Empathy—the was selectively recruited when subjects read

Empathy—the ability to understand and relate to the perspectives of others—is an essentialhuman act. As it requires conceptualizing and identifying with the mental states of other people whilemaintaining the distinction between self and other, empathy necessitates complex social cognition.

Thereis a general consensus in the literature that empathy is comprised of several subordinate processes,including affective empathy (or experience sharing) and cognitive empathy (or mentalizing) (Overwalle& Baetens, 2009; Zaki & Ochsner, 2012). Affective empathy involves resonating with another person’semotional experience, and is linked to activation in a circuit that includes the insula, amygdala, andanterior cingulate cortex (ACC) (Decety & Jackson, 2006). Cognitive empathy, on the other hand, refersto intentionally taking the perspective of the other person, and has been shown to engage midlinestructures such as the superior temporal sulcus (STS) and the temporoparietal junction (TPJ) (Frith , 2006; Spunt & Lieberman, 2012). The TPJ in particular has been emphasized as a key region forempathetic cognition.

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Many studies have found evidence implicating it in comprehending another’sviewpoint while maintaining the distinction between self and other. However, the TPJ has also beendemonstrated to be active during more basic cognitive tasks, prompting the question of whether there isanything particularly “social” or “empathy-driven” about this brain region. This paper will address thedebate over whether the TPJ is inherently social by reviewing the literature on its role in mentalizing,discussing its function in cognition more broadly, and examining the evidence in the context of a networkperspective. The TPJ is consistently recruited during tasks that involve contemplating the mental state ofanother person. In one study, the TPJ was selectively recruited when subjects read stories about aprotagonist’s thoughts and beliefs (Saxe & Powell, 2006). In another experiment using positron emissiontomography (PET), researchers found a hemodynamic increase in the TPJ when participants were Soderberg 1 instructed to take a third person perspective compared to taking their own first person view (Ruby , 2004). As well as mediating these explicitly empathetic acts, the TPJ has been shown to beengaged during the identification of the goals of others’ actions. In a meta-analysis of functional imagingstudies on social cognition, tasks that required the determination of another actor’s goals and perspectiveconsistently engaged the TPJ (Overwalle, 2009).

Along with this imaging evidence, a lesion studyprovides a stronger causal link between TPJ activity and perspective-taking. Three patients with damageto the left TPJ did no better than chance on a task that required the understanding of another person’sfalse belief; importantly, they performed similarly to controls on tasks matched for cognitive demand thatdid not involve false beliefs (Samson, Apperly, Chiavarino, & Humphreys, 2004). This finding impliesthat the left TPJ is necessary for generating representations of someone else’s belief, and that this effect isnot attributable to the cognitive load such a task requires. These results and others provide evidence thatthe TPJ is essential for comprehending the mental states of others. While it has been established that the TPJ is an important component of the mentalizing system,there remain differences in the interpretation of its specific function. Broadly, cognitive empathy isunderstood to entail the act of grasping the mindset of another while maintaining the distinction betweenoneself and that other (Overwalle & Baetens, 2009).

One view emphasizes the TPJ’s role in preserving anindividual’s agency as a perceiver (Decety & Jackson, 2006). Empathy would be disorienting rather thanvaluable if the boundary between these two perspectives was eroded. As this could potentially lead toemotional contagion and maladaptive behavior, it is likely that during the course of evolution, processesthat kept a strong partition between self and other would be selected for. Another interpretation focuseson the temporal domain of empathic thinking. Van Overwalle proposes that the TPJ is responsible for”transient mental inferences” about the other person (i.

e., their present thoughts and goals), as opposed tomore permanent traits and characteristics (Overwalle, 2009). It seems that the TPJ is a site of empatheticprocessing in real time, and as such, has the capacity to dynamically react to changing social input.

Although the TPJ is clearly implicated in perceiving the mental states of others, neglecting itsinvolvement in other realms of cognition would be detrimentally selective. In the domain of attention, the Soderberg 2 TPJ has been classified as an element of the ventral frontoparietal network, which is engaged during thereorienting of attention to a salient stimulus (Corbetta, Patel, & Shulman, 2008). This ventral network actsas a “circuit breaker” for the top-down dorsal attentional system, signaling the presence of behaviorallyrelevant stimuli and directing attention towards them. It is consistently recruited in “oddball” tasks, inwhich subjects are instructed to respond to abnormal stimuli in a stream of repetitive inputs (Marois,Leung, & Gore, 2000). Although much of the work on attentional control has been conducted in thevisual domain, there is evidence to suggest that the TPJ functions in its reorienting capacity independentof sensory modality (Corbetta & Shulman, 2002). In addition to its role in the ventral frontoparietalattention network, the TPJ has also been implicated in the perception of one’s own body relative to theexternal environment.

Electrically stimulating the TPJ can lead to the feeling of being extracorporeal,known as an out-of-body experience (Blanke & Arzy, 2005). An experiment analyzing evoked potentialsfrom electroencephalography readings mapped potentials to the TPJ when subjects were instructed toimagine themselves in the position of a human figure presented on a screen (Arzy, Thut, Mohr, Michel, &Blanke, 2006). Taken together, these findings suggest that the TPJ is responsible for maintaining thedistinction between a corporeal self and the external world. Although these low-level cognitive processes seem far from the complicated cognition ofempathy, their computational mechanisms may be crucial for the complexity of social cognition.Throughout the course of evolution, as understanding the intentions and emotions of conspecifics becameincreasingly adaptive, it is possible that the TPJ’s reorienting and body-perception functions were co-opted to serve social cognition.

Its capacity to draw attention to behaviorally relevant information seemsparticularly suited to empathy, as recognizing and interpreting the perspectives of others in socialinteractions is extremely pertinent to one’s behavior in those interactions. Its role in separating the selffrom the environment could be extended to maintain an individual’s sense of self while they areempathizing with another person. Several theorists have taken this perspective, positing that attentionsignals in the TPJ are central to the act of switching between an internal perspective and another’sviewpoint (Corbetta et al.

, 2008). In this view, the TPJ is defined as carrying out a “domain-general Soderberg 3 computational mechanism:” one that is essential for the higher-order social process of mentalizing(Decety & Lamm, 2007). This compelling interpretation provokes the question of whether the temporoparietal junction isespecially attuned to social perspective taking, or if it is simply performing its low-level cognitivefunction in the social domain. This dichotomy can be difficult to tease apart, as findings that might seemto prove the uniquely social character of the TPJ can be interpreted to support the opposite theory.

Forexample, an fMRI experiment found a greater BOLD signal when subjects read stories about characters’false beliefs than when they read about false physical representations, while this difference was not seenfor expected versus unexpected stories (Young, Dodell-Feder, & Saxe, 2010). The authors conclude thatTPJ response is selective for content about mental state, over and above its function for reorientingtowards salient stimuli. However, if the TPJ’s functional domain is seen as responding to a continuum ofbehavioral relevance, and perspective-taking is recognized as requiring a greater cognitive load thanobject perception, the finding that it is more activated during the mental state condition could beexplained by the behavioral relevance and complexity of understanding that mental state. In addition, thisstudy did not examine TPJ activation during a low-level attention paradigm, so it cannot rule out thepossibility that the TPJ’s functional domain encompasses both attention shifting and perspective-taking.Recognizing that omissions of this kind were an issue in the literature, another experiment compared asimilar belief task with an attention reorienting task within one group of participants, and found that thesame region was modulated by both tasks (Mitchell, 2008). This result indicates the need for a conceptionof TPJ function that integrates across attentional, social, and agential domains, rather than claimingselectivity for any one task. Although not directly concerned with taking the perspective of another person, a study wasconducted that proposed a uniquely social function for the TPJ by examining social versus non-socialdecision-making in a simplified poker game. Participants received a card, saw an image of their opponent(either a human face or a computer), and decided whether to bet or fold.

When comparing the BOLDsignal using a calculation of unique combinatorial performance (UCP), which measures the degree of a Soderberg 4 brain region’s predictive performance, the authors found a strong independent contribution of the TPJ forhuman opponent versus computer opponent trials. When a “social bias” metric was computed using UCPfor regions across the entire brain, the TPJ emerged above all other regions in preferential processing ofinformation concerning a social agent. On the basis of this strikingly different social bias, the authorsconclude that the TPJ has a unique sensitivity to the “perceived behavioral relevance of other agents”(Carter, Bowling, Reeck, & Huettel, 2012). Applying the continuum model discussed above, one couldsimilarly attribute this increased social responding solely to the behavioral relevance of the socialstimulus, thus maintaining that the TPJ’s processing of social information is quantitatively, but notqualitatively, different from its attentional function. However, the fact that out of the entire brain, thetemporoparietal region alone stood out on the social bias metric is hard to ignore.

Presumably, the low-level computational mechanisms of some other brain regions are at play in social decisions, so why dothey not exhibit a significant level of social bias? Perhaps there is something about the particularintegrative power of the TPJ that makes it distinctly suited to conceptualizing another person’s point ofview. This dilemma exemplifies a larger controversy in the field: whether any region (or network ofregions) in the brain is inherently “social.” Although there is an intuitive tendency to search for specificneural locations for our psychological concepts, it seems increasingly likely that there is no one-to-onematch of function to region.

It is probable that the quest for regions that are completely selective forsocial input will turn up no results. So while it might be tempting to look for the “seat of empathy,” it islikely that empathetic processing is accomplished through the combined efforts of many different regions(including the TPJ), each performing more basic functions which are then integrated to generate a unifiedpercept. This aligns with the growing emphasis in the field on prioritizing network interactions overindividual structures (Kennedy & Adolphs, 2012).

Although it is undoubtedly useful to elucidate thespecific contributions of each brain region, attention should be paid to how they operate in functionalnetworks. 


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