The Evolutionary Basis of Empathy

Derived from the Greek empatheia (where en = "in" and pathos = "feeling"), empathy is defined as "the capacity to be affected by and share the emotional state of another, assess the reasons for the other's state, and identify with the other, adopting his or her perspective". Scholars first conceptualized empathy as a cognitive process (e.g., perspective-taking) before describing it in affective terms (e.g., distress), and gradually began approaching it as a multifaceted construct. The evolution of empathy has been compared with a Russian matryoshka doll where the inner core, believed to have developed first, is nested within a series of outer layers. Emotional contagion, the most primitive core of empathy, involves the emotional synchrony between people. It is then followed by sympathetic concern, which is the concern one has about another's state and any subsequent attempts made to ameliorate that state (i.e., consolation). Finally, perspective taking, or the capacity to understand another's situation and needs as separate from one's own, is considered to be the most outer layer of empathic development and the most recent one to have evolved.

What explains how and why empathy evolved in humans? Ever since Aristotle presented his "Four Causes," the question of causality has been central to scientific inquiry. Inspired by Aristotelian logic, ethologist Nikolaas Tinbergen devised a system to better comprehend behavior, one that rested on four basic questions about causality known commonly as the Four Why's. These questions fall into two categories, one dealing with proximal explanations of behavior and the other addressing ultimate ones. The logical starting point when using evolutionary principles begins by observing behaviors to understand how each behavior is structured, and why it was selected for. Whereas proximate causation relates to how a behavior operates, ultimate causation refers to why a behavior exists in the first place. More specifically, proximate explanations deal with the structural machinery underlying the behavior (mechanism or causation) and with the behavior's developmental change over time (ontogeny). Ultimate or evolutionary explanations, on the other hand, are concerned with the benefits the performer receives of having engaged in the behavior (adaptation or function) and the description of the behavior's history in a particular species (phylogeny).

Researchers in evolutionary psychology have been divided on the basis of which category of Four Why's to pursue when studying human behavior. Whereas some have focused primarily on (mostly cognitive) proximate mechanisms, others endorse a holistic view that behavior is best understood by considering both proximate mechanisms and ultimate function. Adherents of the former approach treat the mind not as a blank slate or tabula rasa but instead as an adaptive toolbox that evolved under ancestral environments. Its intellectual spirit drew in part from the human sociobiology movement of the late 1960s and mid-1970s which explained that differences and similarities in behavioral patterns were related to fitness costs. Like sociobiologists, evolutionary psychologists of this school contend that natural selection worked on the effects of behavior on individuals who interacted repeatedly over time. During their course, however, selection pressures designed a multitude of specialized neural circuits or modules. Housed in the brain, these domain-specific modules act like information processing units whose operations are integrated to produce behaviors that help humans solve countless adaptive problems each affecting human survival and reproduction. Therefore, whereas the level of explanation in sociobiology is behavior, the one in evolutionary psychology is the psychological mechanism.

Recent work by de Waal suggests that empathy may have evolved as a proximate mechanism for altruism. Individuals across cultures engage in cooperative behaviors, sometimes risking bodily harm or incurring other costs, such as time and financial resources. Throughout history, helping within collective groups had been a function of the degree of genetic relatedness between helpers and recipients, with participants preferring to help their closest biological kin. For this reason, empathy is believed to have originated from parental care and is as phylogenetically ancient as humans themselves. Infants worldwide communicate their affective states through smiling and crying, and these signals act as vital cues for caregivers to respond. Therefore, parents who possessed an intuitive understanding of the feelings, emotions, and intentions of their infants were more likely to transmit their "empathy genes" to future generations, while those who were indifferent or unsuccessful at soliciting their needs risked losing them. Indeed, the evidence on parental responses to infant distress points to a conclusion that empathy triggers parental sensitivity and subsequent caregiving behaviors.

Natural selection may also account for the evolution of empathy among individuals who share no familial ties. According to the theory of reciprocal altruism, selection acted on the recurring interactions between non-kin members who forged long-term exchange relationships to facilitate each individual's fitness. Thus, psychological mechanisms for offering help and assistance to non-relatives were able to evolve so long as support was mutually reciprocated in the future. Should the survival and reproductive benefits that one party received be larger than the costs the other sustained in providing help, then those who engaged in this type of reciprocation out-reproduced those who did not, causing this kind of helping design to proliferate throughout a population. According to Trivers, among the parameters that are required for reciprocally altruistic behaviors to be selected for, a given species must depend on one another and interact repeatedly over time to assist in a myriad of duties, such as caring for the offspring of non-kin and enlisting each other's help in combat. Natural selection therefore seems to have favored altruism among familiar individuals and previous cooperators - both kin and non-kin - and was likely biased against previous defectors. Such altruism in response to another's distress would not have developed in the absence of empathy. It remains to be seen, however, how evolution can explain sex differences in empathy and how we evolved to perceive and come to expect women to be more empathic than men and, allegedly, better suited to handle crises.


Sex Differences in Empathy

Inasmuch as evolution through natural selection shaped the mind in relation to survival, evolutionary pressures also exercised their influence on the mind through sexual selection. According to this process, men and women ought to exhibit differences in domains where each sex has been faced with distinct adaptive problems. Reproduction is one such domain where the reality of one sex is different from that of the other; across cultural and temporal contexts, compared to men, women have experienced more hardship in childbirth and in ensuring the survival and safety of their children. A subtheory of sexual selection, parental investment theory, maintains that the sex that invests the most in the survival and subsequent raising of offspring should be more discriminating in mate choice, whereas the sex that invests less should compete more aggressively for mating opportunities with the higher-investing one. In other words, behavioral differences between women and men can be explained in part by the roles that each sex faced with respect to resources needed in parental care. Whereas men can choose to invest minimally in the future of their offspring (i.e., a simple act of intercourse), women must bear greater costs extending over a lengthier period of time (i.e., pregnancy sickness, gestation, morbidity, and mortality during childbirth, lactation). In light of their disproportionate costs in raising children and incurring greater costs with the selection of a poorly suited mate, women have evolved a capacity to be generally more sensible in their mating choices compared with men. Therefore, one could argue that women share a Darwinian heritage in being adept at imagining oneself in another's situation and understanding another's feelings, desires, and intentions. Because women typically invest more parental resources than men, having empathy would have been a key asset in this regard. By comparison, women deficient in empathy were more likely to experience interactions with partners and kin who failed to provide them with assistance needed in child rearing. To say that empathy was favored exclusively in women, however, is misleading. Empathy was selected for not only through parent-infant interactions involving mothers as primary caretakers, but also through associations with other group members to facilitate forgiveness among exchange partners and to ensure future cooperation. Men have benefitted and continue to benefit from having empathy as parents and non-parents. The ability to understand others and experience their thoughts and feelings is tantamount in developing genuine social relations irrespective of sex or civil status. However, the mother-infant relationship appears to play a special role in fostering empathy as "infants are emotionally affected by the state of their mothers and mothers are emotionally affected by the state of their offspring".

There now exists a panoply of studies from numerous disciplines on sex differences in empathy. For example, over 40 years ago, Mehrabian and Epstein found that women were more empathic than men (d = 0.98), and suggested that separate statistics be used for each group. More than two decades later in a meta-analysis on sex differences in personality (N > 105,000), Feingold reported that women scored higher than men on tender-mindedness (d = 1.07), a construct known to overlap with empathy. Sex differences in empathy have also been observed indirectly in research on the Big Five model of personality. For example, in a study of 26 cultures (N > 23,000), Costa et al. found that women were more agreeable than men (d from 0.05 to 0.55). Big Five measures of agreeableness, as Nettle noted later, were equivalent to measures of empathy using the Empathy Quotient with women scoring higher than men on both traits (d = 0.60 on agreeableness and d = 0.63 on EQ). These values are consistent with those of Lippa (d = 0.56, N > 250,000) although larger than the ones reported by Schmitt et al. (N > 17,500; mean d = 0.15). Finally, even when using a shortened version of the EQ scale and risking to compromise reliability, Andrew et al. found that women scored higher than men in empathy (d = 0.83).

Other studies not relying on traditional self-report questionnaires have also pointed to a female empathy advantage. For example, Connellan et al. presented newborns with a human face and with a mechanical object and found that, whilst male neonates showed a stronger interest in the object, females showed greater interest in the face. Other studies have revealed that women's empathic superiority reflects the sexes' different exposure to testosterone while in the womb. Longitudinal studies have shown that fetuses exposed to higher testosterone levels in utero make less eye contact as infants in their first year, resort to a smaller vocabulary in their second year, and socialize less with their fellow kindergarten classmates in their fourth year. In another longitudinal study, Udry found that women subjected to more testosterone during embryonic development exhibited masculinized behaviors as adults despite their parents' feminine-oriented socialization efforts. These findings complement others showing that females significantly outperform males in a host of behavioral and cognitive measures of empathy, such as sharing and turn-taking, responding to others' distress, showing sensitivity to facial expressions, inferring what people might be thinking or intending, and accurately recalling information about another person. In light of the theoretical reasoning and empirical evidence for the claim that women possess an empathy advantage, let us consider how this advantage translates to the realm of leadership.