See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/330528693

Herding Brains: A Core Neural Mechanism for Social Alignment

Article  in  Trends in Cognitive Sciences · March 2019

DOI: 10.1016/j.tics.2019.01.002

CITATIONS READS

0 131

4 authors, including:

Inbar Marton Hila Gvirts

University of Haifa University of Haifa
1 PUBLICATION   0 CITATIONS    15 PUBLICATIONS   77 CITATIONS   

SEE PROFILE SEE PROFILE

All content following this page was uploaded by Inbar Marton on 18 February 2019.

The user has requested enhancement of the downloaded file.

Opinion

Herding Brains: A Core Neural Mechanism

for Social Alignment
Simone G. Shamay-Tsoory,1,* Nira Saporta,1 Inbar Z. Marton-Alper,1 and Hila Z. Gvirts1

When we clap our hands in synchrony, feel the sadness of a friend, or match our Highlights
attitudes to peer norms, we align our behavior with others. We propose here a Traditionally, the diverse behaviors that
involve social alignment have been
model that views synchronized movement, emotional contagion, and social
considered individually.
conformity as interrelated processes that rely on shared neural networks.
Building on the predictive coding framework, we suggest that social alignment We synthesize models of collective
action across species with an emer-
is mediated by a three-component feedback loop – an error-monitoring system ging body of neuroscience, neurocom-
that reacts to misalignment, an alignment system, and a reward system that is putational, and psychology research to
activated when alignment is achieved. We describe herding-related syndromes propose that different manifestations
of social alignment are actually linked,
(autism, loneliness) and call for innovative research to investigate the links with motor synchrony, emotional align-
between the levels of alignment. ment, and conformity influencing one
another in a reciprocal manner.

From Herds to Conformism Building on the predictive coding fra-

Herding is ubiquitous throughout the animal kingdom. Birds fly together in flocks, schools of fish mework, we argue that these different
move in a coordinated manner, and swarming ants exhibit collective behavior [1]. Although group levels of alignment reflect the workings
living has various benefits including foraging efficiency [2] and reduction of predation risk [3], of a prototype feedback-loop model.

individuals may benefit from sociality only if they remain connected to other members [4], indicating The social alignment feedback loop
that achieving connectedness is a key motivator of behavior across species. In humans, collective includes three core components.
action may be observed in various everyday situations including unintentional synchronized walking One system is in place to react to
alignment, and another system reacts
[5], rocking [6], and synchronized body-posture sway when conversing [7]. Notably, it is possible to
to misalignment. Based on the misa-
observe diverse behavioral displays of group alignment that go beyond joint movement. These lignment detected, a further system is
‘higher-level’ behaviors may include emotional contagion and social conformity. Given these responsible for aligning to the point of
various forms of group alignment, it has been proposed that herding may be conceptualized as perceived alignment.
the tendency of an organism to align with other organisms in a group in terms of behaviors and
cognitions [8]. Despite a long history of scientific investigation, we are only now beginning to
decipher the different aspects of social alignment. We synthesize here disparate recent findings on
movement synchrony, emotional contagion, and conformity under a single conceptual umbrella: a
social alignment model. We propose that these seemingly different behavioral manifestations of
herding are actually linked, influencing one another in a reciprocal manner and adhering to similar
principles. We advocate in favor of a common set of brain networks underlying social alignment and
hypothesize that, at its core, this system enables connectedness between individuals.

The Various Levels of Herding are Behaviorally Linked

Interpersonal motor synchrony involves the alignment in time of the behavior of two or more
interacting individuals. Close to the constructs of mimicry or imitation, motor synchronization
also introduces the importance of coordination in the timing of the action [9]. Synchronization 1
Department of Psychology, University
can occur implicitly (coordination of walking pace in humans) but can also occur explicitly of Haifa, Haifa 3498838, Israel
(soldiers marching [10]).

*Correspondence:
Emotional contagion represents the alignment of one’s emotions with the emotions expressed sshamay@psy.haifa.ac.il
by those around you, whether consciously or unconsciously [11]. Emotional contagion may (S.G. Shamay-Tsoory).

174 Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3 https://doi.org/10.1016/j.tics.2019.01.002
© 2019 Elsevier Ltd. All rights reserved.
occur implicitly in all types of emotions (e.g., sadness, happiness) during social interactions [12],
as well as in virtual networks [13], or even in situations of mere copresence without directly
attending to each other [14].

Social conformity is the act of fitting in with the group, manifested in the tendency of people to
modify their opinions or judgments according to those of the group [15]. This form of ‘cognitive’
alignment with the group can appear in various domains such as food preference [16], long-
term memory [17], visual perception [18], and moral judgments [19].

Although motor synchrony, emotional contagion, and conformity are different behaviors with
different emphases, evidence demonstrates that these behaviors are related (Figure 1). This
idea is compatible with the embodied cognition theory, according to which bodily experiences
play an integral role in cognition and emotions [20]. Moreover, this framework is largely
supported by the theory of shared reality that views dyadic interactions as units that progress
through shared feelings, shared coordination, and shared identity [21].

In line with the suggestion that the three levels of alignment are linked, it was found that, when
confederates synchronize with participants by mimicking their limb movement, the participants
are likely to conform to stereotypes [22]. Notably, the reverse effect was described in a different
study that shows how team conformity facilitates motor coordination and synchrony [23]. In
addition, previous research has reported a connection between perceived emotional alignment
and endorsement of social beliefs and values, providing a further link between emotion

Alignment in
movements Motor Emoonal Alignment in emoonal
structure, direcon, synchrony contagion states and expressions
and pace

Core
alignment
mechanism

Cognive
synchrony

Alignment in
thoughts, beliefs,
percepons, intenons,
and atudes
Alignment level
Outgroup Ingroup
People we dislike People I like
People I do not feel close to People I feel close to

Figure 1. The Multilevel Model of Social Alignment. The three levels of social alignment (motor synchrony, emotional
contagion, and conformity) are linked, influencing one another in a reciprocal manner. For example, several studies show
that motor synchrony increases emotional alignment and social conformity. By contrast, conformity increases motor
coordination. The alignment level is impacted in all types of alignment by similar principles, with alignment being higher with
ingroup members, people I like, and people I feel close to. The color scale ranges from red (higher alignment) to green
(lower alignment).

Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3 175

contagion and conformity [24]. Likewise, in a variety of studies researchers have demonstrated
the relationship between movement synchrony and emotional contagion. For example, in one
study, inhibition of congruent facial actions impaired listener experiences of a concordant
subjective state, showing that there is less emotional contagion when motor synchrony is
blocked [25]. Moreover, there is evidence that positive emotional contagion (e.g., contagions of
happiness) results in improved motor coordination [26].

Given the theorized link between synchrony, emotional contagion, and conformity, it is not
surprising that the three types of behaviors adhere to a similar set of principles. Indeed, a major
feature of synchrony, emotional contagion, and conformity is the issue of who we align with.
Individuals are more likely to synchronize their movements [27] or conform [28] with others they
like and appreciate [29].

Similarly, the issue of group membership is important to all three levels of alignment. It is known
that people prefer to synchronize in movement with ingroup as opposed to outgroup members
[30]. Likewise, sharing emotions [31] and conforming to the opinion of others [32] have been
shown to be markedly affected by group membership. Notably, movement synchrony, emo-
tional contagion, and social conformity were all reported to lead to greater feelings of closeness.
Several studies have shown a positive relationship between motor synchrony and feeling close
to others [33,34]. Correspondingly, experiencing the same emotion as others, regardless of its
valence, is positively related with higher closeness ratings [35]. Similarly, greater cohesiveness
in groups has been found to lead to greater conformity [36]. Thus, it appears that the link
between the different levels of alignment, likeability, and closeness is bidirectional: liking and
closeness influence all types of alignment, and alignment enhances liking and closeness. It is
possible that the three manifestations of social alignment represent a core mechanism of
connectedness, and that triggering one behavior activates all other behaviors and encourages
likeability, closeness, and connectedness. Collectively, motor, emotional, and cognitive align-
ment appear to be linked together, facilitating one another and being affected by similar social
factors (e.g., level of psychological/physical closeness), suggesting that herding may involve
shared mechanisms (Figure 1).

The Neural Mechanisms Underlying Herding

A well-accepted model of herding in animals argues that there are three basic principles that
guide most types of herds [37]. These principles include: approaching other individuals, while at
the same time moving away from very nearby neighbors, and aligning with the direction of
movement of those that are close by. These principles indicate that social alignment involves a
mechanism that is responsible for detecting the gap between the self and others to evaluate the
level of approach versus avoidance necessary to create an optimal distance from group
members, and a mechanism that generates alignment.

Although these principles were studied in models of collective motion, they may be relevant for
the understanding of higher levels of alignment and its neural underpinnings. For example, both
emotional contagion and conformity require detection of the gap between the emotional state
and cognition of the individual and that of others to allow optimal emotional/cognitive alignment.

Building on these principles, we propose that the different levels of alignment reflect the working
of a prototype feedback loop that includes three core components. The first component is a
gap-monitoring system that detects the levels of gap between the self and others. Another
component is an observation–execution (OE) system that regulates alignment. Finally, a reward
system is responsible for signaling that the gap is optimal and that the individual is aligned with

176 Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3

others. The circular nature of this loop permits a description for each stage of the cycle, at
multiple levels of analysis. The model is mainly informed by findings from traditional single-brain
neuroimaging approaches, which provide us with evidence regarding intrabrain responses to
controlled social stimuli. In addition, we address the ‘second-person neuroscience’ approach
[38], and report findings from interbrain studies (hyperscanning) that show how the core
systems of social alignment may be coupled between multiple brains.

The Neural Systems Involved in Social Alignment

Accumulating evidence indicates that the gap monitoring, reward, and OE systems play similar
roles across various types of alignment – motor, emotional, and cognitive. It is noteworthy that,
until now, the different types of alignment were studied separately, and therefore the evidence
presented should be regarded only as suggesting a common neural mechanism (Figure 2).
Given that the three types of alignment may share similar mechanisms, it is possible that
engaging in one type of alignment (e.g., synchronized walking) would activate the shared
alignment loop, thus activating the two other types of alignment (emotional cognation,
conformity).

The Misalignment Detection System

Studies on misalignment with the group show that deviation from social group norms often
evokes the activity of the dorsal medial prefrontal cortex (dmPFC) and dorsal anterior cingulate
(dACC) [39,40]. The dACC as well as the anterior insula (AI) were shown to be activated during
gap monitoring, and during error/conflict detection in both social and non-social contexts
[39,41]. Emerging theoretical accounts indicate that the dACC primarily operates as a monitor
of conflicts, errors, and, more generally, of differences between actual and predicted or desired
states, which require adjustments [42]. In the context of predictive coding, the dACC was found
to be driven by unexpected outcomes [40,43].

Activity in the dACC was found in a multitude of studies on social influence and conformity, and
it was generally associated with response to a gap between the cognition of an individual and
that of the group [44,45]. The dACC and dmPFC were also found to be active in studies that
examine being ‘out of sync’ in movement [46,47]. Likewise, in the context of emotional
synchrony, the dmPFC was activated when individuals felt that they were not being emotionally
understood [48] and when they attempt to align emotionally with a coattending individual [49].

Figure 2. The Three Core Brain Sys-

tems Underlying Social Alignment.
The gap-detection system (yellow) com-
prises the dorsomedial prefrontal cortex
(dmPFC), the dorsal anterior cingulate
cortex (dACC), and the anterior insula
(AI). The alignment system (red) com-
prises the inferior frontal gyrus (IFG), the
inferior parietal lobule (IPL), and the pre-
motor cortex. The alignment reward cir-
cuit (green) comprises the orbitofrontal
cortex (OFC), the ventromedial PFC
(vmPFC), and the ventral striatum (VS).
Alignment reward
Misalignment detecon
Alignment/observaon–execuon

Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3 177

Activations in the dACC were also reported in studies that observed affective incongruence
between modalities [50]. Therefore, it appears that the role of the gap-detection system is to
estimate the amount of distance (in movement, emotion, and cognition) between the individual
and the group, and once this system is activated it triggers activity in the alignment system.

Alignment and the OE System

The need to align in movement, emotion, or cognition may trigger activity in neural networks
responsible for OE. It has been repeatedly shown that observing a behavior in another may
automatically activate one’s own representations for the behavior, and output from this shared
representation automatically proceeds to motor areas of the brain where responses are
prepared and executed [51]. In humans, the OE system has been identified in the inferior
frontal gyrus (IFG), the inferior parietal lobule (IPL) [52,53], and the superior temporal sulcus
(STS) [54], as well as in the premotor cortex [55]. It was argued that within the OE system the
IFG contains motor representations of actions, whereas the IPL is more relevant for the actual
sensory-to-motor mapping of the visual input and own-body versus other coordinates [56]. It is
also increasingly acknowledged that the OE system is implicated not only in action observation
but also in understanding emotions [57] and beliefs [58].

There is ample evidence indicating that the OE system is involved in social alignment [59]. It was
reported [46] that when participants experience synchrony (e.g., when someone synchronizes
with their motor responses) they show greater activity in the IPL. Similarly, using a paradigm of
finger tapping with a virtual partner, it was demonstrated that increases in perceived synchro-
nization difficulty correlated with greater activation of the right IFG [47]. The IFG was also found
to be active during synchronous speaking [60].

Given that the OE system is engaged not only in understanding movement but also in higher-
level decoding of emotion and cognition [58], it is not surprising that emotional contagion was
found to engage the OE system, particularly the premotor cortex and IPL regions [61]. Lesion
studies further support the role of the OE in emotional contagion [62], emotional empathy [63],
and vicarious pain [64]. Furthermore, in a study on social influence it was reported that the IFG is
activated during social conformity [16]. In line with this, increased activity was found in the IFG
during preference change towards the preference of others [65].

Notably, hyperscanning studies using functional near-infrared spectroscopy (fNIRS) show

interbrain coupling in premotor cortices during a paced finger-tapping imitation task [66].
Furthermore, an increase in interbrain coupling in the left IFG during coordinated face-to-face
dialogue between partners was found [67], as well as interbrain coupling of the left IFG during
synchronized singing [68]. Interbrain coupling in the OE system was also demonstrated to be
higher during tasks that required coordination [69].

Electroencephalography (EEG) studies have found that behavioral synchrony correlates with
interbrain coupling in several oscillatory bands in dyads [70], as well as in groups [71], particularly in
the alpha/mu band, which is regarded as the electrophysiological marker of the OE system. A
study of interactive finger tapping found suppression of alpha and low-beta oscillations over motor
and frontal areas [72]. Finally, interbrain coupling in the alpha/mu band correlated with empathy for
pain (or shared pain), indicating that interbrain coupling underlies emotional contagion [73]. Thus, it
appears that the OE system is not only active in one participant during social alignment, but it is also
coupled between participants. In line with our model, it appears that during social interaction the
feedback loops of the interacting partners are coupled, creating a closed loop between the
participants (Figure 3). Collectively, this line of evidence supports the view that the OE system

178 Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3

 Sensory informaon:

Other Self

Gap was
detected Observaon–execuon
IFG, IPL, premotor

Gap/error detecon
dACC, dmPFC, Al
Predicon about alignment

Other Me

Connectedness reward
Gap was OFC, PFC, VS
not detected
Keep the predicon

Figure 3. The Extended Integrative Model of Alignment. According to the predictive coding framework of alignment, the brain constantly generates predictions
on alignment based on previous information about alignment (‘priors’). When the incoming sensory information of motion, emotion, or cognition about oneself and about
others is different from the prediction (gap detection), that constitutes a prediction error. Detecting a gap activates the alignment system, whereas detecting that there is
no gap activates the reward system. Therefore, the feedback-loop model of social alignment comprises three systems: (i) the gap-monitoring (prediction-error) system
monitors the level of approach/avoidance necessary to align with the group, and includes the dACC and dmPFC. (ii) The alignment system includes the IFG, IPL,
premotor cortex, and STS. (iii) The reward system includes the VS, OFC, and vmPFC, and is activated when alignment is achieved. Although the core model provides
the basis of social alignment, an extended system may be activated depending on the specific requirements of the situation. Abbreviations: AI, anterior insula, dACC,
dorsal anterior cingulate cortex; dmPFC, dorsomedial prefrontal cortex; IFG, inferior frontal gyrus; IPL, inferior parietal lobule; OFC, orbitofrontal cortex; STS, superior
temporal sulcus; vmPFC, ventromedial prefrontal cortex; VS, ventral striatum.

mediates alignment with others in motor synchrony, emotional contagion, and conformity. It is
thus possible that the role of the OE system in social alignment is to observe a behavior in another
(movement emotion, cognition) and to automatically activate one’s own representations for this
behavior to prepare a congruent response.

The Alignment Reward System

Motor, emotional, and cognitive synchrony activates brain areas associated with reward,
potentially related to the sense of satisfaction that occurs when people experience connect-
edness. Neuroimaging studies of reward processing have identified a set of regions compris-
ing, among others, the ventral striatum (VS), ventromedial prefrontal cortex (vmPFC), and
orbitofrontal cortex (OFC) [74] (Figure 2).

In line with recent studies showing that synchrony is enjoyable (e.g., [75,76]), it was found that
the striatum was activated when individuals experienced synchrony during drumming. Simi-
larly, in a hyperscanning study it was reported that the striatum was active during coordinated
movement [77]. Interestingly, activity in the VS was also found in studies on social conformity
that examined how individuals align their beliefs with others [78,79]. A meta-analysis of studies
that examined the neural signature of conformity revealed that stimuli endorsed by others
consistently evoked VS activations [80]. Finally, it was shown that emotional facial synchrony
(having similar facial expressions to those of a target) leads to positive feelings towards the
target and activates the OFC [35].

A Predictive Coding Account of Herding

The feedback-loop model proposed here can be cast in the framework of neurocomputational
accounts of predictive coding (Box 1). According to the predictive coding theory, the brain

Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3 179

 Box 1. Multilayered Social Alignment – A Predictive Coding Approach
Predictive coding holds that the brain produces predictions of input signals (a top-down signal), and then constantly
collects information (a bottom-up signal) that is weighed in relation to the current prediction. Discrepancies between the
top-down and bottom-up signals constitute a prediction error, which the brain strives to minimize by adapting the
models generating the predictions that lead to change in behavior [81].

Time-difference reinforcement learning can be used as basic formulation of a prediction based generative model [111].
In this formulation, the value (Q) of each state of the world is updated when new value information (V) about this world
state is received. Update rate is governed by either a fixed learning rate or according to the new evidence reliability of
precision (W) [112]. The updated world model then informs action. Generally, this can be described by the following
formulation:
Qt ðaÞ ¼ Qt1 ðaÞ þ W  ½V t  Qt1 ðaÞ [I]

Where Qt-1(a) is the prior prediction, V is the new input received. The difference between them [Vt  Qt1(a)] is the
prediction error. W is the weight given to the prediction error, which can be based on the uncertainty of the prediction
and the reliability of the received sensory input.

In our model, the prediction focuses on social alignment (represented by the letter a). For example, when walking next
to a partner synchronizing walking speed may signal alignment and social affiliation. In this case, matching walking
speed is not a mere motor control problem, but a social interaction task. In the predictive coding formulation, different
speeds of walking may represent different world states. The social value of walking speed a will be captured by Q1. If I
walk too fast, I will exceed my partner and there will be a gap between us, resulting in negative social signal V1. This
gap will be used to lower my valuation of walking speed Q1, leading me to choose another walking speed with higher
social value (e.g., Q2).

It has been suggested [113] that, when we use predictive coding to understand social processes, the prior prediction
may encompass beliefs, goals, emotional states, and preceding social experiences. In our context, this means that
Qt1(a) is based not only on the current social alignment process that is taking place but also on all prior social alignment
experiences, across all levels of social alignment (motion, emotion, and cognition). We need to be able to generate
predictions about the social consequences of our actions, evaluate, compare them to our predictions, update our
model, and adapt our future behavior. These components are used in multiple levels and across levels of social
alignment. It is also possible that the prior prediction may be different about an ingroup compared to an outgroup, about
physically close people versus distant people, etc. This may help to explain why people tend to synchronize selectively.

constantly generates predictions based on previous information (top-down signals) to antici-

pate sensory information from within the body and in the environment. When the incoming
sensory information about oneself and about the environment (bottom-up signal) is different
from the prediction, this constitutes a prediction error. The brain can then either take action to
change the incoming sensory information, or alternatively update the prediction based on the
bottom-up information, such that over time the brain would minimize prediction errors [81].

The predictive coding approach is increasingly implicated in models of social processes. It

suggests that processing social information is associated with a nonlinear and probabilistic
combination of social signals and internal signals. For example, social comparison of perfor-
mance in a task involves monitoring one’s own achievements to those of others, comparing
them, updating the relative performance whenever new evidence becomes available, and
changing one’s behavior accordingly [82]. This may suggest that alignment helps the brain to
minimize prediction errors during social interactions [83,84].

We propose here that, during social interactions, a predictive coding mechanism is geared to
maximize alignment in movement, emotion, and cognitions with others. This propensity
towards alignment may be rooted in the reward associated with social alignment which
has evolved either through hereditary genetic predisposition [85] or through reinforcement
learning about the association between connectedness and rewards [86,87]. Therefore, the
individual continuously monitors self and others, and creates predictions regarding the

180 Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3

expected alignment (Figure 3). Detecting misalignment increases prediction error (gap-detec-
tion system) and drives alignment (the OE system). Achieving alignment fulfils the predictive
model goal and activates the reward system (the alignment reward system). This indicates that
alignment with others serves as the optimization principle, and achieving alignment reduces
prediction errors, and makes the environment more predictable, thereby lowering the compu-
tational cost of social processing (also referred to as ‘minimizing free energy’) [84].

Crucially, we propose that the same predictive coding principle operates across all levels of
social alignment, in a manner that each level affects the other. For example, motor signals may
possibly be used to infer cognitive or emotional misalignment, and emotional signals can be
used to update motor predictions (Box 1). A predictive coding framework that spans all the
levels of social alignment may explain why the various levels of herding are behaviorally linked.
Finally, the predictive coding model can also help in explaining why the different levels of
alignment are all affected by social factors such as group membership as well as level of
psychological closeness and liking. In our model, the sensation of reward is central, and if over
the years aligning with outgroup members, people I do not like, or with people I do not feel close
to did not result in a positive outcome, I would learn not to do so. Moreover, as detailed in Box 1,
prior predictions are the heart of predictive coding. This is another crucial mechanism that can
potentially explain why we tend to synchronize selectively: as a result of a learning process, our
prior predictions about an ingroup compared to an outgroup, or about physically close people
versus distant ones, etc., may be different.

Herding-Related Psychopathology
Although herding is a ubiquitous and naturally occurring phenomenon, there are some con-
ditions where some impairment leads to a reduced capacity and/or motivation participate in
herding.

For example, individuals with autism spectrum disorders (ASDs) show aberrant performance at
all levels of social alignment, including motor synchronization [88], emotional contagion [89],
and conformity [90]. Recent studies have used Bayesian predictive coding approaches to
explain the social interaction deficits that individuals with ASD suffer from. They point to the
potential role of attenuated priors, also referred to as hypo-priors [91], or to an aberrant
precision associated with sensory input compared with prior beliefs [92]. In the context of
social alignment, this divergent predictive style was conceptualized as part of a dialectical
misattunement process [93].

It is therefore possible that individuals with ASD demonstrate a low reliance on the prior
prediction geared towards alignment maximization. Interestingly, although numerous studies
show dysfunction in various brain structures, as well as in activity and connectivity patterns in
ASD, to date no single atypical brain structure was found in individuals with ASD [94].
Nonetheless, previous work suggests that abnormal activation in regions related to the
gap-detection system during the processing of social stimuli among individuals with ASD
might underlie their behavioral difficulties in predictive coding of the intentions of others [95].

Another condition that may involve herding related pathology is loneliness. Loneliness is defined
as a form of subjective social pain that evolved to signal that social connections are weak, and
motivate their repair [96]. Lonely people demonstrate poorer empathy [97], indicative of
impaired emotional contagion. Findings with regard to conformity seem to be conflicting, with
some studies showing that lonely individuals are high in conformity [98] and other studies
showing opposite results [99]. Studies of brain structure and function showed that lonely

Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3 181

people have reduced grey matter density in the posterior STS [100], dmPFC, and IPL [101]. The Outstanding Questions
reward circuitry including the VS was also found to be relevant to loneliness [102,103], with a How can we measure social alignment
diminished level of activation in response to social stimuli. Not being able to connect with others in real life? Designing experimental set-
ups to study alignment in groups of
at various levels can result in a lowered sense of affinity and connection, as well as in less humans is challenging. Accessible
engagement and satisfaction from the interaction. Future research will be necessary to paradigms for social alignment in eco-
determine whether lonely people suffer from deficiencies in their capacity to achieve or enjoy logical environments need to be devel-
oped. New methods based on virtual
alignment.
reality environments can also allow
measuring social alignment in semi-
Concluding Remarks and Future Directions realistic contexts, where experiment-
Building on emerging research from multiple fields, the present social alignment model offers an ers can systematically manipulate the
behavior of other participants.
integrative framework for understanding the mechanisms underlying herding. We demonstrate
how the different behavioral manifestations of herding are linked, adhere to similar principles,
What are the neurochemical bases of
and rely on a feedback loop of three networks. Although the three ‘core’ systems of the loop herding? We need better means to
participate in all types of social alignment, each component may play a more dominant role measure different neurotransmitters
depending on the specific type of alignment. For example, it is likely that, because movement involved in herding. Positron-emission
tomography (PET) imaging with radio-
synchrony involves more effort to physically align one’s behavior over time, the OE component
labeled ligands, psychopharmacologi-
is more frequently reported in studies on joint action. cal studies, and behavioral genetic
techniques may further unveil the neu-
It is worth noting that the brain regions detailed here should be considered with caution rochemical bases of social alignment.

because the brain areas suggested to participate in the alignment model have been
What are the neural mechanisms
associated with many distinct functions. For example, the IFG plays a role not only in underlying the negative consequences
alignment but also in language and inhibition [104]. Furthermore, it is likely that regions that of herding? The study of synchrony
mediate self–other distinction (e.g., temporoparietal junction) may also be involved in social mostly focuses on the positive aspect
alignment. In addition, the executive control network would also be highly relevant for of this behavior. Future studies may
examine how herding can negatively
alignment because it involves continuous monitoring of one’s state relative to others [78]. It affect social behavior such as group-
is thus possible that the core systems constitute a feedback loop which is the basis of think and mob behavior.
alignment, whereas an extended system is activated depending on the specific require-
ments of the context. How does dysfunctional herding con-
tribute to the etiology and maintenance
of psychiatric and clinical disorders,
The value of this model derives mainly from focusing on the common denominator between and how can an understanding of
multiple social behaviors. This implies that the multiple levels of alignment inform each other, herding be useful in treating them?
with information about alignment in movement, emotion, and cognition driving further adjust-
ments to achieve alignment across all levels. Future studies may examine patterns of correlation Are the terms herding and alignment
interchangeable? The concepts of
between the degree of synchronization in motor, emotion, and cognitive behaviors for given
‘herding’ and ‘alignment’ need to be
individuals. New directions of studies can expand the model by exploring how this ‘herding better defined.
mode’ enables other behaviors, including cultural assimilation, fashion influences, and eco-
nomic decision making.

It should be noted that the proposed model assumes that interpersonal synchrony is beneficial
to humans and to other social species [85]. Although there is ample support for this view, it also
has some limitations. For example, it was suggested that synchronization may not always result
in improved performance, and that its impact may be selective, based on the task at hand [105].
Considering a related construct, it was recently claimed that mimicry does not always lead to
positive results (such as closeness), and that when mimicry is not expected, it often leads to
opposite reactions [106]. Other possible negative societal consequences of herding may
include influence on risky behaviors (e.g., smoking in adolescence) [107] or mob violence
[108]. Behavioral synchrony might even increase compliance with the request to engage in
aggressive behavior [109]. Therefore, future research should use a variety of ecologically valid
social tasks, take into account the social context in which behavioral synchronization takes
place, and measure multiple outcome variables of synchronization. Additional research will also

182 Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3

 Box 2. Oxytocin (OT) and Herding
Social neuroendocrinology research investigating the influence of OT – a peptide hormone – on affiliation suggests that
the OT system regulates interpersonal synchrony [114], emotional contagion [115], and social conformity [116]. It has
been repeatedly found that both plasma OT and salivary OT in parents are associated with microlevel synchrony of
visuoaffective facial signals between the parent and the infant [117]. In line with a role of OT in synchrony, in a study on
movement synchrony it was found that intranasal administration of OT improved performance of pairs in a task that
required coordinated drawing [118]. Furthermore, it was recently reported [119] that intranasal administration of OT
improves rhythmical counting and enhances interbrain coupling of alpha-band oscillations during the coordination task,
further suggesting that OT enhances brain-to-brain synchrony and thus facilitates social coordination.

Crucially, the OE and the reward systems have been reported to be regulated by the OT system. For example, it was
reported that intranasal OT regulates responses in the IFG and IPL to vignettes describing situations of social synchrony
[120]. Furthermore, numerous studies have found that intranasal OT regulates the activity of the reward mechanism,
including the VS (e.g., Scheele et al. [121]), further suggesting that OT plays a major role in regulating the various types of
social alignment.

be necessary to articulate the conditions in which humans operate against this herding mode.
For example, situations that involve ‘out of the box’ original thinking evidently demand some
type of inhibition of this herding mode to allow the generation of uncommon responses.
Notably, models of creativity postulate that inhibiting core regions of the alignment OE system,
including the IFG, will unleash original thinking [110].

Future research may also use this integrated approach to test the neurochemical mechanisms
underlying herding. For example, the neurohormone oxytocin has been shown to regulate
synchronized movement as well as emotional contagion and social conformity (Box 2).
Furthermore, a computational approach may be used to create simulations of the proposed
model in various contexts that involve social alignment (Box 1). It may be particularly interesting
to examine interbrain networks in ‘real-life’ interactions with a computational approach, and
characterize the unique neural process that allows two or more individuals to mutually adjust
their behavior, cognitions, or emotions as well as the relationship between the different levels of
herding (see Outstanding Questions). This necessary methodological leap will put forward
novel questions on herding in an ecologically natural environment.

Acknowledgments
The authors thank Prof. Rachel Tomer and Dr Uri Hertz for their feedback on an earlier version of this manuscript, and
Rotem Perlmutter for her illustration of Figure 2. Completion of the manuscript was supported by grants 2510/16 from the
Israel Science Foundation (ISF)–Natural Science Foundation of China (NSFC), 959/18 from the ISF, I-142-105.3-2016 from
the German-Israeli Foundation for Scientific Research and Development (GIF), and 2015068 from the US-Israel Binational
Science Foundation (BSF) awarded to S.G.S.T.

References
1. Belz, M. et al. (2013) Spontaneous flocking in human groups.
Behav. Processes 92, 6–14
2. Kułakowska, K.A. et al. (2014) Using an individual-based model
to select among alternative foraging strategies of woodpigeons:
data support a memory-based model with a flocking mecha-
nism. Ecol. Modell. 280, 89–101
3. King, A.J. et al. (2012) Selfish-herd behaviour of sheep under
threat. Curr. Biol. 22, R561–R562
4. Conradt, L. and Roper, T.J. (2005) Consensus decision making
in animals. Trends Ecol. Evol. 20, 449–456
5. Sylos-Labini, F. et al. (2018) Human–human interaction forces
and interlimb coordination during side-by-side walking with
hand contact. Front. Physiol. 9, 179
6. Demos, A.P. et al. (2012) Rocking to the beat: effects of music
and partner’s movements on spontaneous interpersonal coor-
dination. J. Exp. Psychol. Gen. 141, 49–53
7. Varlet, M. et al. (2011) Social postural coordination. J. Exp.
Psychol. Hum. Percept. Perform. 37, 473–483
8. Raafat, R.M. et al. (2009) Herding in humans. Trends Cogn. Sci.
13, 420–428
9. Chartrand, T.L. and Lakin, J.L. (2013) The antecedents and
consequences of human behavioral mimicry. Annu. Rev. Psy-
chol. 64, 285–308
10. Keller, P.E. et al. (2014) Rhythm in joint action: psychological and
neurophysiological mechanisms for real-time interpersonal coor-
dination. Philos. Trans. R. Soc. Lond. B Biol. Sci. 369, 20130394
11. Prochazkova, E. and Kret, M.E. (2017) Connecting minds and
sharing emotions through mimicry: a neurocognitive model of
emotional contagion. Neurosci. Biobehav. Rev. 80, 99–114
12. Anderson, C.L. et al. (2018) Emotion in the wilds of nature: the
coherence and contagion of fear during threatening group-
based outdoors experiences. Emotion 18, 355–368

Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3 183

13. Krämer, N. et al. (2014) Let the weakest link go! Empirical
explorations on the relative importance of weak and strong ties
on social networking sites. Societies 4, 785–809
14. Golland, Y. et al. (2015) The mere co-presence: synchronization
of autonomic signals and emotional responses across co-pres-
ent individuals not engaged in direct interaction. PLoS One 10,
e0125804
15. Coultas, J.C. and van Leeuwen, E.J.C. et al. (2015) Conformity:
definitions, types, and evolutionary grounding. In Evolutionary
Perspectives on Social Psychology (Zeigler-Hill, V., ed.), pp.
189–202, Springer
16. Charpentier, C.J. et al. (2014) The brain’s temporal dynamics
from a collective decision to individual action. J. Neurosci. 34,
5816–5823
17. Edelson, M. et al. (2011) Following the crowd: brain substrates
of long-term memory conformity. Science 333, 108–111
18. Germar, M. et al. (2014) Social influence and perceptual decision
making. Personal. Soc. Psychol. Bull. 40, 217–231
19. Kundu, P. and Cummins, D.D. (2013) Morality and conformity:
the Asch paradigm applied to moral decisions. Soc. Influ. 8,
268–279
20. Meier, B.P. et al. (2012) Embodiment in social psychology. Top.
Cogn. Sci. 4, 705–716
21. Rossignac-Milon, M. et al. (2018) Epistemic companions:
shared reality development in close relationships. Curr. Opin.
Psychol. 23, 66–71
22. Leander, N.P. et al. (2010) Mind your mannerisms: behavioral
mimicry elicits stereotype conformity. J. Exp. Soc. Psychol. 47,
195–201
23. Seth Kaplan, L.B.-S. et al. (2009) Thinking inside the box: how
conformity promotes creativity and innovation. Creat. Groups
12, 229–265
24. Páez, D. et al. (2015) Psychosocial effects of perceived emo-
tional synchrony in collective gatherings. J. Pers. Soc. Psychol.
108, 711–729
25. Hawk, S.T. et al. (2012) Face the noise: embodied responses to
nonverbal vocalizations of discrete emotions. J. Pers. Soc.
Psychol. 102, 796–814
26. Barsade, S.G. (2002) The ripple effect: emotional contagion and
its influence on group behavior. Adm. Sci. Q. 47, 644
27. Miles, L.K. et al. (2009) Too late to coordinate: contextual influen-
ces on behavioral synchrony. Eur. J. Soc. Psychol. 40, 52–60
28. Gommans, R. et al. (2017) Popularity, likeability, and peer con-
formity: four field experiments. J. Exp. Soc. Psychol. 73, 279–
289
29. Qi, S. et al. (2018) A collaborator’s reputation can bias decisions
and anxiety under uncertainty. J. Neurosci. 38, 2337–17
30. Cirelli, L.K. (2018) How interpersonal synchrony facilitates early
prosocial behavior. Curr. Opin. Psychol. 20, 35–39
31. Han, S. (2018) Neurocognitive basis of racial ingroup bias in
empathy. Trends Cogn. Sci. 22, 400–421
32. Howard, L.H. et al. (2015) Infants’ and young children’s imitation
of linguistic in-group and out-group informants. Child Dev. 86,
259–275
33. Tarr, B. et al. (2016) Silent disco: dancing in synchrony leads to
elevated pain thresholds and social closeness. Evol. Hum.
Behav. 37
34. Rabinowitch, T.-C. and Knafo-Noam, A. (2015) Synchronous
rhythmic Interaction enhances children’s perceived similarity
and closeness towards each other. PLoS One 10, e0120878
35. Kühn, S. et al. (2011) Neural correlates of emotional synchrony.
Soc. Cogn. Affect. Neurosci. 6, 368–374
36. McKelvey, W. and Kerr, N.H. (1988) Differences in conformity
among friends and strangers. Psychol. Rep. 62, 759–762
37. Couzin, I.D. (2009) Collective cognition in animal groups. Trends
Cogn. Sci. 13, 36–43
38. Schilbach, L. et al. (2013) Toward a second-person neurosci-
ence. Behav. Brain Sci. 36, 393–414
184 Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3
39. Chang, L.J. and Sanfey, A.G. (2013) Great expectations: neural
computations underlying the use of social norms in decision-
making. Soc. Cogn. Affect. Neurosci. 8, 277–284
40. Somerville, L.H. et al. (2006) Anterior cingulate cortex responds
differentially to expectancy violation and social rejection. Nat.
Neurosci. 9, 1007–1008
41. Lieberman, M.D. and Eisenberger, N.I. (2015) The dorsal ante-
rior cingulate cortex is selective for pain: results from large-scale
reverse inference. Proc. Natl. Acad. Sci. U. S. A. 112, 15250–
15255
42. Mansouri, F.A. et al. (2017) Monitoring demands for executive
control: shared functions between human and nonhuman pri-
mates searching for neural substrates of context-dependent
control of goal-directed behavior. Trends Neurosci. 40, 15–27
43. Alexander, W.H. and Brown, J.W. (2015) Hierarchical error
representation: a computational model of anterior cingulate
and dorsolateral prefrontal cortex. Neural Comput. 27, 2354–
2410
44. Deuker, L. et al. (2013) Playing nice: a multi-methodological
study on the effects of social conformity on memory. Front.
Hum. Neurosci. 7, 79
45. Tomlin, D. et al. (2017) The integration of social influence and
reward: computational approaches and neural evidence. Cogn.
Affect Behav. Neurosci. 17, 784–808
46. Cacioppo, S. et al. (2014) You are in sync with me: neural
correlates of interpersonal synchrony with a partner. Neurosci-
ence 277, 842–858
47. Fairhurst, M.T. et al. (2013) Being and feeling in sync with an
adaptive virtual partner: brain mechanisms underlying dynamic
cooperativity. Cereb. Cortex 23, 2592–2600
48. Morelli, S.A. et al. (2014) The neural bases of feeling understood
and not understood. Soc. Cogn. Affect. Neurosci. 9, 1890–
1896
49. Golland, Y. et al. (2017) Neural dynamics underlying emotional
transmissions between individuals. Soc. Cogn. Affect. Neurosci.
12, 1249–1260
50. Wittfoth, M. et al. (2010) On emotional conflict: interference
resolution of happy and angry prosody reveals valence-specific
effects. Cereb. Cortex 20, 383–392
51. Thornton, I.M. and Knoblich, G. (2006) Action perception: see-
ing the world through a moving body. Curr. Biol. 16, R27–R29
52. Iacoboni, M. et al. (2001) Reafferent copies of imitated actions in
the right superior temporal cortex. Proc. Natl. Acad. Sci. U.S.A.
98, 13995–13999
53. Lestou, V. et al. (2008) Neural substrates for action understand-
ing at different description levels in the human brain. J. Cogn.
Neurosci. 20, 324–341
54. Mainieri, A.G. et al. (2013) Differential role of the mentalizing and
the mirror neuron system in the imitation of communicative
gestures. Neuroimage 81, 294–305
55. Wade, S. and Hammond, G. (2015) Anodal transcranial direct
current stimulation over premotor cortex facilitates observational
learning of a motor sequence. Eur. J. Neurosci. 41, 1597–1602
56. Pilgramm, S. et al. (2009) The role of own-body representations
in action observation: a functional MRI study. Neuroreport 20,
997–1001
57. Jabbi, M. et al. (2008) A common anterior insula representation
of disgust observation, experience and imagination shows
divergent functional connectivity pathways. PLoS One 3,
e2939
58. Rizzolatti, G. and Sinigaglia, C. (2016) The mirror mechanism: a
basic principle of brain function. Nat. Rev. Neurosci. 17, 757–
765
59. Hasson, U. et al. (2016) Mirroring and beyond: coupled dynam-
ics as a generalized framework for modelling social interactions.
Philos. Trans. R. Soc. Lond. B Biol. Sci. 371
60. Jasmin, K.M. et al. (2016) Cohesion and joint speech: right
hemisphere contributions to synchronized vocal production.
J. Neurosci. 36, 4669–4680
61. Nummenmaa, L. et al. (2008) Is emotional contagion special? An
fMRI study on neural systems for affective and cognitive empa-
thy. Neuroimage 43, 571–580
62. Dvash, J. and Shamay-Tsoory, S.G. (2014) Theory of mind and
empathy as multidimensional constructs. Top. Lang. Disord. 34,
282–295
63. Shamay-Tsoory, S.G. (2011) The neural bases for empathy.
Neuroscience 17, 18–24
64. Lamm, C. et al. (2011) Meta-analytic evidence for common and
distinct neural networks associated with directly experienced
pain and empathy for pain. Neuroimage 54, 2492–2502
65. Izuma, K. and Adolphs, R. (2013) Social manipulation of prefer-
ence in the human brain. Neuron 78, 563–573
66. Holper, L. et al. (2012) Between-brain connectivity during imita-
tion measured by fNIRS. Neuroimage 63, 212–222
67. Jiang, J. et al. (2012) Neural synchronization during face-to-face
communication. J. Neurosci. 32, 16064–16069
68. Osaka, N. et al. (2015) How two brains make one synchronized
mind in the inferior frontal cortex: fNIRS-based hyperscanning
during cooperative singing. Front. Psychol. 6, 1811
69. Hu, Y. et al. (2017) Brain-to-brain synchronization across two
persons predicts mutual prosociality. Soc. Cogn. Affect. Neuro-
sci. 12, 1835–1844
70. Dumas, G. et al. (2010) Inter-brain synchronization during social
interaction. PLoS One 5, e12166
71. Dikker, S. et al. (2017) Brain-to-brain synchrony tracks real-
world dynamic group interactions in the classroom. Curr. Biol.
27, 1375–1380
72. Konvalinka, I. et al. (2014) Frontal alpha oscillations distinguish
leaders from followers: multivariate decoding of mutually inter-
acting brains. Neuroimage 94, 79–88
73. Goldstein, P. et al. (2018) Brain-to-brain coupling during hand-
holding is associated with pain reduction. Proc. Natl. Acad. Sci.
U. S. A. 115, E2528–E2537
74. Knutson, B. and Cooper, J.C. (2005) Functional magnetic res-
onance imaging of reward prediction. Curr. Opin. Neurol. 18,
411–417
75. Kokal, I. et al. (2011) Synchronized drumming enhances activity
in the caudate and facilitates prosocial commitment – if the
rhythm comes easily. PLoS One 6, e27272
76. Noy, L. et al. (2015) Being in the zone: physiological markers of
togetherness in joint improvisation. Front. Hum. Neurosci. 9,
187
77. Krill, A.L. and Platek, S.M. (2012) Working together may be
better: activation of reward centers during a cooperative maze
task. PLoS One 7, e30613
78. Prehn, K. et al. (2015) The neural correlates of emotion align-
ment in social interaction. Soc. Cogn. Affect. Neurosci. 10, 435–
443
79. Campbell-Meiklejohn, D.K. et al. (2010) How the opinion of
others affects our valuation of objects. Curr. Biol. 20, 1165–
1170
80. Wu, H. et al. (2016) Neural signatures of social conformity: a
coordinate-based activation likelihood estimation meta-analysis
of functional brain imaging studies. Neurosci. Biobehav. Rev.
71, 101–111
81. Spratling, M.W. (2017) A review of predictive coding algorithms.
Brain Cogn. 112, 92–97
82. Hertz, U. et al. (2017) Neural computations underpinning the
strategic management of influence in advice giving. Nat. Com-
mun. 8, 2191
83. Bolis, D. and Schilbach, L. (2017) Beyond one Bayesian brain:
modeling intra-and inter-personal processes during social inter-
action: commentary on ‘Mentalizing homeostasis: the social
origins of interoceptive inference’ by Fotopoulou & Tsakiris.
Neuropsychoanalysis 19, 35–38
84. Koban, L. et al. (2017) Why do we fall into sync with others?
Interpersonal synchronization and the brain’s optimization prin-
ciple. Soc. Neurosci. 14, 1–9
85. Duranton, C. and Gaunet, F. (2016) Behavioural synchronization
from an ethological perspective: overview of its adaptive value.
Adapt. Behav. 24, 181–191
86. Atzil, S. et al. (2018) Growing a social brain. Nat. Hum. Behav. 2,
624–636
87. Cacioppo, S. and Cacioppo, J.T. (2012) Decoding the invisible
forces of social connections. Front. Integr. Neurosci. 6, 51
88. Cheng, M. et al. (2017) Gender and autistic traits modulate
implicit motor synchrony. PLoS One 12, e0184083
89. Adler, B.A. et al. (2015) Drug-refractory aggression, self-injuri-
ous behavior, and severe tantrums in autism spectrum disor-
ders: a chart review study. Autism 19, 102–106
90. Yafai, A.-F. et al. (2014) Social conformity and autism spectrum
disorder: a child-friendly take on a classic study. Autism 18,
1007–1013
91. Pellicano, E. and Burr, D. (2012) When the world becomes ‘too
real’: a Bayesian explanation of autistic perception. Trends
Cogn. Sci. 16, 504–510
92. Lawson, R.P. et al. (2014) An aberrant precision account of
autism. Front. Hum. Neurosci. 8, 302
93. Bolis, D. et al. (2017) Beyond autism: introducing the dialectical
misattunement hypothesis and a Bayesian account of intersub-
jectivity. Psychopathology 50, 355–372
94. Waterhouse, L. et al. (2016) ASD validity. Rev. J. Autism Dev.
Disord. 3, 302–329
95. von der Lühe, T. et al. (2016) Interpersonal predictive coding, not
action perception, is impaired in autism. Philos. Trans. R. Soc.
Lond. B. Biol. Sci. 371, 20150373
96. Cacioppo, J.T. et al. (2006) Loneliness within a nomological net:
an evolutionary perspective. J. Res. Pers. 40, 1054–1085
97. Beadle, J.N. et al. (2012) Trait empathy as a predictor of indi-
vidual differences in perceived loneliness. Psychol. Rep. 110, 3–
15
98. Mehrabian, A. and Stefl, C.A. (1995) Basic temperament com-
ponents of loneliness, shyness, and conformity. Soc. Behav.
Personal. Int. J. 23, 253–263
99. Hansson, R.O. and Jones, W.H. (1981) Loneliness, cooperation,
and conformity among American undergraduates. J. Soc. Psy-
chol. 115, 103–108
100. Kanai, R. et al. (2012) Brain structure links loneliness to social
perception. Curr. Biol. 22, 1975–1979
101. Nakagawa, S. et al. (2015) White matter structures associated
with loneliness in young adults. Sci. Rep. 5, 17001
102. Cacioppo, J.T. et al. (2009) In the eye of the beholder:
individual differences in perceived social isolation predict
regional brain activation to social stimuli. J. Cogn. Neurosci.
21, 83–92
103. Inagaki, T.K. et al. (2015) Yearning for connection? Loneliness is
associated with increased ventral striatum activity to close
others. Soc. Cogn. Affect. Neurosci. 11, 1096–1101
104. Liakakis, G. et al. (2011) Diversity of the inferior frontal gyrus: a
meta-analysis of neuroimaging studies. Behav. Brain Res. 225,
341–347
105. Wallot, S. et al. (2016) Beyond synchrony: joint action in a
complex production task reveals beneficial effects of decreased
interpersonal synchrony. PLoS One 11, e0168306
106. Hale, J. and Hamilton, A.F. (2016) Cognitive mechanisms for
responding to mimicry from others. Neurosci. Biobehav. Rev.
63, 106–123
107. Liu, J. et al. (2017) The influence of peer behavior as a function of
social and cultural closeness: a meta-analysis of normative
influence on adolescent smoking initiation and continuation.
Psychol. Bull. 143, 1082–1115
108. Russell, G.W. (2004) Sport riots: a social–psychological review.
Aggress. Violent Behav. 9, 353–378
109. Wiltermuth, S.S. (2012) Synchronous activity boosts compli-
ance with requests to aggress. J. Exp. Soc. Psychol. 48,
453–456
Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3 185
 110. Mayseless, N. et al. (2015) Generating original ideas: the neural
underpinning of originality. Neuroimage 116, 232–239
111. Sutton, R.S. and Barto, A.G. (1998) Reinforcement Learning: An
Introduction, MIT Press
112. Knill, D.C. and Pouget, A. (2004) The Bayesian brain: the role of
uncertainty in neural coding and computation. Trends Neurosci.
27, 712–719
113. Otten, M. and Pinto, Y. (2017) A social Bayesian brain: how
social knowledge can shape visual perception. Brain Cogn. 112,
69–77
114. Feldman, R. and Bakermans-Kranenburg, M.J. (2017) Oxytocin:
a parenting hormone. Curr. Opin. Psychol. 15, 13–18
115. Boysza, M. (1961) Children’s rheumatism sanatorium in Wie-
niecspa, a state health resort. Reumatologia 4, 149–153
116. Stallen, M. et al. (2012) The herding hormone. Psychol. Sci. 23,
1288–1292
186 Trends in Cognitive Sciences, March 2019, Vol. 23, No. 3
View publication stats
117. Gordon, I. et al. (2010) Oxytocin, cortisol, and triadic family
interactions. Physiol. Behav. 101, 679–684
118. Arueti, M. et al. (2013) When two become one: the role of
oxytocin in interpersonal coordination and cooperation. J. Cogn.
Neurosci. 25, 1418–1427
119. Mu, Y. et al. (2016) Oxytocin enhances inter-brain synchrony
during social coordination in male adults. Soc. Cogn. Affect.
Neurosci. 11, 1882–1893
120. Levy, J. et al. (2016) Oxytocin selectively modulates brain
response to stimuli probing social synchrony. Neuroimage
124, 923–930
121. Scheele, D. et al. (2013) Oxytocin enhances brain reward sys-
tem responses in men viewing the face of their female partner.
Proc. Natl. Acad. Sci. U. S. A. 110, 20308–20313