Whither Queer Biology? On Richard O. Prum’s “Performance All the Way Down”

By Banu SubramaniamNovember 15, 2023

Whither Queer Biology? On Richard O. Prum’s “Performance All the Way Down”

Performance All the Way Down: Genes, Development, and Sexual Difference by Richard O. Prum

INTERDISCIPLINARY WORK across the humanities and sciences is inherently tricky and fraught. While “the body” mostly lives within the biological sciences, its key descriptive categories—like sex/gender, natural/unnatural, and health/pathology—are not inevitable “scientific” or quasiscientific categories but are very much shaped by particular histories, including patriarchal and Eurocentric ones, and doubtless plenty of others we can’t even name yet. One enduring problem is that in translating colonial politics onto the natural world from the 17th century on, bio-superiority was naturalized and scientized through and in the language of biology. Politics, in other words, became invisible within biology. A central project of feminism is to reveal the politics, but this is no easy task: scientists claim they leave cultural biases behind when they enter their laboratory or field site. Their critics show that this is not true, but the showing, as it were, is a veritable process of whack-a-mole—a scientific study is refuted only for another to reclaim so-called categorical “difference.” Despite the now-long list of tarnished studies—racial differences in skull size, sex differences in scientific ability, correlations between finger size and homosexuality, IQ differences across nations—the obsession with category-creating biological differences continues unabated. During the first two years of the COVID-19 pandemic, for instance, all kinds of claims about racial differences in the incidence, morbidity and mortality rates of COVID-19 were made. These differences were couched in the scientific language of comorbidity rather than, as they ought to have been, in that of economic inequality or unequal access to health care.

Adults, it’s now clear, are not “preformed” as the preformationists, as they were called in the 18th century, believed. We now know they emerge from complex interrelations (entanglements, feedback loops, interactions, intra-actions) between genes and environment, nature and nurture; there is no being but always a becoming. In gender scholar Angela Willey’s evocative framing, life opens up biopossibilities, complexly mediated capacities that cannot be relegated to a “pure” biology or nature but which embody socially salient traits and differences. Characteristics such as height, weight, hair length, and texture are, for example, shaped by natural and cultural factors. Likewise, categories such as sex, gender, race, class, nationality, and behavioral designations such as temperament, intelligence, abilities, and wealth exist within histories of science and politics. How we define these categories themselves constitute biopolitical projects. Biology and society are not separate entities. Some scientists have incorporated this knowledge—but have done so by entirely enfolding the social realm into their disciplines. In particular, versions of sociobiology popularized by E. O. Wilson in 1975 attempt to explain human sociality through biology: humans are the product not only of individual human biology but also of a collective social biology. Rather than nature and culture, everything here is nature and “natural.” Its critics argued, quite rightly in my view, that the early sociobiology looked a lot like the old science—it rationalized an unjust and hierarchical world as natural. A second set of responses have come from feminist, queer, critical race, intersex, trans, and disability rights scholars who refuse to cede biology to the sciences, instead insisting that any meaning-making must be centered on both nature and culture—or, more precisely, on biology, culture, and political contexts. In critiquing the biological determinism of science, they have advocated for new epistemologies, methodologies, and methods that bring the “ands” in the sentences above, in all their entangled complexity, to the fore. Neither response, sociobiological or feminist et al., is homogeneous or without internal debate.

This is where evolutionary ornithologist Richard Prum—a professor of ornithology at Yale University, and author of The Evolution of Beauty: How Darwin’s Forgotten Theory of Mate Choice Shapes the Animal World—and Us (2017)—picks up in his new book Performance All the Way Down: Genes, Development, and Sexual Difference. He combines the two strains—sociobiological and critical feminist—to propose that biology, and sex in particular, is a choreographed performance not just in terms of behaviors or the organism as a whole but all the way down to genes, cells, tissues, and hormones. I am reminded of Charis Thompson’s evocative framing of Making Parents: The Ontological Choreography of Reproductive Technologies (2005). Splicing these two strains into one theory makes for provocative reading—I’ll unpack its problematic nature later in this essay.


Where Prum succeeds is in his reprisal of a well-established critique of the normative binary sex/gender/sexuality system where binary sex (male/female) corresponds to binary gender (masculine/feminine), which together undergird a complementary union of men and women. In the human world, the claim of biological sex as innate translates into powerful intellectual, social, and political infrastructure—birth and death certificates, passports, school and medical forms, laws, sex-segregated spaces and practices. When transposed onto nature, the alleged innateness of biological sex ends up anthropomorphizing animal and plant sexual lives and sexualities into a universal grammar of sex. In short, sexed bodies are everywhere shoring up a foundational infrastructure. Under careful examination, its claims fall apart. As Prum points out, an abundance of exceptions undermines the notion of binary sex and gender, including in the human world. They are traits on a continuum rather than constituting a binary. Take any supposed marker of sex—genes, chromosomes, hormones, genitals, secondary sexual characteristics, behaviors—and there are exceptions. Humans aren’t just XX males and XY females; there are XY females and XX males, as well as XXY and XYY individuals. And there aren’t in actuality discrete sex chromosomes X and Y, but something far more complex. Our understanding is still unfolding.

The point here is that something called “science” manufactures “exceptions,” and society pathologizes these exceptions in an endless feedback loop. Similarly, no single gene, hormone, or behavior can claim to be the root of all sexual difference. Moving beyond the human species to animal and plant worlds, the amount of variation explodes. Many organisms have a chromosomal structure that is contrary to ours. In birds, females are ZW and males ZZ. In some organisms, like turtles, for instance, temperature can help determine sex. Hundreds of fish species regularly change sex as adults (and then sometimes revert). In some lizards, females are parthenogenetic (eggs develop into embryos without fertilization). Biologists Joan Roughgarden (in Evolution’s Rainbow: Diversity, Gender, and Sexuality in Nature and People) and Bruce Bagemihl (in Biological Exuberance: Animal Homosexuality and Natural Diversity) chronicle the astonishing diversity of queer possibilities. Over 85 percent of flowering plants do not have distinct male and female parts—yet even here we are obsessed with binary sex: we label flowers “male” and “female.” If flowers aren’t binary, then we mark flower structures as “male” and “female.” These binary categories then unfold within biology into sexual scripts, featuring Darwin’s “coy females” and “aggressive males.” “Male” and “female” aren’t innocent terms, however—they can generate misogynist scripts. In short, despite resplendent diversity, “science” has created a monotonous world modeled on Victorian gentlemen and ladies! As Prum summarizes, the deviations in sexual development are not failures or aberrations but “evidence that the individual sexual binary does not exist.” It’s easy to agree with Prum’s analysis thus far.

It gets more complicated though. Prum builds on this critique to create a grand unifying theory. The book cover features a blue-and-pink chromosome bathed in pastel shades of pink and blue. Prum is not refusing a binary—rather, he is refusing simple bio-determinism in favor of something more interesting that draws in part on Judith Butler’s theory of gender as performance. In a nutshell, gender, they argue, is socially constituted through everyday speech acts and communication styles that are performative. Gender is a performance, a process, a doing by an individual in a social world. Prum’s extrapolation: Performance is a “playing out of an historically derived role in a social context” (an enactment of Darwinian evolution), and a becoming through doing, which provides a role for individual action and thus for the agency of organisms. Here he also draws inspiration from feminist philosopher Karen Barad and their epistemology of intra-action and agential realism. Refusing the distinction between matter and discourse, agency is not enacted by inter-actions between discrete organisms, Barad argues, but rather through intra-actions: the dynamism of forces in which all “things” constantly exchange, diffract, influence, and work inseparably. In agential realism, both matter and discourse are mutually constitutive.

Prum does not challenge or critique feminist or queer theory, but instead folds them wholesale into biological theories of evolution and animal behavior. That would be fine up to a point—but he misunderstands their core ideas. He insists, for instance, on core differences between sex categories as vital for evolution. Sex, he argues, is relational and dyadic within populations: i.e., there are two sexes even though individuals may not be one or the other. And while critical of sex difference research, he nonetheless finds the notion of sexual difference productive; it is “productive” to study individual “sexual” variation in the absence of a priori binary categories of individual sex. In his theorization, a phenotype, such as the bird’s brilliant plumage or mating dances, is a performative enactment by an individual organism. Hormones don’t cause effects—rather, individual bodies employ hormones to enact desirable phenotypes. Rather than adhering to a story where hormone levels are biologically determined and produce particular behaviors, hormones themselves are open to social and evolutionary forces. In this way, his theory of performance helps outline a choreography of molecular, cellular, and organismal intra-actions.

The book is not actually interdisciplinary but rather multidisciplinary. Two bodies of work—namely biological work on sex on the one hand and feminist and queer work on the other—are juxtaposed, quite literally in separate chapters and appendices. The chapters on feminist and queer theory cite key texts and explain important concepts. These are not simplistic or simplified but elaborated with care and some depth. Yet, reading about them was an odd experience for me. For much of my academic life, I’ve bemoaned the refusal of most scientists (I myself am trained as a scientist) to take feminist and queer theory seriously. And I’ve noticed that when scientists do discover for themselves the complexity of sex and gender, they rarely credit feminist scholarship. So, kudos to Richard Prum for reading this work, citing it, and recommending it to his peers. And yet, the book makes me uneasy. It is as though someone took an advanced course in feminist and queer theory but entirely skipped Feminism or Queer Studies 101! For all its grand theory and terminology, the book is entirely devoid of politics or a theory of power, which is after all the raison d’être of feminist and queer theories. This obliviousness to politics leads to strange conclusions—like arguing that a feminist notion of gender/sex is “precisely congruent” with Richard Dawkins’s concept of extended phenotype! To glibly claim feminist congruence with a figure decidedly hostile to feminism (and vice versa) is jarring, to say the least. The claim demands at least some exegesis. Similarly, he frames his work as extending Butler to Lewis Thomas’s The Lives of a Cell: Notes of a Biology Watcher (1974); he squishes Barad and Bruno Latour together as though they were analogous, and extends Foucault unproblematically to theories of the evolution of sexual reproduction itself. These easy analogies between the two fields may be unnerving for readers conversant with both. One is likely to get whiplash.

Interdisciplinarity is difficult because knowledge formation is organic and accumulates over time, becoming ever more nuanced, debated, and detour-prone. Knowledge is deeply contextual within fields; words and terminology aim to be precise and must be understood within disciplinary histories. Practicing feminist and queer theory isn’t just about using the right words but also about understanding and engaging with power and politics. Much can be lost in translation, and here it most certainly is. Ultimately, for a book engaging with queer theory, it is decidedly unqueer! For example, after the extensive critique of binary sex/gender, Prum holds on to the two categories male and female, but in capitals—which, he claims, makes it easier for him to discuss reproductive biology. But wouldn’t talking about reproduction without Males and Females be productively queer? For example, consider intersexuality. If we support—as we absolutely should—the end of unwanted genital surgeries, then we ought to be able to deal with the continuum of sex, and to do so without othering and stigmatizing nonbinary individuals. Many countries have indeed begun to offer options beyond “male” and “female” in their passports, and social media platforms have embraced a plethora of genders. Workplaces allow individuals to claim their own pronouns (or refuse them). Scholars have proposed models of sex as a continuum. With respect to plants, Madelaine Bartlett and I propose dispensing with the terms altogether, focusing instead on reproductive structures. Yet, despite the proliferation of queer life and possibilities in the world, and despite his own critique of binary sex at an individual level, Prum insists on its significance at higher levels, and especially so for those embodied features that have “evolved for reproductive function.” This means that, in his book, the whole apparatus of reproductive heteronormativity—gender, heterosexuality, family, marriage, monogamy—looms large but unaddressed. In short, Prum offers a theory of biological performances of intra-actions, yet ahistoricized and depoliticized.

Most importantly, the invocation of sex/gender binaries as universal invariably risks reproducing whiteness-related assumptions. For example, many women athletes, especially Black women, are routinely read as masculine. Femininity remains resolutely in the province of delicate whiteness. In many countries, like India, light skin personifies beauty. This is a colonial aesthetic, and a colonial legacy. Colonial hierarchies extended the dehumanized status of animals to nonwhite colonial subjects. In short, Prum may cite work on intersectionality and claim to be “an ornithologist for intersectionality,” but there is evidence to the contrary in his insistence on sex/gender as productive.

Also, by centering reproduction at more meta levels than that of the individual, Prum is able to do something he is clearly keen to do: embrace theories of sexual selection, which also means embracing a gamut of phenomena that some feminists have long critiqued because they are implicated in histories of biological determinism. Those histories, it should be noted, have wrought systematic pain and suffering onto gendered, raced, and colonized bodies deemed marginal. Eugenics is possible because “science” identified some bodies as unworthy of life—e.g., the purportedly feeble-minded, degenerate, perverse, deformed, and promiscuous, as well as epileptics, criminals, alcoholics, paupers, and so on. Eugenics—the late-19th-century science of “good genes,” along with the statistical methods developed by Francis Galton—provided the scientific rationalization for 20th-century sterilization laws in the United States, which then inspired the Holocaust in Europe. Eugenic logics live on in this century, most dramatically in IVF contexts, in who is encouraged to reproduce (wealthy, white), and in who is discouraged or even coerced into long-term contraception or sterilization (people of color and the poor). Some fetuses, as we know, are preferentially terminated. According to the logic of bio-determinism, they can’t be dismantled or undone in the population at large except through the extermination of their germ- or bloodlines. Biology becomes destiny rather than ideology.

On a slightly different note, I find Prum’s easy extension of human phenomena to plants and animals troubling. For example, he advocates the use of the word “rape” in nonhuman organisms because “the term forced copulation in biology has allowed scientists to avoid the recognition that sexual violence is, to paraphrase [journalist and activist Susan] Brownmiller, against the will of the ducks.” For decades, feminists have insisted that we understand rape as an exercise in power rather than a biological imperative. Men rape not because of biological programming but because of male supremacy. Human rape is not an “evolved feature of […] evolutionary histor[y].”

In many ways, Prum proposes a queer sociobiology in much the way that some feminists have posited a feminist sociobiology. Both formulations are oxymorons. I don’t think we need to biologize everything in the world to explain human politics. We don’t need to argue that feminism, civil rights movements, anti–sexual violence, and anti–sexual harassment are biological responses to oppression. We don’t have to wait for our biologies to evolve a countermovement to oppressive histories and actions. We need the language of politics, power, and resistance, not the language of biology. Otherwise, we are left with a scientized humanities, with everything in the world engulfed by a sociobiology that portrays us as the victims of our own biology and evolutionary history (written by and for a colonial and patriarchal science).


A deeper engagement with feminist and queer science and with technology studies opens up other possibilities. First, we might consider the deep asymmetries of power across the humanities and sciences. What counts as science? Prum argues that while feminists and queer scholars have offered critiques, they have been hesitant “to roll up their sleeves and actually do science—try [to] make science work better.” But what does it mean to “do” science? What seems unacknowledged is that while scientists read theory in science journals, they do not read feminist or queer theory as science. So many of us with PhDs in the sciences who continue to work on the sciences outside of formal appointments in scientific disciplines are dismissed as having “left” science. We have not left. We are émigrés of the interior, as it were. We are seeking innovative ways of thinking and perceiving, and new methodologies and instruments for doing science. The common response of scientists, alas, is to more aggressively police their institutional borders.

Second, we might consider that, within the walls of science, a distinguished senior scientist like Prum can embark on this work, but can a graduate student or junior feminist or queer scientist? Will their work be published in scientific journals? Will their work be read as science, credited in decisions regarding promotion and tenure? No, of course not.

Third, while Prum does cite and summarize the role of race in theories of feminist and queer studies, race as a concept drops off repeatedly in his book. He pays a great deal of lip service to intersectionality in feminist and queer studies, but he fails to realize that these fields have not yet fully integrated race or colonialism. In his book, the natural world, for example, is gendered but rarely raced or colonized. Recent work by Bénédicte Boisseron, Zakiyyah Iman Jackson, and Harlan Weaver demonstrates just how much race grounds colonial thinking. Animality and race are transposed not only onto some human subjects but also onto animal and plant worlds. Theorizing gender through the history of colonialism might have helped Prum destabilize the whiteness of a universal sex/gender system.

Finally, there is much work in feminist and queer studies that destabilizes sex and gender and decenters reproduction. Why not do more with horizontal and lateral inheritance, where genetic material moves across organisms—as when bacteria and viruses move genetic material across species—rather than vertically from parent to offspring? While Prum does mention this in passing, he foregrounds vertical inheritance and sexual reproduction as the bedrock of evolution. It’s too bad. Decentering reproduction would have been delightfully queer. Similarly, theories of symbiogenesis queer individualist stories of evolution. Feminist, queer, and trans biologies stress multiplicity in order to open up biopossibilities. Rather than extending human biology to animals and plants, why not go the other way? As Myra Hird argues, given that most plants are intersex, fungi have multiple sexes, other species are transex, and bacteria are devoid of sexual difference altogether, we ought to ground theories of sex/gender in the world that defies sexual logics. Maybe then we can rethink the human beyond its sexual formations.

Whither queer biology? In the early days of institutionalized feminism, scholars debated whether they were working for their own obsolescence. If all disciplines embraced feminist work, then might there be no need for feminist departments? In his conclusion, Prum writes that once the biological sciences more thoroughly engage with queer studies in their research and teaching, they will cease to be queer. Not true! As disciplines engage with feminist work, feminist and queer work doesn’t stand still; it continues to chart new ground, unmoored from the silos of disciplines. Feminist and queer ideas are not static, in other words, but are relational and themselves evolving. By the time biologists embrace this version of queer theory, the fields will have moved on. We don’t need a momentary exchange but, rather, continual scrutinization of the unequal distribution of power across disciplines. And we need to broker new practices of collaboration that open up heretofore unthought-of choreographies of interdisciplinarity.


Banu Subramaniam is the Luella LaMer Professor of Women’s and Gender Studies at Wellesley College. Banu expresses deep gratitude for feedback from Sushmita Chatterjee, Nayanika Ghosh, Rebecca Herzig, Ambika Kamath, Anita Simha, and Angela Willey, and to Michele Pridmore-Brown for her superb editorial input.

LARB Contributor

Banu Subramaniam is the Luella LaMer Professor of Women’s and Gender Studies at Wellesley College and the author of Ghost Stories for Darwin: The Science of Variation and the Politics of Diversity (University of Illinois Press, 2014), Holy Science: The Biopolitics of Hindu Nationalism (University of Washington Press, 2019), and Botany of Empire: Plant Worlds and the Scientific Legacies of Colonialism (University of Washington Press, 2024). Trained as a plant evolutionary biologist, Banu engages the feminist studies of science in the practices of experimental biology. Banu’s current work focuses on decolonizing botany, nativism in plant biology, and the relationship of science and religious nationalism in India.


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