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articles about contagious
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- Abstract
- Spontaneous mimicry appears fundamental to
emotional perception and contagion, especially
when it involves facial emotional expressions.
Here we cover recent evidence on spontaneous
mimicry from ethology, psychology and
neuroscience, in non-human and human animals. We
first consider how mimicry unfolds in non-human
animals (particularly primates) and how it
relates to emotional contagion. We focus on two
forms of mimicry-related phenomena: facial
mimicry and yawn contagion, which are largely
conserved across mammals and useful to draw
evolutionary scenarios. Next, we expand on the
psychological evidence from humans that bears on
current theoretical debates and also informs
non-human animal research. Finally, we cover the
neural bases of facial mimicry and yawn
contagion. We move beyond the
perception/expression/experience trichotomy and
from the correlational to the causal evidence
that links facial mimicry to emotional contagion
by presenting evidence from neuroimaging, direct
manipulation, neuro-stimulation and lesion
studies. In conclusion, this review proposes a
bottom-up, multidisciplinary approach to the
study of spontaneous mimicry that accounts for
the evolutionary continuity linking non-human
and human animals.
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- Yawn contagion
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- Even though spontaneous yawning is
considered a Þxed action pattern, its
description is not Þxed at all.
Spontaneous yawning has been described as
consisting of long, deep inspiration, brief peak
with apnea (acme) followed by a slow expiration
(Walusinski and Deputte, 2004; Guggisberg et
al., 2010) sometimes reported as rapid
(Baenninger, 1997; Krestel et al., 2018) or
shorter (Provine, 2012; Gallup et al., 2016).
When integrating the di_erent descriptions of
the motor patterns deÞning a yawning event
(Walusinski and Deputte, 2004; Guggisberg et
al., 2010; Provine, 2012; Baenninger, 1997), it
is possible to claim that it includes active jaw
gaping, eye closure, con- traction of facial
muscles, and passive jaw closure, accompanied by
neck stretching and head tilting and, in some
cases, by limb and body stretching. Due to its
complicated combination of patterns, Provine
(2012) distinguished di_erent types of yawn
(close-nose yawn; clen- ched-teeth yawn;
sealed-lips nose yawn; the eyes-open yawns)
showing that some features of yawning can be
modulated (Krestel et al., 2017). For all these
reasons, a yawning pattern includes a facial
expression but involves more than just facial
muscles.
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- Once elicited, yawning cannot be completely
suppressed. Therefore, it has been also
deÞned as a stereotyped or
reþex-like pattern (Lehmann, 1979; Provine
and Hamernik, 1986). Barbizet (1958, p.203)
deÞned yawning 'halfway between a
reþex and an expressive move- ment'.
Indeed, the morphological variability
surrounding the yawning display indicates that
this behavioural phenomenon is more articulated
than a simple reþex (Massen and Gallup,
2017). However, despite the di_erent
morphological variants found in some primate
species (e.g. chimpanzees; Vick and Paukner,
2010; geladas: Palagi et al., 2009; macaques:
Zannella et al., 2017; humans:Provine and
Hamernik, 1986, 2012), the basic yawning pattern
is plesiomorphic and well recogniz- able across
species (Baenninger, 1987). This makes yawning
an ex- cellent behavioural marker to investigate
facial display replication in a comparative
perspective.
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- As a physiological response, yawning can be
inþuenced by internal and external factors
(e.g. time of the day: Giganti and Zilli, 2011;
in- tracranial/brain temperature; Gallup and
Eldakar, 2013). Yawning is also socially
modulated: it is more likely to occur in real
social settings, as a result of yawn contagion
(Provine, 1989, 2005). However, in Vir- tual
Reality (VR) trials, the physical presence of a
researcher during testing signiÞcantly
inhibited contagious yawning, even though parti-
cipants were viewing a virtual environment (and
virtual yawns) and were unable to see the
researcher (Gallup et al., 2019).
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- The same factors that a_ect spontaneous
yawning are observed in yawn contagion.
Contagious yawning, although socially modulated,
remains a physiological response. It has been
found in several animal species, including
humans, and it occurs when individuals respond
with a yawn after perceiving a yawning from/in
others (Provine, 2005; Norscia and Palagi, 2011;
Palagi et al., 2009; Demuru and Palagi, 2012;
Campbell and de Waal, 2011, 2014; Romero et al.,
2013).
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- The phenomenon of yawn contagion seems to
rely upon the per- ception-action mechanism also
involving the mirror neuron system (Preston and
de Waal, 2002; Gallese et al., 2004).
Kapitány and Nielsen (2017) suggested
that in Homo sapiens the increased occurrence of
yawning in social conditions - as compared to
solitary - may be due not to contagion but to
the social setting itself. However, their study
showed that the mere presence of others was not
su_cient to increase the probability of yawning.
Instead, in line with what is expected when
contagion is in place, they found that the
yawning rates were sig- niÞcantly higher
in the social non-blind (individuals could see
each other and detect others' yawns) than in the
blind condition (individuals could not see each
other). In both conditions, the auditory
component of yawns was excluded by having the
participants listening to Chopin's Complete
Nocturnes (Kapitany and Nielsen, 2017).
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- Yawn contagion has been found in all the
hominine species: chim- panzees (Pan
troglodytes: Anderson et al., 2004; Campbell and
de Waal, 2011; Campbell and Cox, 2019), bonobos
(Pan paniscus: Demuru and Palagi, 2012; Tan et
al., 2017; but see: Amici et al., 2014 on a vary
small sample size) and humans (Homo sapiens:
Provine and Hamernik, 1986; 1989). Outside the
hominine species but still within the hominid
family, two reports failed to Þnd yawn
contagion in lowland gorillas
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- (Gorilla gorilla; Amici et al., 2014; on a
larger sample including a ba- chelor and a harem
group: Palagi et al., 2019a, 2019b). On the
other hand, yawn contagion has been found in two
cercopithecid monkey species (geladas,
Theropithecus gelada: Palagi et al., 2009;
Tonkean macaque, Macaca tonkeana: Palagi and
Norscia, 2019) but not convin- cingly in others.
SpeciÞcally, yawn contagion has not been
found in the Japanese macaque (Macaca fuscata,
Palagi and Norscia, 2019) and the situation is
unclear for stump-tailed macaques (Macaca
arctoides; Paukner and Anderson, 2005).
Stump-tailed macaques yawned sig-
niÞcantly more while watching a yawn video
than a control video but also showed more
self-directed behaviour like scratching, which
in primates is related to anxiety (Paukner and
Anderson, 2005). Results from stump-tailed
macaques suggests that the yawning response may
be also linked to anxiety and not necessarily to
contagion. Beyond pri- mates, yawn contagion has
been found in wolves (Canis lupus lupus; Romero
et al., 2014) and domestic dogs (Canis lupus
familiaris; inter- speciÞc yawn contagion;
Silva et al., 2012; Romero et al., 2013). In
dogs the yawning response was found not
associated with stress but only with the
perception of a triggering yawn emitted by the
owner (Romero et al., 2013). Therefore, it seems
that phylogenetic closeness is not, per se,
predictive of yawn contagion.
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