Crystals, Fabrics, and Fields Metaphors of Organicism in Twentielh-Century Developmental Biology Donna Jeanne Haraway New Haven and London, Yale University Pros. 1976Sy ‘ Copyright © 1976 by Donna J All rights reserved Jeanne Haraway This book may nwt be reproduced, in whole or in part, in any form (except by reviewers for the public press), without written permission from the publishers, [Sbrary of Congress catalog card number: 75-18) 74 Pmternational standard book number: oon it fy Designed by Susan MeCrillis Kel and set in Baskerville type. Printed in the United States of ‘The Murray Printing Co., ley America by Forge Vill we, Massachusetts Published in Great Britain, Europe, ane Africa by Yale University Press, Ltd. London, Distributed in Latin America by Kaiman & Polon, Inc., New York City; in Aust Services, Artarmon, NSW, sia hy Book & Film » Australia; in India by UBS Publishers’ Distributors Pyt., Lud., Delis in Japan by John Weatherhill, Ine., Tokyo. J vTo G. Evelyn Hutchinson for his inspiration and encouragement at every turn. Joseph Ne Contents Acknowledgments . Paradigm and Metaphor . The Elements of Organicism Ross G. Harrison A Pioncer in the Construction of an Organismic Paradigm ‘Vhe Silliman Lectures tham A Great Amphibian Mature Perspectives Paul Weiss From Fields to Molecular Beology ‘The Living System » Conclusion: Of Paradigms and Scientists References Index oe Gy 64 4 rot tol 138 147 M7 184 188 207 221Acknowledgments Many persons have helped in the writing of this book. ‘Their intellec- tual, emotional, and material support is deeply appreciated. 1 wish especially to thank G. Evelyn Hutchinson, E. J. Boell, F. 1. Holmes, R.S, Brumbaugh, Joseph Needham, Paul Weiss, C. H. Waddington, J. H. Woodger, June G. Goodfield, P. G. Werskey, Carolyn van Heukelem, Richard Stith, and the Danforth Foundation Two special friends are responsible for the warmth and space to work. ‘They are Jaye Miller and Helen Woods. ehParadigm and Metaphor ‘Vo cast off the rotte To cast olf Bacon, Locke, and Newton from Albion's covering To take off his filthy garments and clothe him with imagination; ‘To cast aside from poctry all that is not inspiration, rags of memory by inspiration, ‘That it no longer shall dare to mock with the aspersion of madness Gast on the inspired by the tame high finisher of paltry bloss Indefinite or paltry rhymes, or paltry harmonies Who pretend to poeny that they may destroy imagination by im jon of nature's images drawn yom remembrance William Blak If the history of science were a disinterested, purely descriptive ac- count of objective operations performed to elucidate the workings of a nature no one believed in any’ Similarl more, it would raise few passions. if science were progressive in the sense of an ever enlarging body of true statements about an ever unchanging yet fundamentally inaccessible world, nature would be seen simply as a big puzzle to be solved. Surely there would be no fundamental insights into personal or historical experience to be Simple cumulative science would be a curs xpected from pursuing such a science. -. Those with leisure, money, and arbitrary reason to solve puzzles could devote their time to it They could then present solutions to society, which would translate them into technology in the interests of current power alignments, benign or otherwise. History would be the search into the past for the wuly clever who anticipated the later, better solutions to problems. But at the same time, the hope of approaching nearer and nearer to nature is not surrendered lightly in science. Nor should it be, Wf the views of science espoused both by the old realists and by the more ected, it remains for us to understand in what sense science is progressive and in what way itm recent positivists are to be At lead to anGRYSTALS, FABRICS, AND FIBLDS - 1Cseems that the task of con- appreciation of the structure of natu temporary history and philosophy of sci age a humane sense of scientific research and theory. Vor it is a fact that the history of science docs raise passions, and the basic perspective adopted toward the status of scientific worlds does condition our poetry and our politics. Surely much of our dis-case with modern science comes from appreciation of the inadequacy of the old view of progressive, contro-oriented, objective descriptions of nature (or of meter readings, if nature herself is hiding or dead). ‘Two fundamental problems in understanding science can be noted: How does science change and what are the implications of the view adopted for the theoretical corpus of a science? Second, in what way can a science lead to a sense of the natural? To approach these two s, it is useful to look at a quite restricted arca of one science, developmental biology, .in..the. first-half’ of the. twenticth century, ‘The’ period is a time of basic crisis in which the age-old dichotomy between mechanism and ‘Vilalism was reworked and a fruitful synthetic organicism emerged, with far-reachirig implications for experimental programs and for our understanding of the structure of organisms, Asa key aid in analyzing a period of change in a science, it is instructive to explore the notion of a paradigm as applied to nce is to rediscover for this very broad iss biology. The book by Thomas Kuhn, The Structure of Scientific Revolulions (1962), is of central importance in exploring a change in a biological paradigm. An important aspect of a paradigm is metaphor, and it is suggestive to'itivestigate the use of-metaphor t6 direct research and its, iLGFPFetation. The paradigm. concept is rich with suggestions for ‘analyzing the proper place of metaphor in biology. To begin to under- stand again the place of image in a science should suggest a way in which natural structure can be seen in a post-positivist age.! Visual magery in particular is of critical importance. “draws constantly nearer some goal set by mi. °Cam we not account for 1. Kuhn himself’ does not believe that seience ature in advance.” The progressive aspect of science is different for both sefence’s existence and its suecess in terms of evolution fram [emphasis acted] the comniu- nity’s state of knowledge at any given time? Does it really help te inuagine that there is some one full, objective, true account of nature and that the proper measure of scientific achievement is the extent to which it brings useloser to that ultimate goal?” (1962, p. 170). The view to be developed below is different from Kuhn's but it is through his discussion. of paradigm and paradigm eom- naunities that this study will approach the problem of navural structures, This essay’ will not be able ( disenss adequately many important issues raised by Kuhn, For a relevant eritique sce KordigPARADIGM AND METAPHOR 3 Kuhn raises a host of interesting problems when he introduces the notion of paradigm change. Normal scicnce, he claims, proceeds by exploring the suggestions derived from a dominant paradigm, ‘The paradigm is not a set of explicit rules of procedure but rather “an object for further articulation and specification under new or more stringent conditions” (p. 23). Kuhn simply takes a device common to writers of history, the period, and applies it to a field thought to have developed differently from the rest of human endeavor, that is, in a straight line with occasional Dark Ages and metaphysical sidetrac value lies in refining two aspects of scientific paradigm: paradigm as shared constellation of belief (or disciplinary matrix) and paradigm as model or cxample. Both the communal and the exemplar nature of) paradigms are essential to understanding fundame science ical . But his deeper tal change in a ies: ‘The first aspect leads to a consideration of particular communi in science, their formation and modes of communication, and thei types of interaction with conlli different images and thus different languages. Kuhn's book has been criticized as leading w a highly subjective view of science in which an undefinable central picture conditioned all activity and prevented the be rom talking with any but the converted (Lakatos and Musgrave 1970). ‘Thus the paradigm could beshown but not rigorously defined, The distinction is not far from that made by Wittgenstein in his Tractatuy beoween saying and showing. Science might become more poctic in the mind of the general public if such a belief were widely held, but would it be any more intelligible? howevei cling scientific. communities using ver Kuhn argues, . that his view is more historically accurate for describing what real scientists have done. Further, it is not open to the charge of subjectivity because paradigms and their constitent metaphors are eminently community possessions whose principal value lies in their growing points. In contrast, a view of scientific theory that does not give a large place to metaphor, with its predictive value and potential for development, has trouble accounting for the very pro- gressive aspect of science such views are most often interested: in If one [eels he has a strong support for 1 view of objective science fi.c. separable from the historical context in which the science developed and separable from any heuristic props used in getting to pure formal theory) he will naturally feel that Kulin’s view of science is dangerously relativistic. In the second edition of The Structure af Scientific RevolutionsORYSTALS, FABRICS, AND FIELDS (1970), Kubn answers such charges by refining his meaning of com- munity. More is shared than the articulated theories. A whole system of assumptions, values, and techniques is involved. Paradigm as disciplinary matrix involves sharing (1) symbolic generalizations that give points of attachment for using logical and mathematical techniques in solving puzzles of normal science; (2) belief inthe appropriateness of particular models, such as fields or atoms in a void; (3) values about such things as the place of prediction in an explana- tion; and (4) nplars or conerete typical solutions used in the lite and important to the formation of students. Kuhn recognizes degrees of assent, and discerning the fine structure of paradigm groups would involve constructing a thorough sociology of science A simpler approach to community will be sufficient for the purposes of this essay. In looking at a time of paradigm change in developmental biology, it is important to note the types of association of biologists involved in the revolutionary switch of metaphor, ‘The ‘Theoretical Biology Club in the 19308 in Cambridge, England, is a suggestive cxample. Members of the group were interested in bringing to biology the power of logic and mathematical explanation previously enjoyed in the physical sciences, but they were not merely a modern variant of workers operating on a mechanistic paradigm. They shared also the view that hierarchical organization, form, and development were the central concerns of a new biology. Sensing a crisis and sharing a view of the future, the club self-consciously tried to make explicit among themselves the implications of their views. Its members were importantly involved in the transition to an organismic, nonvitalist paradigm for developmental biology. A second example from biology of an emerging paradigm com- munity of the same period is that of the crystallographers turning tion to biological molecules.* The udes and experimental work were significant in the critical refocusing of the relations of physics to biology and of biochemistry to morphology. A present-day paradigm group sharing the organismic paradigm that was born in its current form in the first half of this century is the Alpbach group. altel 2. John Law of the Seience Studies Unit of the University of Edinburgh is doing an extremely provocative stady of the paradigny community of British erystallographers of the 1gyos aud be- yond, It would be hard te overestimate the Importance of this group for the course of developmsen- Lil and molecular bintogy. ‘The entichment of concepts of struc in malecules was a major ingredient in brewing a nonvitalist organicismyPARADIGM AND METAPHOR 5 Participants in the Alpbach Symposium of 1968 include a former member of the Theoretical Biology Club, CG. H. Waddington." His relationship to the group, however, was tenuous, largely due to divergent views on mechanisms of heredity. Neither the Alpbach group nor the TBC represent tight working groups from closely related laboratories. Instead, they represent scientists groping toward a com mon metaphor appropriate across several previously separate areas of science. Their disagreements are as revealing as their agreements. The nature of the community forming upon the wansformation of the vitalistic-mechanistic controversy will _necd more sensitive attention later. It is enough here to note that a shared paradigm: is more than an aesthetic predisposition pecul concrete, common. picture of the central focus of a science. ‘The picture conditions the problems scen by the community ind the types of solutions admitted as legitimate. The problem for this study is to sketch the picture and to point out subtle but important differences in the paradigm core for various workers. In addition to stressing the community nature of paradigms, Kuhn's thesis is fruitful in another sense. The process of switch of paradigm is compared to a revolution. A major reorientation of fundamental metaphor occurs, leading workers in a field to see new problems and to accept radically different sorts of explanations. Normal sci characterized by the solution of puzzles, a truly cumul: ade possible by “a strong network of commitments conceptual, theoretical, instrumental, and methodological” (Kuhn 1970, p. 42)- By contrast, jar to a few minds. Lt is a nice is tive enterprise, ' the Wansition from a paradigm in crisis to a new one from which a new tradition of normal science can emerge is far from a cumulative process, one achieved by an articulation or extension of the old paradigm. Rather it is a reconstruction of the field from new fundamentals, a reconstruction that changes some of the fieldl’s most elementary theoretical gener many of its paradigm methods and applications. [Kuhn 1979, p. 84] lizations as well as 5. See Koestler and Smythies 19595 the book is based on papers and discuwion of the Alphach symposium, Among the participants were Ladwig von Bertalanfly, Jean Piag Paul Weiss, and W. HL Thorpe. Weiss binlogical issues, and these divisions were also apparent at the hatbel Inshekder nel Warldingion have long been divided on important6 CRYSTALS, FABRICS, AND FIELDS [such is the case, normal science must be particularly good at gencrat- ing periods of crisis in which the switches will occur. Kuhn points out eption of anomaly is the first step in the production of certain expectations. If such that the per full-scale crisis. The” paradigm inspir expectations repeatedly are not met, strains are introduced into the total paradigm system. Kuhn’s historical examples of anomaly and crisis in science are drawn from chemistry and physics. The phlogiston theory led workers to expect metals to behave in a particular way when roasted. Experiments with the red oxide of mercury did not fulfill such anticipations. Perception of the anomaly alone was not sullicient to lead to the discovery of oxygen, but it was critical that the problems presented by the old paradigm were proving resistant to proper solution. ‘The paradigm was failing in application to its own traditional problems (Kuhn 1976, p. 69). ‘There is an illuminating biological parallel to the examples given by Kuhn. The strict atomistic, mechanistic paradigm applied to organisms led Driesch in 1851 to expect the echinoderin gg to behave like a good machiné. That is, its development should have been “mosaic”: the parts should have their fates fixed at the outset and simple interaction of atomic parts according to mechanical laws should be the essence of development. Regulation was not an admissible occurrence within a strict mechanistic paradigm. Driesch formed his expectations ich, allegiance to the paradigm that he was certain his experiment of killing one of the first two blastomeres resulting from the first cleavage of sea urehin eggs would result in halfembryos. The appearance of whole litle animals in his dish precipitated a practical and philosophic crisis, of the first rank in embryology. The old paradigm seals its own fate by the operation of its own dynamic. Two further aspects of paradigm change can be drawn from the ample of Driesch. Kuhn notes that periods of crisis and revolution ence © are marked by concem over the philosophic foundations of the sci not evident in times of normal functioning. Ordinary scientists, used toa certain degree of confidence in their epistemological and metaphy- al commitments, come to feel a need to defend their positions or to evolve new ones. The first third of this century was marked by constant ists, neovitalists, and older brands of each. debates between ncomecha Despite the fact that each camp rested secure in the belief that reason ‘ided with it alone, no resolution could or did occur and experiment re until the machine paradigm common to mechanist and vitalist alikePARADIGM AND METAPHOR 7 was fundamentally altered; the metaphor was of central relevance. Thus the second aspect of Kuhn’s analysis of paradigm change The act of judgment that leads scientists to reject a previously accepted theory is always based upon more than a comparison of that theory with the world. The decision to reject one paradigm is always simultancously the decision to accept another, and the judgment Icading to that decision involves comparison of both paradigms with nature and with ¢ ach other. [Kuhn 1970, p. 77] ‘There is no absolute court of appeal; there are only alternate world views with fertile basic metaphors, This book attempts to trace in some detail a process of paradigm change in metaphor from machine to organic system that took the ground out from under atomism and animism alike in developmental biology. But in the end the essay must also ask how adequate Kuhn's model wuly is in this field of bio- logy. Surely it will be possible to tell the story of qwentieth-century embryology in terms of Kuhn’s pa what depth, in how essenti digm switch. Yet how exactly, to Ja way does to Kulnian scheme apply? So far, the fundamental objection raised against a positivist view of science and history has been that inadequate attention is given to the role of metaphor. Does such an objection reduce theories of scien tific knowledge and progress to a type of formal literary criticism? ‘The question ultimately concerns the nature of language. A neuvral observation language, at least asa goal, is central to positivistic assumptions. Kuhn believes hat the program of op the history of science incomprehensible and the status of a scientific theory sterile.® Operations and manipulations, he feels, are determined by the paradigm and nothing could be practically done in a laboratory ationalism makes |. The argument to follow is directed in particular aystinst the type ol approach taken by Pierre Duhem in bis 114 book La thio ie plysigre, Duhon: allows a certain hearistic role 1 models but denies that theories are essentially embedded in models and lose their coherence and comprehen siveness when uprooted fiom a rich, metaphoric soil, Campbell qzoy provided the ekisiea! qument against Duhem, Campbell's view of model is compatible with the one adopted here But it is necessa ry to explore the diflerences, if any, between physics and biology mvctaphoric systems, in the use of 5- Ernst Much in Te Srismee of Merhanies early stated the positivist, operittionalist perspective Theories do not say anything ante about phenomena or their structured relationships butte eeone= rival systems for restating the content of operations. ‘This radical empiricism has heen attacked for example by FP. PAW, Briddgman’s 1427 Lagiew) Mualern Physics gives the best areonnt of the prowram for eliminating nvey, but it remains influential under various arises, Probably honoperational concepts fhom physics.CRYSTALS, FABRICS, AND FIELDS without one (Kuhn 1970, p. 125}. A pure observation language as the basis of science exactly inverts the order of things. Operationalism might make sense as a postthcorctical exercise in clarification, but it docs not help in the process of planning experiments or in judging the fit of expectation and result, Further, the theory must be “reinte- y, aller positivist analysis if it is to make sense, t grated,” at least tacitly that is, for its structural character to show. The reverse of the positivi claim seems to be the case: the positivist program is the usefiul device but a richer conception is required to generate or understand science As Steven Pepper (1961) observes, structural coordination and empirical and logical data are necessary for adequacy. Positivism admits only the last two clements to science. Metaphoric systems are the core of structural coherence. Kun feels that philosophy of science from the English-speaking world “analyzes the logical structure of a completed body of scientific knowledge (Kuhn 1962, p. 136). History is suppressed in the formalization of a particular system. The formaliza- tion does not codify a pure body of knowledge stable for all future time. The rich languages used in actual science (including any axiom systems complicated enough to generate arithmetic) embody expecta- tions and fundamental views of the structure of nature. Scientific theories are therefore testable and generate crisis and subsequent change. Mary Hesse in her Model and Analogies in Science (1966) gives complete critique of the inadequacies of positivist views of science, She discusses various efforts to avoid the basic pitfalls of pure opera- tionalism, which include versions of the hypothetical-deductive language). The basic program of a more method (dictionary theories, in her positivism and its offspring has been to eliminate from science theoreti- cal terms that lack direct, observable, empirical consequences. ‘The stumbling block has been that success in the program would make all theories (autologies, static restatements of what is already known in a longer form in terms of direct operations and observations (Hesse 1961, p. 8). The theory would lack “growing points.” Loaded terms might be another expression for theoretical concepts pregnant with expectation because of the paradigm metaphor or model in which they are embedded. Hesse gives a number of paradigm model examples from physics such as wave fronts in quantum physics (pp. 8, 17), but as inany can be drawn from biology, one of the most suggestive perhaps being the gene concept. The idea of the genctic particle went simpurPARADIGM AND METAPHOR 9 far beyond what could have been operationally defined, and precise because of that fact, it was fruit in the development of geneti Il is now possible to explore more carefully what is meant in this essay by a metaphor with explanatory power. A metaphor is the vital spirit of a paradigm (or perhaps its basic organizing relation}. Hesse sees metaphor as an intrinsic part of science because metaphor is predictive. According to the formalist view of theories, “it can never be more than a lucky accident that a satisfactory isomorphism is found” between theory and nature.? The number of meanings applicable to theoretical terms is indefinite and there are no systematic criteria for searching out correspondences. Science becomes too arbitrary. Tn the metaphoric view, testing an implied isomorphism is the normal pro- cedure. But metaphor is not just pleasing comparison, Hesse gives a pro- vocative analysis of the logical status of analogical explanation (1966, pp. She secs analogy between two areas from three aspects: posi wal, and negative. Formalist. philosophies of science restrict analogy in science to the positive points of correspondence. Comparison of theory and analogy is then a didactic device and a posttheoretie operation. Other sorts of analogy might have been useful to the scientist but would not be an intrinsic logical part of scientilic knowledge. Analogy would then have a private function only. It would not be part of publi with material analogy, whose essence is the possession of causal implica- tions.* A metaphor is important to the nature of explanation because it leads to the testing of the neutral parts of the analogy. It leads to a searching for the limils of the metaphoric system and thus the anomalies important in paradigm change An important consideration in Hesse’s view is the theory given by cience. Hesse contra sts such formal analogy generates Max Black (1962) of interaction between metaphor and system of 44. ‘The geue also has badd 0 become less atomistic it the process, the particle meramorbesiars system with sirictural Laws somewhere berween 1920 ated 1g6n, the same pevioel that save the deyeloprient of nenvitalist organicism and structuralisny in developmental binlows, But, of course, tht system is firmly rooted in biochemistry 7- Hesse 1966, p46. The terms paradigm and metuphor require further clavitieation, Paradigns, the wider notion, includes techniques, examples, community values, amd the cemval metaphor, metaphor is g nerally related toa sense object such aya machine, aystal. or organism, .V meta phor is an image that gives concrete coherence w even highly abstract thought 1. Ihid., pp. 79-8. decision on the ultimate nature of causality i NOL Necessary to the point The logical status of analouy ist problem ne matter whit view is aduptedd tow rauusality, ‘The findarental assertion is thar scientilic dmacldye is vicher thaw theary anderstond as deductive ssstem with rules of application,10 CRYSTALS, FABRICS, AND FIELDS phenomena under investigation. Again onc is immediately returned to the problem of the nature of language. There is no correct literal description of anything by which an analogy should be judged (Hesse 1966, p. 166). Analogy and primary referent are both altered in meaning as a result.of juxtaposition. The example given by Hesse is that “nature becomes more like a machine in the mechanical philoso- phy, and actual, concrete machines themselves are seen as if stripped down (o their essential qualities of mass in motion” (p. 163). Ic is important to both Black’s and Hesse’s points that metaphor be en_as intelligible, with real impact and consequences explored by communities sharing the language and image. The point is the same m as a community possession, Metaphor as Kuhn’s defense of parad is predictive because it is embedded in a rich system not private to any one nidii. The onditions necessary for metaphor to be explanatory are not loose. ‘They can be summarized as the requirements that the meta- y to be explored, that the metaphor phor have neutral points of analog contain the germ of concrete expectations, and that it give definite limits to acceptable theoretical accounts in science. Metaphor is a pro- perty of language that gives boundaries to worlds and helps scientists using real languages to push against these bounds. Thus one returns to Kuhn’s point that the activity of normal science, puzzle solving within a particular world, itself leads to the anomalies preceding paradigm change. One of the principal foci of this book is what Kuhn calls the qua: metaphysical aspects of a paradigm (1970, p. 41): the sorts of entities ze as the stuf of the universe. Descartes’ immense influence was a major factor in the dominant scientific belief after the mid-seventeenth century that the universe was composed of minute corpuscles. Particular kinds of motion and interaction were part of the world picture of Cartesian atomism. The paradigm functioned like a ings.” The world picture functions to exclude , that as a conse- scientists visual: map to the structure of th other perceptions. It is important to stress, howev quence of the central role of metaphor paradigms operate more as directing tendencies than as clear and tyrannical logical archetypes. 9. This essay does not deal directly with the thorny problenwset for Western dualistic science by Hume and Kant, Terms such as stacture andl ivmorphivn are meant to point 1 a modified reatisan, suggested by A. N. Whitehead in Proress and Reality, ‘The mechanistic dodge of rowel f vicism and positivism by dealing with the world only as ific were made of avonns will be weated latePARADIGM AND METAPHOR u Kuhn treats operation of the paradigm as analogous to the geslall of psychology. However, in science one does not have the option of switch- ing perceptions back and forth, and the community of working scientists is not always aware that its vision has shified. ‘The pedagogical methods of science systematically suppress history, and a person sees basically only his own world and its possibilities. Kuhn gives the example of Galileo’s pendulum versus the Aristotelian’s falling stone (1962, p. 113), but biological examples also abound. A critical one for this essay is the perception of the organism in relation to the crystal. Hone sees the world in atomistic terms (metaphysically and methodologically), the crystal is a smaller, simpler version of the organism in a nearly literal sense. If one sees the world in terms of hierarchically organized levels (the or- ganism becomes the primary metaphor), the crystal becomes an intermediate state of organization. There is no longer a continuum of forms all based on a corpuscular foundation but rather a discontinuous serics of “organisms.” The crystal is still a fruitful metaphor used very seriously in exploring structure. But one sees a different crystal, just as Galileo saw a pendulum not a falling stone. Phillip Ritterbush in his essay The Art of Organic Forms outlines a number of suggestive approaches to the use of metaphor in biology, in particular the use of the crystal analogy (1968). Nehemiah Grew (1628 1712), an early plant anatomist, regarded regularities in natural forms as evidence that the processes of growth consisted of the repetition of simple steps, in which forms might be successfully analyzed.'® Ritter- bush notes that in the nineteenth century Ernst Haeckel, operating on a permutation of the crystal analogy combined with an idea of ideal types, analyzed the radiolarians, His drawings show animals that are “wonders of symmetry and design. Under the influence of his aspira- tions to discover strict symmetry, Hacckel altered his drawings to conform to his belief in the geometrical character of organic form” (Ritterbush 1968, p. 64). Biitschli (1848-1920), still influenced by the crystal metaphor, analyzed protoplasm in terms of'a geometrical space- lattice. Belief in his paradigm led to his seeing structures that could hardly be confirmed today. This fact does not belittle Biitschli’s contri- bution to cell biology but points out the importance of his aesthetic predispositions and the power of a metaphor (p. 67), Thomas Huxley, an untainted empiricist in philosophy, nonetheless made a_ major ar to, Rinerbush tobi p. 8. See alo Grew 1965,CRYSTALS, FABRICS, AND FIELDS an ideal ates by modifying advance in the classification of coelente body plan to explain the place ofa whole group of difficult organisms (p. 36). ‘The illustration of the signifi development of biology raises an important point, Ritterbush develops ive concretization of the idea of organic form. sms, Goethe nce of aesthetic commitments in the the thesis of the progr’ In contrast to the atomistic analysis of the form of orga (1749-1832) stated clearly and powerfully the concept of organic form. Gocthe is not considered primarily a biologist, but his impact on the development of that science is significant.!! Ritterbush’s provocative thesis is that biology can be scen as a concretization of an initial aesthe- tic notion, The metaphor of organic form became progressively more powerful as it became more concrete, never losing its nature as meta- phor. A perfect instance of the thesis is the elaboration of the cell theory The sphere had long been regarded as a st to crystalline in the nineteenth century. perfect form, appropriate to organic nature in contr units. Schleiden went beyond the opposition of the two notions in a great synthetic step, seeing the globular units of organisms as a funda- mental structural basis. The cell theory provided a representation of the whole org: as an assembly of essentially similar structural units which always arose [rom pre-existing cells. This was a scientific representation Decause it referred to structure and function but also because it precisely fulfilled the esthetic requirements of the idea of organic form, In the development of the cell theory we witness the trans- formation of esthetic presuppositions into scientific knowledge in a manner that parallels Kant’s statement that the sense of beau is an aid to the discovery of truth,'® 11, A wsefal diseusion of the relation of Goethe’s seientilie theories to his poetry ist be found in. Salm s97t ta. Ritterbush, 1968, pp. 33. 34. The establishment of the cell theory also meant the triumph of ry biology. But the cell theory has been wansformed in an organismic nt role, Development of the eel] theory’ was an important nodal point in atoms it ninetcenth-cent paradigm and fills differ the history of the mechanist-vitalist convoversy. The crisis in embryology rooted in the Rous: Driesch experiments of the late nineteenth century and resulting in a nonvitalist organicism in de- contest, The debate velopmental biology must be seen in a nuanced and comples: histories Muelley and Schwann was only partly over the former's avowed vitalism, The basie diss ment hinged on the relation of part to whole, the issue that is also central to the laver embryo enw logical debates, Mucller intsisted that the natire of each part is diluenced by its relation t the whole; Schwinn sought eauses only at the level of the smallest organic elements (see Schwann Hii and Mueller 12395PARADIGM AND METAPHOR 13 Ritterbush se es the reduction of ideal form to objective knowledge of biological structure as a fundamental paradigm that makes the “4 cess of biology a coherent sequence from the cell theory to DNA 3.4). The paradigm of the objectification of organic form is not identical to the organismic paradigm that became explicit in this century. Never theless, the Uwo are closely linked and both have been important in transforming our notions of the structure function dichotomy in science in general Several workers of late nineteenth: and early twentieth-century biology are important from the perspective of both paradigms. Por example, the German embryologist Hans Driesch: was instrumental in breaking the limits of a too simple mechanism ini biology and in preci- pitating the crisis Ieading to a nonvitalist organicism, But he violated the mandate of the concretization paradigm in postulating an entelechy as the director of developing form. He relapsed into the ideal, and if anything, weakened the aesthetic appeal of his explanations. D'Arcy Wentworth Thompson (1860 -1948), British biologist and classical scholar, also is relevant to both paradigms. In his weatment of the theory of transformations, he illuminates the geometrical relationships that exist between closely related species. Treating the transformation of fields as. a whole, he coneretized the notion of type and also applied systematic criteria (1917). ‘The focus of this study is a period of crisis and reformulation of basic concepts in experimental embryology and cell biology. The two major concepts sketched above, that of the significance of metaphor in a revo- lutionary change of paradigm and that of the progressive concretization of aesthetic dictates, together form the core of the analysis. Three principal scientilic workers have been chosen for careful consideration: Ross G. Harrison, Joseph Necdham, and Paul Weiss. Ross Harrison 1870 1959) was a pioneer in the construction ofa modern org in distinction to the old vitalisms and mechanisms. Trained at Johns Hopkins with W. K, Brooks and at the University of Bonn with Moritz Nussbaum, Harrison brought experimental embryology to Yale in 1907, where he remained until his death. His extraordinarily rich scien- tific carcer included development of tissue culture, a technique that he used to demonstrate the nature of nerve outgrowth (14175), His major work was an analysis of the establishment of the axes of'symmevy in the pro- cis developing limb bud and inner ear of the newt. His keen interest in problems of pattern and symmetry led him to one of the most beautitialORYSTALS, FABRICS, AND FIELDS 14 serimental treatments of form problems on the “biological level” ever undertaken, Harrison’s experimental work and perceptive analysis of its implications establish him as a primary foundation stone in the edifice of molecular biology, especially as molecular study is concerned with problems of function, structure, and integration of wholes and work spans the years of crisis in develop~ ) ecntury and termi- complex systems. Harrison’s mental biology in the first quarter of the twentic nates as the new paradigm was becoming firmly established in the 1930s and 1940s. His work was rooted in the fertile mechanistic school of Entwicklungsmechanik (developmental mechanics) and developed into a mature tree transcending the old metaphors and programs. Harrison’s work on the limb bud helped lay the foundation for the controversial field concept that informed so much biological work in the 1930s. ‘The concept of the field, considerably modified since the 1930s, still supports the organismic paradigm today. Also, Harrison’s associations with Hans Speman, the German worker on the “organizer” in amphibian development, and with Paul Weiss illustrate the role of paradigm com- munities sketched by Kuhn, From the perspectives of the transforma- tion of the mechanistic crystal analogy to apply to hierarchies and fields, of participation in pivotal paradigm communities, and of his central concern with the form problem in development, Harrison stands as a critical figure for a nonvitalist, experimental approach to modern Yet Harrison also is refractory to analysis in a subtle blend of the old and the organismic framewor terms of paradigm revolution; he new, the continuous and the discontinuous. Joseph Needham (b. 100), a student of the father of British bio- chemistry at Cambridge, F. G. Hopkins, saw as his major scientific task the union of the areas of biochemistry and morphogenesis. He viewed both fields as absolutely essential to a proper understanding of the central problem of developmental biology, that of form and pattern, Following the discovery by Hilde Mangold and Hans Spemann in the 1g20s that a piece of the dorsal lip of the amphibian blastopore im- planted in the blastocoele of another embryo so as to be in contact with ventral ectoderm results in the formation of an organized secondary embryo, Needham worked with C. H. Waddington and Dorothy Need- nical nature of the “organizer.” He remained con- ion, ham on the che! vinced that structure and function, form and chemical composi were inseparable and could be dealt with experimentally without the old dichotomies. Even though the original problem of the organizerPARADIGM AND METAPHOR 1 and induction remains unsolved and even ignored, his vision has been vindicated many times over in the concerns and achievements of modern developmental biology. Necdham was a Christian socialist impressed with the logic and philosophy of dialectical materialism as an alternative to the static systems of mechanism and vitalism. He was also influenced by the organic mechanism of Alfred North Whitehead and perceived that the revolutionary developments in physics and psycho- logy in this century were pregnant with significance for the guiding principles of biology. Organization and organism became the dominant themes of his work. Needham provided one of the first full statements of the organismic paradigm. Beginning with World War IT and a position in Chungking as liaison between Chinese and British scientists, Necdham’'s major interests shifted to the history of Chinese science and technology. He abandoned direct experimental work in the early 1940s to pursue his historical questions, Thus the period of inter ham’s dissertation wor 1 for this thesis extends from Need- in 1925 on the metabolic behavior of inisotol in the developing avian egg to his insightful publications with A. 8. Lawrence and Shih-Ghang Shen in 1944 01 the anomalous viscosity and flow birefringence of protein solutions taken from embryonic issues at critical points in development (Lawrence, Needham, and Shih-Chang). From the beginning of his scientific life, Needhain was uely interested in the philosophical and historical dimensions of science. His writings up to the early 1930s on mechanism and vitalism, culminating in the important Terry Lectures of 1932 (1936), are a valuable record of the nature of the controversies of that period. He has continued since that time to publish papers on the philosophical status of organicism, one of the most interesting being his study (1943) of the significance of Whitehead for biology written in r9q1, He has alo continued to note in provocative papers the progress of his old direct scientific interest, the chemical natu organiz ac © and specificity of the er (1968). The problem is indispensable for a full understanding of the ties between chemistry and morphogenesis If Harrison pioneered in laying the ground work for # biological organicism and Necdham explicitly stated the implication of the par digm, Paul Weiss (b. 1898) has spent a life time constructing a full organismic conception of biology and the organism, Weiss, born and educated in Vienna, studied both engineering and the life sciences. He brought the systems concepts of engineering to the problems posed byDS 16 CRYSTALS, FABRICS, AND FI the developing embryo (19256). Drawing on the fertile analogy from physics, Weiss developed the field concept for developmental biology (1925). His early work on the regeneration of limbs led him to the concept of “resonance,” highlighting a system of interaction between peripheral and central nervous structure and function. He has worked on problems of nerve growth, pattern of outgrowth under various con- ditions in tissue culture, organ formation as a cooperative product of cell and cell matrix interaction, neuroplasmic flow in nerves, and numerous other fundamental problems of morphogenesis and function. His experimental work on cell shape and movement and interaction has been constantly informed by an appreciation of the organism as a system-whole, Weiss exemplifies the biologist who takes organization i¢ problem, rather than as a verbal solution to profound as his ba question: Weiss, like Needham and Harrison, is instruc paradigm communities. His connections with the work and thought of Harrison date from before his 1930 work at Yale. He currently partici- pates in a community of scientists, the Alpbach group, from several fields working out organismic or structural paradigms. Necdham’s connection with the Theoretical Biology Club has already been men- tioned. C. H. Waddington, a member of that ing group of the 1930s and co-worker with Joseph Needham on the chemical nature of the organizer, took part in the Alpbach Symposium with Weiss. Waddington, with his concepts of genes and pattern and of chreods, or structured pathways in development, perhaps more than anyone else has built the foundation for viewing developmental biology as a struc- turalism related philosophically to the psychology of Piaget and anthropology of Levi-Strauss. The organismic paradigm with its associ- aphors has permeated the heart of modern science (Piaget ive on the nature of ated me 19674, 1967b). Harrison, Necdham, and W factors in suggesting experimental work and directing its interpretation, Weiss has explicitly analyzed the relation of art and science in a provo- cative later article, “Beauty and the beast” (1955). Our study will look carefully at the progressive concretization of aesthetic notions resulting from the thought of all three men ILis possible at this point to be more explicit about the questions and plan of the book, Is it legitimate to hold that a major new paradigm developed for embryology in the 1920s and 1930s? If'so, what are the uso demonstrate the role of aestheticPARADIGM AND METAPHOR 17 fruitful metaphors and models and how do they differ from the key images of the previous period? What is the nature of the crisis generated by the normal science preceding the postulated revolution? How do the new paradigm communities develop and fanetion? What are the weaknesses of the paradigm notion for embryology? Finally, what is the relation of the orgar logy to developments in other sciences? It is possible to maintain that branches of physics, mathematics, linguistics, psychology, and anthropology have all es perienced revolutionary and related changes in dominant philosophical perspective. The primary element of the revolution seems to be an effort to deal with systems and their transformations in time; that is, to take both structure and history seriously without reducing wholes to Jeast common denominators. Organization and process become the key cone’ ther than last ditch incantations. But before looking directly at the work and intellectual allegiances of Harrison, of the mechanism--vitalism controversy in order to put it in historical xt. The next section looks at the years from about 1850 to 19 is in biology. Theories of tropisms, physiology, biochemis- iry, developmental mechanies--all illustrate both the uiumphs of work conducted under the mechanistic program and the strains leading to the new paradigm. ‘The major long-standing dualities in biology structure-function, epigenesis~preformation, form-process have all been reformulated as a result of the crisis, Thé section following the historical survey deals with four basic elements of the organismic para- digm., First is the primacy of the form problem, then the roles of symme- uy, polarity, and pattern concepts in the old and new metaphoric systems. Third, the nature of the field particle duality is examined with emphasis on the resolution offered bya nonvitalist organicism, Finally, the relationship of organicism in biology to structuralism as a philosophy is considered with the focus on the fundamental concept of organizing relationships. The following three parts of the book con- sider in detail the work of Harrison, Needham, and Weiss. The con- clusion evaluates the adequacy of the paradigm idea in discussing change in biology and finally differer and Weiss in their d smic paradigm in bi rns r Yeedham, and Weiss, it is necessary to analyze the nature cont 30 as years of eri ates among Harrison, Needham, grce of adherence to the organismic paradigm Brief Historical Sketch In 1850 Wilhelm Roux was bor 1 in Germany. Setting up a program1B CRYSTALS, FABRICS, AND FIELDS. and a method for the experimental analysis of development within a mechanistic world, he was to sce himself as the Descartes of embryology.!* But before considering his program and its effects on embryology, it is necessary to look at the long and complicated interac- Gon of mechanism and vitalism. In this way it will be possible to under= stand of significance of Roux’s school of Entwicklungsmechanik. Joseph Needham in his article “Hunting of the Phoenix” sketches the oscillations of mechanism and vitalism in biology (19314)." His version of the history has been chosen to emphasize his conception of revolutionary change in his own work. In a curious sense Paracelsus in the sixteenth century is the father of both the vitalistic and mechanistic modulations of the machine theme. He gave birth to modem chemistry through the school of iatrochemists that followed him. But side by side with his views on “‘chymical medicines” was his development of the role of “archaei,” or spirits, as governors of the chemical processes. ‘The fundamental ambivalence of Paracelsus was prophetic for the historical and logical development of the mechanism--vitalism controversy. ‘The seventeenth century saw the full flowering of the mechanical corpuscular philosophy. This was Whitehead’s “century of genius,” producing Newton, Leibnitz, Descartes, Harvey, Boyle, Galileo, Francis Bacon, Pascal, Locke. On and on the list grows, including virtually all the names a liberally educated person would recognize in the history and philosophy of science until the mind-shaking develop- ments in contemporary science. There is good reason these are the names remembered, for the seventeenth century articulated and en- throned the paradigin and its metaphoric system, which reigned until this century. In Kuhn’s terms, and consonant with Needham's the intervening 200 years were a time of puzzle solving. The core of the paradigm was summed up by Robert Boyle, author of the Seeptical Chenist of 1680, “I do not expect to sce any principles propos'd more comprehensive and intelligible than the Corpuscularian or Mechani- cal.” The seventeenth century gave science a method, a metaphysic, and a coherent fabric of expectation that was forcibly exploited in the following centuries. [tis difficult to choose a single man from that con- views, 14. Jane M. Oppenheimer drew this analogy in a conversation an July 20, 1970 ty. Much of the amalysis that follows is seriously oversimplified, Mechanisny and vitatism have developed int biology since the carfiest recorded observations of the Greeks, but [feel that che simplification is legitimate im highlighting the particular crisis of dhe le nineteenth and early owentict centuries, Fora recent useful summary af the period, see Coleman 1971PARADIGM AND METAPHOR 19 tury as the most influential, but surely René Descartes would be hard to pass over. He is instructive too for the current of ambivalence in mecha- nism that expressed itself in various vitalistic reactions. Descartes clearly provided for a wanscendent principle to plan and service his machine. His dualism was complete, and his mechanic was as necessary logically as his animal-machine. (Descartes 1955). When his successors tied to banish the substance of mind and to construct a monism of mechanical materialism, the ghost in the machine reappeared just often enough to spook the factory workers and technicians. No argu- ments of logic would banish the vitalists, because the issue was essen lially nota logical one. The metaphor and its rationale led to a vi as surely as it led to a mechanism. The mechanists continually embar rassed the vitalists by solving difficult puzzles on mechanical principles, and the attitudes of the vitalists oficn enough made their suspect status well earned, But the splits of mind and body, structure and function, efficient and final ca rever sutured during the reign of eorpus- cular philosophy. ‘The late seventeenth centur alism use wer aw a wave of reaction to rigid mecha- nism in the anima sensitiva of the chemist- physiologist Georg Ernst Stahl. But the eighteenth century was again dominated by discrete bits of matter in blind motion, Vitalists in this context often had a richer view of the organism. Man a Machine by Julien de ku Metric, physi of the Enlightenment monarch Frederick the Great, was an expression spice ian of full, naive materialism applied to the workings of the finest tin ofall, the human body, [is ironic that current excitement with timing and rhythm in nature might owe a great deal to the mechanical clock metaphor of Enlightenment science. If anything, the anonymous answer Man more than a Mackine only served to emphasize the strength of the machine paradigm. The pair of articles from the eighteenth century provided the images and inspiration for Necdham’s Man a Machine 28, in which he sums up his neomechanism of those years 1928). His own realization of the implications of the new organicism written in 1 dawned graduaily. The nineteenth century was to witness the triumph of mechanistic explanation in biology but also ironically the birth of the strains that were to contribute to a major crisis, Yet the strains are not so surprising 15. Koestler 1967. 11 is instructive to note that Koestler is a friend of Paul Weiss anil was thy prganizer of the Alpbach Symposium, However, Koestler’ and Waddington’s views differ si ficantly, especially on genetics20 CRYSTALS, FABRICS, AND FIELDS own contra- 1c generating i from Kuhn’s perspective of normal scic dictions for the new synthesis. Needham singles out two important developments of the century as critical to the power and acceptance of mechanism: the synthesis of urea, an organic molecule, by purely chemical means by Wohler in 1828 and the demonstration by Atwater and Rosa in 1897 that die Jaw of conservation of energy applies strictly to animal metabolism.’* Also, by 1840 explicit atomism had been triumphantly established in biology by the cell theory. Dalton’s atom did the same for chemistry. By 1859 Darwin had published his Origin of Species, a work that was in many ways suspect to the archmechanists of biology (Sachs, Loch, and Roux). But the Origin represented a kind of thinking very compatible with the corpuscular world view. Natural selection was not unlike the invisible hand of Adam Smith. The free play of forces and the lack of internal constraints and organizing r tions were central, The Enlightenment’s ideal of untrammeled reason, ism was pervasive in the biological as well as in progress, and materi the social order of things. It is instructive to look at the situation in Germany in the mid-nine: teenth century, for it will be in a German context that the neovitalist Driesch will pose a fundamental challenge at the end of the century to Roux’s Cartesian prophecy for embryology. The Mechanistic Quad- rumvirate of Helmholtz, Ludwig, Du Bois-Re: nd Briicke swore a mutual oath in 1845 to explain all bodily processes in mechanistic terms. This group did not make its mechanism a final metaphysic and tolerated various philosophical stances outside biology. But the step from method to metaphysic is an casy one. The medical materialists of the 1850s took the step in no uncertain terms. Jacob Moleschott, Lud- wig Bachner, and Carl Vogt are “remembered for their outrageous dicta that man is what he eats, genius is a question of phosphorus, and the brain sceretes thought as the kidney secretes urine.""7 At about the mond, 16, “Hunting of the Phoenis,” in 19344, p.115. However, full application of the law of conserve tion of energy 10 animals had actually been demonstrated by Max Rubuer in 1893. Fhe essential content of the lave was already widely accepted, and Rubner felt compelled to justily his extensive elect Needham ascribed 10 it elliorts (1294). Abo, the synthesis of urea did not have the crit having more to do with the perplexity over the intricacies of organic eatalysis than with the distine- tion beaween vitalits and mechanists. Liebig interpreted Wobler’s discovery in terms of a clear separation of catalytic and vital processes (sce Samuel 1972). Origin of form, not chemical come fs0e Brooke 168 positions, was the veal isne for vitaismy an mechanisin 17. Loch 1954. The quote is from Fleming's intsoduetion, p. is. For a discussion of important isu in physiology and biochemistry relevant to this eissertation, see the introduction by Frederic 1. Holmes to Justus Liebig’s Animal Chemisty (16, See ale Gallaty (1074 fora diseassion of the nnetecuth century reduetionisn, phitosophical basis of mid