784 consideration. Aristotle (fourth century BC/1985), for example, argued that being a ‘‘good person’’ requires rea- soning about virtues. Kant’s (1785/1993) famous categor- ical imperative was similarly rational, demanding that one should act according to principles that could also become a general law. More recently, Kohlberg et al. (1983) and Kohlberg (1969) advanced a six-stage developmental model acknowledging children’s increasing ability for abstraction and role-taking capacities in moral decisions. In contrast to rational models, the role of emotion in facili- tating moral behavior has been less often emphasized (Haidt 2001). Hume (1777/2006) provides an early notable exception, as he believed in a key role of intuition for recognizing morally good and bad decisions, not requiring willful abstract reasoning. Charles Darwin (1874/1997) further argued that moral decisions are mainly inﬂuenced by emotional drives, which are rooted in socio-emotional instincts already present in non-human primates. Notions of rationality and emotionality also serve as explanations in the contemporary neuroscientiﬁc literature on the psychological processes underlying moral decisions (henceforth: moral cognition). Results from functional magnetic resonance imaging (fMRI) studies by Greene et al. (2004) and Greene et al. (2001) were interpreted as revealing a dissociation between fast emotional responses and subsequent cognitive modulations in moral cognition. FMRI ﬁndings by Moll and Schulkin (2009), Moll et al. (2005a) and Moll et al. (2006), however, were interpreted as revealing a dissociation between group-oriented (i.e., pro-social) and self-oriented (i.e., egoistic) affective drives in moral cognition. Taken together, either abstract-infer- ential or rapid-emotional processing has been emphasized by most previous accounts explaining moral behavior. Rational explanations assumed that moral behavior arises from a conscious weighing of different rules, norms and situational factors, while emotional explanations empha- sized the inﬂuence of uncontrolled emotional states rapidly evoked by a given situation (cf. Krebs 2008). Previous evidence suggests that, the rational and emo- tional facets of moral cognition are likely related to other social skills: theory of mind (ToM) and empathy. ToM refers to the ability to contemplate other’s thoughts, desires, and behavioral dispositions by abstract inference (Premack and Woodruff 1978; Frith and Frith 2003). Indeed, accu- mulating evidence indicates that moral cognition is inﬂu- enced by whether or not an agent’s action is perceived as intentional versus accidental (Knobe 2005; Cushman 2008; Killen et al. 2011; Moran et al. 2011). Empathy, on the other hand, refers to automatically adopting somebody’s emo- tional state while maintaining the self–other distinction (Singer and Lamm 2009; Decety and Jackson 2004). In moral decisions, experiencing empathy was shown to alleviate harmful actions towards others (Feshbach and 123 Brain Struct Funct (2012) 217:783–796 Feshbach 1969; Zahn-Waxler et al. 1992; Eisenberger 2000). Conversely, the deﬁcient empathy skills in psycho- pathic populations are believed to contribute to morally inappropriate behavior (Hare 2003; Blair 2007; Soderstrom 2003). Taken together, moral cognition is thought to involve the representation of intentions and outcomes as well as (vicariously experienced) emotional states (Decety et al. 2012; Leslie et al. 2006; Pizarro and Bloom 2003). This assumption is further supported by the observation that ToM and empathic skills precede mature moral reﬂection in pri- mate evolution (Greene and Haidt 2002) and in ontogeny (Tomasello 2001; Kohlberg et al. 1983; Frith and Frith 2003; Piaget 1932) given that natural evolution tends to modify existing biological systems rather than create new ones from scratch (Jacob 1977; Krebs 2008). Notably, theoretical accounts as well as empirical evi- dence suggest that ToM and empathy are partially overlap- ping psychological constructs. In particular, it has been proposed that embodied representations of affect, which should be relevant for empathic processing, may be further integrated into meta-representational or inferential process- ing (Keysers and Gazzola 2007; Spengler et al. 2009; De Lange et al. 2008; Mitchell 2005). That is, embodied repre- sentation and meta-representation might not constitute two mutually exclusive processes. In other words, more auto- matic, bottom-up driven mapping and awareness of others emotional states in the context of self–other distinction (i.e., empathy) might be modulated by more controlled, top-down processes involved in attributing mind states (i.e., ToM) (Leiberg and Anders 2006; Pizarro and Bloom 2003; Singer and Lamm 2009). Importantly, ToM and empathy differ in the representational content (mental states versus affect), yet both might be similarly brought about by interaction between top-down and bottom-up processes (Singer and Lamm 2009; Lamm et al. 2007; Spengler et al. 2009; Cheng et al. 2007). In line with the theoretical arguments for a partial overlap between ToM and empathy, ventromedial prefrontal cortex lesions associated with ToM impairments were empirically shown to debilitate elaborate forms of empathic processing, while lateral inferior frontal cortex lesions, which leave ToM skills intact, lead to an impairment of basic forms of empathic processing (Shamay-Tsoory et al. 2009). In sum, converging earlier evidence thus suggests that moral cog- nition is subserved by partially interrelated ToM and empathy processes. From a neurobiological perspective, we therefore hypothesized that moral cognition might be subserved by brain areas also related to ToM and empathy, that is, brain networks associated with abstract-inferential and rapid- emotional processing, respectively. Furthermore, we sought to formally investigate to what extent the neurobiological correlates of ToM versus empathy overlap with the neural network associated with moral cognition. Moreover, we