Social cognitive neuroscience is the scientific study of thebiological processes underpinningsocial cognition. Specifically, it uses the tools ofneuroscience to study "the mental mechanisms that create, frame, regulate, and respond to our experience of the social world".[1] Social cognitive neuroscience uses theepistemological foundations ofcognitive neuroscience, and is closely related tosocial neuroscience.[2] Social cognitive neuroscience employs humanneuroimaging, typically usingfunctional magnetic resonance imaging (fMRI). Human brain stimulation techniques such astranscranial magnetic stimulation andtranscranial direct-current stimulation are also used. In nonhuman animals, directelectrophysiological recordings and electrical stimulation ofsingle cells andneuronal populations are utilized for investigating lower-level social cognitive processes.[3][2][4]
The first scholarly works about the neural bases of social cognition can be traced back toPhineas Gage, a man who survived atraumatic brain injury in 1849 and was extensively studied for resultant changes in social functioning and personality.[4] In 1924, esteemed psychologistGordon Allport wrote a chapter on the neural bases of social phenomenon in his textbook ofsocial psychology.[5] However, these works did not generate much activity in the decades that followed.
Inventions in the study of neuronal activity provided the groundwork for field development. In 1924, German physiologist and psychiatrist Hans Berger (1873–1941) recorded the electrical activity of the brain by inventing the first human electroencephalogram EEG.[6] In 1953, Dr. Brownell and Dr. Aronow designed the first clinical positron imaging device, a prototype of a modern Positron Emission Tomography (PET).[7] In 1971, American chemist and physicist Paul Christian Lauterbur invented the idea of MR imaging (MRI).[8] Ph.D. in Chemistry, Michael E. Phelps developed the first PET scanner in 1973, based on discoveries of American scientists David Edmund Kuhl, Luke Chapman, and Roy Edwards, which constructed several tomographic instruments in the late 1950s.[9]
The beginning of modern social cognitive neuroscience can be traced toMichael Gazzaniga's book,Social Brain (1985), which attributedcerebral lateralization to the peculiarities of social psychological phenomenon.[10] Isolated pockets of social cognitive neuroscience research emerged in the late 1980s to the mid-1990s, mostly usingsingle-unitelectrophysiological recordings in nonhuman primates orneuropsychological lesion studies in humans.[4] During this time, the closely related field of social neuroscience emerged in parallel, however it mostly focused on how social factors influencedautonomic,neuroendocrine, andimmune systems.[4][2] In 1996,Giacomo Rizzolatti's group made one of the most seminal discoveries in social cognitive neuroscience: the existence ofmirror neurons inmacaque frontoparietal cortex.[11] The mid-1990s saw the emergence of functionalpositron emission tomography (PET) for humans, which enabled the neuroscientific study of abstract (and perhaps uniquely human, but this issue is still a subject of debate[12][13]) social cognitive functions such astheory of mind andmentalizing. However, PET is prohibitively expensive and requires the ingestion ofradioactive tracers, thus limiting its adoption.[4]
In the year 2000, the termsocial cognitive neuroscience was coined byMatthew Lieberman andKevin Ochsner, who are fromsocial andcognitive psychology backgrounds, respectively. This was done to integrate and brand the isolated labs doing research on the neural bases of social cognition.[1][4] Also in the year 2000,Elizabeth Phelps and colleagues published the firstfMRI study on social cognition, specifically onrace evaluations.[14] The adoption of fMRI, a less expensive and noninvasive neuroimaging modality, induced explosive growth in the field. In 2001, the firstacademic conference on social cognitive neuroscience was held atUniversity of California, Los Angeles. The mid-2000s saw the emergence ofacademic societies related to the field (Social and Affective Neuroscience Society,Society for Social Neuroscience), as well aspeer-reviewed journals specialized for the field (Social Cognitive and Affective Neuroscience,Social Neuroscience).[4] In the 2000s and beyond, labs conducting social cognitive neuroscience research proliferated throughout Europe, North America, East Asia, Australasia, and South America.[15][4][2]
Starting in the late 2000s, the field began to expand its methodological repertoire by incorporating other neuroimaging modalities (e.g.electroencephalography,magnetoencephalography,functional near-infrared spectroscopy),[16] advanced computational methods (e.g.multivariate pattern analysis,causal modeling,graph theory),[17] and brain stimulation techniques (e.g. transcranial magnetic stimulation, transcranial direct-current stimulation,deep brain stimulation).[18] Due to the volume and rigor of research in the field, the 2010s saw social cognitive neuroscience achieving mainstream acceptance in the wider fields of neuroscience and psychology.[4][2][3]
Hyperscanning or inter-brain research is becoming the most frequent approach to studying social cognition.[19][20][21][22] It is thought that exploring the correlation of neuronal activities of two or more brains in shared cognitive tasks can contribute to understanding the relationship between social experiences and neurophysiological processes.[23] In 2024, inspired by research on interpersonal neural synchronization[19][20][21][22] and neuroscience studies on fetal brain responses to auditory stimuli that revealed increased neuronal activity in the fetal brain when it was exposed to an unfamiliar voice stimuli,[24] the mother-fetus neurocognitive model hypothesis has been put forward opening a new research direction.[25][26][27] According to this position, interpersonal neurophysiological processes within the biological system of this dyad provide the fetal nervous system with training for proper reactions to stimuli at the onset of cognition.[25][26][27] The hypothesis has shown that training is successful because of neural synchronization between nervous systems that occurs through the interference of local neuronal oscillations with the low-frequency electromagnetic field of the mother's heart.[27]
Much of social cognition is primarily subserved by two dissociablemacro-scale brain networks: themirror neuron system (MNS) anddefault mode network (DMN). MNS is thought to represent and identify observable actions (e.g. reaching for a cup) that are used by DMN to infer unobservable mental states, traits, and intentions (e.g. thirsty).[28][3][29][30] Concordantly, the activation onset of MNS has been shown to precede DMN during social cognition.[28] However, the extent offeedforward,feedback, andrecurrent processing within and between MNS and DMN is not yet well-characterized, thus it is difficult to fully dissociate the exact functions of the two networks and their nodes.[3][28][30]
Mirror neurons, first discovered inmacaque frontoparietal cortex, fire when actions are either performed or observed.[11] In humans, similar sensorimotor "mirroring" responses have been found in the brain regions listed below, which are collectively referred to as MNS.[11][31] The MNS has been found to identify and represent intentional actions such asfacial expressions,body language, andgrasping.[28][31] MNS may encode theconcept of an action, not just the sensory and motor information associated with an action. As such, MNS representations have been shown to be invariant of how an action is observed (e.g. sensory modality) and how an action is performed (e.g. left versus right hand, upwards or downwards).[32][33] MNS has even been found to represent actions that are described in written language.[34]
Mechanistic theories of MNS functioning fall broadly into two camps: motor and cognitive theories. Classical motor theories posit that abstract action representations arise from simulating actions within the motor system, while newer cognitive theories propose that abstract action representations arise from the integration of multiple domains of information:perceptual,motor,semantic, andconceptual.[35][32] Aside from these competing theories, there are more fundamental controversies surrounding the human MNS – even the very existence of mirror neurons in this network is debated.[36][37] As such, the term "MNS" is sometimes eschewed for more functionally defined names such as "action observation network", "action identification network", and "action representation network".[37]
The hypothesis of the mother-fetus neurocognitive model (MFN) contributes to the theory of the MNS's role in social cognition.[26] It has been shown that MNS mirror neurons are observed only in specific cases of interpersonal dynamics that meet the MFN model conditions.[26] The MFN model is a representation of neurophysiological processes within the biological system of the Mother-Fetus dyad that provides the fetal nervous system with training for proper reactions to stimuli at the onset of cognition.[38] It has been shown that local neuronal oscillations in both nervous systems are synchronized through interference with the low-frequency electromagnetic field of the mother's heart.[38] So that the mother's heart oscillations synchronize the brain oscillations of already excited central and peripheral neuronal ensembles, similar in both organisms, due to physiological entrainment within the shared ecosystem.[38] Therefore, the activation of specific sensorimotor networks in the mother entrains those in the child, and, because of the shared ecosystem, this engagement trains the young nervous system to respond correctly to certain sensory stimuli through statistical mechanisms based on numerous successful and unsuccessful trials. This entrainment provides subliminal perception in the young organism, similar to the Mother's intentional act that initiates the process of cognition.[25][26][27][38]
Mirror neurons were first discovered inmacaquepremotor cortex.[11] The premotor cortex is associated with a diverse array of functions, encompassing low-level motor control, motor planning, sensory guidance of movement, along with higher level cognitive functions such aslanguage processing and action comprehension.[39] The premotor cortex has been found to contain subregions with uniquecytoarchitectural properties, the significance of which is not yet fully understood.[40] In humans, sensorimotor mirroring responses are also found throughout premotor cortex and adjacent sections ofinferior frontal gyrus andsupplementary motor area.[11][31]
Visuospatial information is more prevalent in ventral premotor cortex than dorsal premotor cortex.[39] In humans, sensorimotor mirroring responses extend beyond ventral premotor cortex into adjacent regions of inferior frontal gyrus, includingBroca's area, an area that is critical to language processing and speech production.[41] Action representations in inferior frontal gyrus can be evoked by language, such as action verbs, in addition to the observed and performed actions typically used as stimuli in biological motion studies.[34] The overlap between language and action understanding processes in inferior frontal gyrus has spurred some researchers to suggest overlapping neurocomputational mechanisms between the two.[41][34][33] Dorsal premotor cortex is strongly associated with motor preparation and guidance, such as representing multiple motor choices and deciding the final selection of action.[39]
Classical studies of action observation have found mirror neurons in macaqueintraparietal sulcus.[11] In humans, sensorimotor mirroring responses are centered around the anterior intraperietal sulcus, with responses also seen in adjacent regions such asinferior parietal lobule andsuperior parietal lobule. Intraparietal sulcus has been shown to more sensitive to the motor features of biological motion, relative to semantic features.[31] Intraparietal sulcus has been shown to encode magnitude in a domain-general manner, whether it be the magnitude of a motor movement, or the magnitude of a person's social status.[42] Intraparietal sulcus is considered a part of thedorsal visual stream, but is also thought to receive inputs from non-dorsal stream regions such as lateral occipitotemporal cortex and posterior superior temporal sulcus.[31]
LOTC encompasses lateral regions of thevisual cortex such asV5 andextrastriate body area. Though LOTC is typically associated with visual processing, sensorimotor mirroring responses and abstract action representations are reliably found in this region.[35][43] LOTC includes cortical areas that are sensitive to motion, objects, body parts, kinematics, body postures, observed movements, and semantic content in verbs.[35][43] LOTC is thought to encode the fine sensorimotor details of an observed action (e.g. local kinematic and perceptual features).[43] LOTC is also thought to bind together the different means by which a specific action can be carried out.[35]
Thedefault mode network (DMN) is thought to process and represent abstract social information, such as mental states, traits, and intentions.[3][44][45] Social cognitive functions such astheory of mind,mentalizing,emotion recognition,empathy,moral cognition, and socialworking memory consistently recruit DMN regions in human neuroimaging studies. Though the functional anatomy of these functions can differ, they often include the core DMN hubs of medial prefrontal cortex, posterior cingulate, and temporoparietal junction.[3][44][45][46][29][47] Aside from social cognition, the DMN is broadly associated with internally directed cognition.[48] The DMN has been found to be involved in memory-related processing (semantic,episodic,prospection), self-related processing (e.g.introspection), andmindwandering.[48][49][50] Unlike studies of the mirror neuron system, task-based DMN investigations almost always use human subjects, as DMN-related social cognitive functions are rudimentary or difficult to measure in nonhumans.[50][3] However, much of DMN activity occurs during rest, as DMN activation and connectivity are quickly engaged and sustained during the absence of goal-directed cognition.[50] As such, the DMN is widely thought the subserve the "default mode" of mammalian brain function.[51]
The interrelations between social cognition, rest, and the diverse array of DMN-related functions are not yet well understood and is a topic of active research. Social, non-social, and spontaneous processes in the DMN are thought to share at least some underlying neurocomputational mechanisms with each other.[42][52][53][54][55]
Medial prefrontal cortex (mPFC) is strongly associated with abstract social cognition such as mentalizing and theory of mind.[56][3][29][47] Mentalizing activates much of the mPFC, butdorsal mPFC appears to be more selective for information about other people, whileanterior mPFC may be more selective for information about the self.[56]
Ventral regions of mPFC, such asventromedial prefrontal cortex and medialorbitofrontal cortex, are thought to play a critical role in theaffective components of social cognition. For example, ventromedial prefrontal cortex has been found to represent affective information about other people.[3][44][46] Ventral mPFC has been shown to be critical in the computation and representation ofvalence and value for many types of stimuli, not just social stimuli.[57]
The mPFC may subserve the most abstract components of social cognition, as it is one of the most domain general brain regions, sits at the top of the cortical hierarchy, and is last to activate during DMN-related tasks.[3][50][58]
Abstract social cognition recruits a large area of posteromedial cortex centered aroundposterior cingulate cortex (PCC), but also extending intoprecuneus andretrosplenial cortex.[44][3] The specific function of PCC in social cognition is not yet well characterized,[47][45] and its role may be generalized and tightly linked with medial prefrontal cortex.[44][52] One view is that PCC may help represent some visuospatial and semantic components of social cognition.[59] Additionally, PCC may track social dynamics by facilitating bottom-up attention to behaviorally relevant sources of information in the external environment and in memory.[52] Dorsal PCC is also linked to monitoring behaviorally relevant changes in the environment, perhaps aiding in social navigation.[46] Outside of the social domain, PCC is associated with a very diverse array of functions, such as attention, memory, semantics, visual processing, mindwandering, consciousness, cognitive flexibility, and mediating interactions between brain networks.[60]
Thetemporoparietal junction (TPJ) is thought to be critical to distinguishing between multiple agents, such as the self and other.[47] The right TPJ is robustly activated by false belief tasks, in which subjects have to distinguish between others' beliefs and their own beliefs in a given situation.[29][47][3] The TPJ is also recruited by the wide variety of abstract social cognitive tasks associated with the DMN.[3][44][46] Outside of the social domain, TPJ is associated with a diverse array of functions such as attentional reorienting, target detection, contextual updating, language processing, and episodic memory retrieval.[61][62][63][64][53] The social and non-social functions of the TPJ may share common neurocomputational mechanisms.[65][62][53] For example, the substrates of attentional reorientation in TPJ may be used for reorienting attention between the self and others, and for attributing attention between social agents.[62][65] Moreover, a common neural encoding mechanism has been found to instantiate social, temporal, and spatial distance in TPJ.[42]
Social tasks recruit areas of lateral temporal cortex centered aroundsuperior temporal sulcus (STS), but also extending to superior temporal gyrus, middle temporal gyrus, and the temporal poles.[47][45] During social cognition, the anterior STS and temporal poles are strongly associated with abstract social cognition and person information, while the posterior STS is most associated withsocial vision and biological motion processing.[47][3] The posterior STS is also thought to provide perceptual inputs to the mirror neuron system.[31][28]
There are also several brain regions that fall outside the MNS and DMN which are strongly associated with certain social cognitive functions.[1]
Theventrolateral prefrontal cortex (VLPFC) is associated with emotional andinhibitory processing. It has been found to be involved inemotion recognition from facial expressions, body language,prosody, and more. Specifically, it is thought to access semantic representations of emotional constructs during emotion recognition.[66] Moreover, VLPFC is often recruited in empathy, mentalizing, and theory of mind tasks. VLPFC is thought to support the inhibition of self-perspective when thinking about other people.[1]
Theinsula is critical to emotional processing andinteroception. It has been found to be involved in emotion recognition, empathy, morality, and social pain. The anterior insula is thought to facilitate feeling the emotions of others, especially negative emotions such as vicarious pain. Lesions of the insula are associated with decreased empathy capacity. Anterior insula also activates during social pain, such as the pain caused by social rejection.[67][1]
Theanterior cingulate cortex (ACC) is associated with emotional processing and error monitoring. The dorsal ACC appears to share some social cognitive functions to the anterior insula, such as facilitating feeling the emotions of others, especially negative emotions. The dorsal ACC also robustly activates during social pain, like the pain caused by being the victim of an injustice. The dorsal ACC is also associated with social evaluation, such as the detection and appraisal of social exclusion. Thesubgenual ACC has been found to activate for vicarious reward, and may be involved inprosocial behavior.[67][1]
Thefusiform face area (FFA) is strongly associated with face processing and perceptual expertise. The FFA has been shown to process the visuospatial features of faces, and may also encode some semantic features of faces.[55][1]
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