face, at
90°, 180°, and 270° relative to
one another, replication occurs. The existence
of a susceptibility to contagion among blind
subjects confirms that sight is not the only
trigger. Viewing only part of the face, such as
a widely opened mouth, does not trigger
replication. Therefore, a multimodal perception
of the whole facial configuration and of audible
respiratory moments is necessary for replication
to happen, along with coordinated dynamics
(Provine, 2009). By animals as by humans,
individual yawning in response to perceiving
someone else yawn varies as a function of a host
of variables that have yet to be clarified.
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- The neural basis for contagious yawning
- Contagious yawning is a primitive expression
of cognitive processes involved in
self-awareness and theory-of-mind; that is, the
capacity to build knowledge of mental states and
to attribute them to others. In humans,
contagious yawning first appears around five
years of age, probably linked to the acquisition
of a theory-of-mind. Humans who performed better
at self-recognition and theory-of-mind,
exhibited more contagious yawning (Senju, 2010).
Yawning operates as a low-level form of
nonconscious mimicry, which is necessary for
involuntarily decoding the mental states of
others, notably their emotions. The link between
empathy and contagious yawning has empirical
support but remains controversial (Massen, &
Gallup, 2017). In line with this possible
association, the empathic modeling hypothesis
predicts that species that do not recognize
themselves in mirrors and do not show evidence
of mental state attribution ought to fail to
show evidence of contagious yawning (de Waal
& Preston, 2017; Provine, 2014).
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- Both developmental and neurophysiological
research suggests that resonance between
observed and executed actions is supported by
the mirror neurons system (MNS). Brain areas
that constitute the MNS, namely the premotor
area in the frontal lobe, the inferior parietal
lobule, and the superior temporal sulcus,
participate in the imitation of motor acts and
expressions. These human mirror regions are
homologous with area F5 of the ventral premotor
cortex in macaques (Rizzolatti, 2005). These
regions are also interconnected with the insula
and show activation in functional MRI studies of
imitation, emotional states, and thus, empathy
(Anderson, & Matsuzawa, 2006). The
ventromedial prefrontal cortex also integrates
affective signals to guide emotionally adapted
behaviors. Thus, motor acts and affective states
can transfer from a target to an observer in a
bottom_up, goal-relevant manner through shared
representations for perception and action. In
this way, observing an affective posture or
expression drives feed-back from peripheral
motor representations to activate associated
emotional states. Contagious yawning recruits
brain areas that have been implicated in social
cognition (empathy), self-processing,
theory-of-mind, and emotional contagion
mechanisms (motor mimicry-MNS). Contagious
yawning is the most basic expression of motor
mimicry and emotional contagion, a rudimentary
form of involuntary empathy (Brown et al., 2017;
de Waal, & Preston, 2017). Whatever
underlies contagious yawning, it does not appear
to be based either on conscious imitation or
higher-level, "conscious" empathy. Regardless,
contagious yawning presents a powerful tool to
explore the root of empathy in animal
evolution.
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- An error to avoid
- Equating physiological and communicative
functions of yawning with non-social and social
contexts respectively is inaccurate as both
forms may occur in a social context. Indeed,
ethological studies of non-human primates or
South-African ostriches, for example, show that,
at certain points, an entire group yawns. In
this case, there is no contagious yawning but a
synchronous behavior related to circadian
rest-activity rhythms.
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- Conclusion
- Contagious yawning is an example where,
through evolution, a behavior can be recycled
for different purposes according to the
increasing complexity of the central nervous
system, correlated with the richness of social
interactions. Yawning is managed by the
phylogenetically oldest part of the brain,
namely the diencephalon and brainstem.
Contagious yawning occurs by neocortex
activation, which is necessary for the complex
interactions of social life between individuals.
It remains to be seen whether contagious yawning
has any effect on the activity levels of other
group members. Further research, especially in
wild populations, should examine the regulating
effect of yawning on synchronized group behavior
in order to test its communicative function.
Contagious yawning offers a neglected but
fruitful avenue of investigation in the growing
fields of developmental, affective, and social
neuroscience.
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- A list of related entries from this
volume:
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- Emotional Contagion
- Elisabetta Palagi, Ivan Norscia in
Encyclopedia of Animal Cognition and
Behavior
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- Contagion
- Thomas R. Zentall in Encyclopedia of Animal
Cognition and Behavior
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- Grooming
- Richard McFarland in Encyclopedia of Animal
Cognition and Behavior
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- Sleep
- Albrecht P. A. Vorster in Encyclopedia of
Animal Cognition and Behavior
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- Imitation
- Eóin P. O'Sullivan, Christine A.
Caldwell in Encyclopedia of Animal Cognition and
Behavior
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- Empathy
- Garet P. Lahvis in Encyclopedia of Animal
Cognition and Behavior
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- References
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- 1. Anderson J.R., & Matsuzawa T. (2006).
Yawning: an opening into empathy. In T.
Matsuzawa, M. Tomonaga, M. Tanaka (Eds).
Cognitive Development in Chimpanzees.
(pp.233-245) Tokyo , Springer.
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- 2. Brown, B.J., Kim, S., Saunders, H.,
Bachmann, C., Thompson, J., Ropar, D., Jackson,
S.R., Jackson, G.M. (2017). A Neural basis for
contagious yawning. Curr Biol, 27, 2713-2717.e2.
https://doi.org/10.1016/j.cub.2017.07.062
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- 3. Buttner, A.P., & Strasser, R. (2014).
Contagious yawning, social cognition, and
arousal: an investigation of the processes
underlying shelter dogs' responses to human
yawns. Anim Cogn, 17, 95-104.
https://doi.org/10.1007/s10071-013-0641-z
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- 4. Demuru, E., & Palagi, E. (2012). In
Bonobos yawn contagion Is higher among kin and
friends. PLoS ONE, 7, e49613.
http://doi.org/10.1371/journal.pone.0049613
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- 5. Deputte, B. L. (1994). Ethological study
of yawning in primates. I. Quantitative analysis
and study of causation in two species of old
world monkeys (Cercocebus albigena and Macaca
fascicularis). Ethology, 98, 221-245.
https://doi.org/10.1111/j.1439-0310.1994.tb01073.x
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- 6. de Waal, F.B.M.,& Preston, S.D.
(2017). Mammalian empathy: behavioural
manifestations and neural basis. Nat Rev
Neurosci, 18, 498-509. https://doi.org/10
.1038/nrn.2017.72
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sbadiglio dell struzzo. Psicologia e biologia
dello sbadiglio. Bollati Boringhieri,
Torino.
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- 8. Gallup, A.C., Swartwood, L., Militello,
J., Sackett, S. (2015). Experimental evidence of
contagious yawning in budgerigars (Melopsittacus
undulatus). Anim Cogn, 18, 1051-158.
https://doi.org/10.1007/s10071-015-0873-1
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- 9. Gallup A.C., & Massen J.J.M. (2016).
There is no difference in contagious yawning
between men and women. Royal Society Open
science, 3, 160174.
https://doi.org/10.1098/rsos.160174
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- 10. Gupta, S., & Mittal, S. (2013).
Yawning and its physiological significance.
International Journal of Applied and Basic
Medical Research, 3, 11&endash;15.
http://doi.org/10.4103/2229-516X.112230
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- 11. Harr, A.L., Gilbert, V.R., &
Phillips, K.A. (2009). Do dogs (Canis
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12, 833-837.
https://doi.org/10.1007/s10071-009-0233-0
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- 12. Huber, L., & Wilkinson, A. (2011).
No evidence in contagious yawning in the
red-footed tortoise (Geochelone carbonaria).
Current Zoology, 57, 477-484.
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- 13. Krestel, H., Bassetti, CL., &
Walusinski, O. (2018). Yawning-Its anatomy,
chemistry, role, and pathological
considerations. Prog Neurobiol, 161, 61-78.
https://doi.org/10.1016/j.pneurobio.2017.11.003.
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- 14. Kubota, N., Amemiya, S., Yanagita, S.,
Nishijima, T., & Kita, I. (2014). Emotional
stress evoked by classical fear conditioning
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http://doi.org10.1016/j.neulet.2014.02.064.
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- 15. Massen, J.J.M., & Gallup, A.C.
(2017). Why contagious yawning does not (yet)
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Smell facilitates auditory contagious yawning in
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https://doi.org/10.1007/s10071-014-0798-0
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http://doi.org/10.1073/pnas.0910891106
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Yawn contagion in humans and bonobos: emotional
affinity matters more than species. PeerJ 2:e519
https://doi.org/10.7717/peerj.519
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https://doi.org/10.1111/j.1439-0310.1986.tb00611.x
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E.L., & Hare, B. (2016). No evidence for
contagious yawning in lemurs. Anim Cogn, 19,
889-898.
https://doi.org/10.1007/s10071-016-0986-1
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system and imitation. In S. Hurley & N.
Chater, (Eds). Perspectives on Imitation: from
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Un
comportement recyclé au cours de
l'Evolution
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- Le bâillement contagieux est un
exemple dans lequel, au fil de
l'évolution, un comportement peut
être recyclé pour des
finalités différentes illustrant
la complexité croissante du
système nerveux central,
corrélée à la richesse des
interactions sociales.
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- Le bâillement émane de la
partie la plus ancienne,
phylogénétiquement, du cerveau
à savoir le diencephale et le tronc
cérébral. La réplication du
bâillement, alias contagion,
résulte de l'activation du
néocortex, nécessaire aux
interactions complexes de la vie sociale entre
les individus. Il reste à savoir si le
bâillement contagieux a un effet sur les
niveaux de vigilance des autres membres du
groupe.
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- Des recherches complémentaires, en
particulier dans les populations animales
sauvages, devraient explorer l'effet
régulateur du bâillement sur le
comportement d'un groupe afin de tester sa
fonction de communication.
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- La réplication du bâillement
offre une voie d'investigation
négligée mais fructueuse dans les
domaines en pleine croissance de la neuroscience
du développement, de l'affectif et du
social.
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