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Dolly Mixtures: The Remaking of Genealogy / Edition 1 available in Paperback
While the creation of Dolly the sheep, the world's most famous clone, triggered an enormous amount of discussion about human cloning, in Dolly Mixtures the anthropologist Sarah Franklin looks beyond the much-rehearsed controversy to some of the other reasons why the iconic animal's birth and death were significant. Building on the work of historians and anthropologists, Franklin reveals Dolly as the embodiment of agricultural, scientific, social, and commercial histories that are, in turn, bound up with national and imperial aspirations. Dolly was the offspring of a long tradition of animal domestication, as well as the more recent histories of capital accumulation through selective breeding, and enhanced national competitiveness through the control of biocapital. Franklin traces Dolly's connections to Britain's centuries-old sheep and wool markets (which were vital to the nation's industrial revolution) and to Britain's export of animals to its colonies- particularly Australia-to expand markets and produce wealth. Moving forward in time, she explains the celebrity sheep's links to the embryonic cell lines and global bioscientific innovation of the late twentieth century and early twenty-first. In Dolly Mixtures, Franklin uses the renowned sheep as an opportunity to begin developing a critical language to identify and evaluate the reproductive possibilities that post-Dolly biology now faces, and to look back at some of the important historical formations that enabled and prefigured Dolly's creation.
About the Author:
Sarah Franklin is Professor of Social Studies of Biomedicine and Associate Director of the Bios (Centre for the study of bioscience, biomedicine,biotechnology, and society at the London School of Economics and Political Science
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Dolly MixturesTHE REMAKING OF GENEALOGY
By Sarah Franklin
Duke University PressCopyright © 2007 DUKE UNIVERSITY PRESS
All right reserved.
It is not enough simply to say that sexual reproduction has become the dominant mode of propagation among organisms. One must go further. Cross-fertilisation, either continuous or occasional, is the really successful method of multiplication everywhere. -Edward M. East and Donald F. Jones, Inbreeding and Outbreeding
I began to envisage how to advance the project that I had been thrust into, while also satisfying my own desire for original research in developmental biology. Answer: don't just add DNA to one cell embryos. Add it to plates of cultured cells, and then make embryos from the cells that had taken up the DNA most effectively. In other words, as the 1980s wore on I began to see that the future of genetic engineering in animals lay through cloning. -Ian Wilmut, Keith Campbell, and Colin Tudge, The Second Creation
Clone is a term from botany, derived from the Greek klon, for twig. It refers primarily to regeneration via cultivation and propagation, in which reproduction is asexual and regeneration is of a part from a whole-as in the creation of a new plant from acutting. Clone is both a noun and a verb, and has many figurative as well as practical usages. In all of these senses, cloning is synonymous with copying, its primary synonym. In this very general sense, it could be said that most reproduction occurs through a version of cloning to the extent that asexual reproduction, mitosis, or fission-creating multiple "copies" of an organism-is the most common, or standard, form of replication in living things-most of which are microorganisms and prokaryotes. A common distinction between reproduction and replication is the association of the former with sexual difference and the latter with more primitive copying abilities. However, a counterview would hold that microbes and bacteria, which have a famously loose ability to exchange genes, are equally recombinant to the higher organisms with their much-vaunted abilities of sexual reproduction and the capacity to produce completely new individuals. Through cloning, even a single cell can produce a limitless population, and sterile lines can be reproduced, or grown, in perpetuity. The optimum all-terrain reproductive option of combining asexual division, or cloning, with the capacity to reproduce sexually exists among most species, including vertebrates such as birds and amphibians.
Sexual reproduction is associated with the higher species because it can be seen as more complex and advanced, whereas replication by mere division is associated with less developed life-forms. The perceived advantage of sexual reproduction is that it combines replication with the production of genetic variation, or mix. Sex, in the sense of sexual reproduction, increases mix, which is seen to offer the advantage of maximizing adaptive capacity, or fitness, through variation. Mix, it could be said, adds flex-the ability to change.
In the account of making Dolly, The Second Creation (2000), Ian Wilmut argues that compared to asexual replication, sexual reproduction is expensive, dangerous, and inefficient: "Asexual reproduction is the obvious way to replicate: start with one individual, split down the middle, and then you have two, or many. By contrast sexual reproduction is a bizarre and even perverse way to replicate. Two protozoans that seek to reproduce sexually must first fuse, to produce one.... Sex, in short, is anti-replication. Replication implies that one individual divides to become two or more. But with sex, two combine to produce one" (Wilmut, Campbell, and Tudge 2000, 62). From this point of view, sexual and asexual reproduction are binary opposites: whereas asexual reproduction amplifies one cell into two, sexual reproduction constricts two cells into one. Together, the two processes offer opposite functions. Whereas asexual reproduction is efficient in terms of multiplication, sexual reproduction is a faster way to mix. The ability to combine sexual and asexual reproduction is consequently seen to offer the greatest advantage, and this appears to be the overriding pattern in the majority of species, in which the two mechanisms occur with roughly equal frequency and are often combined. Significantly, this mixed approach to sex is exactly what Wilmut and his team sought to achieve technologically through the series of experiments that led to Dolly's birth. Dolly's birth was also mixed in that she was designed to fulfill specific practical purposes, while being an experimental animal who would contribute to the understanding of basic questions in "pure" science. She was thus typically agricultural in that the project of her creation combined basic questions of genetics, or selective breeding, with commercial and industrial applications. As Wilmut writes of Dolly's significance, she is the animal model who confirms the viability of a technique that constitutes "the third player" completing a "trio of modern biotechnologies [that] taken together [take] humanity into a new age-one as significant, as time will tell, as our forebears' transition into the age of steam, of radio, or of nuclear power" (Wilmut, Campbell, and Tudge 2000, 18). Indeed, it would be cliché, but accurate, to describe Dolly as the proof of a new form of nuclear power-the power of nuclear transfer to advance the project of transgenesis that was Wilmut's overall aim. As he summarizes, the value of the trio the Dolly technique completed, by becoming its means of application, lay in the facilitation of gene transfer.
The point here is that the three technologies together-genetic engineering, genomics, and our method of cloning from cultured cells-are a very powerful combination indeed. Genetic engineering is the conceptual leader: transfer of genes from organisms to organism, and the creation of quite new genes, makes it possible in principle to build new organisms at will. Genomics provides the necessary data: knowledge of what genes to transfer-where to find them and what they do. Cloning of the kind that we have developed at Roslin and PPL [Pharmaceutical Proteins Limited] makes it possible in principle to apply all of the immense power of genetic engineering and genomics to animals. (Wilmut, Campbell, and Tudge 2000, 21)
This "powerful combination" of technologies is thus the metamix for transgenic sex, to which Dolly provided "the gilt on the gingerbread," according to Wilmut (Wilmut, Camble, and Tudge 2000, 20). She and the flock of Roslin sheep who were her contemporaries together confirmed the viability of a means of combining reproductive mechanisms that Wilmut argues "will take humanity into the age of biological control" (Wilmut, Campbell, and Tudge 2000, 24). This is the possibility "to grow animal cells in a dish, as if they were bacteria or cultured plant cells; and then transform these en masse; and then-as is already carried out with bacteria and plants-grow whole new animals from the cells that had taken up the genes most efficiently" (Wilmut, Campbell, and Tudge 2000, 20).
This metamix of sex, in which the reproductive possibilities of plants, animals, and microorganisms are conjoined with biotechnological expertise in order to grow animals from cultured transgenic cells is what offers the totipotency Wilmut describes as bringing into being a new age-"the age of biological control."
What is significant about sex after Dolly, then, is not that existing definitions and formations of it have been transformed, but that they have been, in a sense, sampled, remixed, resequenced, and provided with a novel means of amplification. Sex, which was never pure to begin with, is further hybridized through technological assistance to create a form of mixed-sex, known as the Dolly technique, somatic cell nuclear transfer (SCNT), or the second creation. Sex, in the sense of a reproductive mechanism, has been disassembled and rearranged through processes of reversal, recapacitation, switching, imitation, and transfer that allow it to be redeployed and redirected. The languages and practices of the remaking of sex that accompanied the creation of Dolly thus pose many questions for which sexual activity has become, as it were, a preliminary. That the questions her mixed-sex and multipurpose ancestry pose for normative conceptual regimes of sexual difference, biological reproduction, and animal kinds are inseparable from their ramifications in changing definitions of capital, nation, life, genealogy, and health is the main reason Dolly's mixed sex matters.
Dolly's sexual significance is also notable in the way her own ability to reproduce sexually confirms the scientific legitimacy of her vitality -by authorizing the scientific team that made her in the act of bearing her own viable offspring. In so doing, she models the technique used to create her and in a sense doubly-her own reproductive capacity confirming the viability of the technique by which she was created in a kind of variant on the progeny test (the progeny proof). As the model organism of the Dolly technique, her importance is at one level simply to embody it and to confirm its viability. Her body cells have been repeatedly analyzed to provide confirmation of her genetic provenance, and so Dolly is the technique that made her in some importantly literal senses. Technically, however, Dolly's complete viability was further confirmed by her ability to reach sexual maturity and breed naturally. This second reproductive proof confirmed both her ability to function sexually and the entirety of her sexual function-much in the same way as, but extending, the tradition in which only entire, or sexually complete, animals are allowed into official purebred lines.
The Difference Dolly Makes
Given how she was made, by merging two cells, in what sense is Dolly a clone? The word does not appear in the short letter to Nature announcing Dolly's birth, "Viable Offspring Derived from Fetal and Adult mammalian Cells" (Wilmut et al. 1997). Although Wilmut frequently refers to Dolly as a clone, he and his scientific partner Keith Campbell both acknowledge that the term is not strictly accurate. As Campbell states, "In the strict sense of the meaning, the animals produced by nuclear transfer are not true clones. Account must be taken of possible changes that occur in the genome during embryo and fetal development or while the cells are in culture.... Differences in the components of the egg cytoplasm would result in differences in the offspring. For example differences in the mitochondrial genome" (qtd. in Klotzko 2001, 10-11). In other words, clone is being used to describe Dolly in the absence of a more accurate term. She is not, technically, a clone in terms of a part regenerated from a larger whole, for her origins lie in two cells that were merged, or mixed, to make her. She was also thus not descended from one parent, but from two. She was not created through "mere" division, and she is not even genetically identical to her "clonal mother" (the Roslin term for the sheep whose nuclear DNA was used to make her). Dolly is a genetic mixture-her nuclear DNA derived from a Finn Dorset ewe, her mitochondrial and other cellular DNA provided by a Scottish Blackface. Ironically, Dolly is a clone because she is different (from other sheep)-and indeed unique (among higher mammals). She is, as Campbell points out, also the product of her environmental influences, which in her case included being cultured, passaged, incubated, frozen, thawed, recultured, biopsied, electrified, and gestated by a series of surrogate sheep before she was even born.
Dolly is referred to as a clone because the technique used to make her belongs to the scientific history of experimentation in embryology and genetics associated with cloning in a very general and imprecise way. Clone is the best way to describe Dolly more in terms of what she is not, rather than what she is: she is not an offspring of the usual method of mammalian reproduction, but of an unprecedented process of reproductive recombination. Hence, cloned signifies what sets Dolly apart, which, as we have seen, is both her path of ancestry, or origin, and her ability to embody this difference successfully, unlike all of the other attempted animals in her nonviable sibling cohort. Before she was born, the cell line that would become Dolly, and with which she shares her nuclear genetic identity, was cultured through several passages, meaning that the cells multiplied through division and were transferred, or replated, into petri dishes. This phase of Dolly's emergence is an essential component of the Dolly technique because it is during the period the cells are in culture that they can not only be multiplied but also modified through gene targeting. The fundamental key to the Dolly technique, then, is the ability to switch back and forth between sexual and asexual forms of reproduction.
The difference Dolly makes can thus be seen at several levels-which together express some of the paradoxical features of Dolly's life as a unique clone, a cloned singleton, and a sheep who was distinctive because she was "normal." Ironically, what the word clone means in relation to Dolly is that she was a unique individual because of the means by which she was created. Dolly's sex thus belongs less to a familiar economy of sexual difference but rather to a new scale of clonal difference. The difference cloning makes is not so much sexual as technical: it is a means to change sex in order to achieve specific technical goals, such as the more rapid amplification of flocks of transgenic sheep, for which somatic cell nuclear transfer, or remixed sex, is the most efficient mechanism.
The technical sense of cloning that refers to the production of an identical organism (e.g., a vertebrate) from a parent organism has an unusually controversial history in modern biology, ironically intertwined with the same taint of fraud evident in uses of the term clone to refer to illegitimate copying (e.g., Gucci clones), transgressive sexuality (gay clones), and reverse engineering ("cloned" PCs). Something of a holy grail for twentieth-century developmental biology, claims to have successfully cloned animals have been subject to intense criticism and scrutiny in the past, and have even ended prominent scientific careers. Long before the faked results of the South Korean scientist Woo Suk Hwang's cloning experiments were exposed, an almost implicit association of the science of cloning with fraud and deception already prevailed. David Rorvik's fictional account of cloning published in 1978, in which he so skillfully imitated scientific expertise as to fool even his publisher, famously tarred the science of cloning with the brush of scandal and deceit. The figurative senses of cloning as "imitation" and "simulation" are thus paradoxically exemplified by the history of the science of cloning-so much so that even the word clone sets off alarm bells of various kinds.
The dangerous illicit clone, its negativity doubled by both its figurative and historical associations, is generically and traditionally an abject embodiment of a particular kind of genealogical shame. Suspected of being a fake, a derivative, a copy, or a mere replicant, the clone is diminished by lack of a proper genealogy-and thus identity, substance, or origin. The pedigree of the clone is subaltern, in the sense of inferior and subordinate, because it lacks separation from the original, and thus a distinct identity. It is, rather, the identity with, in the sense of being identical to, its progenitor that makes the clone synonymous with a lack of the most fundamental kind-of individually defining substance. These pejorative associations with clone are combined and reproduced in its usages to refer to illegitimate sexuality based on narcissistic identification (gay clones) and slavery (either as "slavish imitation," or in the association of clones with a worker class of slaves or drones).
Excerpted from Dolly Mixtures by Sarah Franklin Copyright © 2007 by DUKE UNIVERSITY PRESS. Excerpted by permission.
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