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Performance All the Way Down: Genes, Development, and Sexual Difference
An award-winning biologist and writer applies queer feminist theory to developmental genetics, arguing that individuals are not essentially male or female.
The idea that gender is a performance—a tenet of queer feminist theory since the nineties—has spread from college classrooms to popular culture. This transformative concept has sparked reappraisals of social expectations as well as debate over not just gender, but what it is, what it means, and how we know it. Most scientific and biomedical research over the past seventy years has assumed and reinforced a binary concept of biological sex, though some scientists point out that male and female are just two outcomes in a world rich in sexual diversity.
In Performance All the Way Down , MacArthur Fellow and Pulitzer Prize finalist Richard O. Prum brings feminist thought into conversation with biology, arguing that the sexual binary is not essential to human genes, chromosomes, or embryos. Our genomes are not blueprints, algorithms, or recipes for the physical representation of our individual sexual essences or fates. In accessible language, Prum shows that when we look closely at the science, we see that gene expression is a material action in the world, a performance through which the individual regulates and achieves its own becoming. A fertilized zygote matures into an organism with tissues and organs, neurological control, immune defenses, psychological mechanisms, and gender and sexual behavior through a performative continuum. This complex hierarchy of self-enactment reflects the evolved agency of individual genes, molecules, cells, and tissues.
Rejecting the notion of an intractable divide between the humanities and the sciences, Prum proves that the contributions of queer and feminist theorists can help scientists understand the human body in new ways, yielding key insights into genetics, developmental biology, physiology. Sure to inspire discussion, Performance All the Way Down is a book about biology for feminists, a book about feminist theory for biologists, and a book for anyone curious about how our sexual bodies grow.
The idea that gender is a performance—a tenet of queer feminist theory since the nineties—has spread from college classrooms to popular culture. This transformative concept has sparked reappraisals of social expectations as well as debate over not just gender, but what it is, what it means, and how we know it. Most scientific and biomedical research over the past seventy years has assumed and reinforced a binary concept of biological sex, though some scientists point out that male and female are just two outcomes in a world rich in sexual diversity.
In Performance All the Way Down , MacArthur Fellow and Pulitzer Prize finalist Richard O. Prum brings feminist thought into conversation with biology, arguing that the sexual binary is not essential to human genes, chromosomes, or embryos. Our genomes are not blueprints, algorithms, or recipes for the physical representation of our individual sexual essences or fates. In accessible language, Prum shows that when we look closely at the science, we see that gene expression is a material action in the world, a performance through which the individual regulates and achieves its own becoming. A fertilized zygote matures into an organism with tissues and organs, neurological control, immune defenses, psychological mechanisms, and gender and sexual behavior through a performative continuum. This complex hierarchy of self-enactment reflects the evolved agency of individual genes, molecules, cells, and tissues.
Rejecting the notion of an intractable divide between the humanities and the sciences, Prum proves that the contributions of queer and feminist theorists can help scientists understand the human body in new ways, yielding key insights into genetics, developmental biology, physiology. Sure to inspire discussion, Performance All the Way Down is a book about biology for feminists, a book about feminist theory for biologists, and a book for anyone curious about how our sexual bodies grow.
403 pages, Paperback
Published November 16, 2023
About the author
Richard O. Prum
7 books75 followersRichard O. Prum is William Robertson Coe Professor of Ornithology, and Head Curator of Vertebrate Zoology at the Peabody Museum of Natural History at Yale University. Prum describes himself as "an evolutionary ornithologist with broad interests in diverse topics," including phylogenetics, behavior, feathers, structural coloration, evolution and development, sexual selection, and historical biogeography. He has conducted field work throughout the world, and has studied fossil theropod dinosaurs in China. He received a MacArthur Fellowship in 2010.
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March 21, 2025I was curious to read a book that seeks to synthesize a scientific field (evolutionary developmental biology) with an academic tradition that is explicitly political and very often critical to the natural sciences, namely feminist queer theory. Within the latter tradition you do find academics that are explicitly anti-science; who interpret scientific concepts as purely racist, misogynous instruments developed to maintain political status quo and oppress ethnic and sexual minorities. This seemed to me like an odd partner to invite in for cross-diciplinary research, but I decided to give the book a try.
My curiosity was soon replaced with disappointment, however. Several times the author uses historic examples of misuse of scientific concepts as arguments against that science. It is strange to read a trained scientist handle the distinction between descriptive science and normative interpretations so carelessly. Eugenics was a bad political idea inspired by misunderstood concepts from population genetics. The problem was obviously politics and not science. On the contrary, population genetics has helped us to better understand many important principles in biology, like how processes such as genetic drift and natural selection shapes heritable variation in natural populations. (NB! I do not imply, however, that queer theory and other critical theories are without merit in the philosophy of science).
Even though the motivation for this cross-diciplinary investigation at least in part appears to have been politically motivated, it is not given that there can't be useful synergies anyway. Prum himself is convinced that his cross-diciplinary exercise has been fruitful. He utilizes concepts that have been used to describe cultural change and reinterprets them applied to biological development (embryology and beyond). Concepts like "discourse", "agency" and "performativity" are given new meanings when applied to the developing body. However, it is hard to judge how fruitful this exercise actually has been. The text is quite hard to access. The author alternates between advanced scientific terminology and equally inaccessible jargon from philosophy/humanities. Very few readers would be familiar with both. Are we supposed to feel stupid or impressed by the authors intellectual broadness? This is really not effective communication. I guess many readers will be overwhelmed and give up long before finishing the book.
Development is a process. The relationship between genotype and phenotype is a dynamic one. Traditional evo-devo obviously takes this into account. I therefore struggle to see what the dynamic concepts derived from queer theory adds to the table. Prum mentions a few examples of hypotheses that he claims was inspired by his novel "performative science". Although I agree that these hypotheses are interesting I am unable see that they couldn't equally well have been formulated in the framework of classical evolutionary biology. For instance, one hypothesis is about signal interference between the gestating parent and the embryo across the placenta - a scenario that has been repteatedly analyzed in classical evolutionary biology, for instance in the case of the immunity reaction to the Rhesus factor during human pregnancies.
In conclusion, this book is full of problematic mingling of politics and science.
My curiosity was soon replaced with disappointment, however. Several times the author uses historic examples of misuse of scientific concepts as arguments against that science. It is strange to read a trained scientist handle the distinction between descriptive science and normative interpretations so carelessly. Eugenics was a bad political idea inspired by misunderstood concepts from population genetics. The problem was obviously politics and not science. On the contrary, population genetics has helped us to better understand many important principles in biology, like how processes such as genetic drift and natural selection shapes heritable variation in natural populations. (NB! I do not imply, however, that queer theory and other critical theories are without merit in the philosophy of science).
Even though the motivation for this cross-diciplinary investigation at least in part appears to have been politically motivated, it is not given that there can't be useful synergies anyway. Prum himself is convinced that his cross-diciplinary exercise has been fruitful. He utilizes concepts that have been used to describe cultural change and reinterprets them applied to biological development (embryology and beyond). Concepts like "discourse", "agency" and "performativity" are given new meanings when applied to the developing body. However, it is hard to judge how fruitful this exercise actually has been. The text is quite hard to access. The author alternates between advanced scientific terminology and equally inaccessible jargon from philosophy/humanities. Very few readers would be familiar with both. Are we supposed to feel stupid or impressed by the authors intellectual broadness? This is really not effective communication. I guess many readers will be overwhelmed and give up long before finishing the book.
Development is a process. The relationship between genotype and phenotype is a dynamic one. Traditional evo-devo obviously takes this into account. I therefore struggle to see what the dynamic concepts derived from queer theory adds to the table. Prum mentions a few examples of hypotheses that he claims was inspired by his novel "performative science". Although I agree that these hypotheses are interesting I am unable see that they couldn't equally well have been formulated in the framework of classical evolutionary biology. For instance, one hypothesis is about signal interference between the gestating parent and the embryo across the placenta - a scenario that has been repteatedly analyzed in classical evolutionary biology, for instance in the case of the immunity reaction to the Rhesus factor during human pregnancies.
In conclusion, this book is full of problematic mingling of politics and science.
February 18, 2025
Whereas Prum’s book The Evolution of Beauty - which is one of my favorite books of all time- was certainly written for the masses- this important work was written for the more hard science minded. I was swimming in chromosomes and hormones and proteins and signaling pathways - but fortunately he writes- and summarizes his points so well- that my beliefs about gender/sex are fully shook!
“In response to the questions posed by Judith Butler in Gender Trouble, “What is sex, anyway?” and “Does sex have a history?” we can now offer the biological answer that sex is a history—a hierarchical history of coevolved bodily traits with intra-active reproductive functions, and the iterated, individual histories of the material realization of those reproductive possibilities. Sex is not a prior or given fact about any body, because the genetic-discursive process of becoming an organism affords no inputs for any such essence, predetermined truth, or antecedent fact about sex. The sex of the individual is a self-realization, the product of a performative becoming. It is “turtles all the way down.” Or, as I prefer, turtles all the way up.”
“In response to the questions posed by Judith Butler in Gender Trouble, “What is sex, anyway?” and “Does sex have a history?” we can now offer the biological answer that sex is a history—a hierarchical history of coevolved bodily traits with intra-active reproductive functions, and the iterated, individual histories of the material realization of those reproductive possibilities. Sex is not a prior or given fact about any body, because the genetic-discursive process of becoming an organism affords no inputs for any such essence, predetermined truth, or antecedent fact about sex. The sex of the individual is a self-realization, the product of a performative becoming. It is “turtles all the way down.” Or, as I prefer, turtles all the way up.”
May 18, 2026
Performance All the Way Down argues that biology is not the unfolding of fixed essences encoded in genes, but an ongoing, performative process in which organisms actively become themselves through developmental, physiological, evolutionary, and social interactions. Drawing from developmental biology, evolutionary theory, queer feminist theory, and philosophy, Richard Prum challenges adaptationist and binary understandings of sex and gender, proposing instead that phenotypes (including bodies, behaviors, and sexual identities) are enacted through dynamic relationships among genes, cells, environments, and culture. Across examples ranging from feather evolution and bird aesthetics to embryology and intersex variation, the book reframes life as a “performative continuum” in which agency, contingency, and history matter at every level of biological organization.
-
Prologue Summary
The prologue introduces two central themes: biological complexity emerges through developmental processes rather than simple adaptation, and organisms possess forms of agency that shape evolution itself. Prum begins with feathers as an example of evolutionary innovation that cannot be understood solely through natural selection. Feather complexity emerged through “hierarchical modularity,” where structures evolved new dimensions and developmental possibilities over time. He argues that focusing too narrowly on adaptation historically prevented scientists from understanding how feathers actually evolved.
The discussion of structural color in bird feathers expands this argument. Bluebird feathers create color not through pigment but through nanoscale self-organization of keratin and air bubbles. Organisms do not directly engineer these structures; instead, cells create conditions under which materials self-assemble through physical processes like phase separation. Biology therefore depends on emergent interactions between matter and development rather than top-down design.
The prologue then transitions to aesthetics and sexuality in birds, arguing that animal subjectivity, especially desire and attraction, has profoundly shaped evolution. Sexual selection is reframed not as reproductive optimization but as the consequence of individual sensory preferences and aesthetic agency. This perspective leads Prum toward queer feminist theory, which he adopts as a conceptual framework capable of transforming biology itself rather than merely critiquing it from outside.
QUOTES:
“Thinking about adaptation was an intellectual hindrance to progress on the question of the origin of evolutionary innovations, like feathers.”
“The aesthetic view of mate choice and sexual selection reframes reproduction as a downstream consequence of animal subjectivity.”
“This effort is not a scientific accommodation to contemporary culture or politics; rather, it constitutes a genuine intellectual and empirical contribution to biological science.”
-
# Chapter 1: Performance All the Way Down
This chapter establishes the book’s core thesis: the phenotype is not a static biological fact but a performance or enactment of the self. Prum critiques the traditional genotype-to-phenotype model that treats genes as deterministic blueprints. Instead, he argues that organisms continuously become themselves through developmental and environmental interactions. The phenotype includes not just anatomy, but physiology, behavior, consciousness, gender, and sexuality.
Prum introduces the concept of the “performative continuum,” which links molecular biology, organismal development, evolution, and culture. Biology is not separate from culture; scientific concepts themselves are shaped by cultural assumptions and metaphors. He specifically critiques adaptationism for dominating biology and marginalizing alternative frameworks that better account for variability, contingency, and agency.
The chapter also frames queer biology as both an intellectual and political intervention. Prum argues that biology has historically contributed to violence against queer people by naturalizing binary and essentialist ideas about sex and gender. Queering biology therefore means rethinking foundational assumptions about what bodies are and how they develop.
Quotes:
“The phenotype is best understood—materially, biologically, and culturally—as the performance, or enactment, of the self.”
“We are performance all the way down.”
“Queering biology is an appropriate way for science to reckon with the historical and ongoing contributions of biological science” to harms against queer people.
-
# Chapter 2: Critical Concepts
This chapter redefines sex as a historical and developmental phenomenon rather than a fixed binary essence. Prum argues that categories like “male” and “female” are based on reproductive functions and homologous anatomical clusters, but these categories do not apply neatly to all individuals. Infertility, aging, childhood, and developmental variation all expose the instability of essentialist definitions.
Instead of treating sex as an innate property, Prum proposes that sex should be understood historically—as “clusters of embodied reproductive homologies” produced through evolution. Human reproduction requires a binary reproductive process, but that does not mean every individual must fit into one of two natural categories.
The chapter also develops the idea of biological agency. Organisms and cells respond contingently to evolutionary history and current environments. Variation is not necessarily failure; sometimes it is compensatory or stabilizing. Prum’s concept of the “extended phenotype” further blurs the line between reproductive and non-reproductive traits.
Quotes:
“Like many organisms, adult human bodies vary in ways that are related to reproductive functions. These reproductive variations have been traditionally used to refer to individual organisms and their bodies as either male or female. To many people, including many scientists, these terms imply the existence of natural, binary categories that apply to all individuals. …But this definition is not really ac curate because we also use the categories male and female to refer to many individuals that cannot, or do not, sexually reproduce, such as children, the aged, or the infertile.”
-
# Chapter 3: Gender Performativity
This chapter brings Judith Butler’s theory of performativity into dialogue with biology. Prum argues that gender is iterative and citational: every enactment of gender references prior enactments. Gender therefore exists relationally and historically rather than emerging from inner essence.
He extends this insight into molecular biology itself. Communication and signaling among cells constitute forms of biological discourse. Biological systems are therefore performative not just metaphorically but materially: organisms emerge through networks of communicative intra-actions.
The chapter challenges the division between “cultural” performativity and “biological” fact. Instead, performativity operates continuously from cellular signaling to human social identity.
Quotes:
as a lover of citations, i love this “Another crucial aspect of performativity is the iterative property that Butler (drawing on literary theorist Jacques Derrida) refers to as citational-ity. Like academic citations of previous literary, artistic, or intellectual ex-pressions, citationality captures the idea that individual gender realizations exist in reference to, or as citations of, other previous or contemporaneous enactments of gender. Citationality describes the relation between individual instances of gender in different individuals at different places and times.”
-
# Chapter 4: The Enactment of the Biological Self
This chapter explains how developmental biology supports Prum’s performative framework. Genes do not directly determine bodies; rather, organisms emerge through dynamic signaling networks among cells. The genome is not a blueprint but “an informational resource.”
Prum carefully explains endocrine and paracrine signaling, showing how cells communicate through hormones, receptors, and signaling cascades. Cells interpret and respond selectively to signals, which means development involves contingency and agency. Cells can “listen” or fail to listen to one another.
Prum revisits Waddington’s epigenetic landscape, reframing it as a “performative developmental landscape.” Development is canalized—certain outcomes are more likely—but not predetermined.
Quotes:
- thesis of whole book: “The most concise way to express the thesis of this book is that the phenotype is a performative enactment of the individual organism. Each individual organism begins with an inherited genotype and some (modest or extensive) material contribution from its parents. The resulting organism is the outcome of a performative process realized in interaction with the organismal environment.”
-
# Chapter 5: How Do Our Sexual Bodies Develop?
This chapter examines mammalian sexual development and critiques simplistic models of sex determination. Prum argues that sexual phenotypes are realized through developmental processes rather than encoded in genes from the outset.
He explains that mammalian females develop through a “default” pathway shared by all embryos, while Y-linked genes can act as developmental “difference makers.” However, these genes are not master switches; they depend on complex interactions with many other molecular systems.
Prum also compares mammalian and avian sex systems to show that sex determination mechanisms are evolutionarily variable and historically contingent.
Quotes:
- “Sex is an historical constellation, or cluster, of iteratively realized, co-occurring anatomical and physiological homologies that have evolved over evolutionary history through selection for sexual reproduction (see chapter 2).”
- “However, a bimodal distribution in reproductive anatomy and physiology within species does not justify the existence or assignment of an inherent, individual sexual essence or identity to the zygote or genome prior to, or independent of, the sexual phenotype's material realization.”
-
# Chapter 6: Variations in Our Sexual Development
This chapter focuses on intersex and variations in sexual development. Prum argues that developmental diversity demonstrates the absence of an essential binary sex system.
Human sexual development is strongly canalized toward bimodal outcomes, but the complexity of developmental pathways creates many intermediate possibilities. Traditional scientific and medical frameworks pathologized these variations because they assumed a rigid binary that biology itself does not support.
Prum reframes intersex variation not as developmental failure but as evidence that sexual development is inherently variable and contingent.
Quotes:
- “In other words, sex is not an innate property of individual genomes or embryos waiting to be represented in the material world, but a suite of reproductive possibili ties realized, or enacted, by those individuals through development. The sexual phenotype is an active, assertive developmental achievement by the organism. This reframing is critical to understanding the substantial variability in human sexual development. Anatomical variation in human sexual anatomy is strongly, bimodally distributed-that is, the vast majority of human individuals develop the anatomically and physiological capacities to sexually reproduce either through the production and delivery of sperm, or through the production of eggs, pregnancy, lactation, and so on. This bimodal distribution is evidence of the strong canalization (i.e., evolutionary reduction of variation) of sexual development, but it is not sufficient to imply, support, or create an essential (i.e., inherent and indispensable) scientific fact about the sex of a genome, a zygote, or its phenotypic fate.”
- “Rather than regard differences in sexual development as pathological failures of a binary sexual development, we should realize that the developmental variations in the sexual body are actually evidence that the individual sexual binary does not exist in the first place.”
-
# Chapter 7: How Evolution Generates Sexual Variability
This chapter expands the argument evolutionarily, showing that sex determination systems differ radically across species. Sexual differentiation mechanisms are evolutionarily unstable and constantly changing.
Prum argues that this diversity proves there is no universal biological essence of sex. Instead, evolution repeatedly generates new ways of realizing reproductive possibilities. He also discusses placental evolution and immune interactions during pregnancy to show how reproductive biology emerges through evolutionary rewiring.
---
# Chapter 8: The Future of Performative Biology
This chapter explores future implications for developmental biology, medicine, and reproductive theory. Prum examines pregnancy as a performative interaction between gestating parent and embryo, arguing that gestational environments actively shape development.
He proposes hypotheses about why gestating mammals are homogametic (XX) and links performative biology to medical understanding of infertility and pregnancy complications.
The chapter concludes by redefining illness itself as a failed or disrupted self-performance rather than simply mechanical breakdown.
“Thinking about illness and disease as performative mis-realizations of the self…focuses our attention on the fact that there are indeed a greater variety of ways to go wrong than right.”
-
# Chapter 9: Performance All the Way Up
The final chapter broadens performativity beyond biology into culture and communication. Human culture emerges from deeper biological performative systems rather than standing apart from them.
Prum highlights examples of animal culture, especially birds and whales, where vocal and social behaviors are learned and transmitted across generations. He argues that gender and sexual culture are among the earliest and most transformative forms of culture in evolutionary history.
The chapter closes the loop on the book’s central argument: performativity operates continuously from molecules to cells to organisms to societies.
-
Prologue Summary
The prologue introduces two central themes: biological complexity emerges through developmental processes rather than simple adaptation, and organisms possess forms of agency that shape evolution itself. Prum begins with feathers as an example of evolutionary innovation that cannot be understood solely through natural selection. Feather complexity emerged through “hierarchical modularity,” where structures evolved new dimensions and developmental possibilities over time. He argues that focusing too narrowly on adaptation historically prevented scientists from understanding how feathers actually evolved.
The discussion of structural color in bird feathers expands this argument. Bluebird feathers create color not through pigment but through nanoscale self-organization of keratin and air bubbles. Organisms do not directly engineer these structures; instead, cells create conditions under which materials self-assemble through physical processes like phase separation. Biology therefore depends on emergent interactions between matter and development rather than top-down design.
The prologue then transitions to aesthetics and sexuality in birds, arguing that animal subjectivity, especially desire and attraction, has profoundly shaped evolution. Sexual selection is reframed not as reproductive optimization but as the consequence of individual sensory preferences and aesthetic agency. This perspective leads Prum toward queer feminist theory, which he adopts as a conceptual framework capable of transforming biology itself rather than merely critiquing it from outside.
QUOTES:
“Thinking about adaptation was an intellectual hindrance to progress on the question of the origin of evolutionary innovations, like feathers.”
“The aesthetic view of mate choice and sexual selection reframes reproduction as a downstream consequence of animal subjectivity.”
“This effort is not a scientific accommodation to contemporary culture or politics; rather, it constitutes a genuine intellectual and empirical contribution to biological science.”
-
# Chapter 1: Performance All the Way Down
This chapter establishes the book’s core thesis: the phenotype is not a static biological fact but a performance or enactment of the self. Prum critiques the traditional genotype-to-phenotype model that treats genes as deterministic blueprints. Instead, he argues that organisms continuously become themselves through developmental and environmental interactions. The phenotype includes not just anatomy, but physiology, behavior, consciousness, gender, and sexuality.
Prum introduces the concept of the “performative continuum,” which links molecular biology, organismal development, evolution, and culture. Biology is not separate from culture; scientific concepts themselves are shaped by cultural assumptions and metaphors. He specifically critiques adaptationism for dominating biology and marginalizing alternative frameworks that better account for variability, contingency, and agency.
The chapter also frames queer biology as both an intellectual and political intervention. Prum argues that biology has historically contributed to violence against queer people by naturalizing binary and essentialist ideas about sex and gender. Queering biology therefore means rethinking foundational assumptions about what bodies are and how they develop.
Quotes:
“The phenotype is best understood—materially, biologically, and culturally—as the performance, or enactment, of the self.”
“We are performance all the way down.”
“Queering biology is an appropriate way for science to reckon with the historical and ongoing contributions of biological science” to harms against queer people.
-
# Chapter 2: Critical Concepts
This chapter redefines sex as a historical and developmental phenomenon rather than a fixed binary essence. Prum argues that categories like “male” and “female” are based on reproductive functions and homologous anatomical clusters, but these categories do not apply neatly to all individuals. Infertility, aging, childhood, and developmental variation all expose the instability of essentialist definitions.
Instead of treating sex as an innate property, Prum proposes that sex should be understood historically—as “clusters of embodied reproductive homologies” produced through evolution. Human reproduction requires a binary reproductive process, but that does not mean every individual must fit into one of two natural categories.
The chapter also develops the idea of biological agency. Organisms and cells respond contingently to evolutionary history and current environments. Variation is not necessarily failure; sometimes it is compensatory or stabilizing. Prum’s concept of the “extended phenotype” further blurs the line between reproductive and non-reproductive traits.
Quotes:
“Like many organisms, adult human bodies vary in ways that are related to reproductive functions. These reproductive variations have been traditionally used to refer to individual organisms and their bodies as either male or female. To many people, including many scientists, these terms imply the existence of natural, binary categories that apply to all individuals. …But this definition is not really ac curate because we also use the categories male and female to refer to many individuals that cannot, or do not, sexually reproduce, such as children, the aged, or the infertile.”
-
# Chapter 3: Gender Performativity
This chapter brings Judith Butler’s theory of performativity into dialogue with biology. Prum argues that gender is iterative and citational: every enactment of gender references prior enactments. Gender therefore exists relationally and historically rather than emerging from inner essence.
He extends this insight into molecular biology itself. Communication and signaling among cells constitute forms of biological discourse. Biological systems are therefore performative not just metaphorically but materially: organisms emerge through networks of communicative intra-actions.
The chapter challenges the division between “cultural” performativity and “biological” fact. Instead, performativity operates continuously from cellular signaling to human social identity.
Quotes:
as a lover of citations, i love this “Another crucial aspect of performativity is the iterative property that Butler (drawing on literary theorist Jacques Derrida) refers to as citational-ity. Like academic citations of previous literary, artistic, or intellectual ex-pressions, citationality captures the idea that individual gender realizations exist in reference to, or as citations of, other previous or contemporaneous enactments of gender. Citationality describes the relation between individual instances of gender in different individuals at different places and times.”
-
# Chapter 4: The Enactment of the Biological Self
This chapter explains how developmental biology supports Prum’s performative framework. Genes do not directly determine bodies; rather, organisms emerge through dynamic signaling networks among cells. The genome is not a blueprint but “an informational resource.”
Prum carefully explains endocrine and paracrine signaling, showing how cells communicate through hormones, receptors, and signaling cascades. Cells interpret and respond selectively to signals, which means development involves contingency and agency. Cells can “listen” or fail to listen to one another.
Prum revisits Waddington’s epigenetic landscape, reframing it as a “performative developmental landscape.” Development is canalized—certain outcomes are more likely—but not predetermined.
Quotes:
- thesis of whole book: “The most concise way to express the thesis of this book is that the phenotype is a performative enactment of the individual organism. Each individual organism begins with an inherited genotype and some (modest or extensive) material contribution from its parents. The resulting organism is the outcome of a performative process realized in interaction with the organismal environment.”
-
# Chapter 5: How Do Our Sexual Bodies Develop?
This chapter examines mammalian sexual development and critiques simplistic models of sex determination. Prum argues that sexual phenotypes are realized through developmental processes rather than encoded in genes from the outset.
He explains that mammalian females develop through a “default” pathway shared by all embryos, while Y-linked genes can act as developmental “difference makers.” However, these genes are not master switches; they depend on complex interactions with many other molecular systems.
Prum also compares mammalian and avian sex systems to show that sex determination mechanisms are evolutionarily variable and historically contingent.
Quotes:
- “Sex is an historical constellation, or cluster, of iteratively realized, co-occurring anatomical and physiological homologies that have evolved over evolutionary history through selection for sexual reproduction (see chapter 2).”
- “However, a bimodal distribution in reproductive anatomy and physiology within species does not justify the existence or assignment of an inherent, individual sexual essence or identity to the zygote or genome prior to, or independent of, the sexual phenotype's material realization.”
-
# Chapter 6: Variations in Our Sexual Development
This chapter focuses on intersex and variations in sexual development. Prum argues that developmental diversity demonstrates the absence of an essential binary sex system.
Human sexual development is strongly canalized toward bimodal outcomes, but the complexity of developmental pathways creates many intermediate possibilities. Traditional scientific and medical frameworks pathologized these variations because they assumed a rigid binary that biology itself does not support.
Prum reframes intersex variation not as developmental failure but as evidence that sexual development is inherently variable and contingent.
Quotes:
- “In other words, sex is not an innate property of individual genomes or embryos waiting to be represented in the material world, but a suite of reproductive possibili ties realized, or enacted, by those individuals through development. The sexual phenotype is an active, assertive developmental achievement by the organism. This reframing is critical to understanding the substantial variability in human sexual development. Anatomical variation in human sexual anatomy is strongly, bimodally distributed-that is, the vast majority of human individuals develop the anatomically and physiological capacities to sexually reproduce either through the production and delivery of sperm, or through the production of eggs, pregnancy, lactation, and so on. This bimodal distribution is evidence of the strong canalization (i.e., evolutionary reduction of variation) of sexual development, but it is not sufficient to imply, support, or create an essential (i.e., inherent and indispensable) scientific fact about the sex of a genome, a zygote, or its phenotypic fate.”
- “Rather than regard differences in sexual development as pathological failures of a binary sexual development, we should realize that the developmental variations in the sexual body are actually evidence that the individual sexual binary does not exist in the first place.”
-
# Chapter 7: How Evolution Generates Sexual Variability
This chapter expands the argument evolutionarily, showing that sex determination systems differ radically across species. Sexual differentiation mechanisms are evolutionarily unstable and constantly changing.
Prum argues that this diversity proves there is no universal biological essence of sex. Instead, evolution repeatedly generates new ways of realizing reproductive possibilities. He also discusses placental evolution and immune interactions during pregnancy to show how reproductive biology emerges through evolutionary rewiring.
---
# Chapter 8: The Future of Performative Biology
This chapter explores future implications for developmental biology, medicine, and reproductive theory. Prum examines pregnancy as a performative interaction between gestating parent and embryo, arguing that gestational environments actively shape development.
He proposes hypotheses about why gestating mammals are homogametic (XX) and links performative biology to medical understanding of infertility and pregnancy complications.
The chapter concludes by redefining illness itself as a failed or disrupted self-performance rather than simply mechanical breakdown.
“Thinking about illness and disease as performative mis-realizations of the self…focuses our attention on the fact that there are indeed a greater variety of ways to go wrong than right.”
-
# Chapter 9: Performance All the Way Up
The final chapter broadens performativity beyond biology into culture and communication. Human culture emerges from deeper biological performative systems rather than standing apart from them.
Prum highlights examples of animal culture, especially birds and whales, where vocal and social behaviors are learned and transmitted across generations. He argues that gender and sexual culture are among the earliest and most transformative forms of culture in evolutionary history.
The chapter closes the loop on the book’s central argument: performativity operates continuously from molecules to cells to organisms to societies.
June 18, 2024
This book fundamentally changed my understanding of myself. Performance All The Way Down seeks to establish a practice and model of biology based in the feminist and philosophist concept of performativity and the acknowledgement of agency on the level of every ontological individual. Its thesis deconstructs a wider cultural and scientific ideal of individual sex as binary and deterministic. It argues against the supremacy of genes and chromosomes and master controllers of sex expression and against the flattening of biological processes like development into segmented ideas of phenotype, endocrine, etc Sex Difference, in favor of an understanding of sexual variation in all its forms as a sort of ecology, a web of inter- and intra- actions. This book posits selfhood as a performative act, the act of being actually as the act of becoming. The self is endlessly reiterative all the way down, each cell performing the work of itself all the way up to me, as an individual, performing the work of becoming myself. Sex is not an essence, a quality that the body has before observation, a deterministic quality that can sort animals definitely, or an intrinsic and unchanging trait of an individual. It is an act, a doing and becoming that is part of the body's life's work. I now understand myself and my body as endlessly becoming and performing myself, my transsexual being as part of the myriad of the endless potentiality of a self that doesn't just change but its, itself change.
March 19, 2026
this book is fascinating. writing my review while eating rly spicy salsa so i feel crazy. this book also made me feel crazy as in very alive and full of cells and tissues and ideas in flux. also i do not often read books over a period of 13 months so there is something special about ruminating on similar ideas throughout a phase of great personal and bodily change. would recommend for people with a variety of levels of both developmental biological knowledge and gender theory fluency. however it will take some time maybe not necessarily 13 months but a while. alright goodbye everyone also my entire face is still burning. #spicy and delicious mystery sauce
December 27, 2024
I was very excited to read this book after my science book club chose it. I am super interested in learning more about biological sex, determinism, differences in presentation for sex, and how this affects gender and our society. I will say that Prum references many interesting and thought-provoking studies and this was hands down my favorite part of the book. Prum dances around a lot of ideas but I am not sure he ever comes to any actual points. I was super bothered by the word performative and its use in this book. I know it is likely that it is some extra meaning that I am putting on the word but I struggled to get past that. I also struggled with Prum’s use of the words gender and sex interchangeably. Prum states at the beginning of the book that he wants to make science accessible to social science types and the social science accessible to science types and honestly, I don’t think that he accomplished either of these things. There were so many times I had to go look things up to try to understand the social science. As someone comfortable with developmental and evolutionary biology I found myself backing up to reread sections and feeling like he didn’t explain things well. He didn’t even spend the time to define basic terms. In the last chapter of the book, Prum goes completely off the rails talking about how science shouldn’t bother to do any research based on sex because it is useless after he just spent an entire book talking about how sex is a spectrum and we need to change how we think about it. Mostly I left the book feeling extremely confused by the point he was trying to make and frustrated by his inability to make anything clear.
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