Darwin’s vision of deep time encompasses all living things, including human beings. For the first three-quarters of the twentieth century, most social scientists and humanists repudiated the implications of that vision (Fox 1989; Degler 1991; Tooby and Cosmides 1992; Pinker 2002). In the past few decades, evolutionary ways of thinking have become firmly established in the social sciences. In the humanities, evolutionary thinking remains marginal to the academic establishment. Fewer than a hundred published scholars in the humanities can be plausibly described as evolutionists (Carroll et al. 2017). Undergraduate departments in the arts strongly discourage evolutionary thinking, and very few graduate programs accept students who overtly profess evolutionary views. Much of the early work done in this field came from scholars who already had tenure before taking up evolutionary thinking. In the past decade or so, despite the institutional blockage, evolutionists in the humanities have continued to produce important work, much of it by younger scholars (Carroll 2012b; Carroll et al. 2012; Jonsson 2012, 2013; Carroll, McAdams, and Wilson 2016; Kjeldgaard-Christiansen 2016; Clasen 2017; Carroll 2018; Salmon 2018; Saunders 2018; Høgh-Olesen 2019; Kjeldgaard-Christiansen and Schmidt 2019; Kruger and Jonsson 2019; Larsen 2019; Carroll, Clasen, and Jonsson 2020; Dolack 2020; Jonsson 2020; Larsen 2020; Gottschall 2021; Hye-Knudsen 2021; Jonsson 2021; Larsen 2021; McCrae 2021; Scrivner 2021; Bannan 2022; Carroll 2022; Carroll et al. 2022; Larsen 2022; Saunders 2022; Kjeldgaard-Christiansen and Clasen 2023; Larsen 2023).

The late Edward O. Wilson declared that “the greatest enterprise of the mind has always been and always will be the attempted linkage of the sciences and humanities” (Wilson 1998, 8). Wilson perceived an immense gap—methodological, ontological, and epistemological—between the objective physical sciences and research on the subjective complexities of human cultural and artistic experience. The physical sciences, including biology, have succeeded in reducing complex phenomena to general laws and predictable systemic interactions among relatively closed systems—that is, systems with a finite number of causal factors that can be measured. Many scholars, both scientists and humanists, still believe that the physical world and the human mind are essentially different sorts of substances (Carroll et al. 2017). Among more traditional scholars, this belief relegates the mind to a quasi-spiritual world inaccessible to causal reduction, a kind of vapor or mist vaguely associated with an autonomous, transcendental force such as free will (Eichner 1982; Goodheart 2007, 2009). In the currently dominant episteme in the humanities, that autonomous transcendental force has been transferred from the individual human psyche to the collective psyche designated in the term “culture” (Scholes 2006). Scholars who explicitly reject this kind of dualism sometimes nonetheless affirm that the mind and its cultural manifestations, including the arts, have too many unique elements to be reduced to general laws and closed causal systems (Crews 2008; Carroll, Boyd, and Deresiewicz 2009; Deresiewicz 2009). As Edward Slingerland puts it, “fields such as literary studies tend to concern themselves overwhelmingly with emergent structures and idiosyncratic cultural histories” (Slingerland 2008, 268). From this perspective, the general causal principles in the evolutionary human sciences are so remote from the contingent particularities studied in the humanities that for all practical purposes the two fields are separate and irreducible.

Within the evolutionary behavioral sciences, the daunting character of the challenge presented by the humanities finds one form of relief in shrugging off the arts as trivial side effects of human cognitive dispositions that have evolved to solve the adaptively serious problems of survival, reproduction, and successful social interaction (Pinker 1997, 2007). A second form of relief offers itself in reducing the arts to forms of sexual display—elaborate and beautiful, perhaps, as a peacock’s tail, but essentially meaningless in the details of its form and content (Miller 2000; Dutton 2009; Kenrick 2011). Yet a third form of premature cognitive resolution presents itself in identifying the arts as essentially a vessel for carrying useful information about the more urgent adaptive problems of survival, reproduction, and sociality (Scalise Sugiyama 2005). In all these ways of explaining the arts, or explaining them away, imagination does not significantly alter the basic outlines of human nature.

Until Darwin formulated the principle of natural selection and demonstrated its utility in explaining a diverse array of phenomena ranging from embryology through ecology and the geographical distribution of species, biology lacked a unifying principle that could bring the complex and particular phenomena of natural history within the range of closed causal systems. So far as I can tell, we do not at present have a Darwin in our midst—a scientist with the scope, minuteness of knowledge, and power of sustained inquiry necessary to establish a new paradigm uniting the sciences and humanities—and even Darwin’s theory required some three-quarters of a century before the Modern Synthesis finally confirmed its explanatory validity (Huxley 1942; Bowler 1983; Mayr 1991; Carroll 2003). If we do not yet have a Darwin in our midst, we do, I think, have a concept that can serve as a unifying explanatory idea. The one feature of our specifically human nature that most distinguishes the human species from all others is imagination: the power to rise above the present moment in sensory awareness, reflect on events, connect present with past and with anticipated futures, intuit the inner experience of other people, connect actions with abstract concepts such as moral ideas and group social identity, project long-term goals that reflect impersonal values, and envision human life as an aspect of a cosmic world order (Carroll 2012b; Taylor 2013). Imagination is the most obvious and immediate psychological source for the arts: artifacts that use media such as sound, visual percepts, or language to depict human experience, evoke subjective sensations, and articulate meaning (Zhu et al. 2017).

Before the first two decades of the 21st century, commentary on the imagination remained largely speculative. Historically, it was most closely associated with the Romantic poets and philosophers of the late eighteenth and early nineteenth centuries—writers who attributed to it some quasi-divine spiritual force at odds with the explanatory reductions of science (Shelley 1909; Lovejoy 1924, 1961; Wellek 1963; Eichner 1982; Coleridge 1983 [1817]; Wilson 1984 [1931]; Abrams 1984 [1965]). In just the past two decades, a massive new research program in neuroscience has focused on the neurological basis for imagination: the Brain’s Default Mode Network (DMN). The DMN is the most integrative neurological network in the brain (Margulies et al. 2016; Carroll 2020; Smallwood et al. 2021). Neuroscientists have convincingly demonstrated that the DMN is responsible for the uniquely human capacities for Theory of Mind (aka mentalizing, perspective taking), mental time travel (envisioning the present in relation to the remembered past and to anticipated futures), constructing hypothetical scenarios, evaluating moral alternatives, and making long-term plans that subordinate impulse to sustained sequences of disciplined action (Buckner, Andrews-Hanna, and Schacter 2008; Suddendorf, Addis, and Corballis 2009; Buckner 2012; Pace-Schott 2013; Kaufman and Gregoire 2015; Margulies et al. 2016; Beaty et al. 2018; Cao et al. 2018; Smith, Mitchell, and Duncan 2018; St. Jacques et al. 2018; Alves et al. 2019; Buckner and DiNicola 2019; Smallwood et al. 2021; Yeshurun, Nguyen, and Hasson 2021; Menon 2023).

The human capacity for creating an imaginative virtual world has been the culminating adaptation of the long human trajectory of gene-culture coevolution. Sometimes now designated “culture gene coevolution” or even, loosely, “cultural evolution,” gene-culture coevolution is the causal interaction of genetically regulated traits and cumulative, collective forms of learned beliefs, skills, and behavior (Lumsden and Wilson 1981; Richerson and Boyd 2005; Wade 2006; Hill 2007; Cochran and Harpending 2009; Wrangham 2009; Henrich 2016; Henrich 2020). The relatively recent and rapid expansion of the DMN is closely associated with the emergence of behavioral modernity over the past 200,000 years—the use of complex multi-part tools and the fabrication of symbolic culture in the arts (Mellars and Stringer 1989; Carroll 2006; Mellars 2007; Klein 2008, 2009; Neubauer, Hublin, and Gunz 2018; Carroll 2020; Vyshedskiy 2020). Since the advent of agriculture some 10,000 years ago, the imaginative capacity for cumulative cultural innovation has been a necessary component in the development of state-level societies with stratified social orders, specialized productive activities, and institutions for the enforcement of norms (Trigger 2003; Fukuyama 2011).

Even before research on the DMN made it clear that imagination has so decisively altered the trajectory of human evolution, some evolutionary humanists have intuitively understood that their central mission is to analyze interactive causal connections between the elements of human nature and the forms of imaginative experience. They have aimed at constructing continuous explanatory sequences linking the highest level of causal explanation—adaptation by means of natural selection—with particular features of human nature and particular structures and effects in specific works of art. In literary study—to take the humanistic field with which I am most familiar and in which the greatest amount of evolutionary work has been done—the most naïve form of evolutionary critique has contented itself with observing that people seek survival, sex, and status and that artistic works depict people seeking those things (Barash and Barash 2005). More comprehensive methods take in imaginative features (tone, style, theme, formal organization), locate these features in a particular cultural context (ecological, economic, political, religious, ideological, etc.), describe that cultural context as a particular organization of the elements of human nature, register the responses of readers, describe the social and psychological functions the work fulfills for artists and audiences, locate those functions in relation to the evolved needs of human nature, and compare the work with other artistic works, using a taxonomy of themes, formal elements, affective elements, and functions derived from a model of human nature (Carroll 2011, 2012a; Saunders 2018; Carroll 2019; Kruger and Jonsson 2019; Jonsson 2021; Saunders 2022).

The concept of human nature is the common ground on which evolutionary social scientists and humanists have most fully converged. Contributors to both areas agree that human nature consists in motives, emotions, and forms of cognition that have evolved through natural selection and that have been conserved through the dramatic changes in environmental conditions that have characterized human evolution over the past few hundred thousand years. The basic features of human nature can be delineated in detail through depictions of human universals (Brown 1991) and can be summarized in an outline of human life history: the species-typical organization of the human life span modulated by its chief phases, motives, and forms of reproductive and social relationships. That outline includes mother-infant bonding, long-term child dependency on adult care, extended childhood and adolescence, pair-bonded dual parenting, bilateral kinship networks, reproductively functional sexual differences in anatomy, physiology, and hormones, male and female cooperative groups, a uniquely human combination of cooperative male groups and monogamous bonding, post-menopausal survival, and the intergenerational transfer of resources (Trivers Robert 1972; Flinn and Ward 2005; Kaplan, Gurven, and Winking 2009; Konner 2010; Muehlenbein and Flinn 2011; Salmon and Shackelford 2011; Low 2015; Buss 2016; Fisher 2016; Chapais 2017; Bogin and Varea 2020; Richerson and Boyd 2020; Crews and Bogin 2023). For social relationships, evolved human dispositions include reciprocity, dominance hierarchies, reverse dominance (egalitarianism), leadership, internalized norms, strong reciprocity (third-party enforcement of norms), legitimacy in the exercise of power, and the introjection of group identity into individual identity (Trivers 1971; Boehm 1999; Tomasello et al. 2005; Nowak 2006; Hill et al. 2011; Boehm 2012; Carroll 2015a, b; Gintis, van Schaik, and Boehm 2015; Boehm 2016).

Humanist intuitions about the importance of imagination have prompted claims about the universal human need to tell stories and engage in other forms of imaginative construction (Dissanayake 2000; Boyd 2009; Carroll 2012b; Gottschall 2012). In their most circumscribed form, such claims identify the need to create imaginative constructs as one of the basic motives in human nature—a motive added on to the motives more directly targeted to survival and reproduction. In their most expansive form, such claims suggest that humans always occupy an imaginative virtual world and that every basic animal motive is altered—constrained and modulated—by being incorporated within an imaginative virtual world. For most of human history, the collective social creation of imaginative virtual worlds has culminated in religious cosmologies. For the past few hundred years, religious visions have increasingly been supplanted by a naturalistic and scientific understanding of the cosmos. The tension between residual religious cosmologies and naturalistic visions has massive geopolitical consequences that strongly influence alliances and conflicts within and among nation states. Consider, for instance, the current polarization in the United States between evangelical Christians on the right and cultural elites on the left.

If it is true that human motives are constrained and modulated by imaginative constructs, those constructs could be used to alter the course of human history. It seems odd that one reaches this conclusion—here, in this essay—only as the final step in a series of theoretical propositions that many evolutionists (to say nothing of non-evolutionary scholars) would regard as highly doubtful: problematic, speculative, or perhaps simply absurd and implausible. It seems odd because the common, practical sense of governments and individuals alike has always understood the importance of propaganda and didactic messages conveyed through the arts. Some didactic messages, Aesop’s fables, for instance, take a very simple form: the moral of the story is. Others, like Uncle Tom’s Cabin, take elaborate symbolic and allegorical forms. But propaganda and didactic messaging are not limited to such overt efforts at changing hearts and minds. Virtually all art is imbued with some ethos, some set of values and beliefs. Insofar as artists aim to have readers share their own imaginative experiences, and occupy their own imaginative space, all the arts, all the time, aim at affecting human values and motives.

The moral character of artistic influences varies as widely as human personalities and cultural systems. Major influences on imaginative culture have included abolitionists such as Harriet Beecher Stowe (Gottschall 2012) and racialists such as D. W. Griffiths, director of the silent film Birth of a Nation (aka the Klansman). They have reached on one side of the ideological spectrum Hitler’s Mein Kampf and on the other to Mao with his Little Red Book. A grid for the ideological stances of artists would be richly populated on the right, left, and center. Even within a specific set of religious or scientific concepts about human nature and society, ideological commitments can vary all across the grid. Christians can thump that part of the Bible that emphasizes smiting one’s enemies or the part that emphasizes turning the other cheek. Darwinists can emphasize nature red in tooth and claw or the gradual expansion of communitarian sympathy (Hawkins 1997).

In itself, imagination is a force neither for good nor for evil. It is not so much beyond good and evil (Nietzsche’s formula) as before good and evil. Like other fundamental aspects of human nature, it is a framing condition that encompasses what, at any given time, humans might regard as good and evil. Our moral systems are constructed within the boundaries of our own nature. Moral systems that conflict too strongly with basic elements of human nature limit their own effectiveness. They also distort the potentialities of human nature, putting a strain on the possibilities for fulfillment and well-being. Whatever moral vision we might have, prudence suggests that we should aim to gain as clear as possible an understanding of human nature. For the past half-century or so, the evolutionary human sciences have gone a long ways toward advancing our understanding of human nature. In the past two decades, the neuroscience of imagination has taken another major step forward. As evolutionists in both the social sciences and humanities incorporate that new knowledge, they will be coming much closer to uniting the sciences and humanities.


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