Evolution, Literature, and Film:

A Reader


edited by


Brian Boyd, Joseph Carroll,

and Jonathan Gottschall








Sources of the Essays in This Collection


Part 1: Evolution and Human Nature


1.1.        Historical Overview


1. David Buss. Evolutionary Psychology: The New Science of the Mind (2008)


1.2. The Theory of Evolution


Adaptation by Means of Natural Selection


2. Charles Darwin. On the Origin of Species: Recapitulation and Conclusion. (1859)


The Gene as the Unit of Replication


3. Richard Dawkins. The Digital River. (1995)



1.3. Evolution and Humankind


4. Charles Darwin. The Descent of Man: General Summary and Conclusion. (1871)


5. Edward O. Wilson. Man: From Sociobiology to Sociology. (1975)


6. Donald E. Brown. The Universal People. (1991)


7. Edward O. Wilson. Sociobiology at Century’s End. (2000)


8. Steven Pinker.  Evolution and Explanation (2005)


9. David Sloan Wilson. Evolution and Social Constructivism. (2005)


Part 2: The Riddle of Art


10. Steven Pinker. Art and Adaptation (1997)


11. Edward O. Wilson. The Arts and Their Interpretation. (1998)


12. Ellen Dissanayake. Art and Intimacy: How the Arts Began. (2000)


13. Geoffrey Miller. Arts of Seduction. (2000)


14. John Tooby and Leda Cosmides. Does Beauty Build Adapted Minds? Toward an Evolutionary Theory of Aesthetics, Fiction, and the Arts. (2001)


15. Denis Dutton. The Uses of Fiction. (2009)



Part 3: Literature, Film, and Evolution: Theory


16. Brian Boyd. Getting It All Wrong: Bioculture Critiques Cultural Critique. (2006)


17. Joseph Carroll, Jonathan Gottschall, John Johnson, and Daniel Kruger.  Imagining Human Nature.


18. Edward Slingerland. Two Worlds: The Ghost and the Machine. (2008)


19. Marcus Nordlund. Consilient Literary Interpretation. (2002)


20. Robin Headlam Wells. Humanism and Human Nature in the Renaissance (2005)


21. Joseph Anderson. The Reality of Illusion. (1996)


22. Murray Smith. Darwin and the Directors. (2003)


23. David Bordwell. What Snakes, Eagles, and Rhesus Macaques Can Teach Us. (2008)



Part 4: Interpretations


24. Jonathan Gottschall. Homeric Women: Re-imagining the Fitness Landscape. (2008)


25. Michelle Scalise Sugiyama. New Science, Old Myth: An Evolutionary Critique of the Oedipal Paradigm. (2001)


26. Daniel Nettle. The Wheel of Fire and the Mating Game: Explaining the Origins of Tragedy and Comedy. (2005)


27. Marcus Nordlund. Jealousy in Othello. (2007)


28. Nancy Easterlin.  Wordsworth, Psychoanalysis and the ‘Discipline of Love.’ (2000)


29. William Flesch. Vindication and Vindictiveness: Oliver Twist. (2007)


30. Joseph Carroll. The Cuckoo’s History: Human Nature in Wuthering Heights. (2008)


31. Brett Cooke. Human Nature, Utopia and Dystopia: Zamyatin’s We. (2002)


32. Judith P. Saunders. Paternal Confidence in Zora Neale Hurston’s ‘The Gilded Six Bits.’


33. Joseph Anderson. Character in Citizen Kane. (1996)


34. David Bordwell. Convention, Construction, and Cinematic Vision. (1996/2008)


35. Brian Boyd.  Art and Evolution: The Avant-Garde as Test Case: Spiegelman in The Narrative Corpse. (2008)



Part 5: Literature as Laboratory


36. Jonathan Gottschall. Literature, Science, and a New Humanities. (2008)


37. Catherine Salmon and Donald Symons. Slash Fiction and Human Mating Psychology. (2004)


38. Michelle Scalise Sugiyama. Cultural Variation is Part of Human Nature: Literary Universals, Context-Sensitivity and ‘Shakespeare in the Bush.’ (2003)


39. Joseph Carroll, Jonathan Gottschall, John Johnson, and Daniel Kruger. Paleolithic Politics in British Novels of the Nineteenth Century.



Works Cited





Brian Boyd, Joseph Carroll and Jonathan Gottschall






Literary and Film Studies Now: Death or Rebirth?


Adding to a long litany of similar laments, William Deresiewicz recently declared that “The real story of academic literary criticism today is that the profession is, however slowly, dying.” We need not sit passively by, watching as the patient succumbs. Instead, we should take seriously an observation by the philosopher Kwame Anthony Appiah: “In the humanities . . . we are always engaged in illuminating the present by drawing on the past; it is the only way to make a future worth hoping for” (Experiments 1-2). By drawing on the deep past of our evolutionary history, studies in literature and film can ensure their survival and enrich their future.

For at least a decade, many scholars in literature have felt their discipline in crisis. They have been demoralized by falling enrollments and funding, by eroding prestige within and beyond academia, and by a sense of intellectual repetition and exhaustion (see Gottschall, section 36 below). Poststructuralism swept through departments of literature and film in the late 1970s and early 1980s, but its once fresh questions have hardened into habit or dogma. Women’s studies, queer theory, ethnic literatures, cultural studies, post-colonialism, and ecocriticism opened up new subject areas, but these areas have been thoroughly explored through now-familiar research modes. Science studies, under the aegis of Foucauldian discourse theory, have offered another new field but suffer from the same malaise that afflicts poststructuralism in general. The Sokal hoax, in 1996, was only the most spectacular exposé of mortal weaknesses at the heart of this school (Sokal, “Transgressing”; Sokal and Bricmont, Fashionable).

Many scholars working under the aegis of “New Historicism” or “cultural studies” now claim that they are “post-theory” because they focus not on theories but on “empirical” historical data, especially data gleaned from archives. In reality, the archivalists have not left poststructuralist theory behind but have only internalized it. The categories they use derive chiefly from Foucauldian traditions: versions of Marxism and Freudianism filtered through deconstructive epistemology and cast in a programmatically oppositional mode. Simply eschewing explicit theoretical commitment does not invest findings with empirical validity nor insulate them from theoretical critique. It has certainly done little to reverse the declining status of the humanities within the larger world of knowledge.

While the established forms of study in literature and cinema have been drifting into disarray, the evolutionary analysis of human nature has been maturing. It has increasingly placed due emphasis on cooperation, culture, intelligence, and imagination. For over three decades, many observers inside and outside academia have been fascinated to see how evolutionary studies can illuminate human lives, feelings, thoughts, and behavior. Over the past fifteen years or so, evolutionary study in literature and film has emerged as a distinct movement. In the past few years it has gained rapidly in visibility and impact, with many articles and books, and with much attention from the popular and scholarly press, from Nature to the New York Times.

Evolutionary thinking has had revolutionary effects across the human sciences, but it has most dramatically transformed psychology. Those who think that psychological research limits itself to rats running mazes or pigeons pecking levers should think again. Since it began to ask the evolutionary question why—for what benefits—do our minds work as they do, psychology has begun to grapple with much at the heart of literature and film: our core emotions like love, fear, sorrow, happiness; our social and moral emotions like generosity, trust, fairness and indignation; our core relations, parent and child, partner and partner, friends, allies, enemies; Theory of Mind, our capacity to understand other minds; and metarepresentation, our capacity to understand representations as representations. Cognitive, developmental and evolutionary psychology now track empathy in humans and other animals, and our attunement to the emotions of others. Memory researchers study the relationship between our remembered pasts and imagined futures, or between our memory for character traits and our memory for the past. Evolutionary, developmental and cognitive psychology together show why these and other capacities arise, where they emerge from, when they develop in individuals, and how they operate. How could literature and film not seize on all this with excitement?

Although many theorists in literary and film studies have reacted with alarm to claims that biology might help shape culture, acknowledging the reality of human evolution presents no serious dangers and offers immense opportunities. An evolutionary perspective allows us to see ourselves both in the widest angle and with the most precise focus, as individuals solving particular problems within specific contexts, physical and social, using the cognitive equipment—including a predilection for culture—acquired through natural selection. Evolutionary anthropology, biology, economics, and psychology now offer an integrated explanatory framework and a host of new insights we can apply to literature and film.

            Evolutionary theorists of the arts aim not just at offering one more “school” or “approach” to fit within the grab-bag of current theories. We seek to alter the paradigm within which studies in art and culture are now conducted. We have rallied to Edward O. Wilson’s cry for “consilience” among all the branches of learning. We envision an integrated body of knowledge extending from theories of subatomic particles to theories of the arts. Within this consilient world view, evolutionary biology is the pivotal discipline uniting the hard sciences with the human sciences and the arts. Evolutionists seek to investigate how evolution has shaped human bodies, minds and behavior, how culture has emerged out of nature, and how culture has equipped us to modify our behavior.

Virtually all evolutionary theorists of the arts formulate biocultural ideas. That is, we believe that works of art are shaped by our evolved human nature, by culture, and by individual experience. We therefore distinguish ourselves from “cultural constructivists” who effectively attribute exclusive shaping power to culture. We give close attention to “human universals” or cross-cultural regularities that derive from regularities in human nature, but we also recognize the uniquely intense human capacity for culture. We welcome thick descriptions of local context but argue that a true understanding of culture must be rooted in the biological characteristics from which all human cultures grow. Adopting an evolutionary perspective enables us to build theories of literature and film not from near the end of the story but from the start, from the ground up. By building in this way, we can ask altogether new questions and return to older questions with sharper eyes and surer hands.

By insisting on the separateness of humanistic subjects and modes of inquiry, many in the humanities have deprived themselves of the resources discovered in other fields of inquiry. They have also rendered their own research irrelevant to the interests that now most actively engage the minds of the larger educated world. It need not be this way.


What Dangers Does Evolutionary Theory Pose?

The social sciences and humanities have long resisted evolutionary biology, assuming that an evolutionary perspective can only minimize culture, glorify ruthless self-seeking, and deny human uniqueness, diversity, and purpose. In fact evolution now lets us understand the full measure of human uniqueness and human continuity with other life, the power of cooperation, culture, diversity, and the emergence of purpose. Let us quickly allay misgivings about supposed dangers in applying evolution to human minds and behavior.


1. Ignoring human differences: An evolutionary view of human nature does not ignore or deny the enormous cultural differences between peoples, but makes it possible to explain cultural difference in a way that insisting humans are completely “culturally constructed” cannot. (Constructed out of what, in any case?)


2: Biological or genetic determinism: All modern evolutionists recognize that phenotypes (the observable properties of organisms) are not determined solely by genotypes (the genetic recipes in DNA). Behavior is always co-determined by the interaction of genes and the environment. Environments shape, constrain, and elicit the behaviors of organisms. Failing to account for complex interactions between genes and environments is, in fact, profoundly unbiological.


3. Nature versus culture?  Biology is not an alternative to society or culture. Sociality occurs only within living species, and culture only within the social and therefore the biological realm. Moreover, culture is far from being uniquely human. Culture—the non-genetic transmission of behavior, including local customs and even fashions—has been discovered over the last few decades in many social species, in birds as well as mammals.


4. The natural as right: To argue that biology provides a base for human life does not mean it must impose a model for human morality. A particular origin need not predetermine a particular end. Biology in any case offers  a vast array of different models, and culture is itself a part of biology with the power to generate a cascade of new possibilities.


5. Genetic selfishness: Genes are “selfish” in the sense that they prosper according to what benefits them in successive reproductive rounds, but most genes benefit from the health of the whole organism, or even from the success of a whole group of individuals. Richard Dawkins points out that he could just as aptly have called his famous first book not The Selfish Gene but The Cooperative Gene (a title adopted for a more recent book by the biologist Mark Ridley) (Dawkins, Ancestor’s 158). Evolutionary psychology and evolutionary economics concern themselves with the complex mix of cooperation and competition in social life. Indeed, they have placed far more emphasis on generosity, trust, and fairness than non-evolutionary psychology or economics ever had.


6. The supposedly fixed and unchangeable character of human nature: Evolution is change, and species have evolved multiple ways to respond more sensitively to rapid change, including sexual reproduction, nervous systems, flexible intelligence, social learning, and culture. Biologist David Sloan Wilson observes that because of the unique importance of culture in humans, “We have not escaped evolution, as so commonly assumed. We experience evolution in hyperdrive” (“Foreword” 16).


7. Nature and power: Some fear that “Nature, red in tooth and claw” would valorize power and class. In fact evolutionary anthropology and biology increasingly stress that a major difference between humans and other mammals is that humans have found ways to control the urge for dominance. They collaborate to resist being dominated and thus unleash the power of human cooperation (Boehm, Hierarchy). A key concern of evolutionary psychology has been to explain not the redness of our teeth and nails, but why we are, as primatologist Frans de Waal puts it, so “good natured.”


8. Nature versus human singularity: Evolution can explain both our substantial continuity with and our substantial differences from other forms of life. Life has passed through a number of major transitions, each involving new forms of cooperation at one level that enable radically new and more complex possibilities at a higher level. Many biologists now see the cooperation that makes human culture possible as the latest major transition in evolution, as dramatic as the transitions from single-celled to multicellular organisms, or from individual organisms to societies (Wilson and Wilson, “Rethinking”).


What Would We Lose by Adopting an Evolutionary Perspective on Literature and Film?

            Traditional humanists approach literature and film from the perspective of cultivated common sense. The scope of their research extends from scholarship on intellectual and artistic traditions to subtle analyses of tone, theme, style, and form in particular works. At its best, this kind of study embodies in its own perspective the qualities we look for in literary prose and cinematic art: elegance, power, humor, wit, wisdom, passion. The great scholars and critics have assimilated and articulated rich cultural traditions. In constructing and expressing their worldviews, they draw on works of imagination and thus exemplify the value and importance that can attach to the study of artistic culture.

Are any of these values and goals incompatible with an evolutionary perspective on literature and film? We see no reason they should be. Modern science now enters ever more directly into the investigation of what most concerns students of literature and film: human motives and behavior, perception, emotions, cognition, personality and social dynamics. Integrating the new human sciences with humanist literary cultivation presents an immense challenge but also an immense opportunity.

A rapprochement with science need not diminish the creative contribution of the individual literary scholar. The idea that science must mean sterile impersonality is merely a prejudice. In reality, as the physiologist Robert Root-Bernstein explains, the forms of imagination "manifested in styles of scientific creativity” are “just as unique as those of any artist” (“Sciences” 53). Einstein famously affirmed that “Imagination is more important than knowledge”. If creativity is essential to the best work in both the humanities and the sciences, surely it is essential also to work that integrates these domains.

            For the past several decades, studies in literature and film have taken a “theoretical” turn. Without abandoning the ideals of cultivating minds through art, poststructuralist scholars and critics have often rejected the perspective of cultivated common sense. In its place they have looked for theories that open up deeper explanations of the forces shaping human experience and products of the human imagination. Semiotics and deconstructive linguistic philosophy stressed the central role language plays in human consciousness. Freudian psychoanalysis opened up psychosexual symbolism emerging from the most intimate family relationships and the phases of childhood development. Gender theory foregrounded the power conflicts built into human sexual relations. Marxist social theory has inquired into the way works of imagination articulate socioeconomic conditions.

            Is an evolutionary perspective unable to deal effectively with issues that arise out of linguistic philosophy, depth psychology, gender theory, and socio-economic theory? No. Evolutionary human science embraces cognitive neuroscience and cognitive linguistics. Evolutionary psychology concentrates heavily on the often-conflicted relations in the core reproductive relations of families—mothers, fathers, and children—and offers new and penetrating insights into gendered social roles. Evolutionary social theory identifies affiliation and dominance as elemental forces in human social interaction. The way those forces ramify into the complexities of specific social economies provides a rich field of exploration for scholars and scientists.

Culture is part of human nature, and all of the forces we have been describing—linguistic, psychological, and social—manifest themselves in imaginative culture. An adequate understanding of human nature and the products of the human imagination will require the combined and collective work of biologists, psychologists, social theorists, and scholars in the humanities. We need lose nothing of the best that has been thought and said. We need only add to it.


Frequently Asked Questions

Over the years, those who approach art from evolutionary perspectives have met recurrent criticisms, questions, and complaints. Here are some of the most common, along with our responses.


1. Isn’t an evolutionary approach to literature reductive? “Reductive” is a pejorative term signifying that some approach oversimplifies the complex and rigidifies the flexible. “Reductiveness” is a fault, but “reduction” is essential to causal explanation, which necessarily involves reduction to more fundamental principles. Successful reduction connects basic causal principles with the particular features of some specific object. For instance, an explanation of heredity that reduces it to genes and DNA can unify our understanding of diverse phenomena and at the same time bring to light undreamed-of diversity and intricacy. Similarly, evolutionary explanations in the arts can identify elemental causal forces and at the same time open out into a powerfully expansive understanding of particular works or artistic features.


2. There have been many failed attempts to examine literature and film from professedly scientific foundations. Why is this different? Structuralism, Marxist economics, and Freudian psychoanalysis all claimed a scientific basis for studying literature and film. None of these claims survived prolonged empirical scrutiny. The theory of evolution by means of natural selection, in contrast, has been firmly established as the most fertile and robust theory in the life sciences. Evolutionary psychological and social science has emerged as a cumulative, continuously productive research program only within the past forty years. That program now provides the necessary context for evolutionary studies in the arts.


3. How would an evolutionary approach to literature and film avoid the dogmatism and guruism that have characterized much recent literary theory? The theory of evolution by means of natural selection is not a cult or dogma but a scientific research program that took decades to establish on a firm empirical footing. It has to face the continual challenge of consistency with established and emerging evidence from multiple disciplines. Evidence, not assertion or the authority of particular advocates, provides its criterion for validity.


4. Frank Lentricchia claims that if he knows a literary critic's theory he knows in advance how the critic will interpret particular works (64). How do you keep evolutionary approaches from sterile apriorism, if you assume from the outset that all people are ultimately motivated by survival and reproduction? Survival and reproduction are “ultimate” principles that have shaped the characteristics of human nature over evolutionary time. In the case of any individual artist, those characteristics operate through “proximate” mechanisms that are often only tenuously connected to survival and reproduction. Individuals vary widely in their genetic makeup and in the experiences—personal and cultural—that shape their motives. Humans fabricate imaginative worlds that can detach them from survival and reproduction—driving them, for instance, into celibacy or into sacrificing their lives for an abstract cause. Each artistic work challenges us to draw connections from the elemental components of human nature to the particular features of a unique imagined world.


5. Aren't the claims in section two (The Riddle of Art) just-so stories? Isn't this whole approach underpinned by mere storytelling? Evolutionary hypotheses about the adaptive or non-adaptive role of the arts are not final products to be accepted or rejected according to their narrative or rhetorical force. They must be evaluated on the basis of their explanatory and predictive power. The process of testing alternative predictions should—and already has begun to—yield unexpected new information about art and its causes and effects, just as with evolutionary accounts of other familiar but biologically puzzling activities like sleep, dreaming, and play.


6. Isn't it a "category error" to study questions about art in the terms of the sciences? It would certainly be a category error to confuse art itself with science. Art and science have different origins, purposes, effects, and criteria for success. But we can study art as we can human psychology. While we would be wrong to confuse an actual human being with a psychological theory, we can reasonably study both human beings and their cultural products using concepts from the human sciences.


7. Doesn’t analyzing literature and film require us to invoke concepts and methods different from those in biology and the human sciences? Like any other subject area, literature has “emergent” concepts peculiar and appropriate to its own subject. Literary and film scholars necessarily employ categories such as narrative structure, genre, point of view, verse, mise en scène, camera angles, cuts, tone, style, and thematic structure. All such categories engage attention, imagination, cognition and emotion in ways that can be most deeply explored in terms of an evolved and culturally developed human nature.


8. If the paradigm of literary and film study were to shift toward evolution, would everyone have to start reading from the "gene's eye view"? Modern evolutionary analysis requires explanation at multiple levels: at the level of the gene, the organism, the group (pairs, families, alliances), the population, and the species. Recent cultural and literary studies have stressed group-level (class, community, nation) effects, and they have underplayed other levels. The most complete forms of explanation in the arts would connect high-level phenomena like artistic meaning and effect with causal forces at multiple levels.


9. If scholars in the humanities adopt evolutionary perspectives, will there be any room left for imaginative performance—for the kind of work that reveals the depth and richness of the critic’s own mind? The sciences have developed methods like experiment and quantitative analysis to overcome the biases of individuals, cultures and species. Artists, by contrast, have developed methods to appeal to the biases of individuals, cultures, and our species. Critics of literature and film aim at uncovering and explaining facts, but they also respond emotionally and aesthetically to the imaginative qualities in art. Good critics read widely and intensively and write with the boldness and flair that comes from long and deep immersion in great writing. Good criticism needs responsive individual experience, but it also needs the objectivity and impartiality that characterize the scientific ethos.


10. How do evolutionary approaches to literature and film relate to cognitive approaches? Cognition occurs only within evolution. First-generation cognitive psychology derived from early artificial intelligence, from the kinds of information processing involved in computer programming. Second and third-generation cognitive psychology has incorporated a greater recognition of evolved minds: especially of the pre-linguistic bases of cognition, the emotions, and the connections between senses, thoughts, and feelings. Evolutionary and cognitive approaches are only different facets of a common process of inquiry.


Plan of the Volume                                                                                                                                                                                                          

1. Evolution and Human Nature                                                                                                                                                                                        

            Historical Overview                                                                                                                                                                                                                                                          

Part One opens with David Buss’s overview of the historical development of evolutionary biology and evolutionary psychology. In the two sub-sections that follow, we have selected excerpts for their classic status, their representative character, and their power to illuminate central themes in evolution and human nature. The evolutionary ideas delineated here provide the historical and theoretical background for the inquiries into imaginative culture in sections two through five.                                                                        

            The General Theory of Evolution                                                                                                                                                                                                                                    

The most significant scientific theories bring the largest range of phenomena within the smallest compass of causal explanation. Judged by this criterion of significance, Darwin’s theory of adaptation by means of natural selection is one of the most successful efforts at explanation in the history of science, ranking with the theories of Copernicus, Newton, and Einstein.

In the selections we include here from On the Origin of Species, Darwin delineates the core elements in his theory of evolution. Darwin’s theory has not only stood the test of time; it required three quarters of a century before science caught up with his prescience and confirmed in detail the validity of his explanation for evolutionary change. Within just a few years after On the Origin of Species, most scientists had accepted the idea of “evolution,” or, in Darwin’s terms, “descent with modification.” That is, they had accepted that all currently existing life forms had descended from previous forms and changed over the course of geological time. What remained in doubt was the mechanism of change: the theory of “adaptation by means of natural selection.” These elements of Darwin’s theory were not fully established as the unifying principle of evolutionary biology until the nineteen thirties, in the integrative movement now known as “the Modern Synthesis.”

            Darwin himself knew nothing about the specific mechanism of inheritance through which characteristics descend from parents to offspring. When he published On the Origin of Species (1859), genetics as a science did not exist, and it did not begin to achieve scientific maturity until the early twentieth century. DNA, the molecule recording genetic information, was not understood until the middle of the twentieth century. In “The Digital River,” Richard Dawkins formulates the theory of natural selection in modern genetic terms.                                                                                                                                                                                                                                                                               

            The Evolution of Human Nature                                                                                                                                                                                                                                    

            Darwin not only founded the modern concept of evolution but also made the first major contribution to evolutionary psychology. We include key passages here from The Descent of Man, and Selection in Relation to Sex (1871). Natural selection had to wait decades to be fully confirmed by subsequent scientific research. Darwin’s account of the evolved and adapted character of human nature had to wait still longer. In the first decades of the twentieth century, most social science resolutely turned away from evolutionary conceptions, partly in recoil from “social Darwinism,” which in fact owed more to the laissez-faire philosopher Herbert Spencer than to Darwin. Until the last quarter of the twentieth century, most social scientists rejected any appeal to evolutionary biology. The final chapter of E. O. Wilson’s Sociobiology (1975) thus marked a pivotal event in the transition to modern Darwinian social science. We include here excerpts from that chapter and from Wilson’s retrospective and forward-looking commentary, in Sociobiology’s 25th anniversary edition.

The evolutionary understanding of human nature has often concentrated on the “species-typical” or “universal” characteristics of human nature. Anthropologist Donald Brown’s synthetic portrait of “the universal people” offers an engaging introduction to the characteristics shared by humans in all known cultures. In a concise theoretical and historical overview of modern psychology, cognitive psychologist Steven Pinker reveals the explanatory power evolutionary theory brings to the social sciences. And finally, biologist David Sloan Wilson argues that culture reshapes human motives, minds and environments and that “cultural constructivism” should, accordingly, be included within an evolutionary understanding of human nature.


2. The Riddle of Art                                                                                                                                                                                                           

Homo sapiens is a strangely artistic ape. Most humans spend much of their time outside work and sleep lost in landscapes of make-believe. Across the whole breadth of human history, across the wide mosaic of world cultures, there has never been a society where people have not devoted much of their time to seeing, creating, and hearing fictions—from folktales to film, from theatre to television. Nor has there ever been a culture in which people have not devoted precious resources, in time, skill, and material, to producing song, music and dance, pictures and carvings, ornaments, decorations, and designs.

Other animals have adaptations for cooperation in social groups with specialized functions and status hierarchies. Other animals engage in play, produce technology, and share information. Humans alone produce imaginative artifacts designed to provide aesthetic pleasures, evoke subjective sensations, express emotions, depict nature or human experience, or delineate through symbols the salient features of their experience. Dispositions for producing and consuming art constitute uniquely human, species-typical characteristics, and the arts offer rich insights into the human mind. No understanding of human nature that leaves out the arts could possibly give an adequate account of its subject.

Art poses an evolutionary riddle. Why are we storytelling apes? Why do we compulsively concoct and consume fictional stories, stories we know to be untrue? Why do we spend (waste?) so much time telling tales and shaping objects for aesthetic effects—time that could be devoted to activities that produce obvious biological benefits: securing resources, courting mates, or caring for offspring and other kin?

The selections here show some of the range of positions on this contentious and complex issue. Steven Pinker argues that aesthetic aptitudes arose as side effects of other adaptively significant mental powers. Geoffrey Miller offers the most influential version of the idea that the arts serve the purposes of sexual display. E. O. Wilson, Ellen Dissanayake, John Tooby and Leda Cosmides, and Denis Dutton provide overlapping versions of arguments that the arts fulfill definite adaptive functions. Brian Boyd, in Part 4, also addresses the adaptiveness of art in tandem with a close reading of a single avant-garde work. Carroll, Gottschall, Johnson, and Kruger, in Part 5, offer empirical evidence for story’s adaptive function.


3. Literature, Film and Evolution: Theory                                                                                                                                                                          

Evolutionary and cognitive approaches to the arts have been strongly at odds with recently prevailing assumptions in the humanities. There have been two core disagreements: evolutionists claim (1) that we humans share a great deal of our mental and motivational makeup, that differences across cultures and ideologies are dwarfed by similarities; and (2) that our minds have been shaped to provide us with mostly valid information about our world, rather than to operate by arbitrary convention.

            Clashes between bioculturalism and cultural constructivism have sometimes been sharp. For instance, surveying and lamenting the decline of literary studies, critic Louis Menand appealed for renovation but explicitly rejected just one possible form of renovation, the one we regard as most promising: the integration of the humanities with evolutionary biology. "Consilience,” he claimed, “is a bargain with the devil” (“Dangers” 14). Menand’s article prompted Brian Boyd’s rejoinder, “Getting It All Wrong.” While making a sweeping comparison of the key positions in postmodern and evolutionary cultural theory, Boyd delineates the false conceptions that inform Menand’s resistance to consilience. Joseph Carroll and colleagues provide an overview of the literary theory of the last century and contrast it with the emerging evolutionary paradigm. Refuting attempts to present Shakespeare and his contemporaries as deniers of shared human nature, Robin Headlam Wells shows that they had an explicit belief in human nature and a desire to illuminate it. Writing from outside literature (he is a scholar of early Chinese thought), Edward Slingerland argues that a biologically grounded cognitive science can reconcile the humanities and the sciences. Marcus Nordlund incisively formulates a naturalistic basis for a literary hermeneutics alert to the unpredictable complexities of specific interpretive problems.

Reviewing the history of film theory, Joseph Anderson suggests that the accessibility of film depends on its appeal to evolved human modes of “ecological” perception—that is, our perception of the environment that we negotiate as mobile creatures. Beginning with what his American students can understand of an undubbed, un-subtitled Taiwanese martial arts movie—a great deal, it turns out—David Bordwell proposes a naturalistic approach to film response that explains both the considerable convergence and partial divergence in our responses to art.


4. Interpretations                                                                                                                                                                                                                

            Despite first the “theoretical” and then the “archival” turn in much recent scholarship, readings of individual works still dominate the activity of criticism. To vindicate its paradigmatic claims, evolutionary study in literature and film must also demonstrate that it can give compelling interpretive accounts of particular works. Part Four showcases evolutionary readings that display both explanatory power and sensitivity to particular meanings and effects.

If evolutionary approaches work for any art—if evolution has indeed shaped all artists and audiences—they should work for all. The readings here move chronologically from Homeric epic to avant-garde comics. Although many readings focus on English literature, others discuss ancient or modern European literature or cinema from Africa, the Middle East and Asia, as well as America and Europe. Modes include prose, drama, verse, film, and comics. Genres range from epic, comedy, tragedy, realistic novel and story, to science-fiction, film, and serially-composed comic. While most readings focus on a single work, others consider multiple examples as they focus instead on genre, like Daniel Nettle discussing tragedy and comedy; on technique, like David Bordwell discussing shot-reverse shot composition in film; or on response, like Murray Smith discussing emotion in film.

            Some critics stress what might seem like naturally evolutionary topics: anxieties about paternity certainty in Judith Saunders’ essay on Zora Neale Hurston, for instance, or Jonathan Gottschall’s discussion of mating strategies in Homer’s Iliad. While Saunders focuses on a human universal, Gottschall attempts to explain the peculiar ecological and reproductive pressures promoting conflict in the society Homer depicts. For those with no prior acquaintance with evolutionary approaches to literature, other contributions might be more surprising, like William Flesch on altruistic punishment in Oliver Twist, Joseph Carroll’s analysis of Wuthering Heights in terms of human life history theory, or Brian Boyd’s discussion of dominance and counter-dominance in Spiegelman’s contribution to The Narrative Corpse.

Many of the critics here address the question of human nature and its relative persistence across time and circumstance. Marcus Nordlund considers local legal and cultural conditions that modify male sexual jealousy in early seventeenth-century Europe, but he argues strongly against the idea of Othello’s jealousy as a predominantly local construct. Brett Cooke shows how Evgeniy Zamyatin, in his science-fiction dystopia, We, critiques Soviet communism’s attempts to ignore and re-engineer human nature. Others set Freudian against evolutionary accounts of human nature. Michelle Scalise Sugiyama reads Oedipus Rex in the light of evolutionary evidence. Drawing on the developmental theories of John Bowlby and Daniel Stern, Nancy Easterlin revises feminist and psychoanalytic accounts of mother-infant bonding in Wordsworth’s Prelude.    

While evolutionary criticism has a deep interest in human nature, it should not neglect local or individual differences or the effects of artistry. Indeed, just as life itself is a problem-solving process, an evolutionary and cognitive approach to the arts should focus on the partly unique and partly shared problem-solution landscape within which individual artists work. Essays that place particular emphasis on form and individual artistry include Nordlund on Shakespeare, Easterlin on Wordsworth, Bordwell on shot-reverse shot composition, Anderson on Orson Welles, and Boyd on Spiegelman.


5. Literature as Laboratory                                                                                                                                                                                               

Most evolutionary literary scholars believe that mainstream literary studies could do a much better job of generating reliable and cumulative knowledge. We all concur in seeking better sources of knowledge in the biologically grounded human sciences, but some of us have taken a further step: using empirical methods to study subjects in the humanities.

In taking this step we distance ourselves sharply from two salient features of postmodernist literary theory. We reject both the radical skepticism that denies the possibility of ever gaining reasonably objective knowledge, and the practice, so regrettably common in literary study, of looking only or mainly for evidence in support of our hypotheses, especially those that we deem politically desirable.

Science has established many ways to avoid being misled by selective evidence, confirmation bias, and a priori thinking. We believe many of these methods can be adapted to questions about art. We cannot do without reading, thinking, and writing well, but we want to extend the scholarly toolkit by adding scientific methods, which can test, tease apart, and extend intuitions about literature.

      Literature offers an unparalleled source of thick descriptions of imagined human behavior and actual human preferences over thousands of years—evidence that can be used to inspire, enrich or challenge hypotheses about human nature. In this section we include examples of humanities scholars and scientists flowing back and forth across the divide between the two cultures of the arts and sciences. Their goal is not to promote a conquest of humanistic culture by scientific culture. Rather, they are seeking to establish a Third Culture on the fertile ground between the humanities and sciences. We include selections suggesting that (1) stories are a valuable and much-neglected resource in the scientific study of mind and behaviour, and (2) that humanities scholars can draw on the methods, as well as the theories and findings, of the sciences. 




The essays in this volume offer a chance to reshape the landscape of scholarship. The contributions from evolutionary biology, the human sciences, and the humanities provide complementary and convergent perspectives on a single grand subject: human nature in the works of the human imagination. Despite productive disagreements, the authors represented here embody a set of shared attitudes and beliefs. We all believe that evolutionary theory promises the deepest, widest and most reliable knowledge about humankind and all its works. Still more broadly, we believe in science; we believe that the world can be known in a reasonably objective way, that knowledge in any field can accumulate, and that knowledge in diverse fields can be causally connected. We are eager to extend scientific knowledge into the fields of the humanities and to make use of the tools available from both the humanities and the sciences. From the humanities, we incorporate the skills of historical scholarship and close reading. We respect the sensitivity critics display in responding to the artistry of literature and film, and we are committed to preserving and celebrating the rich heritage of the humanities. From evolutionary biology and the human sciences, we incorporate a knowledge of human universals and of the cognitive mechanisms underpinning art. We recognize that the building blocks of human nature combine in different ways and under different conditions to form unique structures in any given culture. We also recognize the rich genetic and experiential differences between individuals. Whether or not we adopt empirical, quantitative methods, we share a profound respect for the spirit of impersonal, disinterested inquiry. We recognize that a passionate responsiveness to the arts is natural to scholarship in the humanities, but we are also determined to have access to the impersonal, objective scrutiny of science.

If the impulses behind this anthology were to become active across the humanities, that would constitute an epistemic revolution expanding the scope of both science and the humanities. In the short term it would open the products of the human imagination to the human sciences and the methods and results of the human sciences to the humanities. In the long term, it would enable humanists to join with scientists in contributing to the continuous development of more reliable and durable knowledge. The revolutionary impulses implicit in this volume are thus fundamentally creative and constructive. We have no illusions that our formulations are fixed and final, but we have felt the excitement of making new discoveries and look forward confidently to more. We have all been inspired by the exhilarating sense that we are joining together in an intellectual adventure of great scope. We invite you to join us.


Sources of the Essays in this Collection:


            Most of the journal articles and excerpts from books included in ELF have been abridged, and their various forms of citation have been standardized. In condensing the articles, the editors have aimed at shaping coherent essays and have not indicated omissions through ellipses. Sources for all the essays are included in the bibliography. Essays that were originally published as journal articles or as book chapters in edited collections can be located in the bibliography by their titles in this volume.

Below, we list the source for essays excerpted from monographs or otherwise titled in ways that are not identical with titles in the bibliography. The sources listed below are sequenced in parallel with the sequence of essays in this volume:


Dawkins’ “The Digital River” is from River out of Eden.

Edward O. Wilson’s “From Sociobiology to Sociology” is from Sociobiology: The New Synthesis, and his “Sociobiology at Century’s End” is from the 25thanniversary                                          edition of Sociobiology.

Donald Brown’s “The Universal People is from Human Universals.

Steven Pinker’s “Evolution and Explanation” is the Foreword to David Buss’s Handbook of Evolutionary Psychology.

Steven Pinker’s Art and Adaptation is from How the Mind Works.

Edward O. Wilson’s “The Arts and their Interpretation” is from Consilience: The Unity of Knowledge.

Ellen Dissanayake’s “Art and Intimacy: How the Arts Began” is from Art and Intimacy.

Geoffrey Miller’s “Arts of Seduction” is from The Mating Mind.

Denis Dutton’s “The Uses of Fiction” is from The Art Instinct.

“Imagining Human Nature” by Carroll, Gottschall, Johnson, and Kruger is from Graphing Jane Austen.

Robin Headlam Wells’s “Humanism and Human Nature in the Renaissance” is from Shakespeare’s Humanism.

Joseph Anderson’s “The Reality of Illusion” is from The Reality of Illusion.

David Bordwell’s “What Snakes, Eagles, and Rhesus Macaques Can Teach Us” is from Poetics of Cinema; the final segment, though, is a condensed version of the                                               “Foreword” to Moving Image Theory, edited by Anderson and Anderson.

Jonathan Gottschall’s “Homeric Women: Re-imagining the Fitness Landscape” is from The Rape of Troy.

Marcus Nordlund’s “Jealousy in Othello” is from Shakespeare and the Nature of Love.

William Flesch’s “Vindication and Vindictiveness” is from Comeuppance.

Brett Cooke’s “Human Nature, Utopia and Dystopia: Zamyatin’s We” is from Human Nature in Utopia.

Joseph Anderson’s “Character in Citizen Kane” is from The Reality of Illusion.

David Bordwell’s “Convention, Construction, and Cinematic Vision” is from Poetics of Cinema.

Jonathan Gottschall’s “Literature, Science, and a New Humanities” is from Literature, Science, and a New Humanities.



Contributor Biographies

David Bordwell is Jacques Ledoux Professor Emeritus of Film Studies, University of WisconsinMadison. He has written several books on the aesthetics and history of cinema, including Narration in the Fiction Film, On the History of Film Style, and, most recently, Poetics of Cinema (2007).


Brian Boyd is University Distinguished Professor of English, University of Auckland. His Nabokov work (biography, criticism, editions) has been translated into twelve languages. His On the Origin of Stories: Evolution, Cognition and Fiction (2009) proposes art and storytelling as adaptations, explains the evolved cognitive mechanisms underpinning fiction, and shows the implications for reading classic fictions. He has also published many evolutionary literary essays, theoretical and interpretive.


David Buss is Professor of Psychology at the University of Texas, and Past President of the Human Behavior and Evolution Society.  He is author of more than 200 scientific publications, as well as a number of books, including The Evolution of Desire:  Strategies of Human Mating; The Dangerous Passion; and Evolutionary Psychology:  The New Science of the Mind.


Donald E. Brown is professor emeritus of anthropology at the University of California, Santa Barbara.  His research focuses on the peoples and cultures of Southeast Asia, social structure, ethnohistory, human universals and ethnicity/ethnocentrism.  His principal publications include Brunei: The Structure and History of a Bornean Malay Sultanate; Principles of Social Structure: Southeast Asia; Hierarchy, History, and Human Nature: The Social Origins of Historical Consciousness; and Human Universals.


Joseph Carroll is Curators' Professor of English at the University of Missouri, St. Louis. His Evolution and Literary Theory (1995) integrated traditional humanist theory with evolutionary psychology and set both into sharp opposition with poststructuralist theory. The essays collected in Literary Darwinism: Evolution, Human Nature, and Literature (2004) took in new developments in the field and worked toward a comprehensive theory of human nature and literature, and he has written many subsequent essays. His annotated edition of Darwin's On the Origin of Species (2003) is widely used in classes on the history of science.


Brett Cooke is Professor of Russian at Texas A&M University. In addition to Darwinist studies of opera, Western science fiction, Irish art, ballet, and, naturally, Russian literature, he is the author of Human Nature in Utopia: Zamyatin's We (2002). He is also co-editor (with Frederick Turner) of Biopoetics: Evolutionary Explorations in the Arts (1999) and (with Jan Baptist Bedaux) of Sociobiology and the Arts (1999).


Charles Darwin, independent scholar, biologist, discoverer of the theory of evolution by natural selection. Author of On the Origin of Species by Means of Natural Selection, The Descent of Man and Selection in Relation to Sex, The Expression of Emotions in Animals and Man, and many other works.


Richard Dawkins retired in 2009 as the first Charles Simonyi Professor of the Public Understanding of Science at Oxford. Dawkins is the author of many books including The Selfish Gene, The Blind Watchmaker, The Ancestor's Tale, The God Delusion and, most recently, The Greatest Show on Earth. He is a Fellow of the Royal Society and the Royal Society of Literature. His prizes include the International Cosmos Prize, the Kistler Prize, the Lewis Thomas Prize, the Shakespeare Prize, and the Nierenberg Prize.


Ellen Dissanayake is an independent scholar, writer, and lecturer, whose writings about the arts over 35 years synthesize many disciplines and draw upon fifteen years of living and working in non-Western countries. Author of three books and numerous scholarly and general articles, she is currently Affiliate Professor in the School of Music, University of Washington.  Her book, Homo Aestheticus (1992), has been translated into Chinese and Korean.


Denis Dutton is Professor of Philosophy at the University of Canterbury in New Zealand. He edits the website "Arts & Letters Daily" and the journal Philosophy and Literature,
published by the Johns Hopkins University Press. He is also author of The Art Instinct: Beauty, Pleasure, and Human Evolution (2009).


Nancy Easterlin is Research Professor of English at the University of New Orleans. She is coeditor with Barbara Riebling of After Poststructuralism: Interdisciplinarity and Literary Theory (Northwestern,1993) and author of Wordsworth and the Question of ‘Romantic Religion’ (1996) as well as numerous essays on biocultural approaches to literature. She is currently completing a book entitled What Is Literature For? Biocultural Theory and Interpretation.


William Flesch teaches English, film, and sometimes philosophy at Brandeis.  In addition to Comeuppance (2007), he is the author of Generosity and the Limits of Authority: Shakespeare, Herbert, Milton, and The Facts on File Companion to Nineteenth Century British Poetry.


Jonathan Gottschall is an adjunct in the English Department at Washington & Jefferson College.  He is the author of The Rape of Troy: Evolution, Violence, and the World of Homer (2008) and Literature, Science, and a New Humanities (2008). He is co-editor of The Literary Animal: Evolution and the Nature of Narrative (2005).


Robin Headlam Wells is Professor Emeritus of English Literature at Roehampton University, London. His most recent book is Shakespeare’s Humanism (2005). He is currently writing A Short History of Human Nature.


John A. Johnson is Professor of Psychology, at Pennsylvania State University, DuBois. He has published many articles on the personality and evolutionary psychology of moral and educational development, career choice, and work performance. He is currently co-editing a book, Advanced Methods for Conducting Online Behavioral Research.


Daniel J. Kruger is an assistant research professor at the University of Michigan, where he is affiliated with the Prevention Research Center in the School of Public Health and the Life Course: Evolutionary and Ontogenetic Dynamics program at the Institute for
Social Research. His evolutionary research interests include: altruism, cooperation, competition, risk taking, mortality patterns, mating strategies, and applications for social and ecological sustainability. Many of his research projects are grounded in evolutionary life history theory.


Geoffrey Miller teaches evolutionary psychology and human sexuality, and does research on mate choice, sexual selection, intelligence, creativity, art, music, personality, psychopathology, consumer behavior, and behavior genetics.  His books include The Mating Mind (2000), Mating Intelligence (co-edited with Glen Geher 2008), and Spent: Sex, evolution, and consumer behavior (2009).  Following a B.A. from Columbia and a Ph.D. from Stanford, he worked at University of Sussex, University College London, and London School of Economics, and is currently associate professor of psychology at University of New Mexico.


Daniel Nettle is a Reader in the Centre for Behaviour and Evolution, Newcastle University, where he studies the application of evolutionary theory to human behaviour in many domains. He is the author of several books including, most recently, Personality: What Makes You the Way You Are (2007). He also spent several years as a professional actor and theater director, and still retains an interest in theater in practice as well as theory.


Marcus Nordlund is an associate professor of English at the University of Gothenburg, Sweden, and specializes in English Renaissance literature and biocultural literary theory. He has published two books -- The Dark Lantern: A Historical Study of Sight in Shakespeare in Shakespeare, Webster, and Middleton (1999) and Shakespeare and the Nature of Love: Literature, Culture, Evolution (2007) -- and articles on a variety of literary subjects from Shakespeare to Toni Morrison.


Steven Pinker is Harvard College Professor and Johnstone Family Professor of Psychology at Harvard University, and the author of seven books on language, cognition, and human nature, most recently The Stuff of Thought.


Catherine Salmon is an associate professor of psychology at the University of Redlands. She is the author (with Don Symons) of Warrior Lovers and the co-editor (with Charles Crawford) of Evolutionary Psychology, Public Policy and Personal Decisions and  (with Todd Shackelford) Family Relationships: An Evolutionary Perspective. Her research interests include female sexuality and pornography, eating disorders, and birth order and family relationships. When not in the office, she is usually found working with pit bull terriers or horseback riding.


Judith P. Saunders is Professor of English at Marist College in New York State.  Her articles take in a wide range of literary figures, including Edgar Allan Poe, Henry Thoreau, Stephen Crane, Sherwood Anderson, Gertrude Stein, Virginia Woolf, Gwendolyn Brooks, Elizabeth Bishop, and others.  She is the author of a book-length study of the British poet Charles Tomlinson. She has undertaken Darwinian analysis of a variety of literary works, most recently in Edith Wharton and Evolutionary Biology: Reading Her Fiction through a Darwinian Lens (2009).


Michelle Scalise Sugiyama is Research Associate at the University of Oregon Institute of Cognitive and Decision Sciences, and founder and Director of the Cognitive Cultural Studies Division at the Center for Evolutionary Psychology, UC Santa Barbara.  Her research focuses on cognitive adaptations for cultural transmission, with an emphasis on narrative and art behavior in foraging societies.  She has written numerous articles on the origins of storytelling, the role that folklore plays in foraging societies, and the cognitive foundations of narrative.


Edward Slingerland is Associate Professor of Asian Studies and Canada Research Chair in Chinese Thought and Embodied Cognition at the University of British Columbia. His first book, Effortless Action: Wu-wei as Conceptual Metaphor and Spiritual Ideal in Early China (2003), won the American Academy of Religion’s award for the Best First Book in the History of Religions. His most recent monograph, What Science Offers the Humanities: Integrating Body and Culture (2008), argues for the relevance of the natural sciences to the humanities.


Murray Smith is Professor of Film Studies at the University of Kent, Canterbury, UK. He is the author of Engaging Characters: Fiction, Emotion, and the Cinema (1995), Trainspotting  (2002), and co-editor of Film Theory and Philosophy (1997), Contemporary Hollywood Cinema (1998) and Thinking through Cinema: Film as Philosophy (2006). His research interests include the psychology of film viewing and the place of emotion in film reception, as well as the philosophy of film, music and of art more generally. He is currently working on the implications of evolutionary theory for film culture.


Don Symons received a B.A. in psychology and a Ph.D. in biological anthropology from the University of California at Berkeley.  He has been a faculty member of the anthropology Department at U.C. Santa Barbara since 1970, and is currently Professor Emeritus.  He studied social play among free ranging rhesus monkeys (Play and Aggression, 1978) and was an early contributor to the field of evolutionary psychology (The Evolution of Human Sexuality, 1979).


John Tooby and Leda Cosmides are best known for their work in pioneering the new field of evolutionary psychology. They are professors of anthropology and psychology at the University of California, Santa Barbara, where they co-direct the Center for Evolutionary Psychology.  Both were educated at Harvard and Stanford (postdoctoral). Awards for their research include the NIH Director’s Pioneer Award, the American Association for the Advancement of Science Prize for Behavioral Science Research, the American Psychological Association’s Early Career Award; a National Science Foundation Presidential Young Investigator Award, and J. S. Guggenheim Fellowships; John Tooby has been president of the Human Behavior and Evolution Society.


David Sloan Wilson is SUNY Distinguished Professor of Biology and Anthropology at Binghamton University. He applies evolutionary theory to all aspects of humanity in addition to the rest of life, both in his own research and as director of EvoS, a unique campus-wide evolutionary studies program. He is known for championing the theory of multilevel selection, which has implications ranging from the origin of life to the nature of religion. His books include Darwin’s Cathedral: Evolution, Religion, and the Nature of Society (2002) and Evolution for Everyone: How Darwin’s Theory Can Change the Way We Think About Our Lives (2007). His next book is titled Evolving the City: An Evolutionist Contemplates Changing the Word—One City at a Time


Edward O. Wilson is University Research Professor, Emeritus, at Harvard University. His 25 books include On Human Nature (1978) and The Ants (1990, with Bert Hölldobler), which won Pulitzer prizes, Consilience: The Unity of Knowledge (1998), and most recently The Future of Life (2002), and again with Bert Hölldobler, The Superorganism (2009).