A growing alienation?

A growing alienation?

Stud. Hist. Phil. Biol. & Biomed. Sci. 39 (2008) 374–377 Contents lists available at ScienceDirect Stud. Hist. Phil. Biol. & Biomed. Sci. journal ho...

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Stud. Hist. Phil. Biol. & Biomed. Sci. 39 (2008) 374–377

Contents lists available at ScienceDirect

Stud. Hist. Phil. Biol. & Biomed. Sci. journal homepage: www.elsevier.com/locate/shpsc

Essay Review

A growing alienation? Tim. J. Horder Department of Physiology, Anatomy and Genetics (Le Gros Clark Building), South Parks Road, Oxford OX1 3QX, UK

When citing this paper, please use the full journal title Studies in History and Philosophy of Biological and Biomedical Sciences

From embryology to evo-devo: a history of developmental evolution Manfred D. Laubichler & Jane Maienschein (Eds.); MIT Press, Cambridge, MA, 2007, pp. vii+569, Price $55.00, ISBN-13: 978-0-26212283-2. The volume here under review is a doubly welcome new contribution to the defining of the differing perspectives of historians of science and practising scientists in the context of a newly central focus of interest in biology, namely ‘evo-devo’. What exactly this term covers will be a theme we will need to come back to. The book raises interesting questions such as: What is a scientific discipline? How does a new one emerge? What role do historians, and other non-scientists, play in establishing it? The book is welcome, first, because it attempts to survey the history behind a recently emerging research discipline that is gaining much scientific attention; so recently in fact that it has as yet hardly been touched by historians. ‘Evo-devo is clearly taking center stage among modern biological disciplines’ (p. 123). And yet Laubichler and Maienschein (the editors) refer to ‘the largely neglected history of twentieth-century embryology’ (p. 2); ‘next to nothing was known of what happened . . . in the period between the 1920s and the 1970s’ (p. 4). Secondly, it is welcome because, as the outcome of two meetings, the first organised as the annual Dibner Institute Seminar in the History of Biology in 2001 at Woods Hole and a follow-up workshop in 2002 at MIT, it sets out to show the potential for collaborative interactions between historians and scientists. The editors claim a new departure in the involvement at the meetings of working scientists. ‘An unusual and most effective aspect of the workshop was the inclusion of three individuals involved day to day in the execution of evo-devo’ (p. 468) who had ‘expressed an interest in the history of their discipline’ (p. 2). ‘The way history is reconstructed by practitioners reflects a lot about their assumptions concerning the current status of the field’ (p. 27). The practising scientists invited to the workshop (Hall, Müller and Wagner; all well known and already much published evo-devo devotees) contribute ‘Reflections’ in the final part of the book. In the end the exercise raises far more questions

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than it answers, many of them important and interesting for the nature and future of biology. It also points, in concrete form, to the potential for both scientists and historians of science to enrich each others work. A key issue underlying the book as a whole is the following: is this a new discipline within biology or is it some sort of delayed continuation of the much older tradition of relating embryology and evolution particularly associated with Ernst Haeckel’s speculations on recapitulation 150 years ago? Part One is designed to cover the period of the early twentieth century whereas Part Two covers later periods. As this structure implies, there is general acceptance in the book of an hiatus in the history of this subject, and as the editors put it, ‘Exploring what happened in between . . . (1920–1970) . . . was the goal’ (p. 4). Given the general agreement that there was an historical discontinuity, the question arises as to why the subject fell out of favour. Churchill documents the ‘internal’ critique of Haeckel that started in the 1880s: his paper offers a typically insightful appraisal of Haeckel’s idea of recapitulation and its early critics and defenders. The best known explanatory viewpoint is that there was a ‘revolt from morphology’ around the end of the nineteenth century, mainly the result of the rise of experimental embryology and eventually of genetics. Allen’s wide-ranging account offers a number of explanations (while avoiding that catchphrase which he himself coined). He emphasises the disciplinary ‘splitting’ according to proximate, as against ultimate, biological causal factors; and within the former, of genetics as distinct from experimental embryology, driven by the ‘dialectics’ of epistemological levels (historical/sociological, and biological). This account nicely illustrates the effects of the constant historical tendency for disciplinary sub-specialisms to emerge based on concepts and methods. By recalling charges of vitalism levelled against the key contribution of Hans Spemann, Allen picks out only one of the many complex reasons why embryology failed as a discipline, as compared with the conspicuous successes of genetics. Gerson and Griesemer reinforce Allen’s themes. Richmond’s contribution represents the most direct coverage in the book of the gap period; her fascinating analysis of Richard Goldschmidt (the figure who most kept alive the linkages of interests in genetics, embryology and evolution in the period 1920–1950)

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brings out the overriding cause of his marginalisation: the dominance of the Morgan school in American biology, an assessment endorsed by Allen. Maienschein’s paper offers Bonner as another (even less successful) contributor to the attempts to link the three domains. A second, clichéd, account of the history of evo-devo has it that embryology was actively excluded from the Modern Evolutionary Synthesis of the 1930–1940s. But, again: why? This thesis is reiterated at several points in the book, particularly in the papers by Love and Gerson. Using the examples of morphologist D. D. Davies and palaeontologist W. K. Gregory, Love shows how they both used embryologically based concepts (such as allometry, gradients, larval forms) in their thinking. (He could well have added G. G. Simpson as a similar example.) Love argues that morphology was excluded from the Modern Synthesis alongside embryology, but this is hardly borne out by the list of contributors to the landmark Princeton meeting (Jepson et al., 1949). Rather, his paper indicates that in so far as morphology and palaeontology, alongside population genetics, were key disciplines in the Modern Synthesis, embryology nonetheless played a behind-the-scenes role in providing crucial linking concepts, especially for the morphologists. Four of the papers (Wourms, Love, Maienschein, Churchill) serve to remind us that the morphological research traditions that triumphed in the Haeckelian period have continued to be practised uninterruptedly. This is not only a measure of the historical inertia of research traditions: it suggests (as Griesemer argues) that disciplines become perceived, and labelled, as passé (and failing) only by comparison with other disciplines which are seen at the time as new and ‘leading-edge’. Novelty is an important driving force in science. But in order to address the question of whether the recent new interest in evo-devo is a rediscovery of the older tradition or is better seen as a new discipline, one has to have reasonably clear agreement about what actually characterises the field and delimits it in its manifestation today. Many of the papers allude to the problem of defining the subject, particularly Chapter Two. Allen’s definition of the field’s aims is a good starting point: ‘a synthetic paradigm or research program in which developmental biology is combined in some ways with both genetics and evolutionary theory. Some versions of ‘‘evo-devo” in the past, as well as the present, emphasize the genetic, while others stress the evolutionary aspect, but the common element is that in some way all three components are intertwined’ (p. 123). The editors suggest that ‘By the late 1990s evolutionary developmental biology had all the markings of a new scientific discipline’ (p. 3), a judgement spelled out by several authors. It has the ‘institutional hallmarks of a successful scientific discipline’ (Wagner, p. 526) including dedicated international symposia, new or renamed journals, its own textbooks, and funding through specialised panels at grant agencies. A desirable requirement for any newly emerging discipline is a new, identifying name. The term ‘evo-devo’ seems to be the winner among a number of contending alternatives variously suggested throughout the book (Müller, p. 501). And yet this faltering over an emblematic title is perhaps a pointer to a lack of conviction about the reality of the supposed novelty of the discipline, evident among a number of the authors. Even the editors sometimes appear uncertain: ‘it is still unclear whether the Evo-Devo focus can succeed in providing new perspectives that go beyond what would be possible within other explanatory schemata’ (p. 5). In setting out the agenda of the book, Laubichler remarks that ‘Now . . . after roughly thirty years of gestation, evolutionary developmental biology is solidly entrenched within the conceptual framework of modern biology . . . However, today’s evolutionary developmental biology is far from being a uniform scientific discipline with a single research agenda and a well-defined methodology’ (p. 13). Müller’s daunting Table 15.1

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provides ample evidence of the disunity and complexity at the conceptual level. ‘What, exactly, is the new and genuine product of this synthesis?’ (p. 14). ‘An integrative epistemic space that would unite all these different ‘‘Evo-Devo” applications within a genuine new ‘‘synthesis” has not yet emerged’ (p. 24). ‘Clearly the new discipline is still in a process of consolidation, searching for unifying principles, but already a number of institutional attributes foreshadow its future importance in the biosciences . . . surprisingly different accounts of the origins, the contents, the aims and the significance of the nascent discipline are being told. Some even question whether the field exists at all’ (p. 499). Of course such judgements depend on what one means by a ‘discipline’ in the first place, a topic briefly addressed by Gerson. The organisers of the two meetings retrospectively refer to a ‘growing skepticism about what the current emphasis on a ‘‘synthesis” of evolutionary and developmental biology actually entails’ (p. 5). These uncertainties about the nature of the nascent discipline not only explain the book’s hesitant character, but also pose obvious difficulties for the historian. How can historians do good history on a subject that is so ill defined, and which may not really exist? It is not surprising that the professional historians tend to stick to the relative safety of the earlier period, even though it has already been relatively well documented, without comment on the historical gap or any possible new manifestations. The three representative evo-devo scientists perhaps have the inherent right to claim to define their own subject, on the basis of first-hand experience and with automatic authority. It could be argued that, through his numerous and pioneering publications, Brian Hall almost single-handedly delimited the new subject. All three seem to share a vision that sees evo-devo as encompassing virtually the entire range of biological considerations, from molecular genetics all the way up to ecology, adaptation, behaviour and even environmentalism (pp. 511–512). And yet Wagner’s concluding paper is distinctly guarded about the ultimate prospects for success, and even pessimistic: ‘it is much too early to say whether this challenge (i.e., the necessary analytic power to gain scientific knowledge on a major evolutionary transformation) can be answered successfully’ (p. 530). Whereas Müller in particular urges the need to go beyond the molecular level, Wagner shows this may not be that easy. I turn now to the second theme of the book: the implications of the interactions of the scientists with the lay observers, and interpreters, of science. ‘What do birds (i.e., the working scientists) have to do at a meeting of ornithologists’ (p. 524)? Overall the meetings were judged ‘a tremendously successful experience’ (p. 2). Distilling from the potted biographies provided, participants comprised three professional historians, three philosophers, one sociologist, eight biologists (four or five of whom are lab-based; the remainder might be considered theoretical biologists actively involved in the evo-devo field). The potential meeting of minds across a wide spectrum of academics adds greatly to the interest of this book: what was going on in the original meetings and how is this manifested across the pages of the book? If, through history, one can get an understanding of what has controlled the direction of travel in past science, then one might be better able to anticipate and rationally select future directions. ‘[The] historical approach gives a unique perspective on, and informs, current developments; in that way, historical awareness can actually improve scientific practice’ (p. 25). Wary though historians will be about considering contemporary, let alone future, events, understanding the progression of science may, and should, benefit from the historian’s expertise because, to an exceptional degree compared to other fields of human experience, science is explicable (and even predictable) given that it advances according to known and unvarying methodological and cognitive principles. However, as the editors say: ‘Of late, writing the history of science with a long-range perspective in

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order to get at the underlying persisting traditions . . . has become unfashionable . . . [This] new orientation in the historiography of science (focused . . . on the local, immediate, contingent, and particular aspects of the scientific enterprise) has led to important new insights, but it has also contributed to a growing alienation between the communities of historians of science and scientists’ (p. 5). Why should this be? Does it matter? Does this volume help us to see how to reverse the trend?. Almost all papers in the book say something about the history of evo-devo, but the wide range of approaches taken is in itself of considerable interest. They reveal something about the motivations (perhaps unacknowledged) of the authors and about the forms of communication that are possible in such an interactive exercise. Wourms (a biologist) takes essentially a listing approach to history: he presents a chronology of key events, institutions and personnel (backed up by useful listings of expeditions, surveys, journals, classic books, research students, universities, museums and marine stations, all in the USA). This appears to be documentation for its own sake, although his Fig. 8.2 attempts the big picture in summarising lines of intellectual descent and of splitting among the disciplines. In such an approach explanation and causality are merely implied. An alternative form of history (seen here in varying degrees with Hall, Wimsatt, Love, Newman, Maienschein) is to attempt microhistory on some highly selected corner of the wider subject, involving minute study of an obscure, peripheral figure or institutional arcana (as distinct from the microhistory of seminal figures as so well illustrated by Churchill and Richmond). Thus Wimsatt devotes considerable length to Rupert Riedl and Lancelot Law Whyte, two distinctly idiosyncratic figures: he freely admits that they had very limited influence in the Anglo-American biological tradition. Or again, Hall devotes four pages to microtomes and the Cambridge Instrument Company. This approach can amount to an antiquarianism that, in a book tackling the big picture, surely needs explanation. Another type of distraction from useful, valid explanatory history is at times in evidence: Newman, Gilbert and Griesemer could be said to have raised the spectre of Whiggism, that is, of finding interest in an earlier historical episode apparently on the grounds that it seemed to have anticipated later developments. Even Richmond seems uncertain in her avoidance of this trap: ‘it would certainly be wrong to view Goldschmidt as a precursor of contemporary research programs in Evo-Devo’ (p. 173), but ‘Goldschmidt’s approach to problems of heredity, development, and evolution continues to resonate today, even if those currently engaged in attacking these problems are armed with a different set of empirical data and theoretical assumptions’ (p. 199). It hardly needs insisting that historians avoid following along with the fashions and prejudices of the contemporary scientists and, even worse, backing them up by creating retrospective historical myths. An additional and familiar handicap to successful communication is also represented in the book. Externalist history, or history that emphasises, for example, social contexts, techniques, institutions, technoscience or material cultures, is difficult for practising scientists to engage with wholeheartedly. Since such approaches frequently want to equate science and technology, they seem to deny the centrality or even relevance of what is ultimately most important to scientists: namely the ideas that motivate them. Gerson perhaps comes nearest to representing a science studies input: although he attempts to explain what was going on in the Modern Synthesis in institutional and organisational terms, his paper is simply too short. Finally, one can point to the contrasting roles of laboratorybased scientists as compared to the ‘theoretical biologists’. Are their respective contributions to evo-devo science necessarily of an equivalent standing or equally and mutually beneficial? Wimsatt identifies the missing element in the Modern Synthesis with

‘internal factors in evolution’ and ‘burden’; his frank critique revealingly highlights many of the shortcomings of the theoretical biologists’ contributions. Of the two other biological theoreticians (both with philosophical backgrounds), Love’s is a brief and useful historical piece, but Griesemer’s is long and, for this particular labbased scientist, his attempt to see Mendel as a developmentalist and Spemann as a hybridist was baffling. The wide range of uses of history and contextualisation of science that we see in this book leads me to the following observation. The possibility of fruitful interactions and exchange between scientist and non-scientist contributors presupposes clarity of shared (or at least overlapping) styles, languages and aims. Without this, ships merely pass in the night. It seems likely that scientists would find it easiest to engage with historical treatments that help to explain the situation they find themselves in: central to their concerns and ways of working are hypotheses, theories, thought patterns, concepts and ideas. Any usefully explanatory historical contribution must focus on the historical continuities up to the present: it must explain where we are by accounting for how we got here historically and, in particular, it cannot leave unexplained, arbitrary gaps. Hence the editors’ plea for the ‘longrange perspective’ on ‘persisting traditions’. Such history cannot be done briefly. The three spokesmen scientists in fact say very little about how and why they think interest in evo-devo has reemerged in the way it has. Indeed this historical question is barely touched on anywhere in the book; the nearest approach we get to it is in Laubichler (pp. 22–24) and, very briefly, in Allen and also in Wimsatt. ‘There is still no consensus about how one should interpret these recent events: many of the (still living) participants have their own interpretations, some of them collected in this volume’ (Laubichler, p. 22). However a number of papers make it perfectly clear how the new discipline is in reality practised; Hall, Newman, Gilbert, Wimsatt and especially Wagner illustrate how, in the typical research department, molecular genetics defines the way the subject is actually pursued. And yet Wagner states that: ‘devo-evo as a scientific movement was well articulated many years before molecular developmental biology joined the act’ (p. 526). There are two broad explanations of the rediscovery of evo-devo and they could not be more different. The first explanation is the 1977 publication of Stephen J. Gould’s first book, Ontogeny and phylogeny. It arrived almost entirely out of the blue. There can be little doubt that Gould’s book served to educate a new and unsuspecting generation of biologists about these areas of interest, and to make it respectable and safe again to discuss such subjects as recapitulation. The alternative explanation would hinge on the discovery of Hox genes around 1985 and the rapid recognition of their importance in linking species phylogenetically and explaining embryogenesis. Although Gould had briefly speculated about the molecular basis of development and heterochrony, almost nothing was known at that time: when, soon after, Hox genes were identified, their discovery arose from a quite different research tradition, molecular biology. This present book has largely missed out on the opportunity to explore these contrasting and distinct historical backgrounds. The implications of the two explanations are remarkably different in kind and important: that is, the impact of a technical outcome of a technique-driven tradition as against a largely historical exercise in the history of ideas. Of course, measuring degrees of causal influence at earlier historical periods of time presents very severe problems. Gerson, Wimsatt, Wourms and Allen all refer to Gould’s importance, but only briefly and without any analysis. Perhaps the best measure we have is the judgement of witnesses who lived through the relevant episodes. Overall Gould’s thesis amounted to a reformulation of de Beer’s earlier notions of heterochrony. (Gould also owed

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something to Simpson whose approach would fit well into the pattern of influences discussed by Love.) Part of the explanation for the impact of Gould’s book lay in its most unusual two-part structure; Part One was purely historical, Part Two was theoretical biology. His two-pronged presentation perhaps had the effect of doubling-up his authority as a new, important, though still relatively young, author; it bestrode both history and an apparently novel contribution to biological theory. The historical section allowed him the freedom to discuss the Haeckelian notions that had earlier become so discredited: he could then re-legitimise them in using them, in part, in his scientific contribution. In sum, this is in many respects a well produced book and it will have obvious appeal for people close to the subject of evo-devo. However, it has lapses: for example ‘John Burdon and Sanderson Haldane’ (p. 173) (one and the same); the running title of Richmond’s article is cut short; de Beer did not work on the molecular nature of the organizer (p. 144). The referencing system is unusual, unnecessarily cumbersome and inconsistent; for most papers, text references refer you to numbered notes which only then lead you on to a separate alphabetical list of source references, but some papers refer you directly in the text. As with any multi-author book, there is ultimately no coordinated consensus and there are the usual repetitions, redundancies and gaps detracting from the balanced coverage of the material. The following quotation captures the overall message of the book well. ‘[W]e do not seem to be dealing with a single set of questions and methods; its distinctness from other areas of biology is still under debate; and we do not know how the story will turn out . . . The intellectual and scientific promise of Evo-Devo is so exciting, however, that we cannot not tell the story’ (p. 15). In other words, it is not yet clear how much new there is in it. Essentially it may be merely a rediscovery; it may amount to the latter-day recognition by evolutionists (and especially geneticists) of the relevance and significance of the classic issues of embryology. Important classic themes do not disappear: they just become unfashionable. However, the book does forcefully remind us that biology and history face very similar problems. Intrinsically, and distinctively among the sciences, biology is characterised by its complex, integrative, multi-level nature. History suffers, like biology, from disci-

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plinary fragmentation and overspecialisation: more and more about less and less. As scientists and/or historians we require perspective and integration in the face of these ever increasing and inevitable tendencies. Scientists, and perhaps historians, ought to be far more worried than they appear to be at their disunity. The work of historians can help scientists in a number of ways: their work challenges the clarity of the scientists’ concepts (whose origins and meaning so often are obscured by historical accretion); it alerts scientists to assumptions that have become unconscious; it reminds them of forgotten, alternative ways of thinking and neglected data sources (so avoiding repetition of past mistakes); it helps them detect trends; and above all it broadens their perspectives generally. A strong case could be made that the very best scientists tend to be those with wide horizons, which often correlates with having a sense of their subject’s history. If indeed we face ‘a growing alienation’, this book shows that it does not have to be so and that a solution must surely lie in combining the respective skills of scientists and historians: having first acknowledged that their aims, though different, must have much in common. In an ideal world, one could hope for many more meetings of the kind that resulted in this book. They could not fail to be important in helping to steer scientists in the most productive directions. The case of Gould’s influence on evo-devo stands as an outstanding example of the proposition that the recall of historical material and perspectives can, under appropriate circumstances, have very significant effects on scientific priorities. Here the history of ideas appears to have mattered in a way that dwarfed the historical blips and diversions associated with technical breakthroughs or institutional and financial structures. At their best, independently minded historians of science can indeed serve to reorientate the autopilot that usually guides scientists. References Gould, S. J. (1977). Ontogeny and phylogeny. Cambridge, MA: Harvard University Press. Jepson, G. L., Mayr, E., & Simpson, G. G. (Eds.). (1949). Genetics, paleontology and evolution. Princeton: Princeton University Press. Müller, G. B. (2007). Six memos for evo-devo. In M. D. Laubichler & J. Maienschein (Eds.), From embryology to evo-devo: A history of developmental evolution (pp. 499–524). Cambridge, MA: MIT Press.