(Gingerich, Raza, Arif, Anwar, & Zhou, 1994). While such events are rarely predictable, they can be reliably explained by Charles Darwin's and Alfred Russel Wallace's great idea, natural selection. Four-legged whale ancestors that could swim better underwater had more offspring; over thousands of generations, their descendants gradually became superb aquatic athletes.Happenstance occurs in intellectual evolution as well. After finishing The Descent of Man, and Selection in Relation to Sex (Darwin, 1871), Darwin realized that materials he had long collected on emotions could be organized to refute Charles Bell's earlier claim that the elaborate musculature of the human face was evidence of Divine design. He quickly wrote The Expression of the Emotions in Man and Animals (Darwin, 1872(Darwin, /1965, emphasizing the phylogenetic consistency of emotional expressions from animals to humans. The book is, as advertised, about expression, and it says little about the selective forces that produced emotions, leaving a persisting anti-Darwinian legacy for emotions research (Fridlund, 1992).However, Darwin clearly recognized that evolution shaped not only the physical characteristics of an organism but also its mental processes and behavioral repertoires. The knowledge that natural selection shaped the brain mechanisms that mediate motivation and emotions offers a solid foundation on which a modern theory of emotions is being built.Although current psychological theories of emotion differ widely in many particulars, almost all now agree that emotions are adaptive responses that arise from mechanisms shaped by selection (Plutchik, 2003). It is now hard to imagine that just four decades ago emotions were generally seen as products of learning unrelated to natural selection. It took Ekman's, Izard's and Eibl-Eibesfeldt's studies of cross-cultural consistency in emotional expression to overthrow that view (Eibl-Eibesfeldt, 1983;Ekman & Davidson, 1994;Izard, 1991). In retrospect, it is obvious that learning cannot be the whole story and that emotions would not exist unless they were useful. Evolution is not an alternative to other theories of emotions; it is the common foundation for all. Many of its contributions are so simple that they are not always recognized. To highlight the continuing importance of Darwin's theory of natural selection for emotions, we consider its implications for several classic questions.
What Emotions AreDefinitions of emotions typically describe proximate aspects such as physiology, subjective experience, or facial expression, often emphasizing one or another component (Ekman & Davidson, 1994;Izard, 2007). An evolutionary approach defines what emotions are in terms of how they came to exist. Emotions are modes of functioning, shaped by natural selection, that coordinate physiological, cognitive, motivational, behavioral, and subjective responses in patterns that increase the ability to meet the adaptive challenges of situations that have recurred over evolutionary time (Nesse, 1990). They are adapta...