Special Section
Daphnia and Apollo: An Epigenetic Fable
San Francisco State University
mkenney@sfsu.edu
Abstract
Environmental epigenetics is a field of molecular biology that studies how signals from the environment (e.g., food, toxicants, and even our social milieu) affect gene expression. Although, for some, environmental epigenetics promises a new and dynamic account of the relationship between organisms and their environments, current research using model organisms often relies on stereotypical assumptions about gender, race, class, and sexuality in humans (Kenney and Müller 2017). In this article, rather than simply critiquing dominant narratives, I tell a different story—a feminist fable about small crustaceans called Daphnia, who display remarkable epigenetic responses to their environments. For example, many species of Daphnia are made up of clonal females who reproduce asexually, but when resources are scarce they produce males and practice sexual reproduction. In the presence of predators, they grow helmets and neckteeth, passing these adaptations along to their clonal daughters—a potentially epigenetic inheritance. Working within the metaphoric tissue of technoscience, I bring recent research on Daphnia epigenetics together with the story of Daphne and Apollo from Ovid’s Metamorphoses in order to activate new possibilities for understanding the relations between humans and animals, organisms and their environments.
Keywords
environmental epigenetics, chemical ecologies, critical fabulation, model organisms, metaphors, comparison
Introduction
Donna Haraway describes technoscience as “a practice of materializing refigurations of what counts as nature, a practice of turning tropes into worlds” (1994, 60). She argues that it is imperative that we contest visions of nature that are predicated on dominant values such as “military combat, sexual domination, security maintenance, and market strategy” (60–61). Feminist science studies scholars have a long history of challenging the narratives and tropes of scientific inquiry, especially when they harbor harmful norms and stereotypes about gender, race, sexuality, disability, immigration status, and other categories of difference (e.g., Haraway 1989; Martin 1991; Subramaniam 2014; Willey 2016). However, the purpose of this critique has always been intimately linked to another project—the project of creating a feminist lexicon for a “successor science” (Harding 1986), generating more promising tropes for alternative knowledge-making practices (see, e.g., Hustak and Myers 2012; Kenney 2019). “Queering what counts as nature,” Haraway writes, “is my categorical imperative” (1994, 60).
In this article, I take up this intellectual and political tradition as I consider the stories emerging from the new field of environmental epigenetics. Rather than simply critiquing dominant narratives in the field, I work within the metaphoric tissue of technoscience to tell a feminist fable about Daphnia—small crustaceans who display remarkable epigenetic responses to their environments. I bring recent research on Daphnia epigenetics together with the story of Daphne and Apollo from Ovid’s Metamorphoses in order to activate new possibilities for understanding the relations between humans and animals, organisms and their environments. Throughout, I pay special attention to the role of metaphor and analogy in model organism experiments, offering the technique of “speculative comparison” as a theoretical incitement to interrupt dominant epigenetic narratives and imagine more radical ecological futures. I begin the article with a discussion of dominant epigenetic narratives, followed by a description of “speculative comparison,” before introducing the Daphnia and their lifeworlds. Overall, this article is an exploratory attempt to use the practice of storytelling to open social, scientific, and political questions about environmental epigenetics and multiply the possibilities for figuring life in the postgenomic era.
The Politics of Environmental Epigenetics
In recent years, research from the burgeoning field of environmental epigenetics has captured the imaginations of STS scholars concerned with creating more complex, socially embedded, and justice-oriented accounts of human health and biology. Environmental epigenetics is a branch of molecular biology that studies how signals from the environment affect gene expression via mechanisms such as DNA methylation, histone modification, and chromatin remodeling. In humans, specific environmental stimuli such as foods, toxicants, and social experiences are believed to have epigenetic effects, which contribute to health and well-being throughout the life course. For feminist STS scholars, environmental epigenetics beckons because it offers molecular explanations for how social and environmental exposures become embodied—exposures that are often uneven and socially stratified (e.g., Bullard 2000).
For example, environmental epigenetics has the potential to explain how experiences of trauma, deprivation, and oppression get under the skin, causing harm to our physical and mental health. Those who have experienced violence and oppression know that trauma has corporeal effects; Claudia Rankine, for example, describes how everyday experiences of racism take up residence inside bodies: “You can’t put the past behind you. It’s buried in you; it’s turned your flesh into its own cupboard” (2014, 63). Whereas the genetic determinism of the era of the Human Genome Project relegates these experiences to the realm of the social or psychological, research in environmental epigenetics recasts these experiences as biosocial, capable of affecting human health and well-being in significant ways. Cohort studies (Yehuda et al. 2005; Heijmans et al. 2008; Lehrner et al. 2014) support the hypothesis that trauma leaves molecular traces—on bodies, in utero, and potentially across generations—that modulate our gene expression, offering scientific validation for our innate awareness that bodies remember the past and that trauma can be passed down from our parents and grandparents.
Feminist STS scholars have turned to environmental epigenetics not only to account for the biosocial effects of structural and interpersonal violence but also to theorize the relationship between organisms and their environments.1 As biologist and feminist science studies scholar Lisa Weasel argues, “the science of epigenetics holds…revolutionary implications for re-imagining the relationships entwined and emerging out of naturecultures, and the political potentials of such” (2016, 109). Environmental epigenetics appears to confirm many of the feminist critiques of genetic determinism where DNA is cast as the master molecule and the blueprint of life (see, e.g., Keller 2000; Fausto-Sterling 2004; Lee 2011), while offering more complex accounts of embodiment that contextualize our bodies in relation to our social and material environments. In interviews, molecular biologists sometimes read like Deleuzian feminists as they draw out the radical ontological implications of epigenetics research. UCLA professor of medicine Steve Cole, for example, uses epigenetics research to reconceive the human as a radically porous organism:
We think of our bodies as stable biological structures that live in the world but are fundamentally separate from it. That we are unitary organisms in the world but passing through it. But what we’re learning from the molecular processes that actually keep our bodies running is that we’re far more fluid than we realize, and the world passes through us. (Cole quoted in Dobbs [2013] 2017)
Cole offers a vision of biology where the relations between inside and outside, organism and environment are fundamentally reworked. The world passes through us and we are not unchanged. This is a vision of molecular biology that seems amenable to the relational ontologies we see in recent work in new feminist materialism (Alaimo and Heckman 2008; Coole and Frost 2010; Van der Tuin and Dolphijn 2012; Pitts-Taylor 2016): to speak of worlds and bodies as vital, and in flux. We are far more fluid than we realize. These epigenetic imaginaries have the potential to account for lively and deadly encounters between molecules and tissue, how bodies are made in and through these encounters, often in surprising ways. As Brian Dias, assistant professor of pediatrics at the University of Southern California, attests, “If science has taught me anything, it is not to discount the myriad ways of being and becoming” (Dias quoted in Hughes 2014).
However, despite the ontological and political promise of environmental epigenetics, we know that no knowledge project is inherently emancipatory. Feminist science studies scholars have also found harmful assumptions about gender, race, sexuality, and class in the scientific and popular literature on this research. Specifically, because environmental epigenetics researchers often look at gestation and early life as key windows of developmental plasticity, mothers have become a central focus of epigenetics research and proposed public health interventions (e.g., Richardson 2014; Richardson et al. 2014; Lappé 2016; Kenney and Müller 2017; Valdez 2018).
In “Of Rats and Women: Narratives of Motherhood in Environmental Epigenetics,” Ruth Müller and I analyze peer-reviewed journal articles and media coverage of a series of influential studies at McGill University on the epigenetic effects of maternal care in rats. The McGill group found that the amount a mother rat licks and grooms her pups regulates the expression of a glucocorticoid receptor gene in the pups, which affects stress response throughout their lives. Pups who were licked and groomed frequently by their mothers grow up to be calm adult rats, whereas pups who were licked and groomed less grow up to be anxious and difficult for researchers to handle. While these results compellingly demonstrate that the social environment has biological consequences, the McGill experiments focus solely on the behavior of the mother to the exclusion of all other environmental factors. In accounts of this research, the mother is figured paradoxically as both responsible for the health of her offspring and simultaneously having very little agency, as her behavior is also believed to be influenced by her environment. Ultimately, this explanatory framework results in a new kind of environmental determinism, where mothers’ bodies become “vectors of developmental or epidemiological risk” (Richardson 2014, 227; see also Waggoner and Uller 2015).
Much of the research on the epigenetic effects of maternal care is conducted using model organisms such as mice and rats. While model organism experiments can offer insight into biological mechanisms shared across species (Nelson 2018), there are important differences between rodents and humans that researchers often fail to highlight in publications and press releases. For example, rats (unlike humans) do not practice bi-parental care and their cage environment is deliberately simplified to exclude any other environmental stimuli that could affect their development. Despite obvious differences, these research findings are translated into the human context, with articles that demand parents “Cuddle Your Kid” (Kristof 2012), or else you could be responsible for a lifetime of anxiety and poor health. While researchers often blame the media for these hasty translations, we found that the peer-reviewed literature also moves between model organisms and humans without considering the social, political, and epistemological implications of these comparisons (Kenney and Müller 2017).
For example, Steve Cole, who said that “we are far more fluid that we realize,” contributed to a peer-reviewed study on the biological effects of maternal care entitled “Maternal Warmth Buffers the Effects of Low Early-Life Socioeconomic Status on Pro-inflammatory Signaling in Adulthood” (Chen et al. 2011). In this article, the authors use the McGill group’s model organism experiments as the basis for framing their hypothesis about health disparities in humans. They hypothesize that if licking and grooming has a protective effect for the rat pups, it is possible that “maternal warmth” in humans can buffer against the biological effects of living in poverty. They test this hypothesis by measuring systemic inflammation in people who grew up in low-socioeconomic status households, dividing the study population into two groups: those who self-reported experiencing low maternal warmth and those who self-reported experiencing high maternal warmth. They found that the high maternal warmth group exhibited less systemic inflammation than the low maternal warmth group—supporting their hypothesis that maternal warmth buffers against the negative health effects of poverty.
Here, the narrow focus on the mother and the stereotypical feminine quality of “warmth” emphasizes maternal responsibility to the exclusion of all other social and environmental factors. Although this study was conducted on human subjects, the legacy of the model organism experiments remains; despite the fact that human children are often cared for by people other than mothers (fathers, queer, trans, and non-binary parents, aunties, grandparents, childcare workers, etc.) “maternal warmth” remains the salient category, which significantly shapes the article’s recommendations. Although the researchers acknowledge that health disparities are caused by economic disparities, the mother nevertheless becomes the site of intervention. They write:
Working to alleviate poverty, as lofty and important a goal as this is, has remained an intractable problem in our society. Complementing this effort, encouraging and teaching parenting behaviors that facilitate warm emotional climates, even in the face of adversity, might prove to be a supporting, effective target of intervention (as suggested by cross-fostering and environmental manipulation studies in previous animal research). (Chen et al. 2011, 735)
In recommending parenting classes rather than economic policy, Chen et al. participate in a long history of ineffective and condescending social policies that disproportionally target poor women of color. In the 1980s and 1990s, Black feminists such as Angela Davis (1990), Patricia Hill Collins (1990), and Dorothy Roberts (1997) effectively critiqued the legacy of US policy interventions that fail to address the structural effects of racism and poverty on the lives of Black people, families, and communities. Angela Davis and Fania Davis write:
While the difficulties besetting the family should by no means be dismissed, any strategies intended to alleviate the prevailing problems among poor Black people that methodologically target the family for change and leave the socioeconomic conditions perpetuating Black unemployment and poverty intact are doomed to failure at the outset. (1990, 81)
While Chen et al. state that their goal is to improve the health of children in low-socioeconomic status households, by framing poverty as an intractable problem, mothers again become the target of public health interventions while the conditions that produce health disparities remain intact.2 Environmental epigenetics has the potential to strengthen social justice claims by explicating the biological mechanisms by which racism and poverty affect health and well-being; however, when researchers ask a narrow set of questions about bodies and environments, they can reinforce harmful assumptions about gender, race, class, and sexuality and offer policy recommendations that pathologize those they desire to help. Here we see how the ontologically imaginative proposition that “we are far more fluid than we realize” can quickly transform into racist calls for parenting classes that “facilitate warm emotional climates, even in the face of adversity.”
The narrow policy recommendations in Chen et al. demonstrates both the need for interdisciplinary inquiry—for example, Black feminist theory should be required reading for molecular biologists—and the need to take care when translating model organism experiments into policy recommendations. Claims about the epigenetics of maternal care are buttressed by analogies that are made too quickly between rats and humans, between the lab and the world. Differences are collapsed in sameness. Rats in highly controlled cage environments become proxies for humans in complex material and social environments, resulting in stories that overemphasize maternal responsibility and narrate kinship only through pathology. However, rather than continue to highlight and critique these hidden translations, in this article I offer a different approach to the politics of epigenetic storytelling—an approach that begins with a technique I call “speculative comparison.”
Speculative Comparison
Speculative comparison is designed to intervene in the politics of comparability by calling naturalized analogies into question and to open up different possibilities for thinking and doing relations. An analogy makes a comparative claim: “A is like B.” A speculative comparison, on the other hand, asks, “How is A like or unlike B”? In turning assertions into questions, speculative comparisons are “occasions for theorizing; occasions for telling differences and samenesses in new ways” (Verran 2002, 729). When we ask the question how is A like or unlike B, there is not a right or wrong answer, but many possible avenues of narration, imagination, and empirical investigation. Speculative comparison is particularly good for teaching or thinking together in a group; through collective brainstorming you quickly find that some ways of telling samenesses and differences conflict with one another, that some comparisons are more generative that others, and that different analogies do different epistemological, pedagogical, and political work.
For example, in the class Science, Technology, and Sexuality, my MA students and I used speculative comparison to analyze a public health campaign for PrEP (pre-exposure prophylaxis drugs for the prevention of HIV/AIDS) that targeted gay men in San Francisco. I presented them with an ad that depicts a group of multiracial gay men dancing in a club superimposed with the phrase “It’s our sexual revolution.” To explore the politics of sexuality, risk, and pharmaceuticals at work in the campaign, we asked, “How is PrEP like or unlike ‘the pill’?” The poster asks the passerby to draw an analogy between (A) the so-called (heterosexual) sexual revolution of the 1960s catalyzed, in part, by the advent of hormonal birth control, and (B) the novel use of PrEP to prevent HIV transmission among gay men. The public health campaign uses this analogy—A is like B—to present PrEP as a pharmaceutical that offers personal sexual empowerment by reducing risk. However, as we discussed in class, risk of HIV transmission and risk of pregnancy are materially, socially, and historically different. Exploring the unstated differences between HIV and pregnancy led to an important class discussion about the history of HIV/AIDS activism, social determinants of health, and the ongoing role of homophobia, racism, and stigma in the HIV/AIDS epidemic. As we increasingly politicized HIV/AIDS risk, we also began to reframe hormonal birth control in the context of reproductive justice and efforts to end population control, rather than through the ideologies of choice, empowerment, sexual freedom, and reproductive rights. Tacking back and forth between PrEP and the pill, asking how A was like or unlike B, allowed us to critique the framework of the public health campaign and explore different ways of telling stories about these two classes of pharmaceuticals. Of course, a speculative comparison exercise such as this one requires skills in intersectional and social justice storytelling—however, the activity of making and unmaking analogies helps us in these efforts by destabilizing what we take for granted and creating opportunities for alternative stories to emerge.
Speculative comparison challenges us, much like science fiction, to make the familiar strange and, in doing so, open new ways of approaching the politics of everyday life. This kind of comparison is speculative rather than definitive, because it involves experimenting with relations rather than making ontological claims. It figures comparison as a practice, rather than as a given, in a way that draws attention to what comparisons do as they move through the world. The speculative, in Isabelle Stengers’s terms, is about activating possibilities, generating “narratives that populate our worlds and imaginations in different ways” (2007, 9). For example, Anna Tsing uses a speculative comparison between humans and mushrooms to teach us to see mushrooms in a new way: “Like an inside-out stomach, fungi secrete enzymes into the soil around them, digesting organic material and even rocks, and absorbing nutrients released in the process” (2010, 191). Comparing two organisms that seem very different, turning one of them inside out, Tsing gives us a far-fetched simile designed to stimulate our imaginations and draw us into the life worlds of mushrooms. Tsing gives us comparison as sf worlding (Haraway 2016, 14), where sf stands for a multiplicity of speculative practices: “science fiction, speculative fabulation, string figures, speculative feminism, science fact, so far ” (Haraway 2016, 2). Tsing’s sf worlding uses analogy as invitation to imagine and inhabit worlds otherwise.
By contrast, the research on the epigenetic effects of maternal care relies on a literalized analogy: rats are like humans. The purpose of using model organisms is to generate situated comparability between species, to provide insights that can be generalized or extrapolated to the biology of other organisms, especially humans in the context of biomedical research. However, in the analogy between rats and humans in the case of the McGill experiments, the comparison has been asserted rather than investigated. It relies on the assumption that we will recognize a human tenderness in the licking and grooming of the mother rats and not stop to ask if it is relevant that humans tend not to lick their young. In Primate Visions (1989), Haraway argues that when researchers seek answers to social questions with model organism experiments they translate “metaphor into hardware” (231–43), materializing contemporary biopolitical questions in the experimental design—the cages, the experimental protocols, the enrichments, the stressors, the deprivations. The results of these experiments then circulate as if they provide direct insight into the epigenetic effects of maternal care across species.
One way to destabilize the naturalized analogy produced by these experiments would be to ask which samenesses and differences between species matter here. There are serious scientific questions we could ask, such as whether social structures and molecular pathways are similar in rats and humans, and how so? Biopolitical questions about whether the lack of aggressiveness in rats is an indicator of mental health, or if instead, docility is privileged by the laboratory workers, not the rats themselves. Or more peculiar questions like, if the human lab technicians licked the rat pups with their own tongues, would the pups grow up to be calm adults? However, in the remainder of this article, I will leave the rats behind, to take up a very different model organism that is currently being used for epigenetics research: small crustaceans called Daphnia, planktonic organisms that are barely visible to the naked eye.
Daphnia
There are over 150 species of Daphnia living in lakes, ponds, streams, swamps, pools, and rivers worldwide. Daphnia are highly responsive to the environment, transforming their bodies in response to the conditions of their habitat. For example, although most Daphnia populations are typically made up of females who reproduce asexually, in times of environmental fluctuation, such as change in temperature or scarcity of food, they lay eggs that hatch into males and then reproduce sexually, though some species rarely, if ever, switch to sexual reproduction. Cyclical parthenogenesis, switching from asexual to sexual reproduction, is an example of environmentally induced sex determination (Ah-King and Nylin 2010). Female and male Daphnia have the same chromosomes; it is the epigenetic modification of their DNA that regulates sexual difference. Daphnia also display dramatic “predator induced-polyphenisms,” growing defensive helmets, neckteeth, or tail spines when exposed to chemicals called kairomones released by nearby predators. The daughters of the armored Daphnia may also grow helmets and neckteeth even when no predators are present, demonstrating a potentially epigenetic inheritance.
In our current age of increased anthropogenic environmental change, this remarkable environmental sensitivity has made Daphnia into important organisms for ecological research and surveillance. They have become sentinel species used to monitor the health of ecosystems and waterways, studied in their habitats or used in bioassays in laboratories. Living Daphnia are used as biosensors in sophisticated water monitoring devices, such as the Daphnia Toximeter II, because their health, behavior, and morphology reliably shift in relation to the chemical make-up of their water. According to the product’s website, the Daphnia Toximeter II is a sixty-kilogram machine containing a chamber filled with Daphnia used for real-time testing of water from rivers, industrial plants, and drinking water infrastructure. The website explains, “Sample water continuously runs through the measuring chamber containing the daphnia. The live images obtained using a CCD-camera are evaluated online with an integrated PC to analyse changes in the behaviour of the daphnia. If the change is statistically significant, an alarm is triggered” (bbe moldaenke, n.d.). Changes in the Daphnia’s movement, size, distribution, or number signal the presence of a potential toxicant that could affect river ecology or the potability of drinking water. This water monitoring technology can detect minute changes in the water’s chemical make-up that are difficult to detect through other means; for example, in 2004 a Daphnia biomonitor discovered an unknown contaminant in the Meuse River in the Netherlands that took months to identify through chemical analysis (de Hoogh et al. 2006). Here in the Daphnia Toximeter II, the Daphnia’s liveliness and capacity for corporeal plasticity is captured for predicting risk to human health.3
In 2011 Daphnia pulex was the first crustacean to have its genome sequenced and was found to have the most genes of any animal sequenced so far. Studies in ecotoxicogenomics are currently investigating how Daphnia use their large complement of genes to adjust to “environmental stressors like heavy metals, rising temperatures, emerging diseases, and bioreactive organic compounds” (Ebert 2011, 540). Daphnia have also been enlisted as research subjects in the new science of “resurrection ecology,” the study of animals who have been revived from viable eggs that are decades or even centuries old to learn how their morphology has changed in response to their environments over time. The oldest animals ever brought back to life were Daphnia pulicaria, revived from dormant eggs buried for seven hundred years in lake sediment (Frisch et al. 2014). When these ancient hatchlings were compared to their contemporary sisters, they showed dramatically different strategies for storing phosphorous, which the research team believes is a response to the introduction of commercial fertilizer in the nineteenth century. They now store less phosphorous because it is readily available in the fertilizer-polluted lake water. In other words, the bodies of Daphnia pulicaria remember the industrial histories of their ecosystem.
Learning about the Daphnia and their sensitivity to environmental change, I began to wonder how Daphnia are like or unlike humans. Whereas the familial scene of the mother rat licking her pups invites comparison with humans, the Daphnia, who have clones and time travelers among their ranks, are more difficult to approach comparatively. What can we learn from the Daphnia and her brood of clonal daughters? What can we learn about reproduction, sexuality, and motherhood from these cyclomorphic crustaceans, who have one of the largest animal genomes but no tongues with which to lick their young?
Developmental biologist Scott Gilbert gives us one possibility for comparing humans and Daphnia in his article “The Genome in Its Ecological Context” (2002). After detailing how Daphnia develop helmets and neckteeth in response to predators, he writes, “Humans have specific predator-induced developmental plasticity on a scale unimaginable in invertebrates or amphibians” (Gilbert 2002, 201). The predators Gilbert invokes are not apex predators like lions or sharks, but microbes. Gilbert explains that our immune system produces specific antibodies depending upon which bacteria and viruses we have previously been exposed to. He notes that identical twins do not have identical immune systems, since their phenotypes develop in relation to their respective environments. Gilbert explains that “we have also evolved nonspecific defenses wherein bacterial components activate genes inside our body that produce nonspecific defensive substances” (210). Here he is referring to enzymes that are secreted by our intestines as a first line of defense against bacteria. Note Gilbert’s strange and wonderful use of language. In this article humans are creatures with “predator-induced polyphenisms” and “nonspecific defense fluids.” These unfamiliar but alluring terms have migrated from the ecologically sensitive world of the Daphnia into the human realm to give us a stranger hominid.
Through speculative comparison, we can indulge our curiosity about our own ecological sensitivity and ask what we might learn about biological agency, phenotypic plasticity, and ecological resilience from these tiny crustaceans who do not elicit easy mammalian identification. Like the Daphnia changing in response to their fertilizer-polluted waters, we are also alterlife, “life already altered, which is also life open to alteration” (Murphy 2017, 498). Telling stories about organisms like Daphnia can help us push against fantasies of ecological purity (Shotwell 2016) and biosecuritization (Chen 2011; Ahuja 2016) to “meet the future organisms we are becoming” (Ah-King and Hayward 2014, 9). Wondering about how the Daphnia are like or unlike humans, I am compelled to write feminist theory as “critical fabulation” (Hartman 2018, 470), “attend[ing] lovingly to stories” (Haraway 2004, 201) as a practice of enacting better relations with a more-than-human world.
Daphne and Apollo
As I write these epigenetic fables about the Daphnia, their name entices me into a second speculative comparison, more mythological than biological, a comparison that I will pursue for the remainder of this article. They are called Daphnia after Daphne, the nymph who transforms into a laurel tree in Ovid’s Metamorphoses. The Metamorphoses is a Latin poem written in dactylic hexameter published in the year 8 CE. It begins, “My soul is wrought to sing of forms transformed to bodies new and strange,” and through 15 books goes on to tell 187 stories of transformation, where characters are turned into swans, and stones, and constellations, rivers, and plants and cicadas. We meet Daphne in Book 1 of the Metamorphoses. We learn that she is not interested in marriage but instead takes delight in wandering the woods, wearing the skins of animals she hunts. Apollo, the god of poetry and medicine, sees her, falls in love, and runs after her. She flees, uninterested in Apollo, uninterested in participating in the heterosexual economy, scared of what he might do if she stops running. He continues to chase her—like a hound chases a rabbit, Ovid tells us. As he’s about to catch her, Daphne prays to her father Peneus, a river god, and she’s instantly changed into a laurel tree. A predator-induced polyphenism. A morphological adaptation she will pass on to her leafy daughters, who bear her name. Daphne is the Greek word for laurel tree.
And indeed, the crustaceans are called Daphnia because of the branch-like antennae they use to smell and to move around in the water. The analogy between crustacean and nymph was originally made in 1785 by naturalist Otto Friedrich Müller and has laid there since, hidden in plain sight. As we learned from Emily Martin (1991), scientific language is replete with calcified metaphor, with signifiers lying dormant, waiting to signify again. With this speculative comparison, I am interested in performing my own resurrection ecology, wresting back some of the poetry, animating the analogy, awakening the dormant signifiers. A re-laureated science. In honor of Daphne.
I begin with this photograph of Gian Lorenzo Bernini’s seventeenth-century sculpture group Apollo and Daphne that stands in the Galleria Borghese in Rome (Figure 1). Taken in the dark, the flash illuminates the marble, capturing the instantaneousness of the transformation. I perform a reading of this photograph, rather than the sculpture itself, because for me, the photograph takes the eternal moment preserved in marble and gives it back its urgency. The story of Daphne and Apollo, made less threatening by familiarity and historical distance, here regains its terror and implicates the viewer.
Figure 1. Bernini, Apollo and Daphne, 1622–1624. Photo: Carol Yoho. 2010.
This is how I imagine it: A chase, moving from the glow of a campfire where Apollo glimpsed Daphne, now running headlong into the darkness. It is all happening very quickly, bodies moving recklessly through the forest, indistinct voices, tree branches snapping. But unmistakably a chase. Daphne then Apollo then the camera. Not a camera camera, but an iPhone, hastily pulled from a back pocket. Apollo screams into the darkness, again and again, half pleading half menacing. Then Daphne’s voice; a fearful prayer. The phone flashes. Too brief to tell what is happening. The moment of transformation caught digitally; but only by chance. Nymph in the dark /flash/ tree in the dark. The metamorphosis becomes terrifyingly clear only on the screen. “Holy shit.” Her hair. Her hands. Leaves where there was flesh. Fingers receding, branches forking and twisting up towards the moonless sky. The photographer is implicated in the scene. Whether complicit in the assault or trying in vain to stop it, or just caught in the commotion attempting to document something. He is left with, we are left with, this image and its obligations. (Haraway 1991, 192).
What this photograph captures for me is this question of how to account for the metamorphosis in the aftermath and how to understand the violence that can accompany feats of radical plasticity. Although it was Daphne’s desire, or rather, her desiring otherwise, that catalyzed her transformation, it is difficult to know how Daphne feels about being a tree—the poem refuses to say. Does it even make sense to speak of Daphne any longer at all or has being nymph already given way to arboreal sensations and desires? Apollo copes with this lacuna through appropriation: “If you cannot be my wife,” he says, “you shall be my tree.” Apollo takes the laurel as his own. A symbolic violence. Her burgeoning liveliness appropriated by her pursuer. This is why Apollo always wears laurels on his temples and why we have poet laureates. Apollo (and by extension Ovid’s narrator) would like to believe that Daphne’s consent can be read in the movement of the laurel’s branches. But this is a belated analogy. This tree is Daphne, this tree is still like Daphne, this tree at least used to be Daphne, right? We do not know if Daphne gives her consent; she only seems to nod with her leafy crown. As comparative literature scholar Heather James writes, “Ovid’s tale dramatizes the ambiguity of signs and exposes the motives for variable readings: power and passion affix meanings to words, bodies, and gestures” (2007, 46).
As a modern reader of the Metamorphoses, the ambiguity of signs and the ambivalence of transformation are two themes that stand out for me. Although Roman literature has been appropriated by the dominant historical narrative about the continuity of “Western civilization,” the stories that Ovid tells in the Metamorphoses are two thousand years strange and their moral universe is populated with unfamiliar stars. Ovid makes it difficult to classify Daphne’s transformation as triumph or tragedy. She is safe from Apollo; but her swift feet are now stuck fast by slow growing roots. Since her voice is gone, her subjectivity is lost to us. Like Apollo we can only search the poem and the sculpture for signs. We are all just holding up our iPhones in the dark. However, this moment of uncertainty might also spark a feeling of hope. The ambivalence of the transformation gives ways to polyvalence. A host of stories churn below the surface. Which story can do justice to this metamorphosis? What does it mean to honor Daphne?
Daphnia and Apollo
After this reading of Daphne and Apollo, I bring these questions back to bear on the small translucent crustaceans—a new model organism for the postgenomic age. What story can do justice to their epigenetic metamorphosis? What could it mean to honor the Daphnia? On the one hand, it’s possible to tell a determinist story about their bodies optimally adapting to their environment, a story of an evolutionary arms race in a mechanistic nature that is red in tooth and claw. However, Ovid’s story of Daphne becoming tree opens up new narrative possibilities for the Daphnia’s becoming armored. We might think of their predator-induced polyphenisms not as an epigenetic program determined by natural selection and triggered by their environment, but as a pattern of ongoing signals and responses, where what counts as a signal can only be determined in the context of response. A kairomone is only a kairomone because it induces a helmet. In this way, Daphnia grow helmets and neckteeth to protect themselves and their daughters from predators they have encountered for hundreds of thousands of years, in ongoing epigenetic metamorphosis. But like Daphne, they have also changed quite quickly in response to the dramatic and unprecedented increases in chemicals like phosphorous. Here, we might think of the Daphnia as organisms “learning how to attune their sensory bodies to the ongoing improvisational rhythm of differences that make up the world” (Hustak and Myers 2012, 105). When we imagine “species are sensuous responses” (Hayward 2010, 233), we can begin to see how the environmental sensitivity that makes the Daphnia vulnerable to toxicity and other environmental change is also a source of ecological resilience and transformation.
It is this capacity to attune to their environment that is captured by the Daphnia Toximeter II—a cyborg device, simultaneously animal and machine (Haraway 1991). Although changes in behavior and morphology have meaning and function in the lifeworlds of the Daphnia themselves, here, like in many bioassay techniques and model organism research (see, e.g., Hayden 2003; Nelson 2018), their value lies in their ability to predict harm to humans. As Apollo appropriates Daphne’s transformation for himself, so too, can researchers use model organism as stand-ins for humans, rather than beings in and of themselves. Here they are figured as enough like humans to be effective sentinels, but different enough to avoid ethical consideration (see Hayden 2003, 201). As we unmake the dominant comparative practices of technoscience through speculative comparison, it is important to ask whether the purpose of epigenetic knowledge is surveillance, control, and the manipulation of life or whether other values, such as consent and reciprocity, are actively being centered (Simpson 2014; Benjamin 2016; Liboiron 2021).
Although Daphnia’s patterns of ecological response are reliable enough for their bodies to be made into chemical detectors in sophisticated water monitoring devices, the Daphnia continue to surprise. For example, evolutionary biologists believe that sexually reproductive species of animals are always more successful than asexual species, not only because of the genetic diversity created through recombination but because asexual species are believed to build up deleterious mutations in their DNA, which threaten their survival, a theory known as Müller’s Ratchet (see, e.g., Roughgarden 2009, 77–78). These long-standing evolutionary theories have been challenged by an asexual population of Daphnia that was found to be successfully displacing populations that cycle between sexual and asexual reproduction on the African continent. The article that reports these findings—“Invasion of an Asexual American Water Flea Clone throughout Africa and Rapid Displacement of a Native Sibling Species” in Proceedings of the Royal Society B (Mergeay, Verschuren, and De Meester 2006)—is thick with troubling associations: ecological concern is expressed through alien invasion, colonialism, sororicide, and a reproductive heterosexual panic about being unseated by aggressive clones. The article tells the story of a North American asexual subspecies of Daphnia—a Daphnia pulex / Daphnia pulicaria hybrid—that was introduced to Lake Naivasha in Kenya in the 1920s and has since completely displaced the Daphnia pulex subspecies that previously lived in the lake—a species that switched between sexual and asexual reproduction. The article continuously refers to the North American subspecies as an “invading clone” (2841) or “asexual pest species” (2842)—a monolithic colonizing force that is stripping the lake of its natural biodiversity.4
As Banu Subramaniam points out in Ghost Stories for Darwin (2014), this kind of xenophobic rhetoric is pervasive in invasion biology. Rather than focus on habitat degradation caused by colonialism, imperialism, and global capitalism, ecologists often pathologize invasive species as intrinsically harmful, in need of policing and eradicating. “Invasion of an Asexual American Water Flea” traffics in the moral economy of invasion biology where native plants and animals are under constant threat from invasive, alien, and exotic species. Referring to the Daphnia pejoratively as “water fleas” adds to this effect—stripping them of their mythological mantle and insisting on referring to them only through a crude analogy with a common pest, like Woody Allen calling pigeons “rats with wings.”
The assumption at the heart of Mergeay, Verschuren, and De Meester (2006) is that clonal species ought to be less evolutionarily successful than those that practice sexual reproduction. They argue that “sexual recombination allows rapid spread of favourable genetic mutations through populations, maintaining recurrent investment in sexual reproduction promotes adaptation to new habitat conditions” (2842), thus conferring an evolutionary advantage when compared to a subspecies that shares a single genome. The authors continuously express their surprise that the genetically diverse native population of Daphnia could be “outcompeted by a single asexual genotype with intrinsically lower evolutionary potential” (2843). They are unable to reconcile the evidence of displacement with mainstream evolutionary theories that posit asexuality as an evolutionary dead-end. They write, “How this obligate asexual exotic species could eradicate an indigenous sibling species that optimally combines the advantages of sexual and asexual reproduction is puzzling as well as alarming” (2843). While it’s true that, in animals, all clonal species are relatively recently derived from sexual species (Roughgarden 2009, 74–75), these asexual Daphnia present an opportunity for us to reconsider the received assumptions about sexual reproduction in evolutionary theory. Sexual recombination is thought to confer evolutionary advantage because it allows species to adapt to new environments through natural selection. However, this ignores the fact that in nature “nonsexuated microorganisms literally outweigh all sexuated life” (Schaefer 2021, 530); when we center bacteria rather than animals, we can see that evolution does not inherently privilege sexual reproduction.5 Environmental epigenetics helps us to understand that asexual reproduction is not reproduction of the same, but clones, like twins, differ from one another through their dynamic relationship with their environment. It is possible that these asexual Daphnia do not require genetic diversity because they are continuously responding epigenetically to their environments.
In other cases, asexual populations of Daphnia have been known to convert previously cyclical parthenogens to asexuality. So-called contagious asexuality has been demonstrated throughout the Daphnia pulex species-complex when different populations come into contact with one another. Contagious asexuality challenges the imagined opposition between sexuality and asexuality because, “paradoxically, contagiously asexual lineages…spread asexuality through sex” (Decaestecker, De Meester, and Mergeay 2009, 302). Daphnia from asexual populations can lay eggs that grow into males, who then reproduce sexually with females from sexual populations of Daphnia. This results in offspring, 50 percent of whom are asexual females. Thus, surprisingly, as sexual recombination occurs, asexuality spreads through previously sexual populations. And yet, despite the complexity of this phenomenon, biologists continue to speak about sexuality as the norm and asexuality as the condition to be explained: “Why switch to asexuality when you can be a cyclical parthenogen?,” they ask (Decaestecker, De Meester, and Mergeay 2009, 304). These evolutionary stories that figure clones as dangerous invaders and asexuality as an evolutionary dead-end not only narrow our biopolitical imagination, but, as Angela Willey argues, they participate in the “pervasive cultural privileging of sexual relationships over other types of relationship” (2016, 72)—the same privileging of sexual relationships that threatened Daphne’s desire to turn away from marriage. I am reminded, here, of James’s reading of the Daphne and Apollo story in Ovid’s Metamorphoses. Here too, in the field of invasion biology, “power and passion affix meanings to words, bodies, and gestures.”
However, rather than seek to scrub scientific language clean of stories, of passions, of mythological and science fictional valences, this is a place where we can join in as scholars who care about the politics of life and the politics of reading, writing, and storytelling. To interrupt the dominant accounts, not only through critique but through composition. To tell stories of Daphnia cucullata growing helmets in response to threats, like Daphne becoming laurel to protect herself from Apollo. Stories of the Daphnia pulicaria with corporeal memories of their fertilizer-polluted waters, scattering their eggs in times of scarcity, the dormant embryos buried in the sediment, awaiting more plentiful times, even, perhaps, anticipating the advent of resurrection ecology, dreaming of hatchings to come. Stories of Daphnia magna conscripted into acting as a sentinel species, whose sensitivity to toxicity and ability to predict ecological harm for larger organisms has entrapped them in biomonitors that capture their vital capacities for purposes not their own, like the precogs in Phillip K. Dick’s Minority Report. Stories of contagious asexuality spreading from one population of Daphnia pulex to the next, part lesbian separatist agenda, part attack of the clones, using their epigenetic sensitivity to respond to unfamiliar environments, making a mess of our best evolutionary theories.
Of course, some of these comparisons might strike the reader as far-fetched or downright improper. I would argue, however, that the speculative mode is always a bit improper. It is not about what is but what might be. Because it is not in accordance with accepted reality, speculation therefore also feels dubious, embarrassing, and unseemly. However, risking speculative impropriety, I argue, might be important in getting ourselves unstuck from the usual patterns of biological storytelling and activate different possibilities for attending and responding within our more-than-human world. The “impoverished realm of realism” (Hartman 2019, 261; see also Mitchell 2002, 123–52) is often silent on these most vital matters. We might need the help of what I call “fables of response-ability” (Kenney 2019)—didactic, efficacious (Stengers 2008) stories that can generate and strengthen care on a damaged and profoundly unjust planet (Tsing et al. 2017).
Conclusion
In an era when our own bodies are experiencing unprecedented chemical exposures, it might be savvy to continue to wonder how Daphnia are like or unlike humans. Comparing ourselves to the Daphnia, learning about our own predator-induced polyphenisms, we can see how we are already becoming with our environments, changing in the presence of microbes we encounter. As Malin Ah-King and Eva Hayward write, “We…are living in a time of intensified exposure to toxicity where life requires reinvention (if it can) or risks extinction and disease…Nonhumans and humans are vulnerable, but also exuberant, adaptable, resilient and constantly changing in interaction with environment” (2014, 9). It is this ambivalent formulation of biological agency, phenotypic plasticity, and ecological resilience that seems like a promising response to determinist stories of mother rats programming their offspring for lives of endless stress and fear. Reading the Daphne and Apollo story through the ecologically responsive morphology of the Daphnia, we arrive at an agential and relational biology where species are made through sensuous response. A more radically and politically engaged biology where it becomes possible to ask how environmental epigenetics can become Daphne’s science, not Apollo’s. It is here that we can begin to feel the force of the proposition from epigenetics researcher Steve Cole that I invoked at the beginning this article: “we’re far more fluid than we realize, and the world passes through us.” Looking at the Daphnia, their translucent bodies pulsating under the microscope, we can begin to tell new stories of epigenetic metamorphoses, stories that can keep up with the myriad ways of being and becoming, stories that populate our worlds and imaginations in different ways.
Acknowledgements
This article benefited from feedback over many years in different settings. The author wishes to thank Donna Haraway, Scott Gilbert, Ruth Müller, Micaela Janan, Priscilla Wald, Stephanie Clare, Nicole Archer, the Department of Gender, Sexuality, and Feminist Studies at Duke University, the STS Department at the University of San Diego, the participants of the We Are Compost Not Posthuman colloquium at the University of Leiden, the participants of the Ways of Reading workshop organized by Kadist and e-flux, her colleagues at the Women and Gender Studies Department at San Francisco State University, and the editors and reviewers at Catalyst.
Notes
1 As ecofeminist Val Plumwood (1993) has argued, dualisms such as nature/culture and organism/environment are part of a centuries-old Euro-American logic of domination. There are many human cultures, however, that do not subscribe to these dualisms, understanding the environment not as background but as animate, as having personhood, or as kin (see, e.g., Kimmerer 2002; Todd 2017). Environmental epigenetics is only one epistemology among many that we can turn to in order to unlearn these dualisms. It is important to consider why environmental epigenetics is given more authority in this regard than Indigenous and non-Western epistemologies that have long challenged this Euro-American logic (see Harding 2008; Todd 2016).
2 Furthermore, without carefully differentiating between the McGill study and their human subjects, they run the risk of making docility rather than health equity a central value in the study. Since the McGill group was unable to measure the mental health of rats directly, the rats’ aggressiveness and difficulty to work with in a laboratory setting became a stand-in for mental health. When researchers translate these results to the human world, more work must be done to show that biosocial interventions are designed to address health inequity rather than to control “unruly” racialized populations.
3 See Landecker (2010) for a discussion of the consequences of harnessing the living potential of cells for human purposes. Here the Daphnia’s potential for epigenetic change is harnessed for human purposes without their consent; they become components within human devices, defined only by their use value.
4 The Daphnia’s presence in the lake is a direct result of colonialism: “Most likely, the invading water flea clone was introduced accidentally to Lake Naivasha in 1927–1929, during stocking of largemouth bass (Micropterus salmoides) from the USA, following the suggestion by the then US President T. Roosevelt in 1910 that the sport fishing in British East Africa needed improvement” (Mergeay, Verschuren, and De Meester 2006, 2842). In this article, human politics are displaced onto the Daphnia, narrowing the stories we can tell about these organisms and their genetic/epigenetic capacities.
5 Donovan O. Schaefer writes, “From a biological standpoint, dimorphic heterosexuality, is something that some organisms do, an evolutionary detour that branched off from nonsexual reproduction—the main superhighway of life” (2021, 543). From this perspective, the success of the asexual Daphnia is not “puzzling as well as alarming,” even if it is rarer in animals than in plants or bacteria.
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Author Bio
Martha Kenney is an Associate Professor in Women and Gender Studies at San Francisco State University. Located in the tradition of feminist science studies scholar, her research examines the poetics and politics of biological storytelling. Her latest project on environmental epigenetics uses speculative fiction to interrupt dominant biological narratives and imagine more radical ecological futures.