PeRSpeCTIveS

time. Building on this model, we make

The default mode network: three related arguments. First, we propose
that activity in the DMN is modulated by

where the idiosyncratic self meets incoming external information (top arrow
in Fig. 1a), which is integrated over many

the shared social world seconds to many minutes (horizontal arrow

in Fig. 1a). Therefore, the DMN should
not be considered an intrinsic system
Yaara Yeshurun, Mai Nguyen and Uri Hasson active only during internally oriented or
stimulus-​independent processes. Second, we
Abstract | The default mode network (DMN) is classically considered an ‘intrinsic’ argue that activity in the DMN is nonetheless
system, specializing in internally oriented cognitive processes such as sensitive to intrinsic information (long-​term
memories, conditional responses, beliefs,
daydreaming, reminiscing and future planning. In this Perspective, we suggest that
emotions and so on), which influences
the DMN is an active and dynamic ‘sense-​making’ network that integrates and interacts with the interpretation and
incoming extrinsic information with prior intrinsic information to form rich, context-​ processing of incoming external information
dependent models of situations as they unfold over time. We review studies that (bottom arrow in Fig. 1a). Therefore, the
relied on naturalistic stimuli, such as stories and movies, to demonstrate how an DMN should not be considered an extrinsic
individual’s DMN neural responses are influenced both by external information system, sensitive only to incoming sensory
information. Last, we combine these points
accumulated as events unfold over time and by the individual’s idiosyncratic past to propose that the DMN is an active and
memories and knowledge. The integration of extrinsic and intrinsic information dynamic ‘sense-​making’ network that
over long timescales provides a space for negotiating a shared neural code, which integrates incoming extrinsic information
is necessary for establishing shared meaning, shared communication tools, shared with prior intrinsic information over long
narratives and, above all, shared communities and social networks. timescales to form rich, context-​dependent,
idiosyncratic models of the situation as it
unfolds over time.
Making sense of the world is a dynamic, includes the posterior medial cortex, medial As the activity in the DMN is shaped
ever-​changing process requiring the prefrontal cortex and temporoparietal by our unique history, it is by nature
integration of incoming information with junction1–3, was classically viewed as an idiosyncratic. However, at the same time, our
prior knowledge about the situation over ‘intrinsic’ system, showing decreased knowledge, memories and beliefs are shaped
multiple timescales. For example, while activity during experimental tasks. Later by the people we are connected to and the
listening to a debate, a listener must integrate work updated this view to suggest that the world in which we are immersed (Fig. 1b).
their own prior thoughts and beliefs about DMN is specialized for internally oriented Thus, in the last section of this Perspective,
the debate topics with the current and or non-​stimulus-​induced tasks, such as we argue that the DMN provides a space
past statements of the speakers. Thus, our ‘mind-​wandering’, recall of past events for social ‘others’ (the extrinsic people we
mindsets and our actions at each moment and internal simulation of future events4–7. interact with) to shape the self (our set of
in time are shaped by at least three factors: Although a recent review further divided the intrinsic memories and beliefs), which in
what is happening now in the world DMN into multiple interleaved networks1, turn can enable us to shape the memories
(incoming sensory input); what happened in researchers have often made a distinction and beliefs of others. This unique interplay
the world in the preceding moments (recent between this ‘intrinsic’ system and ‘extrinsic’ between the extrinsic and intrinsic forces
active memories that further contextualize systems that process incoming sensory provides a mechanism for negotiating a
and shape the present); and who we are and information or outgoing motor outputs4,5,8–14 shared neural code to facilitate learning and
what we bring to the moment (that is, our (see Box 1). communication via shared common ground.
long-​term memories, conditional responses, Here, we challenge the view of the Our view of the DMN is informed by
beliefs and emotions that shape the way we DMN as an intrinsic system by pointing to numerous studies using naturalistic stimuli
process the incoming inputs). The ongoing the active and dynamic role of the DMN and inter-​subject analyses that map shared
integration of these three factors happens in synthesizing extrinsic and intrinsic neural responses across participants (Fig. 2).
effortlessly in our brains, almost by default. information over long timescales. In a
In this Perspective, we propose a previous review, we argued that memory Extrinsic responses of the DMN
key role for the default mode network is an integral component of any processing We begin by proposing that the DMN
(DMN) in dynamically integrating these act15 and that the DMN resides at the top of integrates moment-​by-​moment external
three factors over extended timescales in this ‘process–memory’ hierarchy, playing information with prior information over
order to make sense of a novel, constantly a unique and central role in processing seconds to minutes, as events unfold
evolving situation. The DMN, which events as they dynamically unfold over over time.

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Perspectives

DMN activity is influenced by context of visual integration across the visual cor- neural processing of new information.
and incoming inputs. Research in visual and tex is captured by spatial receptive fields: Studies using naturalistic stimuli have
auditory perception has revealed the influ- early visual areas process local parts of the identified a topographical hierarchy of
ence of context on neural activity even in visual field and so have small spatial recep- processing timescales extending from
early sensory areas. In the primary visual tive fields, whereas higher-​order visual areas early sensory cortex to higher-​order areas
cortex of non-​human primates, for example, integrate information over larger parts of including the DMN7,20–23. TRWs are short
the response of a simple cell to a vertical line the visual field and thus have larger spatial in sensory areas and become gradually
will change as a function of lines in its extra-​ receptive fields. longer towards higher-​order areas. The
visual field, or its spatial context16,17. Regions Analogously, temporal receptive windows responses in early sensory areas such as
in higher-​order cortex then integrate these (TRWs) capture the influence of context the early auditory and visual cortex are
responses, resulting in sensitivity over over time. The TRW is the window of influenced by the temporal structure over
larger spatial contexts18,19. This hierarchy time in which past information affects tens of milliseconds20,21. These findings are
consistent with research22 suggesting that
Box 1 | Alternative accounts of default mode network function early sensory areas process fast-​changing,
low-​level stimulus features, such as sound
The function of the default mode network (DMN) is a matter of substantial interest and debate. amplitude or visual edges. By contrast,
In this box, we aim to highlight several prominent lines of studies that expose the richness
neural responses in mid-​level areas, which
and diversity of functions associated with the DMN. Furthermore, we aim to outline how such
diversity of functions can be unified by our theoretical perspective. include linguistic regions along temporal
cortex, are affected by information
Baseline activation for healthy function integrated over a few seconds (for example,
The DMN was initially conceptualized as a ‘task-​negative’ network, active only when the brain is
the preceding words in the sentence).
not occupied by a task4,8,123,124. Later, it was discovered that regions of the DMN show spontaneous
fluctuations in activity that are highly correlated across regions, suggesting that the DMN forms a
At the apex of the processing hierarchy,
functional network11. These findings led to the hypothesis that the activity and connectivity of the the neural response of the DMN to each
DMN have an important role in the offline intrinsic activity that is required to maintain balanced sentence is influenced by prior information
and steady internal states during rest2,125. accumulated over many minutes15,20,21,23.
Mind-​wandering
In addition to the modulation of DMN
Follow-​up studies aimed to further characterize the cognitive functions associated with periods of activity by information accumulated
rest, or when the mind is disengaged from processing external information. As a result, researchers over many seconds, in the next section
found that DMN activity is increased during periods of ‘mind-​wandering’126–129 and that an we discuss evidence for the modulation
individual’s tendency to mind wander is correlated with the DMN response130. Moreover, the of DMN connectivity by information
content of mind-​wandering may modulate DMN activity131 (for example, thinking about what one accumulated over many seconds.
needs to advance their research project versus thinking about how one’s bedroom looks) and the
connectivity of the DMN is different for different components of mind-​wandering (for example, DMN connectivity is influenced by context
memory, social cognition and planning)132. and incoming inputs. To map DMN
Autobiographical memory functional connectivity, analyses sample
The first study to detect DMN responses associated with internally generated autobiographical the temporal response in one node of the
memories in the absence of external stimuli was exploring brain mechanisms underlying personal DMN and use this as a seed to search
memory133. Since then, various neuroimaging studies have demonstrated that DMN activity and for co-​varying fluctuations in neural
connectivity are modulated by tasks that require the retrieval of past events and by spontaneous
activity across other nodes of the network
autobiographical memory128,134–136. It was suggested that autobiographical memory is one of the
within an individual brain (Fig. 3a). This
prominent processes that shows the involvement of the DMN in mental representations generated
using intrinsic information, independent of extrinsic information1. analysis, although effective in outlining
the anatomical boundaries of the DMN,
Prospective memory is unable to distinguish subtle task-​related
In addition to the retrieval of episodic memories, participants are often occupied with thinking
fluctuations in connectivity across various
about future events and planning ahead while resting in the MRI scanner137. Intriguingly, memory
‘for the future’, like memory for the past, activates the DMN and shapes its connectivity131,136,138,139. experimental conditions including, for
example, lying at rest in a scanner, listening
Social cognition to a list of scrambled words and listening to
Many studies have found extensive overlap between the DMN and regions involved in social
an intact story several minutes long24–26
cognition that are collectively known as the ‘social brain’140–148. In addition to being involved in
(Fig. 3a). The stability of the averaged
self-​referential processes such as thinking about self-​mental states149,150, the DMN is involved
in thinking about other people’s beliefs, intentions and motivations53,136,151, and in priming the connectivity maps across numerous
intentional stance152. Importantly, thinking about other people is not confined to moments of conditions has frequently been used to
disengagement but is also, and perhaps even primarily, evident as people are engaged with support the claim of the intrinsic nature of
other social agents in the real world, for example during telling (speaking) and listening to the DMN5,27. Such interpretation, however,
(comprehending) real-​life personal stories33,46,153 as well as watching and listening to fictional was called into question by the realization
stories37,42,47. that within-​brain functional connectivity
Where intrinsic meets extrinsic patterns are heavily influenced by direct
Taken together, the studies described here suggest that, on the one hand, the DMN is active during and indirect anatomical wiring, but are less
internal-​related thoughts that are self-​generated in the absence of external stimuli, as in the sensitive for isolating spontaneous activity
case of mind-​wandering, and therefore can be considered intrinsic; and, on the other, the response (for example, during rest) from stimulus-​
of the DMN is locked to external stimuli, especially during social interactions, and thus can be locked activity that propagates through the
considered extrinsic. Thus, in this Perspective, we combine many of these findings to suggest network as during processing of extrinsic
that the DMN is at the nexus of the interaction between the external and internal worlds.
information26,28.

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 Perspectives

Task-​related connectivity patterns Process memory

can be isolated from spontaneous neural Active integration Active (recurrent) memory
fluctuations by switching to a multi-​brain
perspective that characterizes the shared a DMN in integration of extrinsic and intrinsic information over long timescales
response across brains during the process- Sensory input
ing of temporally extended naturalistic (what is happening now)
stimuli. We recently developed such an Accumulation
over time
approach: inter-subject functional correla­
tion analysis26. Inter-​subject functional DMN
correlation differs from standard functional Active (recurrent) memory
connectivity analysis in one crucial way: Event start Present
it calculates the functional correlation
across brains, as opposed to within brains
(Fig. 2). Intrinsic neural dynamics during Accumulation
over time
rest, as well as non-​neuronal artefacts
LTM
(such as those associated with respiration (prior knowledge)
and head motion), can only influence the
pattern of correlations within each brain,
b Shared or coupled DMN activity as the basis for social cognition
but cannot induce correlations between
Speaker Listener
brains. By contrast, neural processes that are
locked to the structure of the stimulus and
shared across participants can be correlated
across brains. Start of Start of
Using inter-​subject functional Old self conversation conversation
Old self
cor­relation, we observed shared connectivity
patterns within the DMN that are locked
to the stimulus26 (Fig. 3b). Crucially, we
observed these stimulus-​locked connectivity
patterns only during processing of an intact,
coherent and temporally extended spoken
story. There are no shared connectivity LTM LTM
patterns during rest or while participants
listened to temporally incoherent stimuli
such as scrambled words (Fig. 3b). The c Regions of the DMN
finding of stimulus-​locked responses in TPJ dmPFC Prec/PCC MFG IPL
Prec
the DMN only for intact, but not temporally
scrambled, narratives highlights the
importance of integration of information
over long timescales in the DMN. This
temporal dependency over long timescales vmPFC PCC mPFC
MTG
makes the DMN more context-​dependent
than early sensory brain regions, in which RH LH
responses are less affected by temporal
structure of stimuli over such long Fig. 1 | A new view of the default mode network. a | Activity in the default mode network (DMN) is
timescales. These findings also highlight the modulated by incoming external information (top arrow), which is actively accumulated (grey expand-
importance of studying the brain’s dynamics ing triangle) and integrated (red circle) over hundreds of seconds (horizontal arrow) with our intrinsic
information (long-​term memories (LTMs), conditional responses, beliefs and so on, represented by the
under temporally extended, ecological
bottom arrow) to form a rich, context-​dependent, dynamic model of the unfolding situation. b | Our
conditions29. thoughts, feelings and actions are constantly being shaped by the actions, memories and stories of
others. At the same time, our LTMs shape the way we process the external input. This unique interplay
Integration in the DMN depends on the between the extrinsic and intrinsic forces provides a space for negotiating a shared neural code nec-
content, not form, of external input. DMN essary for establishing shared meanings, shared communication tools, shared narratives and, impor-
responses are locked to the content of tantly, shared communities and social networks. c | Regions of the DMN defined by functional
the stimuli over long timescales. At the connectivity analysis. These regions include the posterior cingulate cortex (PCC) and precuneus (Prec),
same time, they are relatively invariant the ventromedial prefrontal cortex (vmPFC) and dorsomedial prefrontal cortex (dmPFC), and the bilat-
to even dramatic changes in low-​level eral temporoparietal junction (TPJ). IPL, inferior parietal lobule; LH, left hemisphere; RH, right
stimulus properties that do not change the hemisphere; MFG, middle frontal gyrus; MTG, middle temporal gyrus. Part c adapted from ref.26,
CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/).
interpretation (meaning) of the stimuli.
Despite substantial differences in low-​
level properties, the DMN shows similar movies versus verbal descriptions32–34; abstract Heidel and Simmel-​like shape
temporal responses to the same narrative different synonyms35; different languages, animation versus concrete verbal description
presented using: different modalities, such as such as English versus Russian36; and of the same story37. A similar pattern of
written text versus speech30,31 or audiovisual different levels of abstraction, such as results was identified in the DMN using

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Perspectives

a Temporal ISC
Voxel xA Voxel xB

Inter-subject functional correlation matrix

0.3

0.2

Participant A Participant B ISC for voxel x 0.1

Correlation (r)
0.0
b Temporal inter-subject functional correlation
Voxel xA –0.1
Voxel xB
Voxel yB
–0.2
Voxel zB

Voxels
–0.3
Voxels
ISC map
(diagonal)
Inter-subject functional
Participant A Participant B correlation map for seed voxel x

c Spatial ISC
Spatial ISCs across time

ti–2 0.3

ti–1
ti 0.2
ti

0.1

Correlation (r)
Spatial ISC at time i 0.0

–0.1
Time points

Participant A Participant B –0.2

ISC of spatially distributed response patterns

–0.3
Time points
d Speaker–listener neural coupling model Inter-subject pattern correlation
Lag –2 for each time point (diagonal)
Lag –1
Lag 0
Lag 1
Lag 2

Speaker Listener

encoding models across written and spoken a different modality. Other researchers have in integrating information across sensory
narratives38. In these studies, a model trained trained decoding models, showing that the modalities40; for example, the precuneus
to predict neural responses to a narrative in specific story a participant is reading can be and bilateral temporoparietal junction have
one modality accurately predicted neural decoded from DMN neural responses, even been shown to integrate sensory and motor
responses to the same narrative presented in across languages39. The DMN also has a role information to represent word concepts41.

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 Perspectives

◀ Fig. 2 | Using inter-subject correlation analysis to map shared responses across subjects. Information acquired prior to events can
Inter-​subject correlation (ISC) analyses allow researchers to measure neural responses that are shared substantially change their interpretations.
across participants and locked to the structure of the external input. ISC can be computed in the In one intriguing study47, two groups of
following ways (for more details, see ref.122). a,b | Temporal ISC within and between brain areas. In this participants were presented with the same
basic form, the neural response to a continuous natural stimulus (as a story or a movie) in a given brain
short story by J. D. Salinger, Pretty Mouth
area is correlated with the responses in the same brain region in another participant (part a). This
simple analysis allows us to measure the consistency of neural responses to complex naturalistic stim-
and Green My Eyes, in which a husband
uli and to identify neural responses shared across individuals. The ISC method can be extended to calls his friend at night asking for the
correlate the response time course between each voxel in one participant’s brain and all other voxels in whereabouts of his wife. Before listening
another participant’s brain to yield a voxel by a voxel inter-subject functional correlation matrix (part b). to the story, the two groups were biased
The diagonal values of this matrix reflect the conventional ISC map in which correlations are com- with different contextual information.
puted only between homologous targets in the participants’ brains. A single value on the diagonal One group was led to believe that the
corresponds to the ISC for a given area. A single column (or row) of this matrix represents the func- wife was unfaithful, whereas the other
tional connectivity map for one seed voxel. c | Inter-​subject pattern correlation. ISCs can also be com- group was led to believe that the husband
puted across spatially distributed response patterns. For a given region, the correlation between was paranoid and jealous. DMN regions
voxels’ response patterns at a given moment (as illustrated by the coloured squares) is correlated
showed stronger alignment of neural
across participants. Inter-​subject pattern correlations can also be computed across time points to
capture the evolution of response patterns over time (for example, if a particular pattern recurs at responses within a group that shared an
multiple time points). d | Speaker–listener neural coupling model. ISCs can also be extended to model interpretation than between groups with
the direct interaction between a speaker’s brain and a listener’s brain during communication (that is, differing interpretations. This suggests
not induced by shared external input). The speaker–listener coupling model uses the speaker’s brain that participants’ prior beliefs (acquired
activity as a model for predicting the brain activity in each listener. To capture the temporal structure just before the story started) changed the
of the speaker–listener interaction, the speaker’s time courses are shifted by time intervals backwards way the DMN processed the same story,
and forwards relative to the moment of vocalization. These shifted speaker time courses are combined which unfolded over many minutes. Similar
with linear weights to build a predictive model for the listener brain dynamics. This method enabled findings have been observed following
investigation of whether the speaker’s neural activity during speech production is coupled over time manipulations of perspective48,49 or focus50,
with the listener’s neural activity during speech comprehension. Adapted with permission from ref.122,
as well as when context has led participants
CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/).
to interpret a situation as ironic51, reflecting
an attempt to ‘save face’52 or intentionally
Nevertheless, a relatively small change By manipulating attention to one or the harmful53. Moreover, a recent study suggests
to the stimulus can produce increasingly other modality, processing of unattended that regions in the DMN are involved in
large differences in neural responses across stimuli was shown to be primarily restricted reinstating the context of a storyline, which
the timescale hierarchy when it changes the to sensory cortex, whereas attended stimuli facilitates the convergence of the plot of two
interpretation of the stimulus. For example, went on to be processed by the DMN and storylines into one54. Together, these findings
manipulation of a small number of words dorsal attention networks. suggest that DMN activity is influenced not
in a spoken story (for example, changing Crucially, although attention can only by the structure of the incoming events,
‘laughing’ to ‘sobbing’), resulting in two modulate responses in the DMN, it cannot, but also by mindset and prior knowledge.
very different stories35, led to differences in isolation, account for the results described
in the neural responses elicited by the two above. Between-​subject multivoxel pattern The DMN is sensitive to long-​term
stories that correlated with the size of the analysis (Fig. 2) within each node of the memories. Context extends to the days and
TRW. As such, early auditory cortex showed DMN revealed that the multidimensional even years before an event, as information
relatively little difference in its response to spatial patterns of responses are aligned gathered throughout a lifetime is used to
the two stories, reflecting the relatively intact across participants watching a movie46. contextualize an event. For example, an
low-​level acoustic properties of the stimuli. Furthermore, using a shared response individual’s experience of watching episode
By contrast, DMN regions (which are on the model, the same study reported at least 5 in season 3 of the series Game of Thrones
top of the timescale hierarchy) showed large 20 orthogonal dimensions in the DMN’s both depends on and is influenced by
differences, reflecting the large difference in spatial response patterns that are shared internally stored knowledge of events and
story meaning. across participants, suggesting that no single characters in episodes that aired years ago.
univariate dimension, such as attention or Indeed, researchers have exploited stimuli
Integration in the DMN is modulated by arousal, can fully account for the complex such as television shows or stories to demo­
attention. Evidence suggests that the DMN multidimensional aligned responses in nstrate how the activity in the DMN is
is also modulated by attention. Stimuli that the DMN46. influenced by information acquired over days.
are more engaging elicit greater shared For example, one study55 used functional
neural responses across participants in Intrinsic responses of the DMN MRI (fMRI) to scan participants as they
DMN regions such as the medial prefrontal We next review research demonstrating watched the second half of an episode of the
cortex42, and DMN regions of listeners how the activity in the DMN is affected Twilight Zone. One group of participants
who are more engaged and attentive show not only by external events unfolding over watched the first half of the episode a
greater shared brain responses across many minutes but also by an individual’s day before they watched the second part
participants43,44. Intriguingly, attention may long-​term memories, schemas and belief (long-​term context); a second group
facilitate propagation of information along systems acquired over the years (Fig. 1a). watched the entire episode without a break
the cortical processing timescale hierarchy. (short-​term context); and a third group
In one study45, two unrelated narratives DMN responses are modulated by prior watched only the second half of the episode
were presented simultaneously, with one beliefs. Context is not narrowly confined (no context). Inter-​subject correlation in
in written form and one in spoken form. by the structure of the unfolding event. the DMN was greater among participants

NaTure RevIewS | NeUroScIence volume 22 | March 2021 | 185
Perspectives

who had the background knowledge to uninterrupted conversations, while showing a mental codification of experience.
interpret the clip than among participants profound impairments in tasks that require Neuroimaging studies suggest that
who did not. Crucially, responses in the the retrieval of long-​term memories60. instantiation and reinstatement of schemas
DMN were similar across the short-​term Consistent with such behavioural results, are mediated by the interaction among
context and the long-​term context groups a recent neuroimaging study involving regions within the DMN — namely, the
(although DMN regions showed increased an individual with hippocampal amnesia ventromedial prefrontal cortex, precuneus,
connectivity to the hippocampus in the suggests that integration of information bilateral temporoparietal junction and
long-​term context group compared with in over several minutes in the DMN can be hippocampus63. One study demonstrated
the short-​term context group). This result sustained even after hippocampal lesion62. the involvement of the DMN in schema
suggests that the responses in the DMN In this study, the participant with amnesia representation by measuring brain responses
can be shaped by information accumulated showed typical response patterns in the to different schemas (‘eating at a restaurant’
across many minutes during uninterrupted DMN while listening to an intact narrative, or ‘going through the airport’) presented as
viewing (short-​term context) as well as by as seen in neurotypical controls. Moreover, as stories that varied widely in terms of their
information gathered over days (as in in neurotypical controls, the DMN activity characters and storylines, and in terms
the long-​term context group) — and, of the participant with amnesia was of format (audiovisual clips or spoken
conceivably, even years. disrupted by temporally scrambling the narration)64. Regions including the posterior
The ability of the DMN to use long-​term paragraphs of the narrative, suggesting that medial cortex and medial prefrontal cortex
context is probably related to its anatomical the DMN otherwise successfully integrated exhibited schema-​associated event patterns
and functional connections to the medial information over windows of at least that generalized across stories, participants
temporal lobe and hippocampus, which 10–30 s. Further research is needed to better and modalities. Moreover, the DMN was
are central to learning and memory56–59. characterize the role of the hippocampus in found to be involved in updating schemas
At the same time, the ability of the DMN to supporting the integration of information when events violated expectations or were
accumulate information over many minutes over long timescales in the DMN. inconsistent with the established schema65,66,
(short-​term context) may explain why as in the case of surprising events during
individuals with amnesia can often sustain The DMN represents acquired schemas. a movie (for example, an unexpected plot
coherent conversations over a few minutes60 The ability to integrate information over twist). A recent review highlights the role
or recall coherent narratives61 during days is necessary for building schemas, of the DMN in representing and reinstating

a Functional correlation using PCC as a seed

Rest Processing single words Listening to a story

0.25 0.5
r
b Inter-subject functional correlation using PCC as a seed

Rest Processing single words Listening to a story

Participant 1 Participant 2

RH LH 0.13 0.5
r

Fig. 3 | Isolating stimulus-locked brain connectivity using inter-subject functional correlation. a | Within-​participant functional correlation maps
between the posterior cingulate cortex (PCC) seed (yellow voxel in the schematic; dashed circles in the brain maps) and the whole-​brain neural activity.
The functional correlation analysis delineates nodes of the default mode network (DMN) in which the activity fluctuates together (co-​varies) in a given
participant, owing to the direct or indirect anatomical connections during rest (left panel), processing of single words (middle panel) and listening to a
coherent story (right panel). b | Inter-​subject functional correlation maps between the PCC seed and the whole-​brain neural activity observed in other
participants. This analysis can filter out spontaneous intrinsic neural facilitation, and as such reveals no substantial stimulus-​locked correlations in the DMN
during rest (left panel) or during the processing of single words (middle panel). By contrast, however, inter-​subject functional correlation exposed
stimulus-​locked shared responses across participants in the DMN as subjects listen to and process a spoken story minutes long (right panel). LH, left
hemisphere; RH, right hemisphere. Adapted from ref.26, CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/).

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 Perspectives

schemas while processing narratives67. Thus, timescales as well as our idiosyncratic prior Reciprocally, the verbal and non-​verbal
the DMN is involved in generating and dispositions, including learned schemas, actions of receivers’ brains can also shape
retrieving schemas in order to dynamically prior knowledge and beliefs that shape our the neural responses of the sender in a
interpret an external situation. responses to a given context (Fig. 1b). In coupled dynamical system84,85. Although we
other words, we suggest that the DMN is an review evidence supporting our claim that
The DMN is sensitive to individual active and dynamic sense-​making network DMN response patterns are shaped by the
differences in interpretation. The finding that integrates incoming extrinsic social actions of other social agents, we note that
that responses in the DMN are shaped by information with prior intrinsic information this notion does not preclude the possibility
our long-​term memories and beliefs raises to form rich, context-​dependent models of that additional genetic factors on longer
the intriguing possibility that individual the situation as they unfold over time. evolutionary timescales also shape responses
differences in the way we perceive the world Perhaps one of the most intriguing in the DMN.
can account for individual differences in findings across these studies is the discovery
DMN responses. Indeed, several studies of neural patterns in the DMN that are Brain–brain coupling between speakers
have linked similarities in spontaneous, shared across participants and aligned to and listeners. Brain–brain coupling was
idiosyncratic interpretations of narratives the abstract structure and interpretation first reported in a study that correlated the
to neural similarities. In one study37, of the external events. In other words, responses in the speaker’s brain recorded
participants were scanned using fMRI while the results suggest that participants who while telling a personal story in the fMRI
watching an ambiguous animation that understand the situation in the same way scanner with the listeners’ brain responses
conveyed a complex social narrative using will have similar neural patterns in nodes while listening to the story86. The speaker’s
only the motion of simple geometric shapes, of the DMN, irrespective of considerable neural responses were coupled (that is,
in the vein of work by Heider and Simmel68. differences in the low-​level stimulus correlated, with or without a short time lag;
Greater similarity in the interpretation of properties of the sensory input. At the same Fig. 2d) to the listeners’ neural responses
the ambiguous shape animation (as reported time, subtle differences in DMN neural in brain regions at various levels of the
by each of the participants) was associated response patterns across participants seem timescale processing hierarchy. Speaker–
with greater similarity in neural response to correlate with subtle differences in their listener coupling in early auditory regions
in DMN regions across participants. interpretation. Together, these studies reflected shared processing of low-​level
Similar studies have reported that increased suggest that, through the interaction between acoustic properties of the stimulus, such as
alignment in neural response across the intrinsic self and the extrinsic world, the the audio amplitude envelope. By contrast, in
subjects in the DMN is correlated with DMN develops a shared neural code. language areas and the DMN, the responses
the similarity in interpretation of spoken in the listeners’ brains lagged behind the
narratives69, similarity in emotional response The DMN in social interaction responses in the speaker’s brain, suggesting
over time during spoken narratives70, A fundamental question is why similar that responses in the speaker’s brain causally
similarity in humour perception over time71, understandings or interpretations of shaped the responses in the listeners’ brains.
similarity in socio-​sexual desire while high-​level concepts produce shared neural Furthermore, speaker–listener neural
watching erotic movies72 and similarity in activity across individuals in the DMN. coupling in higher-​order areas, including
moral decision-​making73,74. It was recently suggested that a crucial aspect the DMN, reflected communication and
Individual differences in behaviour or of interpersonal interactions is a sense of shared understanding of the narrative. This
interpretation of a narrative that are linked dyadic shared reality82. Here, we propose that study along with further studies revealed
to differences in DMN activity have been the shared representation in the DMN arises that speaker–listener coupling in the DMN,
shown to be associated with trait-​level from dynamical social interaction between but not early auditory regions, is disrupted
personality differences, demographic the self and others. In other words, through when communication is disrupted by
factors or cultural differences75,76. Several reciprocal and dynamical social interaction, either speaking nonsense words86,87, telling
recent studies have shown that individual other people’s brains can shape your brain a story in a language that the listener does
differences in personality traits, including responses and, at the same time, be shaped not understand86 or temporally scrambling
levels of paranoia77, memory load75 and a by your brain’s responses to generate a the stimulus88. Moreover, speaker–listener
tendency to hold an analytic versus holistic shared reality (Fig. 1b). Indeed, many studies coupling in the DMN predicted the quality
perspective78, may modulate the level of suggested an extensive overlap between of communication: the better the listener
shared responses in regions of the DMN. the DMN and the ‘social brain’ — the brain understood the speaker, the closer
Several researchers have also identified regions involved in social cognition (Box 1). their brain responses matched the speaker’s
correlations between neural similarity in Throughout the day, DMN neural brain responses86–89. The speaker–listener
regions of the DMN and measures of mental responses are shaped by the actions and coupling was observed in the temporal
health in clinical populations, including gestures of other social agents. Even when fluctuations of the response time courses
depression79,80 and schizophrenia81. isolated at home or lying down in a dark within each node of the DMN86–89 (for
fMRI scanner during experiments, an example, the time course of neural responses
An active sense-​making network individual’s brain responses are still shaped in the speaker’s precuneus was correlated
The evidence reviewed in this Perspective by the actions of others, as conveyed by with that of the listener) as well as in the
suggests that activity in the DMN is neither audiovisual movies, spoken words or within-​area alignment of spatial patterns
solely intrinsic nor solely extrinsic. Rather, written text. The switch from a single-​brain across participants (for example, the spatial
the activity in the DMN is shaped by perspective to a multi-​brain perspective83 voxel pattern in the speaker’s precuneus was
both the unfolding dynamical structure of opens the way to studying how the correlated with that of the listeners)33.
real-​life events (including what other social responses of one brain (the sender) shape These findings demonstrate that
agents say and do in those events) over long the responses of other brains (the receivers). comprehension-​based processes in a

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Perspectives

listener’s brain are driven by, or shaped by, Sender–receiver coupling has also been demonstrated33 (Fig. 4a). In this study,
production-​based processes in the speaker’s reported during non-​verbal communications participants were scanned using fMRI
brain. Speaker–listener coupling has such as gestures96,97 and facial expressions98. while watching an episode of a television
been observed using various stimuli and series. Participants were then scanned while
imaging modalities, including fMRI33,70,88,90,91, The communication cycle. The full circle of describing the episode out loud in their
func­tional near-​infrared spectroscopy experience transmission from the speaker’s own words. A second group of participants,
(fNIRS)92–94 and electroencephalography95. brain to the listener’s brain has also been who had never seen the television episode

a Circle of communication b Between-subject pattern

Movie-viewing
Scene 01 Scene 01
Corr

Scene 02 Scene 02

Condition Y
Condition X
Corr

... ...
Baseline:
... random pairs ...

Scene 22 Scene 22
Spoken recall Listening to Corr
the narrative

c Shared neural patterns across movie-viewing, recalling d Friends exhibit similar neural responses to narratives
the movies and listening to the movie recall
Social distance Social distance Social distance Social distance
LH RH = 1 (friends) = 2 (friends of friends) =3 =4
Prec
dmPFC
LH

PCC

TPJ

TPJ RH

Prec dmPFC
P
LH
0.0089 1.0000 × 10–4

PCC vmPFC

Inter-subject time series similarity (normalized within brain regions)

–0.45 0.45

Fig. 4 | Transmission of experience and the similarity of neural responses among friends. a | An experimental paradigm for measuring how we share
our memories with other brains. First, a participant was scanned while watching an episode of the television series Merlin (encoding information to mem-
ory). Next, the participant was scanned as they used words to recall and share their memories with a new group of participants who had never seen the
television episode before. b | Neural coupling between groups was assessed using a between-​subject pattern similarity spotlight analysis. An average
spatial pattern of activity was calculated for each participant for each scene of the movie (coloured bars). Spatial patterns for each scene were then cor-
related (corr) across participants in different groups and averaged. c | Participants who had never seen the episode before and listened to the description
of the episode had a similar pattern of neural response within the default mode network (DMN) to the participant who recounted the episode from memory.
d | The social network of a cohort of students at a business school was reconstructed based on a survey of the students. From this network, 42 students
participated in a functional MRI (fMRI) study and watched a collection of short movie clips while in the scanner. Similarities in fMRI response while watching
these clips were calculated according to the social distance (defined by the social network) between a dyad. These were calculated by correlating the time
series of response for each participant to that of every other participant in a given brain region and taking the mean of the resulting correlations. Warmer
colours indicate relatively similar responses for a given brain region; cooler colours indicate relatively dissimilar responses for that brain region. Note the
higher degree of inter-​subject alignment of the neural responses in regions of the DMN (posterior cingulate cortex, PCC; precuneus, Prec; ventromedial
prefrontal cortex, vmPFC; dorsomedial prefrontal cortex, dmPFC; and temporoparietal junction, TPJ) and nearby networks to the same movie clips among
dyads that are social closer. LH, left hemisphere; RH, right hemisphere. Parts a–c adapted from ref.33. Part d adapted from ref.117, CC BY 4.0 (https://
creativecommons.org/licenses/by/4.0/).

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 Perspectives

before, were then scanned while listening to coordination among group members102–106, refers to a length of thread, whereas ‘string’
one of the viewer participants’ descriptions. finding increased neural alignment — both in computer science refers to a sequence
Strikingly, participants who had never seen spatial and temporal — during periods of characters. Several recent studies have
the episode before showed spatio-​temporally of the task that required heightened identified teacher–student DMN coupling
similar DMN activity to the speaker both coordination and communication. Another during teaching and learning in both
as the speaker recalled the episode and as electroencephalography study tested neural real-​life classrooms110,111 and simulated
the speaker watched the episode (Fig. 4b,c). alignment between romantic dyads and classrooms88,112–116. Notably, teacher–student
Classification analysis showed that similar­ unacquainted dyads, and found elevated neural coupling predicts student learning
ities in the DMN responses between the coupling in the DMN during mutual outcomes88,110,113,114,116.
movie viewers, the speaker and the listeners gaze and smiling107. It was suggested that
were scene-​specific and invariant to low-​ rudimentary non-​verbal social cues such as Social networks shape shared understanding.
level sensory features (such as differences mutual gaze and smiling, which are used by An old proverb says “Tell me who your
between the audiovisual movie and its caregivers to initially establish coupling with friends are, and I will tell you who you are”.
spoken description), suggesting that these children, can be used later on to facilitate Emerging neuroscientific research provides
shared responses reflected the transmission neural coupling routinely throughout life. evidence that neural alignment across
of information. Moreover, listener com­ subjects is influenced and shaped by the
prehension was better predicted by the The emergence of shared understanding. people with whom we are connected in our
unique alignment between the listeners’ There is no period in life when our brain real and virtual social networks. A striking
and the speaker’s response patterns as the responses are more strongly shaped by recent study117,118 mapped the close-​knit
speaker encoded the movie to memory, others than during the early years of child social network of a class of students at a
relative to the alignment to the responses development. This is especially apparent business school and found that pairs of
of other participants who watched the during the acquisition of a shared linguistic people who have closer social relationships
movie but did not share their experience code in early childhood, which enables have more similar neural responses to
with the listeners. This tight alignment the sharing of semantic representations narratives than do pairs of people who are
between the speaker and their audience may between a child and their caregivers. less close (Fig. 4d).
reveal the mechanism by which we share Emerging fNIRS studies in which caregivers Moreover, recent work from the
our idiosyncratic perspectives and memories and children were simultaneously scanned same group measured brain responses of
with other brains who never directly during communication and interactive play participants from the same social network
experienced them. shows coupled neural responses between as they watched ambiguous movie clips119.
caregivers and children (ages 8–12 years108) The participants then gathered in groups to
One-​way versus two-​way communication. and between caregivers and infants as with the aim of coming to a consensus about
Although unidirectional communication young as 9–10 months old1–3. This coupling the narrative interpretation of each movie
through television, movies, social media was evident in a few DMN regions, mainly clip. Consensus-​building conversation
and even online learning is increasingly in the prefrontal cortex. Infant–caregiver changed the minds of the participants and
common (Fig. 1b), much of communication is coupling was most strongly associated with led to the alignment of their subsequent
bidirectional and requires mutual adaptation mutual gaze (eye contact), infant smiling DMN neural responses as they watched the
and coupling between individuals who act and joint attention to objects109, supporting clip again. The increase in neural alignment
simultaneously as senders and receivers84. a mutual influence of neural dynamics. following the consensus-​building task
Investigating these dynamical processes Together, these findings suggest that that persisted over novel clips sampled from
requires a shift to simultaneous scanning caregiver–child DMN coupling emerges the same movie. Participants who were
of multiple brains, which is sometimes early in development and likely plays a role central in their real-​world social networks
called hyperscanning99. Emerging work in learning and language acquisition. played an outsized role in building group
using hyperscanning is uncovering the The development of shared alignment. Group-​level differences in neural
bidirectional flow of information during free understanding of concepts is also the responses between cultures have also been
dialogue and other naturalistic, complex hallmark of teaching and learning in observed during processing of culturally
tasks. For example, fNIRS was used to formal education. In such contexts, novel, relevant stimuli, such as stories describing
simultaneously scan dyads engaged in free sometimes jargonized, information must protected collectivist values, that may be
dialogue; this approach showed that face-​to-​ be transmitted from an expert’s brain to related to the degree of identification with
face communication elicits greater brain– novices’ brains. Here, we conceptualize the culture120,121.
brain coupling than does back-​to-​back teaching as a process of building a shared
communication100, suggesting a role of non-​ understanding of new information, Conclusions
verbal signals in enhancing communication supported by new or changed neural Our thoughts, feelings and actions are
and neural alignment. A later study representations. For example, taking a constantly being shaped by the actions,
using fNIRS identified conversational computer science class might lead to the memories and stories of others. At the same
‘leaders’ that emerge naturally in group development of a new understanding of time, our actions, memories and stories
discussions and used Granger causality computer-​science concepts and terms such shape and influence the thoughts, feelings
analysis to show that the leader’s neural as ‘recursion’ or ‘multiplexor’, as well as the and actions of the people to whom we are
activity more strongly predicted the contextual adjustment and modification connected. In this Perspective, we argue
followers’ neural activity than vice versa101. of already familiar concepts. It might also that we are able to shape and be shaped by
Other work has focused on groups of require flexibly changing the interpretation others in part because activity in the DMN
individuals completing complex, real-​life of words depending on the context: for is capable of both shaping other brains’
tasks that require substantial goal-​directed example, ‘string’ in common parlance responses and being shaped by the actions

NaTure RevIewS | NeUroScIence volume 22 | March 2021 | 189
Perspectives

of other brains during social interaction. 5

These authors contributed equally: Yaara Yeshurun, 25. Greicius, M. D., Supekar, K., Menon, V. &
Mai Nguyen Dougherty, R. F. Resting-​state functional connectivity
Shared neural activity at the sensory level reflects structural connectivity in the default mode
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