Is there a core neural network in empathy? An fMRI based quantitative meta-analysis

“…the bilateral SMA, bilateral AI, and bilateral putamen — brain areas, which are well known for their involvement in empathy (Carr et al., 2003; Fan et al., 2011; Hooker et al., 2008, 2010; Jabbi and Keysers, 2008; Jabbi et al., 2007; Lamm et al., 2007; Mathur et al., 2010; Singer et al.,…”

“2003; Blair 2008; Decety 2010; Decety and Michalska 2010), the 2 previous metaanalyses in the domain of empathy for pain did not specifically highlight these regions as being consistently activated across studies (Fan et al. 2011; Lamm et al. 2011).”

“…neural network that comprises the bilateral anterior insula, the anterior cingulate cortex, the thalamus, and the medial prefrontal cortex [Fan et al., 2010, 2011; Molenberghs et al., 2011; Singer and Lamm, 2009; Decety et al., 2006; Lamm et al., 2007, 2011] that is activated during the…”

“The anterior cingulate cortex may have a special role in parental empathy. fMRI metaanalyses implicate the anterior cingulate in empathy (Fan et al., 2011), and a recent review concluded that the anterior cingulate gyrus is central to the vicarious experience of both pain and reward (Lockwood,…”

“For an analysis of such emotional stimuli, showing results consistent with the present study, the reader is referred to the meta-analyses by Fan et al. (2010) and Kober et al. (2008).”

“The neural representation of empathy has received considerable attention in recent years (for meta-analyses, see Fan et al., 2011a; Lamm et al., 2011).”

“, 2017), others suggest that the MCC represents a cognitive-evaluative form of empathy, whereas the AI represents an affective-perceptual form (Fan et al., 2011).”

“For example, some early researchers tended to divide empathy into cognitive empathy and affective empathy and defined cognitive empathy as the process of accurately recognizing and understanding others’ feelings and emotions, whereas affective empathy is defined as emotional resonance or empathic accuracy, that is, th…”

“There is increasing evidence from neuroimaging and lesionmapping studies that the insula, and especially its anterior portion, is a key structure within the ‘empathic network’ (Driscoll et al., 2012; Fan et al., 2011; Leigh et al., 2013; Morelli and Liberman, 2013; Singer et al., 2004).”

“…consisting of the anterior insula/inferior frontal gyrus (AI/IFG) and cingulate cortex, i.e. within its rostral-ventral (ACC) and midcingulate (MCC) subdivisions, corresponding to Brodmann areas 24 and 32 (e.g. (Fan et al., 2011; Morrison et al., 2004; Morrison et al., 2007a; Singer et al., 2004).”

“For internally guided decision-making, the clusters in MPFC, pACC, PCC, and superior temporal gyrus (STG) were revealed.”

“Implication of inferior frontal and inferior parietal mirror neuron areas is not a universal finding in the empathy literature, and some have suggested that it may reflect specific socially relevant tasks or stimulus types, not empathy in and of itself (Fan et al., 2011).”

“In addition to its role in social reward and the integration of affective and social-cognitive processes, a recent meta-analysis of imaging studies on the neural basis of empathy described a dACC–SMA axis that was observed across all studies of human empathy, whether to the pain, social exclusion, anxiety or happiness…”

“Key neural regions of interest were retrieved from three recent meta-analyses on neural activations associated with empathy (Seitz et al., 2006; Fan et al., 2011; Lamm et al., 2011), which include the ventromedial prefrontal cortex (VMPFC), ventral anterior cingulate cortex (VACC), dorsol ACC…”

“…expressions (e.g. Singer et al., 2004; Jackson and Decety, 2005; Lamm et al., 2007; Gu and Han, 2007; Sheng et al., 2014; Cui et al., 2015; Luo et al., 2015; see Lamm et al., 2011; Fan et al., 2011; Jauniaux et al., 2019; Ding et al., 2020; Fallon et al., 2020 for a review and meta-analysis).”

“A meta-analysis of studies has demonstrated overlapping networks for what the authors referred to as affective-perceptual and cognitive–evaluative empathy (Fan et al., 2011).”

“Despite its role in empathy (Lamm et al., 2010; Fan et al., 2011) and in emotion understanding (Zaki et al., 2012), hyperactivity in the AI has long been implicated in anxiety and anxiety disorders (Critchley et al., 2004; Simmons et al., 2006, 2008, 2011).”

“The embodied-simulation/empathy network was defined on the basis of recent meta-analyses on empathy (Fan et al., 2011) and mirror properties (Molenberghs”

“We found the typical neural empathic response to painful versus non-painful touch, involving activation bilaterally in the insula cortex, as found in many previous studies (Singer et al., 2004; Lamm et al., 2011; Fan et al., 2011).”

“For related examples involving such analyses showing results consistent with the present study, see [121,122].”

“A recent meta-analysis, in fact, included this region in an empathy network, including insular, cingular and supplementary motor cortices.(78) Another frontal region, Brodmann area 6, according to our multimodal analysis, showed an overlapping modulation by oxytocin and mother–infant stimuli.”

“…motor area (SMA) (Singer et al., 2004; Jackson et al., 2005; Saarela et al., 2007; Gu and Han, 2007; Han et al., 2009; Gu et al., 2010, 2012; Fan et al., 2011; Lamm et al., 2011), recent research has revealed increased ACC/SMA activity to perceived painful stimuli applied to (Xu et al.,…”

“Also included were the anterior insula, superior temporal sulcus (STS), and temporal pole (TP) that have been repeatedly implicated in human empathy research (32, 35).”

“Patient studies (Adolphs, 2009; Blair, 2005) and human neuroimaging experiments (Fan et al., 2011; Lamm et al., 2011) have contributed substantially during the past decades in characterizing the neural correlates involved in these processes (Gonzalez-Liencres et al., 2013).”

“Facial synchrony (Light et al., 2015) Occurrence of positive and negative emotions from facial electromyography recorded from corrugator (frowning) and zygomatic (smiling) Motion synchrony (Varni, Camurri, Coletta & Volpe, 2009) Phase synchronization and reoccurrence quantification approach to measure nonverbal cues by…”

“Furthermore, fMRI research suggests that the MCC is part of a neural network involved in empathy (Fan et al., 2011).”

“…meditators, practicing mindfulness meditation activates the prefrontal cortex, anterior cingulate cortex, and anterior insula (Chiesa & Serretti, 2010), areas that are part of a core network activated during empathy (Fan, Duncan, de Greck, & Northoff, 2011; Masten, Morelli, & Eisenberger, 2011).”

“…in the anterior middle cingulate cortex (aMCC), dACC, supplementary motor area, and anterior insula/IFG, with the aMCC more frequently reported in studies of cognitiveevaluative empathy, where subjects are explicitly instructed to evaluate others’ emotional or sensory states (Fan et al. 2011).”

“…that empathy and compassion are multifaceted constructs, and that measuring both affective and cognitive processes (i.e., feelings and attributions) will be important for understanding compassion behavior (Ashar et al., in press; Fan et al., 2011; Shamay-Tsoory et al., 2009; Zaki & Ochsner, 2012).”

“The embodied simulation network enables parents to respond to infant pain and emotions by representing them in the self (Fan et al., 2011; Feldman, 2015a), ground emotional experiences in the present moment on the basis of interoceptive representations (Craig, 2009; Gallese, 2014), and afford perceptual-motor coupling…”

“Physiologically, it was discovered how practicing paying attention to one’s inner experience develops the prefrontal cortex, anterior cingulate cortex, and anterior insula, the same regions in the brain that are needed to improve empathy [59,60].”

“Indeed, while prior work has identified a core neural network that appears to underlie both components of empathy (Fan et al., 2011), studies have also identified distinct neural regions for cognitive and affective empathy.”

“We also calculated contrast values in the ACC and AI defined in a meta-analysis (Fan et al., 2011).”

“…around the rostrum of the corpus callosum, is generally involved in emotional awareness (Devinsky et al. 1995), a recent meta-analysis of neuroimaging research on empathy found that the left dorsal anterior mid-cingulate cortex is specifically pertinent to cognitive empathy (Fan et al. 2011).”

“Likewise, the AI has also been identified in a variety of tasks associated with cognition, emotion or the interaction between them [Gu et al., 2012, 2013a,b; Fan et al., 2011].”

“Empathy is also involved in social decisions, emotional contagion, helping behaviors and group belongingness [47,48,49].”

“Therefore, our data support a dissociation between cognitive empathy and affective empathy in modulating decision-making in moral dilemmas, in line with the evidence of a functional distinction between these two dimensions, which appear to involve different processes and to rely on partly distinct neural networks [40,…”

“These brain regions are known to be largely implicated in emotional processing (Fan et al., 2011; Fusar-Poli et al., 2009; Kurth et al., 2010; Lamm et al., 2011).”

“It is well known that the ability to share the physiological and emotional states of others is a basic dimension of empathy [4], which is broadly defined as the experiencing of an affective or sensory state similar to that shown by another individual [5].”

“…consistently activated in empathy, including anterior and mid-cingulate cortex, insula, dmPFC, mPFC, septum and amygdala (Decety and Grezes 2006; Fan et al. 2011; Morelli and Lieberman 2013; Seitz et al. 2006; Singer and Lamm 2009) that have been consistently shown to also become activated in…”

“…insula has a strong connection with the amygdala (Naqvi and Bechara, 2009; Meyer- Lindenberg and Tost, 2012; Sescousse et al., 2013) and is involved in emotional processing and empathy (Vogt, 2005; de Vignemont and Singer, 2006; Fan et al., 2011; Morita et al., in press; Ponz et al., in press).”

“The ACC is probably the highest brain region in the hierarchy of affective responses closely related to the limbic system.(35,36) The 2 subgroups of patients with positive and negative BOLD responses are too small to detect a reliable difference.”

“, 2004; Zaki, Wager, Singer, Keysers, & Gazzola, 2016), particularly in response to close others or in-group members (Beckes, Coan, & Hasselmo, 2012; Hein, Silani, Preuschoff, Batson, & Singer, 2010), and has been implicated in affective empathy and nonreciprocal altruism more generally (Fan et al., 2011; Kanske, Bock…”

“Owing to the requirements of neuroscientific methods and their experimental approach of replicable and standardized measurement conditions and stimuli, SN has predominantly exposed participants to abstract and standardized situations, such as pictures or videos of other persons in aversive states (for recent overview…”

“…the anterior insula, dACC and IFGOp, structures consistently reported to be involved in face processing as well as in empathic processing of pain and disgust (Fan et al., 2011; Lamm et al., 2011); and also the amygdala, a region associated with responding to emotional saliency (Adolphs, 2010).”

“Interestingly, this network seems independent of whether spontaneous or instructed empathizing is tested (Fan et al. 2011).”

“Further, tasks tapping cognitive empathy versus emotional empathy have been found to result in different patterns of activation (Fan et al., 2011; Shamay-Tsoory, 2011).”

“The five ROIs defined here were drawn from a recent meta-analysis that identified brain regions associated with empathy for pain (Fan et al. 2011).”

“A whole-brainbased quantitative meta-analysis of recent fMRI studies identified that the dorsal anterior cingulate cortex (dACC), anterior midcingulate cortex (aMCC), supplementary motor area (SMA) and bilateral anterior insula are consistently activated during empathy [22].”

“Additionally, it would be important to keep in mind that structures lying deeper in the brain such as the insula or cingulate cortex are also likely important for mentalizing [49,50].”

“Previous research has identified distinct clusters of brain regions involved in affective empathy: medial/anterior cingulate cortex (MCC, ACC), anterior insula (AI) (Fan et al., 2011; Lamm et al., 2011), and supplementary motor area (SMA) (Lamm et al., 2011).”

“Among nonparents, extensive research on the neural mechanisms of perceptual affective empathy induced by imitating others’ emotions (Abu-Akel, 2003; Fan et al., 2011) has highlighted the functional importance of the right frontoinsular (RFIC) and rolandic operculum (ROC) cortices in the neural operations of action rep…”

“ies that elicit empathy during fMRI (Fan et al., 2011; Lamm et al., 2011) find consistent activation of the insula bilaterally, as well as the anterior”

“Both cognitive empathy and affective mentalizing recruit similar neural networks [Fan et al., 2011; Gonzalez-Liencres et al., 2013; Hooker et al., 2008; Smith et al., 2015], and arguably describe synonymous behaviors [Walter, 2012].”

“, 2015]); these regions are the middle cingulate cortex (MCC), the supplementary motor area (SMA) [Fan et al., 2011], and the dlPFC [Lieberman, 2007 for a review; Weissman et al.”

“A multitude of imaging experiments in humans in the domain of empathy for pain, disgust, taste, and touch revealed that, in contrast to mirror neuron networks, in the domain of motor actions, sharing sensations and feelings with others engages somatosensory cortices, as well as anterior parts of the insula and anterio…”

“Though not consistently activated by many of the empathy-forpain tasks utilized by functional neuroimagers (Fan et al., 2011; Lamm et al., 2011), the amygdala is arguably a core structure that subserves empathy and compassion.”

“These results replicate the previous neuroimaging findings of activations in the empathy and theory-of-mind networks in response to perceived pain in others (Fan et al., 2011; Lamm et al., 2011; Shamay-Tsoory, 2011), and of enhanced neural responses to ingroup rather than outgroup pain (Xu et al.”

“…studies reported an involvement of the insula in the first-hand experience of pain and vicarious pain induced by observing another person in a painful situation or expressing a painful facial expression (Bzdok et al., 2012; Corradi-Dell’Acqua et al., 2016; Fan et al., 2011; Singer et al., 2004).”

“The cingulate cortex is considered a key correlate of both alexithymia and empathy (for meta-analyses of fMRI studies, see Fan et al., 2011; Lamm et al., 2011 for empathy; van der Velde et al., 2013 for alexithymia).”

“In studies localizing functional correlates of empathy for pain, the dorsal anterior insula (dAI) and anterior midcingulate cortex (aMCC), in particular, have been shown to play a pivotal role in first-hand and vicarious experience of pain (Fan et al. 2011; Lamm et al. 2011).”