Social cognitive neuroscience

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Social cognitive neuroscience is the scientific study of the biological processes underpinning social cognition. Specifically, it uses the tools of neuroscience to study "the mental mechanisms that create, frame, regulate, and respond to our experience of the social world". [1] Social cognitive neuroscience uses the epistemological foundations of cognitive neuroscience, and is closely related to social neuroscience. [2] Social cognitive neuroscience employs human neuroimaging, typically using functional magnetic resonance imaging (fMRI). Human brain stimulation techniques such as transcranial magnetic stimulation and transcranial direct-current stimulation are also used. In nonhuman animals, direct electrophysiological recordings and electrical stimulation of single cells and neuronal populations are utilized for investigating lower-level social cognitive processes. [3] [2] [4]

Contents

History and methods

The first scholarly works about the neural bases of social cognition can be traced back to Phineas Gage, a man who survived a traumatic brain injury in 1849 and was extensively studied for resultant changes in social functioning and personality. [4] In 1924, esteemed psychologist Gordon Allport wrote a chapter on the neural bases of social phenomenon in his textbook of social psychology. [5] However, these works did not generate much activity in the decades that followed. The beginning of modern social cognitive neuroscience can be traced to Michael Gazzaniga's book, Social Brain (1985), which attributed cerebral lateralization to the peculiarities of social psychological phenomenon. Isolated pockets of social cognitive neuroscience research emerged in the late 1980s to the mid-1990s, mostly using single-unit electrophysiological recordings in nonhuman primates or neuropsychological 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 influenced autonomic, neuroendocrine, and immune systems. [4] [2] In 1996, Giacomo Rizzolatti's group made one of the most seminal discoveries in social cognitive neuroscience: the existence of mirror neurons in macaque frontoparietal cortex. [6] The mid-1990s saw the emergence of functional positron emission tomography (PET) for humans, which enabled the neuroscientific study of abstract (and perhaps uniquely human) social cognitive functions such as theory of mind and mentalizing. However, PET is prohibitively expensive and requires the ingestion of radioactive tracers, thus limiting its adoption. [4]

In the year 2000, the term social cognitive neuroscience was coined by Matthew Lieberman and Kevin Ochsner, who are from social and cognitive 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 first fMRI study on social cognition, specifically on race evaluations. [7] The adoption of fMRI, a less expensive and noninvasive neuroimaging modality, induced explosive growth in the field. In 2001, the first academic conference on social cognitive neuroscience was held at University of California, Los Angeles. The mid-2000s saw the emergence of academic societies related to the field ( Social and Affective Neuroscience Society, Society for Social Neuroscience ), as well as peer-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. [8] [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), [9] advanced computational methods (e.g. multivariate pattern analysis, causal modeling, graph theory), [10] and brain stimulation techniques (e.g. transcranial magnetic stimulation, transcranial direct-current stimulation, deep brain stimulation). [11] 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]

Functional anatomy

Much of social cognition is primarily subserved by two dissociable macro-scale brain networks: the mirror neuron system (MNS) and default 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). [12] [3] [13] [14] Concordantly, the activation onset of MNS has been shown to precede DMN during social cognition. [12] However, the extent of feedforward, feedback, and recurrent 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] [12] [14]

Mirror neuron system (MNS)

Mirror neurons, first discovered in macaque frontoparietal cortex, fire when actions are either performed or observed. [6] In humans, similar sensorimotor "mirroring" responses have been found in the brain regions listed below, which are collectively referred to as MNS. [6] [15] The MNS has been found to identify and represent intentional actions such as facial expressions, body language, and grasping. [12] [15] MNS may encode the concept 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). [16] [17] MNS has even been found to represent actions that are described in written language. [18]

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, and conceptual. [19] [16] 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. [20] [21] 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". [21]

Premotor cortex

Mirror neurons were first discovered in macaque premotor cortex. [6] 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 as language processing and action comprehension. [22] The premotor cortex has been found to contain subregions with unique cytoarchitectural properties, the significance of which is not yet fully understood. [23] In humans, sensorimotor mirroring responses are also found throughout premotor cortex and adjacent sections of inferior frontal gyrus and supplementary motor area. [6] [15]

Visuospatial information is more prevalent in ventral premotor cortex than dorsal premotor cortex. [22] In humans, sensorimotor mirroring responses extend beyond ventral premotor cortex into adjacent regions of inferior frontal gyrus, including Broca's area, an area that is critical to language processing and speech production. [24] 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. [18] The overlap between language and action understanding processes in inferior frontal gyrus has spurred some researchers to suggest overlapping neurocomputational mechanisms between the two. [24] [18] [17] Dorsal premotor cortex is strongly associated with motor preparation and guidance, such as representing multiple motor choices and deciding the final selection of action. [22]

Intraparietal sulcus

Classical studies of action observation have found mirror neurons in macaque intraparietal sulcus. [6] In humans, sensorimotor mirroring responses are centered around the anterior intraperietal sulcus, with responses also seen in adjacent regions such as inferior parietal lobule and superior parietal lobule. Intraparietal sulcus has been shown to more sensitive to the motor features of biological motion, relative to semantic features. [15] 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. [25] Intraparietal sulcus is considered a part of the dorsal visual stream, but is also thought to receive inputs from non-dorsal stream regions such as lateral occipitotemporal cortex and posterior superior temporal sulcus. [15]

Lateral occipitotemporal cortex (LOTC)

LOTC encompasses lateral regions of the visual cortex such as V5 and extrastriate body area. Though LOTC is typically associated with visual processing, sensorimotor mirroring responses and abstract action representations are reliably found in this region. [19] [26] LOTC includes cortical areas that are sensitive to motion, objects, body parts, kinematics, body postures, observed movements, and semantic content in verbs. [19] [26] LOTC is thought to encode the fine sensorimotor details of an observed action (e.g. local kinematic and perceptual features). [26] LOTC is also thought to bind together the different means by which a specific action can be carried out. [19]

Default mode network (DMN)

The default mode network (DMN) is thought to process and represent abstract social information, such as mental states, traits, and intentions. [3] [27] [28] Social cognitive functions such as theory of mind, mentalizing, emotion recognition, empathy, moral cognition, and social working 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] [27] [28] [29] [13] [30] Aside from social cognition, the DMN is broadly associated with internally directed cognition. [31] The DMN has been found to be involved in memory-related processing (semantic, episodic, prospection), self-related processing (e.g. introspection), and mindwandering. [31] [32] [33] 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. [33] [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. [33] As such, the DMN is widely thought the subserve the "default mode" of mammalian brain function. [34]

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. [25] [35] [36] [37] [38]

Medial prefrontal cortex (mPFC)

Medial prefrontal cortex (mPFC) is strongly associated with abstract social cognition such as mentalizing and theory of mind. [39] [3] [13] [30] Mentalizing activates much of the mPFC, but dorsal mPFC appears to be more selective for information about other people, while anterior mPFC may be more selective for information about the self. [39]

Ventral regions of mPFC, such as ventromedial prefrontal cortex and medial orbitofrontal cortex, are thought to play a critical role in the affective components of social cognition. For example, ventromedial prefrontal cortex has been found to represent affective information about other people. [3] [27] [29] Ventral mPFC has been shown to be critical in the computation and representation of valence and value for many types of stimuli, not just social stimuli. [40]

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] [33] [41]

Posterior cingulate cortex (PCC)

Abstract social cognition recruits a large area of posteromedial cortex centered around posterior cingulate cortex (PCC), but also extending into precuneus and retrosplenial cortex. [27] [3] The specific function of PCC in social cognition is not yet well characterized, [30] [28] and its role may be generalized and tightly linked with medial prefrontal cortex. [27] [35] One view is that PCC may help represent some visuospatial and semantic components of social cognition. [42] Additionally, PCC may track social dynamics by facilitating bottom-up attention to behaviorally relevant sources of information in the external environment and in memory. [35] Dorsal PCC is also linked to monitoring behaviorally relevant changes in the environment, perhaps aiding in social navigation. [29] 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. [43]

Temporoparietal junction (TPJ)

The temporoparietal junction (TPJ) is thought to be critical to distinguishing between multiple agents, such as the self and other. [30] 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. [13] [30] [3] The TPJ is also recruited by the wide variety of abstract social cognitive tasks associated with the DMN. [3] [27] [29] 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. [44] [45] [46] [47] [36] The social and non-social functions of the TPJ may share common neurocomputational mechanisms. [48] [45] [36] 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. [45] [48] Moreover, a common neural encoding mechanism has been found to instantiate social, temporal, and spatial distance in TPJ. [25]

Superior temporal sulcus (STS)

Social tasks recruit areas of lateral temporal cortex centered around superior temporal sulcus (STS), but also extending to superior temporal gyrus, middle temporal gyrus, and the temporal poles. [30] [28] 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 with social vision and biological motion processing. [30] [3] The posterior STS is also thought to provide perceptual inputs to the mirror neuron system. [15] [12]

Other regions

There are also several brain regions that fall outside the MNS and DMN which are strongly associated with certain social cognitive functions. [1]

Ventrolateral prefrontal cortex (VLPFC)

The ventrolateral prefrontal cortex (VLPFC) is associated with emotional and inhibitory processing. It has been found to be involved in emotion recognition from facial expressions, body language, prosody, and more. Specifically, it is thought to access semantic representations of emotional constructs during emotion recognition. [49] 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]

Insula

The insula is critical to emotional processing and interoception. 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. [50] [1]

Anterior cingulate cortex (ACC)

The anterior 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. The subgenual ACC has been found to activate for vicarious reward, and may be involved in prosocial behavior. [50] [1]

Fusiform face area (FFA)

The fusiform 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. [38] [1]

Notable figures

See also

Further reading

Related Research Articles

In psychology, theory of mind refers to the capacity to understand other people by ascribing mental states to them. A theory of mind includes the knowledge that others' beliefs, desires, intentions, emotions, and thoughts may be different from one's own. Possessing a functional theory of mind is crucial for success in everyday human social interactions. People utilise a theory of mind when analyzing, judging, and inferring others' behaviors. The discovery and development of theory of mind primarily came from studies done with animals and infants. Factors including drug and alcohol consumption, language development, cognitive delays, age, and culture can affect a person's capacity to display theory of mind. Having a theory of mind is similar to but not identical with having the capacity for empathy or sympathy.

<span class="mw-page-title-main">Brodmann area 44</span> Brain area

Brodmann area 44, or BA44, is part of the frontal cortex in the human brain. Situated just anterior to premotor cortex (BA6) and on the lateral surface, inferior to BA9.

A mirror neuron is a neuron that fires both when an organism acts and when the organism observes the same action performed by another. Thus, the neuron "mirrors" the behavior of the other, as though the observer were itself acting. Mirror neurons are not always physiologically distinct from other types of neurons in the brain; their main differentiating factor is their response patterns. By this definition, such neurons have been directly observed in humans and primate species, and in birds.

<span class="mw-page-title-main">Prefrontal cortex</span> Part of the brain responsible for personality, decision-making, and social behavior

In mammalian brain anatomy, the prefrontal cortex (PFC) covers the front part of the frontal lobe of the cerebral cortex. The PFC contains the Brodmann areas BA8, BA9, BA10, BA11, BA12, BA13, BA14, BA24, BA25, BA32, BA44, BA45, BA46, and BA47.

<span class="mw-page-title-main">Orbitofrontal cortex</span> Region of the prefrontal cortex of the brain

The orbitofrontal cortex (OFC) is a prefrontal cortex region in the frontal lobes of the brain which is involved in the cognitive process of decision-making. In non-human primates it consists of the association cortex areas Brodmann area 11, 12 and 13; in humans it consists of Brodmann area 10, 11 and 47.

<span class="mw-page-title-main">Posterior cingulate cortex</span> Caudal part of the cingulate cortex of the brain

The posterior cingulate cortex (PCC) is the caudal part of the cingulate cortex, located posterior to the anterior cingulate cortex. This is the upper part of the "limbic lobe". The cingulate cortex is made up of an area around the midline of the brain. Surrounding areas include the retrosplenial cortex and the precuneus.

The simulation theory of empathy holds that humans anticipate and make sense of the behavior of others by activating mental processes that, if they culminated in action, would produce similar behavior. This includes intentional behavior as well as the expression of emotions. The theory says that children use their own emotions to predict what others will do; we project our own mental states onto others.

<span class="mw-page-title-main">Temporoparietal junction</span> Area of the brain where the temporal and parietal lobes meet

The temporoparietal junction (TPJ) is an area of the brain where the temporal and parietal lobes meet, at the posterior end of the lateral sulcus. The TPJ incorporates information from the thalamus and the limbic system as well as from the visual, auditory, and somatosensory systems. The TPJ also integrates information from both the external environment as well as from within the body. The TPJ is responsible for collecting all of this information and then processing it.

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<span class="mw-page-title-main">Default mode network</span> Large-scale brain network active when not focusing on an external task

In neuroscience, the default mode network (DMN), also known as the default network, default state network, or anatomically the medial frontoparietal network (M-FPN), is a large-scale brain network primarily composed of the dorsal medial prefrontal cortex, posterior cingulate cortex, precuneus and angular gyrus. It is best known for being active when a person is not focused on the outside world and the brain is at wakeful rest, such as during daydreaming and mind-wandering. It can also be active during detailed thoughts related to external task performance. Other times that the DMN is active include when the individual is thinking about others, thinking about themselves, remembering the past, and planning for the future.

<span class="mw-page-title-main">Biological motion</span> Motion that comes from actions of a biological organism

Biological motion is motion that comes from actions of a biological organism. Humans and animals are able to understand those actions through experience, identification, and higher level neural processing. Humans use biological motion to identify and understand familiar actions, which is involved in the neural processes for empathy, communication, and understanding other's intentions. The neural network for biological motion is highly sensitive to the observer's prior experience with the action's biological motions, allowing for embodied learning. This is related to a research field that is broadly known as embodied cognitive science, along with research on mirror neurons.

<span class="mw-page-title-main">Vittorio Gallese</span> Italian physiologist (1959–)

Vittorio Gallese is professor of Psychobiology at the University of Parma, Italy, and was professor in Experimental Aesthetics at the University of London, UK (2016-2018). He is an expert in neurophysiology, cognitive neuroscience, social neuroscience, and philosophy of mind. Gallese is one of the discoverers of mirror neurons. His research attempts to elucidate the functional organization of brain mechanisms underlying social cognition, including action understanding, empathy, language, mindreading and aesthetic experience.

Cultural neuroscience is a field of research that focuses on the interrelation between a human's cultural environment and neurobiological systems. The field particularly incorporates ideas and perspectives from related domains like anthropology, psychology, and cognitive neuroscience to study sociocultural influences on human behaviors. Such impacts on behavior are often measured using various neuroimaging methods, through which cross-cultural variability in neural activity can be examined.

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<span class="mw-page-title-main">Marcus Raichle</span> American neurologist

Marcus E. Raichle is an American neurologist at the Washington University School of Medicine in Saint Louis, Missouri. He is a professor in the Department of Radiology with joint appointments in Neurology, Neurobiology and Biomedical Engineering. His research over the past 40 years has focused on the nature of functional brain imaging signals arising from PET and fMRI and the application of these techniques to the study of the human brain in health and disease. He received the Kavli Prize in Neuroscience “for the discovery of specialized brain networks for memory and cognition", together with Brenda Milner and John O’Keefe in 2014.

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<span class="mw-page-title-main">Salience network</span> Large-scale brain network involved in detecting and attending to relevant stimuli

The salience network (SN), also known anatomically as the midcingulo-insular network (M-CIN) or ventral attention network, is a large scale network of the human brain that is primarily composed of the anterior insula (AI) and dorsal anterior cingulate cortex (dACC). It is involved in detecting and filtering salient stimuli, as well as in recruiting relevant functional networks. Together with its interconnected brain networks, the SN contributes to a variety of complex functions, including communication, social behavior, and self-awareness through the integration of sensory, emotional, and cognitive information.

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Valeria Gazzola is an Italian neuroscientist, associate professor at the Faculty of Social and Behavioral Sciences at the University of Amsterdam (UvA) and member of the Young Academy of Europe. She is also a tenured department head at the Netherlands Institute for Neuroscience (NIN) in Amsterdam, where she leads her own research group and the Social Brain Lab together with neuroscientist Christian Keysers. She is a specialist in the neural basis of empathy and embodied cognition: Her research focusses on how the brain makes individuals sensitive to the actions and emotions of others and how this affects decision-making.

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