Distributed affective space represents multiple emotion categories across the human brain

“, 2001) and transforming the masks to the native space using FLIRT (Jenkinson et al., 2002) with nine degrees of freedom.”

“, 2002) and non-brain matter was removed using BET (Smith, 2002).”

“Participant-wise gray matter masks were generated by segmenting the T1-weighted images into gray and white matter and cerebrospinal fluid using the FAST segmentation tool (Zhang et al., 2001) and transforming the masks to the native space using FLIRT (Jenkinson et al.”

“Accordingly, the specific number of basic emotions has been debated and the distinction between basic and non-basic emotions has been questioned (e.g. Russell, 2003).”

“Yet, a wide array of other emotions, including ‘secondary’ or ‘social’ emotions (see reviews and proposed taxonomies in Damasio, 1999; Adolphs, 2002a), also serve adaptive survival functions and are characterized by distinctive facial expressions (Baron-Cohen et al., 2001; Shaw et al., 2005), bodily sensations (Nummen…”

“Second, somatomotor areas including premotor cortex, cerebellum (including vermis and the anterior lobe), globus pallidus, caudate nucleus, and posterior insula were activated during most emotions, but according to the cluster visualizations especially during the processing of emotions that have a strong impact on act…”

“tems which are however not specific to emotional processing per se (Kober et al., 2008; Lindquist et al., 2012).”

“Especially, left hemisphere activation in orbitofrontal cortex connected to rewards and punishments (Kringelbach and Rolls, 2004), as well as in inferior frontal cortex and dorsolateral prefrontal cortex, which subserve language and executive control (Poldrack et al., 1999; Kane and Engle, 2002), and in anterior insul…”

“This accords with prior work suggesting that emotion-specific neural activation patterns might explain why each emotion feels subjectively different (Damasio et al., 2000; Saarimäki et al., 2016).”

“Especially, left hemisphere activation in orbitofrontal cortex connected to rewards and punishments (Kringelbach and Rolls, 2004), as well as in inferior frontal cortex and dorsolateral prefrontal cortex, which subserve language and executive control (Poldrack et al.”

“Nevertheless, the psychological and neural mechanisms of these non-basic emotions, as well as their commonalities or differences relative to basic emotions, remain largely unresolved (Ekman, 1999; Ekman and Cordaro, 2011; Adolphs, 2002b).”

“, 2008), emotion regulation (Schutter and van Honk, 2009), and somatomotor processing and action preparation related to emotional processing (Kohler et al., 2002; Wicker et al., 2003; Mazzola et al., 2013).”

“Nevertheless, the psychological and neural mechanisms of these non-basic emotions, as well as their commonalities or differences relative to basic emotions, remain largely unresolved (Ekman, 1999; Ekman and Cordaro, 2011; Adolphs, 2002b).”

“, 2008) and maternal love (Bartels and Zeki, 2004), as well as aesthetic feelings such as wonder or nostalgia (Vuilleumier and Trost, 2015).”

“…and Rolls, 2004), as well as in inferior frontal cortex and dorsolateral prefrontal cortex, which subserve language and executive control (Poldrack et al. , 1999; Kane and Engle, 2002), and in anterior insula linked to processing of social emotions (Lamm and Singer, 2010) was activated…”

“gration then giving rise to the interpretation of the subjective feeling (Northoff and Bermpohl, 2004; Northoff et al., 2006).”

“Yet, a wide array of other emotions, including ‘secondary’ or ‘social’ emotions (see reviews and proposed taxonomies in Damasio, 1999; Adolphs, 2002a), also serve adaptive survival functions and are characterized by distinctive facial expressions (Baron-Cohen et al.”

“, 2008), as well as amygdala, brainstem and thalamus (Adolphs, 2010; Damasio and Carvalho, 2013; Kragel and LaBar, 2014).”

“In turn, constructivist emotion theories suggest that all emotions are generated by a shared set of fundamental functional systems which are however not specific to emotional processing per se (Kober et al. , 2008; Lindquist et al. , 2012).”

“According to the basic emotion theories, emotions have been shaped during the evolution to serve distinct survival functions via distinct neural circuits and physiological systems (Panksepp, 1982; Ekman, 1992, 1999; Damasio, 1999).”

“The organization of human emotion systems is currently a topic of lively debate (Hamann, 2012; Lindquist et al. , 2012; Kragel and LaBar, 2014; Meaux and Vuilleumier, 2015; Saarima¨ki et al. , 2016; Adolphs, 2017; Barrett, 2017).”

“, 2005), bodily sensations (Nummenmaa et al., 2014a), and neural activity patterns (Kassam et al.”

“all emotion categories, likely reflecting the offset of the auditory stimulation followed by mental imagery of the emotionevoking situation (Ganis et al., 2004) and emotion-related modulation of this activity (Holmes and Mathews, 2005; Nummenmaa et al.”

“…using univariate analyses have quantified neural responses to emotions such as regret (Coricelli et al. , 2005; Eryilmaz et al. , 2011), guilt (Wagner et al. , 2011), pride (Takahashi et al. , 2008; Zahn et al. , 2009; Simon-Thomas et al. , 2012), rejoice (Chandrasekhar et al. , 2008) and…”

“Prior studies using univariate analyses have quantified neural responses to emotions such as regret (Coricelli et al., 2005; Eryilmaz et al., 2011), guilt (Wagner et al.”

“, 2004) and emotion-related modulation of this activity (Holmes and Mathews, 2005; Nummenmaa et al., 2012; Kassam et al., 2013).”

“, 1999; Kane and Engle, 2002), and in anterior insula linked to processing of social emotions (Lamm and Singer, 2010) was activated during these emotions.”

“, 2004) and emotion-related modulation of this activity (Holmes and Mathews, 2005; Nummenmaa et al., 2012; Kassam et al., 2013).”

“…include cortical mid-line regions (Peelen et al. , 2010; Chikazoe et al. , 2014; Trost et al. , 2012), somatomotor regions (Adolphs et al. , 2000; de Gelder et al. , 2004; Nummenmaa et al. , 2008, 2012; Pichon et al. , 2008), as well as amygdala, brainstem and thalamus (Adolphs, 2010; Damasio…”

“The organization of human emotion systems is currently a topic of lively debate (Hamann, 2012; Lindquist et al. , 2012; Kragel and LaBar, 2014; Meaux and Vuilleumier, 2015; Saarima¨ki et al. , 2016; Adolphs, 2017; Barrett, 2017).”

“Our results extend previous studies, which have shown classification of specific emotional states usually focusing on the basic emotions only or a subset of these (Ethofer et al. , 2009; Peelen et al. , 2010; Said et al. , 2010; Kotz et al. , 2013; for a review, see Kragel and LaBar, 2014).”

“This activation might reflect the control of autonomic nervous system’s reactions to different emotions (Critchley et al. , 2005; Linnman et al. , 2012) and/or covert activation of particular motor programs (Blakemore et al. , 2016).”

“These emotions may involve more elaborate cognitive representations acquired through experience, education, and social norms (Panksepp and Watt, 2011), and hence, recruit brain systems partly distinct from those implicated in more ‘primitive’ and possibly innate basic emotions.”

“These areas are engaged during emotion perception (Nummenmaa et al. , 2008, 2012; Pichon et al. , 2008), emotion regulation (Schutter and van Honk, 2009), and somatomotor processing and action preparation related to emotional processing (Kohler et al. , 2002; Wicker et al. , 2003; Mazzola et al. ,…”

“Both of these regions are related to salience processing and emotional arousal (Anders et al., 2004; Adolphs, 2010; Damasio and Carvalho, 2013; Kragel and LaBar, 2014) and show specific activation patterns for basic emotions (Wang et al.”

“In turn, constructivist theories consider specific emotions emerging from the interaction of fundamental processes shared across psychological domains and, thus, not unique to emotions alone (see, e.g. Russell, 2003; Lewis and Liu, 2011; Hamann, 2012; Cunningham, 2013).”

“Third, anterior prefrontal cortex was activated especially during positive emotions (happiness, love, pride, gratitude, longing) according with previous research linking anterior prefrontal cortex with positive affect (Bartels and Zeki, 2004; Zahn et al., 2009; Vytal and Hamann, 2010).”

“These include cortical midline regions (Peelen et al., 2010; Chikazoe et al., 2014; Trost et al., 2012), somatomotor regions (Adolphs et al.”

“These regions might code emotional valence (Colibazzi et al., 2010; Chikazoe et al., 2014), participate in self-relevant and introspective processing (Northoff and Bermpohl, 2004; Northoff et al.”

“, 2004) and emotion-related modulation of this activity (Holmes and Mathews, 2005; Nummenmaa et al., 2012; Kassam et al., 2013).”

“These regions might code emotional valence (Colibazzi et al. , 2010; Chikazoe et al. , 2014), participate in self-relevant and introspective processing (Northoff and Bermpohl, 2004; Northoff et al. , 2006), and integrate information of internal, mental, and bodily states (Northoff et al. , 2006;…”

“, 2008) and maternal love (Bartels and Zeki, 2004), as well as aesthetic feelings such as wonder or nostalgia (Vuilleumier and Trost, 2015).”

“The organization of human emotion systems is currently a topic of lively debate (Hamann, 2012; Lindquist et al. , 2012; Kragel and LaBar, 2014; Meaux and Vuilleumier, 2015; Saarima¨ki et al. , 2016; Adolphs, 2017; Barrett, 2017).”

“have shown classification of specific emotional states usually focusing on the basic emotions only or a subset of these (Ethofer et al., 2009; Peelen et al., 2010; Said et al., 2010; Kotz et al., 2013; for a review, see Kragel and LaBar, 2014).”

“…in Damasio, 1999; Adolphs, 2002a), also serve adaptive survival functions and are characterized by distinctive facial expressions (Baron-Cohen et al. , 2001; Shaw et al. , 2005), bodily sensations (Nummenmaa et al. , 2014a), and neural activity patterns (Kassam et al. 2013; Kragel and LaBar, 2015).”

“Narrative-based guided imagery is known to be an efficient emotion induction technique that engages reliably the brain’s emotion circuits (Costa et al., 2010; Nummenmaa et al., 2014b) and results in strong subjective emotional feelings.”

“have shown classification of specific emotional states usually focusing on the basic emotions only or a subset of these (Ethofer et al., 2009; Peelen et al., 2010; Said et al., 2010; Kotz et al., 2013; for a review, see Kragel and LaBar, 2014).”

“These areas are engaged during emotion perception (Nummenmaa et al., 2008, 2012; Pichon et al., 2008), emotion regulation (Schutter and van Honk, 2009), and somatomotor processing and action preparation related to emotional processing (Kohler et al.”

“, 2011), guilt (Wagner et al., 2011), pride (Takahashi et al.”

“, 2011), pride (Takahashi et al., 2008; Zahn et al., 2009; SimonThomas et al., 2012), rejoice (Chandrasekhar et al.”

“This is in line with previous research showing that surprise tends to separate from other emotions in subjective ratings (Toivonen et al., 2012).”

“…accumulating behavioral evidence suggests that other, non-basic emotions are also characterized by distinctive features in facial expressions (Baron-Cohen et al. , 2001; Shaw et al. , 2005), bodily changes (Nummenmaa et al. , 2014a), and physiology (Kreibig, 2010; Kragel and LaBar, 2013).”

“Prior studies using univariate analyses have quantified neural responses to emotions such as regret (Coricelli et al. , 2005; Eryilmaz et al. , 2011), guilt (Wagner et al. , 2011), pride (Takahashi et al. , 2008; Zahn et al. , 2009; Simon-Thomas et al. , 2012), rejoice (Chandrasekhar et al. , 2008)…”

“have shown classification of specific emotional states usually focusing on the basic emotions only or a subset of these (Ethofer et al., 2009; Peelen et al., 2010; Said et al., 2010; Kotz et al., 2013; for a review, see Kragel and LaBar, 2014).”

“…Eryilmaz et al. , 2011), guilt (Wagner et al. , 2011), pride (Takahashi et al. , 2008; Zahn et al. , 2009; Simon-Thomas et al. , 2012), rejoice (Chandrasekhar et al. , 2008) and maternal love (Bartels and Zeki, 2004), as well as aesthetic feelings such as wonder or nostalgia (Vuilleumier and…”

“emotions, it does not prove that these patterns are strictly necessary for the corresponding emotional state (Nummenmaa and Saarimäki, 2017).”

“…such as regret (Coricelli et al. , 2005; Eryilmaz et al. , 2011), guilt (Wagner et al. , 2011), pride (Takahashi et al. , 2008; Zahn et al. , 2009; Simon-Thomas et al. , 2012), rejoice (Chandrasekhar et al. , 2008) and maternal love (Bartels and Zeki, 2004), as well as aesthetic feelings such as…”

“…such as regret (Coricelli et al. , 2005; Eryilmaz et al. , 2011), guilt (Wagner et al. , 2011), pride (Takahashi et al. , 2008; Zahn et al. , 2009; Simon-Thomas et al. , 2012), rejoice (Chandrasekhar et al. , 2008) and maternal love (Bartels and Zeki, 2004), as well as aesthetic feelings such as…”

“, 2008), as well as amygdala, brainstem and thalamus (Adolphs, 2010; Damasio and Carvalho, 2013; Kragel and LaBar, 2014).”

“In turn, constructivist theories consider specific emotions emerging from the interaction of fundamental processes shared across psychological domains and, thus, not unique to emotions alone (see, e.g. Russell, 2003; Lewis and Liu, 2011; Hamann, 2012; Cunningham, 2013).”

“, 2008), emotion regulation (Schutter and van Honk, 2009), and somatomotor processing and action preparation related to emotional processing (Kohler et al., 2002; Wicker et al., 2003; Mazzola et al., 2013).”

“, 2012) and/or covert activation of particular motor programs (Blakemore et al., 2016).”

“In turn, constructivist theories consider specific emotions emerging from the interaction of fundamental processes shared across psychological domains and, thus, not unique to emotions alone (see, e.g. Russell, 2003; Lewis and Liu, 2011; Hamann, 2012; Cunningham, 2013).”

“cific emotion at both psychological and behavioral levels (Meaux and Vuilleumier, 2015).”

“Recent evidence indicates that it is possible to classify more specific emotions also, such as love and pride within the “happiness” basic-emotion category (Saarimäki et al., 2018).”

“Experiences of speci fi c emotions, then, are found to involve multiple interacting systems – situated near DMN and subcortical regions – that enter distinct states in response to perceived threats and opportunities [9,62,80 – 82].”