Clan mentality: evidence that the medial prefrontal cortex responds to close others

“Stimuli were generated on an Apple MacBook Pro (Apple Computer) using MATLAB software (The MathWorks) and the Psychophysics Toolbox extension (Brainard, 1997) and projected onto a screen at the head of the magnet bore.”

“There is considerable evidence that social species develop mechanisms to aid in identifying and evaluating kin, in-group members, and individuals otherwise relevant to one’s survival (Trivers, 1971; Rendall et al., 1996).”

“Bush et al. (2000), in a comprehensive review, noted that regions of the ACC near to our region respond to a range of tasks laden with affective content, many of which do not have explicit social demands.”

“Lesions to human ventromedial PFC result in failure to represent the consequences of choices in nonsocial decision-making tasks (Bechara et al., 1994, 2000).”

“By the age of 5 or 6, children can flexibly process social information, represent mental states of others, and engage with peers and family in a way that maintains cohesive group interactions (Flavell, 1999; Wellman et al., 2001; Hughes and Leekam, 2004; Fehr et al., 2008).”

“Meta-analyses of neuroimaging studies using social content have emphasized the aMPFC (e.g., Amodio and Frith, 2006; Northoff et al., 2006; Mitchell, 2009).”

“Data were spatially normalized to the Montreal Neurological Institute (MNI) standard anatomical atlas space (2 mm isotropic voxels) using a T2-weighted EPI BOLD-contrast atlas in SPM2 (Wellcome Department of Imaging Neuroscience, London, UK) (Friston et al., 1995).”

“Lesions to human ventromedial PFC result in failure to represent the consequences of choices in nonsocial decision-making tasks (Bechara et al., 1994, 2000).”

“…interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al., 2003; Mah et al., 2005; Rudebeck et al., 2006, 2007) [for review, see Damasio et al. (1996), Stuss (2002), and Adolphs (2003, 2009)].”

“Meta-analyses of neuroimaging studies using social content have emphasized the aMPFC (e.g., Amodio and Frith, 2006; Northoff et al., 2006; Mitchell, 2009).”

“Specifically, human aMPFC and rACC may overlap a “medial network” identified in the monkey that is concerned with visceral and affective systems and is largely distinct from an “orbital network” consisting of mainly ventrolateral-orbital structures (Ongür and Price, 2000).”

“It is thus important to note that explicitly nonsocial tasks also engage the frontal midline, in particular those that require calculating expected reward or value of a choice (e.g., Daw et al., 2006; see also Beckmann et al., 2009 for meta-analysis).”

“After the judgment task participants completed 50 min of resting-state scan conditions as reported elsewhere (Van Dijk et al., 2010).”

“The stimulus set consisted of 160 single-trait adjectives selected from a pool of normalized personality adjectives (Anderson, 1968) and either the word “self” or “Bush” in black text against a white background.”

“In experiment 1 (n 31) participants were scanned as they performed an attribute-judgment task used in previous studies (Kelley et al., 2002) that included themselves and former U.S. President George W. Bush as targets.”

“…and behavioral interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al., 2003; Mah et al., 2005; Rudebeck et al., 2006, 2007) [for review, see Damasio et al. (1996), Stuss (2002), and…”

“…of social interpretation and behavioral interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al., 2003; Mah et al., 2005; Rudebeck et al., 2006, 2007) [for review, see Damasio et al.…”

“By the age of 5 or 6, children can flexibly process social information, represent mental states of others, and engage with peers and family in a way that maintains cohesive group interactions (Flavell, 1999; Wellman et al., 2001; Hughes and Leekam, 2004; Fehr et al., 2008).”

“…(rACC) may be critical for certain components of social interpretation and behavioral interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al., 2003; Mah et al., 2005; Rudebeck et al., 2006,…”

“Data were projected onto the Caret-inflated cortical surface (Van Essen, 2005).”

“Participants completed the task in four runs each containing 25 trials that were interleaved with variable-duration fixation events to optimize jitter (Dale, 1999; Miezin et al., 2000).”

“6C), and similar dissimilar, collapsing across the closeness dimension (Fig.”

“…for certain components of social interpretation and behavioral interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al., 2003; Mah et al., 2005; Rudebeck et al., 2006, 2007) [for review, see…”

“By the age of 5 or 6, children can flexibly process social information, represent mental states of others, and engage with peers and family in a way that maintains cohesive group interactions (Flavell, 1999; Wellman et al., 2001; Hughes and Leekam, 2004; Fehr et al., 2008).”

“We intentionally constrained our ROIs to be relatively small to avoid averaging signal across many brain areas, given the convergence of several different architectonic zones and their corresponding brain networks (Ongür et al., 2003).”

“It is thus important to note that explicitly nonsocial tasks also engage the frontal midline, in particular those that require calculating expected reward or value of a choice (e.g., Daw et al., 2006; see also Beckmann et al., 2009 for meta-analysis).”

“This analysis yielded results similar to those reported above.”

“Other studies have investigated the effects of closeness or familiarity of the target to the participant (Gobbini et al., 2004; Lou et al., 2004; Schmitz et al., 2004; Ochsner et al., 2005; Heatherton et al., 2006; Zhu et al., 2007; Vanderwal et al., 2008).”

“Rather, the loss of signal-tonoise ratio (SNR) owing to susceptibility artifacts (Ojemann et al., 1997) precluded making any observations in the most ventral prefrontal regions (see supplemental Fig.”

“Other studies have investigated the effects of closeness or familiarity of the target to the participant (Gobbini et al., 2004; Lou et al., 2004; Schmitz et al., 2004; Ochsner et al., 2005; Heatherton et al., 2006; Zhu et al., 2007; Vanderwal et al., 2008).”

“Other studies have investigated the effects of closeness or familiarity of the target to the participant (Gobbini et al., 2004; Lou et al., 2004; Schmitz et al., 2004; Ochsner et al., 2005; Heatherton et al., 2006; Zhu et al., 2007; Vanderwal et al., 2008).”

“Responses falling within the rACC are also frequently observed in studies of social cognition (Mitchell et al., 2002; Heatherton et al., 2006; Moran et al., 2006; Ames et al., 2008, Vanderwal et al., 2008).”

“By the age of 5 or 6, children can flexibly process social information, represent mental states of others, and engage with peers and family in a way that maintains cohesive group interactions (Flavell, 1999; Wellman et al., 2001; Hughes and Leekam, 2004; Fehr et al., 2008).”

“Under normal circumstances, personally familiar or relevant individuals are also likely to elicit affective responses or trigger relevant past experiences that may contribute to how individuals are evaluated (Bechara et al., 2003).”

“There is considerable evidence that social species develop mechanisms to aid in identifying and evaluating kin, in-group members, and individuals otherwise relevant to one’s survival (Trivers, 1971; Rendall et al., 1996).”

“For instance, thirsty monkeys will forgo juice to view pictures of other monkeys, but only when they are socially relevant (e.g., either of higher status or a possible mate) (Deaner et al., 2005).”

“Responses falling within the rACC are also frequently observed in studies of social cognition (Mitchell et al., 2002; Heatherton et al., 2006; Moran et al., 2006; Ames et al., 2008, Vanderwal et al., 2008).”

“Responses falling within the rACC are also frequently observed in studies of social cognition (Mitchell et al., 2002; Heatherton et al., 2006; Moran et al., 2006; Ames et al., 2008, Vanderwal et al., 2008).”

“Whether the representation is selectively social or related to valuation of a stimulus or an outcome more generally is unclear (Adolphs, 2009, 2010; Rushworth et al., 2007; Beckmann et al., 2009).”

“Other studies have investigated the effects of closeness or familiarity of the target to the participant (Gobbini et al., 2004; Lou et al., 2004; Schmitz et al., 2004; Ochsner et al., 2005; Heatherton et al., 2006; Zhu et al., 2007; Vanderwal et al., 2008).”

“Other studies have investigated the effects of closeness or familiarity of the target to the participant (Gobbini et al., 2004; Lou et al., 2004; Schmitz et al., 2004; Ochsner et al., 2005; Heatherton et al., 2006; Zhu et al., 2007; Vanderwal et al., 2008).”

“Area 10m projects to the anterior and posterior cingulate, superior temporal sulcus, amygdale, and hippocampal formation (e.g., Petrides and Pandya, 2007; Saleem et al., 2008).”

“Lesions to anterior cingulate gyrus in the monkey do decrease interest in social stimuli (Rudebeck et al., 2006), but while social information is both rewarding and incontrovertibly valuable (Anderson, 1998), this is especially true when such information is useful for behavior.”

“Although similarity was not found to have an effect in our experiments, affective evaluations do interact with mentalizing processes (Hornak et al., 2003; Moran et al., 2006; Olsson and Ochsner, 2008).”

“perceived similarity of others to the self (Mitchell et al., 2006; Rilling et al., 2008; Mobbs et al., 2009; Volz et al., 2009).”

“work suggesting that the self is perceived to be a more appropriate reference point for reasoning about similar others than for dissimilar others (Cadinu and Rothbart, 1996).”

“Based on architectonic evidence in the human (Ongür et al., 2003; Palomero-Gallagher et al., 2008), these two regions are certain to be comprised of multiple, distinct areas.”

“…interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al., 2003; Mah et al., 2005; Rudebeck et al., 2006, 2007) [for review, see Damasio et al. (1996), Stuss (2002), and Adolphs (2003, 2009)].”

“Harris and Fiske (2007) found that pictures of “socially undesirable” people such as the homeless result in more reduced MPFC response than pictures of “socially desirable” people.”

“Tomlin et al. (2006) reported rACC activity in an economic exchange game during observation of partner decisions that were relevant to the participant, but not during a control condition with personally relevant outcomes that were not generated by a social partner, i.e., by a computer.”

“Social context can affect perceptions about in-group constituency (Haslam et al., 1992).”

“Neuroscientific data are increasingly being brought to bear on this question (Mitchell, 2009).”

“…interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al., 2003; Mah et al., 2005; Rudebeck et al., 2006, 2007) [for review, see Damasio et al. (1996), Stuss (2002), and Adolphs (2003, 2009)].”

“…(aMPFC) and rostral anterior cingulate cortex (rACC) may be critical for certain components of social interpretation and behavioral interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al.,…”

“…interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al., 2003; Mah et al., 2005; Rudebeck et al., 2006, 2007) [for review, see Damasio et al. (1996), Stuss (2002), and Adolphs (2003, 2009)].”

“Responses falling within the rACC are also frequently observed in studies of social cognition (Mitchell et al., 2002; Heatherton et al., 2006; Moran et al., 2006; Ames et al., 2008, Vanderwal et al., 2008).”

“…and evolutionary arguments for the idea that both of these dimensions, similarity and closeness, should be processed by brain systems in the service of social decisions, for instance in aid of identifying social status and processing kin or in-group relationships (Platek and Kemp, 2009).”

“This analysis yielded results similar to those reported above.”

“perceived similarity of others to the self (Mitchell et al., 2006; Rilling et al., 2008; Mobbs et al., 2009; Volz et al., 2009).”

“perceived similarity of others to the self (Mitchell et al., 2006; Rilling et al., 2008; Mobbs et al., 2009; Volz et al., 2009).”

“…interactions (Glees et al., 1950; Eslinger and Damasio, 1985; Damasio et al., 1990; Rolls et al., 1994; Stone et al., 1998; Anderson et al., 1999; Hadland et al., 2003; Mah et al., 2005; Rudebeck et al., 2006, 2007) [for review, see Damasio et al. (1996), Stuss (2002), and Adolphs (2003, 2009)].”

“, 2006), but while social information is both rewarding and incontrovertibly valuable (Anderson, 1998), this is especially true when such information is useful for behavior.”

“Activity in these clusters has been linked to social cognition when people make social judgments or employ theory of mind to predict the actions, beliefs, and intentions of others (Amodio and Frith, 2006; Lieberman, 2007; Behrens et al., 2008, 2009; Krienen et al., 2010) (Fig.”

“…large portion of the variance in aMPFC activity during introspective tasks (Andrews-Hanna et al., 2010a), and aMPFC activity increases with perceived self-relevancy (Macrae et al., 2004; Mitchell et al., 2006; Moran et al., 2006, 2009; Benoit et al., 2010; Krienen et al., 2010; Denny et al., 2012).”

“about close others with whom participants experience self-other overlap (Mitchell et al., 2006; Krienen et al., 2010).”

“For instance, previous research has implicated MPFC in distinguishing friends from strangers 13 , and a recent study implicated the hippocampus and posterior cingulate cortex (PCC) in tracking social distances between participants and characters in an interactive game 18 .”

“Medial prefrontal and VS activity has been linked to computations of value (Levy and Glimcher, 2012), the degree of closeness with other people (Harris and Fiske, 2007; Krienen et al., 2010), thinking about the self (Denny et al.”

“The ventral mPFC may also be activated when representing the mental states of similar or close others (Mitchell et al., 2006; Jenkins et al., 2008; Krienen et al., 2010) or when judging personal traits of a close other (Zhu et al., 2007; Wang et al., 2012).”

“In summary, we detail the previous idea of ongoing social cognition as a possible neurophysiological baseline (Schilbach et al. 2008, 2012; Krienen et al. 2010) by identifying a characteristic task-rest sub-differentiation in social brain systems.”

“This entices to speculate that the human vPCC mediates self-relevance assessment, potentially extending to the hierarchically higher processes self-monitoring and self-reflection (Bzdok et al., 2013a; Johnson et al., 2002; Kircher et al., 2001; Krienen et al., 2010; Vogt, 2005).”

“Evidence implicates mPFC in various social processes (Kelley et al., 2002; Ochsner et al., 2004; Amodio and Frith, 2006) relating to self and close or similar others (Heatherton et al., 2006; Mitchell et al., 2006; Krienen et al., 2010).”

“Representations of the self and of close others recruit mPFC (Heatherton et al., 2006; Mitchell et al., 2006; Krienen et al., 2010) and, within social interactions, distinct regions of mPFC represent cooperative human partners (McCabe et al., 2001), volatility of the social environment (Beh-rens et…”

“The closer a stimulus or event is to one’s self-concept, the more it matters—and the stronger the vmPFC activity (Krienen et al. 2010, Tamir & Mitchell 2010).”

“(b) Increased responses in the ToM areas and the precuneus were also observed when participants answered regarding personal preferences of their friends (Krienen et al., 2010).”

“present MDP account can parsimoniously incorporate the dominance of social content in human mental activity as high value function estimates for information about humans (Baker et al., 2009; Kampe et al., 2001; Krienen et al., 2010).”

“Furthermore, it is increasingly recognized that self-processing extends to incorporate significant others (123,124).”

“Since psychological constructs such as familiarity, but also similarity and physical proximity, have previously been demonstrated to upregulate mPFC activity (Jenkins et al., 2008; Krienen et al., 2010; Mitchell et al., 2006; Tamir and Mitchell, 2011), this raises the question whether an increase in familiarity might…”

“Thus, DMPFC may have been activated when participants contemplated targets who were dissimilar to themselves, while MPFC may have been activated when thinking about similar targets (Mitchell et al., 2006; Krienen et al., 2010).”

“…when making decisions pertaining to other people we value, including our friends and relatives (Benoit, Gilbert, Volle, & Burgess, 2010; Krienen, Tu, & Buckner, 2010; Murray, Schaer, & Debbane, 2012), as well as those we deem similar (Mitchell, Macrae, & Banaji, 2006; but see Krienen et al., 2010).”

“For example, vmPFC is more active for judgments about the self than about others (D’Argembeau et al., 2008; Krienen et al., 2010), and mediates the selfreference effect in memory (Philippi et al., 2012).”

“Our results support the proposal that the MPFC mediates a function to represent self-relatedness of others (Northoff et al., 2006; Han and Northoff, 2009; Krienen et al., 2010) and can be used as an objective measurement of close relationships, complementing subjective evaluations of close relationships.”

“Medial prefrontal cortex, pACC, DMPFC and PCC are also often active in tasks that involve thinking about the thoughts and feelings of others (Frith & Frith, 2006; Krienen et al., 2010; Lieberman, 2010; Mitchell, 2009).”

“Importantly, the choice of one type of music over another likely reflects limbic, decision-making brain regions, such as the prefrontal cortex (reviewed in 19), which is known to be engaged during judgments of kinship and close others later in life (20).”

“…of people and evaluations of self are associated with medial prefrontal cortex (MPFC) and posterior cingulate cortex (PCC) activity (e.g., Vogeley et al. 2001; Harris et al. 2005; Mitchell et al. 2005; Ochsner et al. 2005; Lindner et al. 2008; Krienen et al. 2010 but see Kelley et al. 2002).”

“…to be similar to the self engaged this ventral mPFC region, whereas social targets manipulated to be dissimilar to the self engaged dorsal mPFC. Krienen et al. (2010) clarified Mitchell et al.’s findings by demonstrating that the driver of activation in mPFC was closeness rather than…”

“The current finding of spontaneous activity in self-sensitive brain areas induced by self-associated objects is in line with behavioral and neural findings suggesting incorporation of close others in one’s selfconcept (Aron et al., 1991; Mashek et al., 2003; Krienen et al., 2010).”

“Prior closeness findings (Krienen et al., 2010) are also consistent with the ToMp account of MPFC, insofar as we typically represent the minds of close others with detailed, idiosyncratic models.”

“However, more recent research has demonstrated that regions within the medial PFC respond more strongly when participants make judgments about friends compared to strangers, regardless of whether the other person is perceived as similar to the self (Krienen et al., 2010).”

“In particular, greater MPFC activation has been associated with interpersonally close others [63–67], and with humanized others [68], compared to those who are more distant or dehumanized.”

“162 Interestingly, pattern similarity within the components of the DMN (i.e., mPFC) also appears to cluster according to the social closeness we feel toward others.”

“, familiar) and/or perceived to be similar in personality by the participant (Mitchell et al., 2006; Krienen et al., 2010), we also expected to find neural differences when otherbased counterfactuals involved either familiar and/or similar characters.”

“Neuroimaging studies show the mPFC is preferentially activated when mentalizing about similar and psychologically close others (Krienen et al., 2010; Tamir and Mitchell, 2011; Denny et al., 2012).”

“Neural evidence of these principles has been detected in cortical and limbic brain circuits (Rilling et al., 2002; Krienen, Tu, & Buckner, 2010; Watanabe et al., 2014) and there is some evidence that the relative strength of these circuits can provide motivation for certain types of altruistic…”

“A paradox is introduced when attempting to measure unconstrained mind-wandering by imposing this behavior on the participants as the task to follow while lying in a brain imaging scanner cf. [58, 59, 60].”

“Consistently, a previous fMRI study had shown that the medial prefrontal cortex responded more strongly when participants made judgments about the personal preferences of friends and close others, which arguably are likely to trigger relevant past experiences, relative to strangers, regardless of whether the other per…”

“Activity in the dorsomedial prefrontal cortex is associated with social cognition both when making judgments about others’ mental state or intentions (Amodio and Frith, 2006; Lieberman, 2007; Behrens et al., 2008; Krienen et al., 2010) and while reflecting on one’s own self, beliefs, intentions, and actions (Amodio an…”

“Neuroimaging evidence also demonstrates that regions of the brain that are reliably active when reasoning about the mental states of others are more active when people think about close others than when people think about more distant others (Krienen et al. 2010).”

“, 2006) and in judging social closeness (Krienen et al., 2008).”

“The vmPFC activation is related to processing similar others (Schmitz et al., 2004; Ochsner et al., 2005; Jenkins et al., 2008; Chen et al., 2010; Krienen et al., 2010), which is consistent with our finding that the vmPFC was activated in response to the self and similar others in both the TD group and the ASD group.”

“In Experiment 1, we modified a well-established experimental paradigm that has been widely used to probe the neural basis of self-knowledge (e.g., Kelley et al. 2002; Krienen et al. 2010; MeyerandLieberman 2018).”

“This is in line with the fact that processing personally relevant information related to close others engages sectors of the mPFC and the PPC/Prec [Buckner et al., 2008; Krienen et al., 2010].”

“, 2006) and close friends in comparison with similar others (Krienen et al., 2010).”

“For instance, my favorite coffee mug may be personally relevant to me, but not necessarily self-relevant in the strict sense of being part of my core self-identity.”

“First, we successfully replicated findings on mentalizing (Heatherton et al. 2006; Krienen et al., 2010; Mar, 2011), as all social conditions and the self-condition engaged regions of the default network including dmPFC, vmPFC and PCC, in addition to lateral PFC.”

“Increased activation of the precuneus and adjacent posterior cingulate cortex has also been linked to increases in retrieval of autobiographic memories (Cavanna and Trimble, 2006; Krienen et al., 2010).”

“between socially close and distant others? The medial PFC, part of the social brain network, has been shown to respond differently when making judgments about close versus unfamiliar others (Krienen et al., 2010) and self-reported social proximity positively correlated with”

“Our study offers novel evidence in favor of the thesis that the vmPFC’s role for racial evaluations is largely confined to the more positively valued and prized groups, potentially through regulating desirability-related emotions or representing values assigned to these social groups (Kringelbach and Rolls, 2004; Mitc…”

“Activations in this region have been associated with making judgments about close others and the self [21,25].”

“Closeness or intimacy between self and other appears to reduce the differentiation observed in this region (Chen et al. 2010; Krienen et al. 2010; Ochsner et al. 2005; Schmitz et al. 2004).”

“…and complex incentives that can broaden our understanding of the social brain beyond what we can learn from simpler games (Güroglu et al. 2008; Krienen et al. 2010; Redcay et al. 2010), and neuroimaging studies that utilize true face-to-face interactions as part of their design will play a…”

“The frontal lobe areas involved in selfreferential processing and empathy, particularly orbital and medial prefrontal cortices have been shown to respond more to in-group members and friends (Volz et al., 2009; Freeman et al., 2010; Krienen et al., 2010) and showed reduced activity in response to dehumanized and stigm…”

“…are familiar and closely related to the self, we hypothesize that adolescents will show more activation to parents than to an unfamiliar peer in self-relevant regions, specifically regions within the mPFC demonstrated to be more responsive to familiar than to similar others (Krienen et al., 2010).”

“Conway, Lowder, & Cabeza, 2011), interpersonal closeness (people who are closer, or more important to the self elicit responses more ventrally, Krienen et al., 2010), or affective content ( " hot " affective states elicit responses more ventrally (Ames, Jenkins, Banaji, & Mitchell, 2008;…”

“…DACC activation appears to represent conflict (Botvinick et al., 1999) and MPFC likely represents mentalizing (Figure 5(c); Hampton et al., 2008; Krienen et al., 2010); therefore, we suggest that the less information about conflict that is online and readily available to the MPFC during a…”

“Interestingly, previous studies in adults indicate a stronger vMPFC involvement for stimuli that are closer or more important to oneself [Krienen et al., 2010; Murray et al., 2012; van der Meer et al., 2010].”