Journal of the History of Biology, Vol. 3, No. 1

Journalof the Historyof Biology SPRING 1970: VOLUME 3, NUMBER 1 THE BELKNAP PRESS OF HARVARD UNIVERSITY PRESS

Editor: Everett Mendelsohn, Harvard University Assistant Editor: Judith P. Swazey, Harvard University e

Copyright 1970 by the President and Fellows of Harvard College

CONTENTS Science and Philosophy in Aristotle's Generation of Animals

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ANTHONY PREUS

Descartes' Physiological Method: Position, Principles, Examples

53

THOMAS S. HALL

Harvey and Fludd: The Irrational Factor in the Rational Science of the Seventeenth Century

81

ALLEN G. DEBUS

Towards a Synthesis: Population Concepts in Russian Evolutionary Thought, 1925-1935

107

MARK B. ADAMS

Historical Aspects of F. W. Putnam's Systematic Studies on Fishes

131

RALPH W. DEXTER

Vertebrate Paleontology, an Early Nineteenth-Century Transatlantic Science

137

PATSY A. GERSTNER

The History of the Naming of the Loblolly Bay

149

EDMUND BERKELEY ESSAY REVIEW:

The History of Embryology as Intellectual History

155

FREDERICK B. CHURCHILL

The J. H. B. Bookshelf

183

JOURNAL OF THE HISTORY OF BIOLOGY is published semiannually in the spring and autumn by the Belknap Press of Harvard University Press, 79 Garden Street, Cambridge, Massachusetts, 02138. Editorial Board: Bentley Glass, State University of New York, Stony Brook; Hebbel E. Hoff, M.D., Baylor University; Ernst Mayr, Harvard University; Everett Mendelsohn, Harvard University; Jane Oppenheimer, Bryn Mawr College. Advisory Editorial Committee: Enrique Beltr6n, Mexico; Georges Canguilhem, France; John T. Edsall, M.D., U.S.A.; A. E. Gaissinovitch, U.S.S.R.; Ralph W. Gerard, M. D., U.S.A.; John C. Greene, U.S.A.; Marc Klein, M.D., France; Vladislav Kruta, M.D., Czechoslovakia; Joseph Needham, England; Dickinson W. Richards, M.D., U.S.A.; K. E. Rothschuh, M.D., Germany; Conway Zirkle, U.S.A. Editorial Correspondence and manuscripts should be sent to Professor Everett Mendelsohn, Editor, Journal of the History of Biology, Holyoke Center 838, Cambridge, Massachusetts, 02138. Subscription correspondence should be addressed to Mrs. W. H. Carpenter, Harvard University Press, 79 Garden Street, Cambridge, Massachusetts, 02138. Subscriptions, which are payable in advance, will start with the first issue published after receipt of the order. Please make remittances payable to Harvard University Press. Subscription rates are $7.50 a year in the U.S.; $8.50 in all other countries; $4.50 for a single copy. Journal Design by David Ford

ScienceandPhilosophyin Aristotle'sGenerationof Animals ANTHONY PREUS Department of Philosophy, Harpur College, State University of New York at Binghamton

In the Generation of Animals, Aristotle tries to solve a group of problems which are central both for biology and for Aristotle's philosophical system. The biologist supposes that he has made some progress toward the understanding of life if he has understood how a new life begins; Aristotle shares this concern with origins, and the explanation of animal generation also has metaphysical importance for him. Individual animals are among his paradigm cases of "entities" (ousiai); if his metaphysical system (which depends so much upon the concept of an entity) is to be validated, then the generation of an entity must be explicable within that system. He sketches the sort of explanation which he supposes must be given, in the Metaphysics: When one inquires into the cause of something, one should, since "causes" are spoken of in several senses, state all the possible causes. E.g. what is the material cause of man? Shall we say "the menstrual fluid"? What is the moving cause? Shall we say "the seed"? The formal cause? His essence. The final cause? His end. But perhaps the latter two are the same.' 1. Eta (VIII) 4, 1044a33-37, translation by W. D. Ross, The Works of Aristotle, vol. VIII, Oxford, 1928. Cf. A (I) 6, 988a5-8; A (V) 1, 1013a30; Z (VII) 8, 1033b23ff; 0 (IX) 7, 1049alff; A (XII) 3, 1070a28; 4, 1070b30; 6, 1071b30; 7, 1072b30ff. (In references to Aristotle's Metaphysics, A=Alpha a=little alpha==II, B=Beta==III, r=Gamma==IV, ?\=Delta=V, -I, E=a Epsilon = VI, Z = Zeta= VII, H = Eta =VIII, g = Theta= IX, I = Iota ==X, K= Kappa =XI, A==Lambda=XII, M=Mu=XIII, N=Nu=XIV.) In putting the GA (Generation of Animals) very near the end of the series of Aristotle's compositions I am not simply following W. Jaeger, (Aristotle, Oxford, 1934, pp. 329-341), but also I. Block ("The Order of Aristotle's Psychological Writings," American Journal of Philology, 82 Journal of the History of Biology, Vol. 3, no. 1 (Spring 1970), pp. 1-52.

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Here the explanation of generation seems to be put forward as a model of explanation; Aristotle shows no fear of the difficulties of his problem. Sketched briefly, the problem is of the change from non-being to being. Since Parmenides, philosophers had had to deal with the problem of change: how is change thinkable? How can a changing world be understood? Roughly speaking, Parmenides had argued that non-being is unthinkable, that all change involves non-being, and therefore that change is unthinkable. Philosophers before Plato had tried to get round this argument by trying to find a way to deny that change involves non-being (perhaps the atomists were an exception). Plato did grapple with the problem in his later dialogues, but Aristotle is clearly dissatisfied with the solution proposed by Plato in the Timaeus (Generation and Corruption II.Iff). Aristotle's theoretical solution is to be found in Physics V-VIII; it depends upon the persistent application of the distinction between potentiality and actuality to each variety of change which may be distinguished, as this distinction was made central in the definition of 'change' given in III.1: "the actualization of that which is potentially, as such." This general definition works out fairly well in the case of change in any of the categories where the "is" of the definition is predicative, that is, except the first. Change in the first category (entity) must, however, involve a radical difference in the sense of the definition itself, because in this case the "is" is existential. Plato had made this problem disappear by analyzing the entities in the phenomenal world into so many attributes, fleetingly found together. This solution seemed to Aristotle no solution at all, since one of the most obvious truths about the phenomenal world was that it was composed of things, of men and animals, and so on. Whatever else was to be done, entities must not disappear in the metaphysical analysis. Aristotle's solution is, none the less, based upon much of the Platonic solution of the problem of existential change. For Plato, the phenomenal attributes had their reality, their being, in some relation to an eidos, which Plato understood as an eternal object. For Aristotle, phenomenal entities have their reality as examples of an eidos, which he understands as an (1961) 50-77) and Pierre Louis, in the introduction to his Aristote: de la GCztration des Animaux (Paris: BudE, 1961). I take this opportunity to express my appreciation of the help of the late Professor Ludwig Edelstein, who first suggested this line of research to me, and of Professors Edward N. Lee and George Boas, who read earlier versions of this essay and offered valuable suggestions.

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Aristotle's Generation of Animals everlasting object existing in and through the individuals. For Plato, only an eidos was an ousia (entity, "real being"); for Aristotle, both the individual and the species (eidos) may be called ousia. However, for both philosophers, everything which is not an ousia has its being in and through some relationship to an ousia. A potentiality (dynamis) is such in relation to the ousia which it can become or can bring about in something else. Potentialities answer the Eleatic problem of non-being to Aristotle's satisfaction, because that which is potentially an X is not an X, but it is not nothing either. Potentiality gives Aristotle a way to save both ex nihilo nihil fit and substantial (existential) change. There need be no absolute non-being (no void of the atomists); everything which can be made into something more complex is matter for such a generation; it is potentially the result of such a generation, but is at the same time not-(some particular) being. The active power belongs to something which already has the eidos (form, species) actually, but which is potentially the generating cause of a (numerically) different individual with the same eidos. While the source of movement is actually a member of the species, it is not (even potentially) another member of the same species; rather, it has the potentiality of causing such an individual. In this way it too generates from a kind of relative non-being. So much may be taken as an all too brief summary of the teaching of the Physics and Metaphysics with regard to the problem at hand. In the Generation of Animals (GA), Aristotle turns to the problem of showing that his metaphysical analysis does indeed account for the phenomena of generation. The application of the metaphysical concepts to the phenomena results in a number of problems in explanation for Aristotle. In this essay I shall try to describe some of these problems, how they arise, and how Aristotle tries to solve them. In GA I, Aristotle uses his concepts of matter and form to provide a starting point, at least, for the explanation of generation. The rigorous distinction between matter and form assists in the refutation of some earlier theories of generation, but it also causes difficulties for Aristotle's own account. In Part A we examine how Aristotle's problem concerning matter and form arises in Book I, and how an attempt is made to resolve the difficulty. As a result of the investigations recounted in Book I, Aristotle is sure that the male provides the moving cause of generation;

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that which provides the moving cause also provides the form. In nearly all species, the male contributes to generation by means of semen; in order to fit the theory to the phenomena, Aristotle must show how semen transfers movement from the male parent to the material for generation, and how semen transfers the form (eidos) and the soul from male parent to offspring. In GA II. 1-3, Aristotle attempts a solution of the problem of the transfer of soul; he juxtaposes explanatory analogies with a subtle development of the implications of the distinction between potentiality and actuality, and he introduces (to the surprise of readers of the de Anima) a special material basis of the soul, which is said to carry the soul from male parent to offspring. In Part B Aristotle's explanation of generation as it appears in IL.1-3is critically examined, and some related problems which arise in the interpretation of these chapters are commented upon-among them, the problem of the origin of mind. Aristotle's explanation of animal generation often utilizes the ramifications of the concept of energeia (activity, actuality). This part of his explanation has the closest relationship with the theoretical philosophical positions expressed in Physics, Metaphysics, and de Anima, and is most typical of the more theoretical sections of the remainder of the GA. Part C discusses the explanation of generation which appears in II.4, and which relies heavily upon the concept of energeia rather than upon the concept of pneuma, the supposed physical basis of soul. Also discussed are the ways in which the concepts of matter, movement, form, and end are related to energeia in the context of the explanation of generation. Finally, we show how the developed concept of energeia is used in the solution of the problem, especially difficult for Aristotle, of the resemblance and lack of resemblance between parent and offspring. A. MALE AND FEMALE, FORM AND MATTER: BOOKI The major problem in explanation which Aristotle faces in Book I-and it plagues him for the rest of the work-arises from an initially too rigid application of the distinction between form and the active power, and matter and the passive power, to the natures of male and female.2 Matter and move2. 1.2, 716a2-bl3, applies the distinction to male and female, and Aristotle seems to be still committed to this rigorous application in IV.1, 765b9-15; it is all-important in the key passage at II.4, 738bl8-27, which summarizes

4

his explanation

of sexual

generation.

Aristotle's Generation of Animals ment are strictly segregated to female and male respectively, the female bringing about no change, and the male providing no material, for that which is produced. In I.17-23 Aristotle defends this thesis through the examination of a series of "problems" which his own theory proposes to solve more adequately than any other. The resolution of these puzzles leads to exposition of individual elements of the Aristotelian solution of the general problem of generation. 1. The refutation of "pangenesis" and preformationism; the male principle as "organizer" (I.17, 721b6, - 18, 724a13). The theory which may be called "pangenesis" held that the spermatic materials were drawn from the entire body of one or both parents, and that the fact that these materials were so drawn should explain the resemblance of offspring to parents.3 The tactic which Aristotle uses to refute this theory is typical, and our understanding of the refutation will help in the understanding of the theory which he develops for himself. He argues that even if parts from every region of the parents' bodies were to mix together, one would still have no reason to expect an organized whole, an organism, to result. If something were to come from a written word, the whole word, then it would have to come from each of the syllables, from each of the letters, and from the synthesis (combination). If flesh and bone are constructed from fire and so on, then the spermatic secretions would come from the elements only; how could they come from the synthesis? But without the synthesis there is no similarity. And if something makes the synthesis later, then this is responsible for the similarity, and not the fact that the spermatic secretions come from the whole.4 3. Compare the Hippocratic Airs, Waters and Places XIV, 19 (W. H. S. Jones, Hippocrates [Cambridge, Mass. 1957], II, 120-123), and IIEp' rovis (E. iUttre, Oeuvres complhte d'Hippocrate, reprinted Amsterdam, 1962, VII, 407ff) I, III, VIII, whose arguments closely parallel those refuted by Aristotle in the present passage. For an analysis of all these theories, and others concerning animal generation and development proposed by ancient Greek authors, including Aristotle, see Erna Lesky, Die Zeugungs- und Vererbungslehren der Antike und ihr Nachwirken (Abhandlungen der Geistes- und Sozial-wissenschaftlichen Klasse, Akademie der Wissenschaften und der Literatur in Mainz, 1950, no. 19). There is a great deal in the present essay which overlaps material in Dr. Lesky's treatise; since it is done from a different direction, and a different point of view, I hope that my essay will be complementary to hers. 4. I.18, 722a29-b4. Cf. Metaphysics Z.17, 1041bll-33.

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The arrangement of parts is not a material thing, but it is the principle of resemblance. Thus the pangenesis theory does not solve the problem which it is meant to solve. This refutation uses the principle upon which Aristotle bases his more general criticisms of reductionist theories of various sorts: they generally reduce any organizing principle out of their analysis. Aristotle's theory is precisely that theory which makes the organizing principle central, for his eidos, as the active power, functions as a kind of "organizer," in something akin to the modern sense of the word.5 The theory thus refuted supposes the individual parts to be scattered in the spermatic secretion, but another popular theory, which we call "preformationism," held that all these parts were connected, in which case it should be "a small animal." 6 The trouble with this theory is that it cannot explain how a female is, an animal with female animal could be generated-that parts rather than with male parts. The theory of Empedocles represents a compromise position between the Hippocratic pangenesis and the homunculus, or preformationist, theory, in that each parent seems to provide half the animal.7 Aristotle's objections to this theory, as he understands it, are perfectly straightforward: a) the parts could not remain healthy and alive if separated; b) it is not reasonable that two halves of animals should grow together to form one animal; c) the parts would have to be separated in more than two parts (up and down, right and left, front and back). In contrast to all these theories, Aristotle argues that one ought to look for some one material which can carry the active principle for making all the others; one should look for a 5. The term "organizer" seems to have been introduced by H. Spemann. L. von Bertalanffy, Modern Theories of Development (Oxford, 1933), gives a bibliography of the relevant works and explains the concept on pp. 121128. 6. 722b5. Aeschylus Eumenides 657ff, and Euripedes Orestes 552, use this theory as part of a defense of Orestes' matricide, that the male parent implants this homunculus as a seed in the female parent. Plato in the Symposium seems to subscribe to this view too (J. S. Morrison, "Four Notes on Plato's Symposium" Classical Quarterly 14 [1964], 42-55); Lesky, Die Zeugungs-, pp. 18-20, argues that Plato is not a preformationist in the Timaeus, at least, since he says at 91d that the seed is "still unformed" when put into the womb. J. Needham, History of Embryology, 2nd ed. (New York, 1959), pp. 34, 43ff, shows how popular preformationism has been. 7. Cf. Empedocles fragments 63-70 DK (= Diels, Kranz, Die Fragmente der Vorsokratiker, 6th ed. [Dublin/Zuirich 1952], I, 336-337); H. Cherniss, Aristotle's Criticism of Presocratic Philosophy (Baltimore, 1935), pp. 250ff; G. E. R. Lloyd, "Right and Left in Greek Philosophy," Journal of Hellenic Studies, 82 (1962), 56ff.

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Aristotle's Generation of Animals spermatic material which is not from all the parts of the body, but for the whole body. Thus the correct explanation must, in Aristotle's view, have an irreducibly teleological character. 2. The nature of the spermatic secretions, sperma. Having disposed, to his satisfaction, of some of the more popular theories of sexual generation, Aristotle must proceed to show how his a priori description of the character of male and female fits with the observed phenomena concerning the contributions to generation of the two sexes (he has described many of these phenomena in 1.3-16). His general name for semen and menstrual fluid is sperma; he proposes a definition of this word for examination: "Sperma wants to be this sort of nature: the first 'from what' in the generation of natural constructions." 8 Characteristically, Aristotle lists a number of senses of the word "from" with an eye to showing which are applicable in this case.9 As we should expect, the two senses which he chooses are "from matter" and "from something in that it is the source of movement." The source of matter and movement for the independent living body is, to a great extent, its food; it is reasonable to suppose that there should be some close relationship between both sorts of sperma and food. Aristotle finds this relationship by arguing that both the menstrual fluid and the semen are residues10 of blood, which is the final food of the body. Thus in a sketchy way (the full discussion is reserved for Book IV) the problem of family resemblance which had given rise to the pangenesis theories and their variations may be solved, for it is reasonable that "there is a resemblance between (the 8. 724a17. 1rphf0ro(first) poses problems of translation. A. L. Peck, Aristotle: Generation of Animals, Loeb ed., (Cambridge, Mass., 1953), takes it with yivcETat, translating "originally formed"; A. Platt, Aristotle de Generatione Animalium (Oxford, 1912 [Oxford trans., vol. V]), and Louis (Aristote: GA) correctly bring it closer to et ov;:"from it as their origin," "le principe d'o6 sortent," but these translations lead us to expect the word apx7 in the text. The above translation was suggested by L. Edelstein in conversation. "First" here, as often elsewhere, has something of the sense of "proximate." On the words which follow, see H. J. Drossaart Lulofs, Aristotelis de Generatione Animalium (Oxford, 1965), at 724a17. 9. The senses distinguished at 724a may be compared with those distinguished at Metaphysics A.24, 1023a26f (the most exhaustive list) and Metaphysics a (II) 994a20. The discussion of the senses of "from" at Physics I.7, 190a22, is closely related to the notion of genesis. 10. See Peck's discussion of nourishment and residues in the introduction to his Loeb edition of the Generation of Animals, pp. lxiii ff.

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blood) which is distributed to the various parts of the body and that which is left over" (I. 19, 726b14). Given Aristotle's notion that the "menstrual fluid" is the female sperma of the mammalia, the most obvious resemblance between blood and sperma is found in the female. The mense is obviously bloody in character; the problem which faces Aristotle is to show that it is in fact the material for generation.11 His argument in support of his notion that the menstrual fluid is spermatic in character is, in part, observational. He argues (727b) that females with no menstrual flow do not conceive, nor usually when the menstrual flow is in progress, but after the flow is over.12 This fits into his system as one more instance of the "mean," of not too much nor too little. Because it fits, and because there is no other candidate on the horizon, Aristotle asserts that the menstrual fluid is the spermatic contribution of mammalian females. He has already argued (726b30ff) that the female is weaker, 11. This is one of the more famous of Aristotle's wrong guesses; it is also one of the more pardonable, given the difficulty of finding a mammalian egg. Cf. Peck, Introduction to GA, p. xii, noting the discovery of the mammalian egg by K. E. Baer in 1827; George Sarton, "The discovery of the mammalian egg and the foundation of modern embryology," Isis, 16 (1931), 315ff, notes that the discovery was by no means easy even with a microscope. "The menstrual fluid," or "mense," is the standard translation of "the A problem arises in the interpretation of Aristotle's theory KaTcra747ta. in that menstruation, properly so-called, occurs only in primates, but Aristotle ascribes a spermatic function to the katamenia in all mammals. The beginning of a solution is simple: Aristotle supposes that the menstrual discharges in primates is the same in character as the estrous discharge in some other mammals. In Historia Animalium (HA) VI.18, 571b3-37, 581a5, Aristotle treats of the phenomena which appear in connection with heat in the mammals; in this passage he identifies the estrous discharge with the katamenia of human females at 572b28 and passim. S. A. Asdell, Patterns of Mammalian Reproduction (Ithaca, N.Y., 1946), p. 23, mentions estrous discharges in several mammals, notably the dog and cow. In the dog, the bleeding is proestrous, while in the cow it is metestrous. Aristotle mentions the proestrous discharge in dogs as a katamenia at HA VI.20, 573b30ff, and he supposes that the time of estrous discharge in the cow is the best time for impregnation VI.18, 573a3ff. 12. "After the flow is over" may seem a peculiar way of describing the period of peak fertility for human females, in whom ovulation occurs halfway between menses. But Aristotle might be thinking of some other mammals, particularly dogs, with proestrous discharges. The fact that he is thinking of mammals generally in this passage is borne out by the fact that he finds it necessary to explain fertilization during the flow, which is highly infrequent if not unknown in human females, but rather common in mammals which exhibit estrous bleeding, such as cows. Peck inserts the word "women" in his translation at 727b; it is not in the text at this point.

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Aristotle's Generation of Animals and therefore likely to produce that which has less form; that which has less form is matter; therefore the female is likely to produce the matter for generation. Combining the two arguments, the menstrual fluid should be that spermatic contribution which serves as matter for generation. 3. How mense serves as matter for generation. Given the assumption that the male provides the form and the female the matter (cf. 729alO-12), how can generation proceed? Aristotle actually gives several separate accounts of generation, each more sophisticated than the last, each from the point of view of a particular problem in his account. At I.20, 729alOff, he gives a provisional account into which mense as spermatic can fit. Specifically, he thinks he understands the role of mense through this analogy: "In the coagulation of milk, the milk is the body, but the fig juice or rennet is that which has the constructing principle. The semen of the male acts in the same way" (729al2-14). Both milk and mense are "residues" of blood; both rennet and semen contain vital heatthese are points made clear elsewhere.13 This analogy gives him a further occasion for the development of the consequences of his definitions of male and female in a "general rational account": There must be that which generates and the "from what"; this is true even if they are the same individual. They at least differ in form (eidos) and in that the definition (logos) of each of them is different; in those which have the powers separated, the bodies and the nature of the active and of x the passive are also different. Now if "male" means "mover" and "doer," and "female" (qua female) means "passive," the female would not contribute semen to that of the male, but matter. This is what is seen to occur, for the nature of y the menstrual fluid is in the class of proximate matter.14 13. HA III.6, 516al; GA I.20, 729alO; II.4, 739b20; 2, 735bl; IV.8, 776al5ff; 4, 772a22, cf. 771b23. 14. I. 20, 729a24-32. Peck translates, at x, "if the male is the active partner . . ." as though Aristotle were talking about individual males and females and the manner in which they have sexual intercourse; Platt's translation, "if the male stands for the effective and active ...," is much better. My translation stresses the account of the meaning of words. Peck and Louis have a quite different interpretation at y, where the text reads "KaT' T74 7pTP V'Xv." Peck translates, "the natural substance of the menstrual fluid is to be classed as 'prime matter.'" Louis translates, "la

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The force of the original definitions is increased by tying them in to the notions of eidos and logos, and the notion of power (dynamis). Male and female each has, per se, its own form and definition, and its own power. When said in that way, it sounds reasonable; the net result is the driving of a further wedge between the characteristics which one may expect to find in actual individuals of each sex. 4. How the male contributes to generation; the carpenter analogy. No ancient Greek author seems to have doubted that in most species the male contributes to generation by means of semen. Aristotle need not argue that this, rather than something else, is the male contribution. He has, however, more difficulty describing how semen contributes to generation, once he has rejected preformation as a possibility. His position is already clear, in one respect: the male provides the form, the dynamic structure. Even though preformation has been rejected, two possible ways of sharing in generation are still open to semen: (a) by mixing with and remaining present in and part of the generated body; (b) not sharing in the body of the embryo, except as a power and movement. The strict interpretation of the original definitions of male and female lead him to reject the first way and to accept the second. This will get him into considerable difficulties, particularly in 11.3; we must examine the argument which he offers in support of his choice. a

To those who examine the problem in general it does not appear that one thing is generated from the passive and the active with the agent immanent in that which is generated, nor in general from the moved and the mover in this way. The female, qua female, is passive, and the male, qua male, is active and the source of movement. If the extremes are taken, on the one hand the active and mover, on the other

nature des menstrues appartient au domaine de la matiere primordiale." v.x'q"as this term is Actually, Peck gives a quite good account of "irpwdirv used in the biological books in his introduction, pp. xi-xv. Aristotle is not referring to the "prime matter" of the Metaphysics, e.g. 0.7, 1049a24, for mense is "bloody," which qualifies as some sort of "thaten" in the sense of "thaten" explained in Metaphysics e. Metaphysics A.6, 1015a8-12, gives two senses of the phrase 7r pOarqU, primary in relation to the thing, and in general first. Menstrual fluid is primary in relation to the thing to be generated, or proximate. Cf. Metaphysics Eta 4, 1044al5-b2, where this proximate matter is called 'oiKfZoV', and Bonitz, Index Aristotelicus, 653b25ff.

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b

c

passive and moved, that one thing which is generated is not "from" them except in the sense that the bed is "from" the carpenter and the wood, or the ball "from" the wax and the form (eidos). It is clear then that it is not necessary for anything to be emitted by the male, nor, if something is emitted, does this mean that that which is generated is from this as immanent in it, but as from the mover and from the form, as one who is cured by the medical art.15

At a Aristotle gives what amounts to a deductive demonstration of his point, based on the original definitions. While the argument may be taken as disclaiming a position like that of certain modern vitalists,16 the question at issue is not precisely that, but whether the semen makes a material contribution to the offspring. Matter is by definition that which is acted on, not that which acts; anything that had both powers would change itself, but the female does not generate without the male.'7 Thus the definitions lead us to the conclusion that the powers are separate not only logically, but also in fact. The analogy of fig juice and rennet which had been used to illustrate the role of mense in generation was admirably suited 15. 729b9-22, partly following Louis rather than Peck. 16. I.e., that there is an internal entelechy which causes or guides the development of the individual living thing. I am thinking of H. Driesch, Philosophie des Organischen, 4th ed. (1928), in particular, and also E. Rignano, Biological Memory (London, 1926). 17. Aristotle recognizes cases of parthenogenesis. Apart from plants, in which sex differentiation is only analogically present, there are cases of animals which generate another animal like themselves without benefit of males: the fishes which he calls erythrinoi and channai (II.5, 741a35; I11.5, 755b21; III.10, 760a8; HA IV.11, 538a19, 567a27). The channa is rather clearly identified with Serranus cabrilla or Serranus scriba (D'Arcy Thompson, A Glossary of Greek Fishes [London, 1947], p. 283); the erythrinos is somewhat more difficult, but the consensus seems to be that Cuvier was right in identifying it with serranus anthias, a brilliantly red (erythrinos means red) fish. Alternatively, it might be any of several members of the Serranidae or Sparidae, like the "rouget," as this word is used in French (see Thompson, pp. 65-67). Some insects, particularly bees (III. 10, 759a8ff), generate parthenogenetically, thinks Aristotle, but sometimes something different and "lower" in kind than themselves. Aristotle never really grapples with the problem which results from observations of parthenogenesis for his general theory; in the case of the generation of bees, he says simply that "like plants, they have within themselves both the female and the male" (759b30). In the case of plants one may isolate the male and female elements in many species (e.g. the pistil and stamen, in many flowering plants), but he does not attempt to isolate the male and female elements in the case of the parthenogenetic (and perhaps hermaphroditic) animals; this was done in the case of the hermaphroditic species of Serranidae beginning in the eighteenth century (see Thompson, 283-284).

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to show the way in which a natural material serves for change, but it was not a close analogy for pointed change, for fig juice or rennet and milk result in cheese, but semen and mense may generate a living being. The analogies used to illustrate the role of semen are not deficient in this respect, for they are frankly teleological in character. This is not an unreasonable difference between the two sets of analogies, for although matter is to be understood teleologically, the end is more truly present in that which has the form and source of movement. The source of movement and end are most closely related through the notion of form, for a movement is a pointed movement in virtue of its form, the structure of the movement itself (which may be per se valuable), and the normal results of that movement in a material, if they are valued results. The use of these analogies at b and c in the passage quoted leads us to a further understanding of the thought process which led to the rejection of the theory that semen participates materially in the generated embryo; they are examples of the general tendency to draw analogies between the workings of art and the workings of nature. These analogies lead Aristotle to suppose that the fact that the artist does not become a material part of his product shows somehow that the natural generator does not become a material part of his product either. The carpenter analogy is explicitly used in this way: he makes the bed, but neither he nor a part of him becomes or becomes a part of the matter of the bed.18 The wax analogy emphasizes the distinction between the form and matter of the particular thing. If it is true to say that the male provides the form, someone might say that he is providing a "part" of the generated thing. Aristotle simply reminds us of the distinctions made elsewhere (de Anima II.1, 412b7-9; Physics VII.3, 245bll, et al.), that the form and matter of the individual are one, and not "parts" of the thing. It is odd to find the form of the wax ball classed with active principles in this place; all I can do to reduce the sense of strangeness which I feel with that notion is to point out that in the case of the male principle, at any rate, the form is thought to be provided as a movement. The analogy of the medical art brings the analogical account closer to the circumstances of the natural event to be ex18. The carpenter analogy was first used in the GA at 1.18, 723b28; it is further developed at I.22, 730b6ff; II.4, 740b25; and 11.6, 743a25. In Parts of Animals (PA) 1.1, 641a, and 11.7, 652b15, there are related uses of this analogy.

12

Aristotle's Generation of Animals plained. We are reminded of the passages in the Metaphysics and Physics in which the medical art is taken as paradigmatic in the understanding of nature, and of many passages in the biological books in which a medical predicate is attached to "nature." 19 Aristotle's trust in the general analogy between natural and artificial generation is one of the causes of his only slightly modified confidence that the matter for a particular generation is totally passive in respect of that generation, and that the source of change provides only form and movement, and no material contribution. I believe that this trust is greater than is strictly demanded by the metaphysical presuppositions which govern his explanatory scheme. In Physics I1.1, 192b24, the doctor who cures himself is said to do so "accidentally." This is taken to be a general truth about the relation of efficient and formal causes on the one hand, and matter on the other, in all the arts. Yet the products of nature are taken to be analogous to the doctor who cures himself, except that in nature this process is essential, not accidental (192b14). If these distinctions are taken into account in relation to the use of the analogy of the medical art at GA I.21, 729b, one notes that not only semen but also mense is a natural product; mense exists physei, and should therefore have a source of movement within itself. Thus Aristotle has a possible alternative position regarding the activity and passivity of these contributions to generation: the mense may have some source of movement, some form, some power, even active, but it may lack something which the male can provide. Under this interpretation, parthenogenetic production of things different in kind becomes comprehensible: if parthenogenetic reproducers did have that which a male could provide, they could produce things like in kind to themselves. Something like this alternative solution does begin to appear in the later books, not only in the case of the generation of bees, but also in the various discussions of the generation of infertile eggs,20 and most of all, in the discussion of the resemblance of offspring to parents, in Book IV. 19. E.g. Metaphysics A (XII) 3, 1070a30; 4, 1070b30; Z (VII) 7, 1032blff; Physics II., 192b24; PA I.1, 639bl5ff. Some of the many examples of Onts 1a&rpeuKe and the like are cited by Bonitz, Index Aristotelicus, 837a5ff. 20. Wind-eggs are discussed at length at 1I.5, 741a; I11.1, 749a5ff; and briefly elsewhere. Aristotle might have concluded that the viviparous animals also produce eggs, by analogy from the oviparous and especially ovoviviparous animals, but the near impossibility of finding a mammalian egg without a microscope effectively prevented this hypothesis; Aristotle is A (v) 12, 1019al5ff;

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However, in Book I, Aristotle remains committed to the radical separation of matter and maker; this commitment may be seen in his attempt to find (at 729b22ff) empirical evidence in support of this separation. Most important to his case, if accurate, is his supposed observation that some insects copulate without transfer of semen from male to female.2' In I.22, 739b27-32, Aristotle fits this case into the analogy of art and nature with evident satisfaction: It is as if someone were to bring the matter to the craftsman. Because of the weakness of this sort of male "the nature" is not such as to act through something else; the movements seem scarcely strong enough with nature sitting right there. Here nature seems to resemble modellers in clay, not builders, for she does not fashion that which is constructed by means of something else, but does it with her own parts.22 In further support of his case, he cites the production of infertile eggs by birds prior to copulation. The implication drawn is that the material is all there; only the cause of growth and development must be missing. He goes on to cite the practice of chicken breeders of putting a second cock to the hen in hope of improving the breed, believing that the brood would resemble the second cock.23 Aristotle seems to think that the second cock adds to the vital force of the offspring, and gives it some of its own valued characteristics. As a final bit of evidence, Aristotle calls attention to the mode of fertilization in oviparous fish, in which the male sprinkles milt over the eggs; those which the milt touches are fertile, while the others are not, "as though the male contributes not to the quantity but to the quality" (730a22). Aristotle is now ready to account for semen, which could readily have been thought to become some part of the gensufficiently empirical in his approach that he must account for the facts as observed. His theory is developed in such a way as to account for the absence of eggs in mammals. 21. He claims that the female inserts a part of herself into the male, and that the material is thus worked up by direct contact with the internal organs of the male insect. This error possibly arises from taking the ovipositor, as seen during oviposition, for the phallus, as seen during copulation, and confusing male and female between the two events (see K. G. Davey, Reproduction in Insects [San Francisco, 19651). 22. 730b27-32, after Peck. Incidentally, the passage counts against, rather than for, the notion that Aristotle's physis is a transcendent individual, for the physis which works here is that of the male animal. 23. Platt's note to 730a18 tells the source of this notion.

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Aristotle's Generation of Animals erated offspring, in terms of his carpenter analogy. In the case of the carpenter, the form is present in his soul, and is generated in the matter by means of the movement, and his soul and knowledge move his hands or some other part in a particular movement, different for different products ... his hands moving the tools and the tools the matter (1.22, 730b13-19). Aristotle here regards carpentry as strictly analogous to the male role in sexual generation in that semen is viewed precisely as a tool, for the tool does not become a material part of the production (not in carpentry at any rate), but serves to transmit the formative movements from the maker to the made. The distinction which Aristotle finds here between semen and the tools of carpentry is this: the semen is a tool which "has the movements in activity," while the tools of the arts have their movements "somehow." Semen has an actual "disposition" 24 to act in a certain way; tools of the arts have certain potentialities, powers, but they are not in themselves sources of movement; semen is a source of movement. 5. The accomplishment of GA I. Aristotle set out, in the first book of the GA, to describe many of the phenomena related to generation, particularly phenomena related to sex differentiation (we have not discussed this aspect of the book), to refute some of those who had proposed explanations differing from his own, and to understand the phenomena of sexual generation in terms of his own philosophical system. The major distinction which Aristotle uses in the explanation of sexual generation in Book I is the distinction between matter and form. I have argued that he gets himself into difficulties through his rigorous application of this distinction to the distinction between male and female; he seems to have been led to this rigorousness as much by the explanatory models, the carpenter and the doctor, as by demands of his philosophical system. The application of the distinction between matter and form and the analogies of art and nature as presented in Book I are understood by Aristotle as substantially correct, since he continues to use them through the GA, but they leave problems in explanation which force him to more drastic measures in 24. "hexis": see his definition at Metaphysics A (v) 20, 1022b3ff.

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Book II. There is a certain plausibility in the view that the menstrual fluid serves as the material for generation, for the body is fed by blood once it is born; it should have been made of blood, or something very like blood, in the first instance. But the notion that the semen becomes no material part of the generated embryo would seem implausible to many members of Aristotle's audience. Thus the climax of the first book is the explication of this relatively unpopular thesis. Aristotle's account of the soul in the de Anima had identified the soul, or all the soul but mind, with the dynamic structure of the body; as a first approximation, it seems to him appropriate to argue that the male has as its function the installation of soul in body. In defending this position it is constantly necessary for him to argue against those positions which identify the soul (psyche) with some material. As far as it goes, the analogy of craftsman and procreator seems appropriate as a defense of the position. Aristotle knows very well, however, that his materialist opponents can persuasively argue that the relevant difference between craftsman and procreator is just that the procreator passes on soul, and the craftsman does not. Aristotle must, therefore, defend the validity of the analogy. In order to do so he will have to explain just how semen functions as a tool for transferring soul from procreator to offspring. This is the major problem of the first part of book II. B. THE PROBLEM OF THE TRANSFER OF SOUL (GA II.1-3) In II.1-3 Aristotle takes up the account of generation from the direction of a teleological analysis. In Book I a teleological theory of generation operated implicitly rather than explicitly; the notions of "point" and "end" are implicit in the notion of form, and essentially involved in the analogy of the craftsman, but Aristotle does not exert himself to make these points crystal clear. In the second book he begins with an account of the teleological principles which must operate in an adequate explanation of animal generation, particularly sexual generation, and then works back to the problem of the role of semen in generation. When the problem has been laid out, he proceeds to examine the "nature" of semen, and finally to give an account of its role in generation which he hopes will be acceptable. 1. The purpose of sexual generation (II.1, 731b18-732a25). The question "why is there sexual generation?" may be analyzed into "why is there animal generation?" and "why are 16

Aristotle's Generation of Animals there sexes?" The answer to the first question is easy enough. Animals (or plants) cannot last forever as individuals, due to the imperfections in their matter, their potentiality for nonexistence. However, everything that exists aims at continued existence. This is possible for a species of animals or plants, through reproduction. Thus the purpose of reproduction is the continued existence of the species. The normal mode of reproduction in animals is sexual, so the distinction of sexes exists for the sake of reproduction. Aristotle explains the distinction between the sexes thus: As the proximate moving cause is better and more divine in nature, in that the logos and eidos are present in it, than the matter, so it is better that the superior be separate from the inferior. That is why the male and female are separate in as many species and to the greatest degree possible; for the male is better and more divine in that it is the source of movement in generated things; the female is the matter.25 Aristotle argues in much the same fashion in de Anima 11.4, 415a26ff, in his explication of the nutritive and generative function of the soul; in Generation and Corruption II.10 there is also a general relation established between the stars, which are everlasting as individuals, and the continuation of species on earth.28 Several commentators have argued that the GA gives a large place to "material necessity" in explanation, instead of and in opposition to teleological explanation.27 The present passage, particularly given its placement in the book, should discourage the view that Aristotle has softened his position on teleology in nature. We note that he relates closely the moving and final causes in this passage. Indeed, he regards the "participation" of the source of movement in the end as sufficient reason for the distinction between the sexes and the purported superiority of male animals over female anrimals. This tack of the argument is by no means accidental, for Aristotle is aiming at a final defense of the notion that the male provides the soul and the female the matter. If the notions of "male," "moving cause," and "better" can be closely 25. 732a2-10. Peck's note to the passage is useful. 26. See Metaphysics A (XII) 10, 1075al2-24, for the ordering of the universe; GA IV.10 discusses the cycles of animal generation in relation to the astronomical cycles; Peck's notes to the passage are valuable. One may treat GA II-IV as an explication of Generation and Corruption II.10. 27. E.g. Pierre Louis, Introduction to the Bude GA, pp. x-xi.

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related in the minds of his audience, he is that much closer to convincing us of the truth of his position. The arrangement of the exposition may owe something to his view of rhetoric (Rhetoric I.1, 1355b27; Topics I.2, 101a30-34), for he seems to begin his argument in Book II with something which his audience, presumably male, will be ready to accept: that the male is superior to the female. He can then proceed to argue that it is better to provide the form than to provide the matter. From these premises it "follows," by a kind of rhetorical move, that the male provides the form rather than the matter. 2. The statement of the problem of semen (733b23ff). "It is necessary for that which is generated to be generated from something and by something, and to become something."28 The "from what" may be treated summarily; it is the matter, whether mense or egg, or the material gotten through the umbilicus, or the later milk, and other food. The new individual, the new entity, is a "this something" of the same species as its parents (as in Metaphysics Z [VIII 7-9, 0 [IX] 8). The problem at hand is the "by what," the moving cause of generation. Aristotle develops and tries to resolve (733b23-735a29) a kind of dilemma in regard to the relationship between semen and the notion of a source of movement. It seems reasonable to suppose that the source of movement for generation, the "by what" of generation, should be either within or external to the semen. It is unreasonable to suppose, Aristotle claims, that something external makes each of the parts of the embryo, particularly in the case of the internal organs, because nothing can be set in movement except by something which touches it.29 So the construction of the internal organs requires the presence in the fetus of something which either as a part of it or as separable from it constructs at least these parts. But it cannot be separate or different from the fetus itself: if there were such a thing, it would either remain or disappear. If it were to remain, it would be distinguishable, a detectable entity within but not a part of the living body, but no such entity 28. 733b25. This principle is familiar from Physics, Generation and Corruption, Metaphysics Z (VII) 7-9, 0 (IX) 8. 29. Cf. irp6s at 730b27; Physics VII.2 notes that air is often the medium of movement. We shall see how Aristotle uses pneuma as an analogous medium of movement in the solution of this problem.

18

Aristotle's Generation of Animals is discoverable.30 Furthermore, it is "absurd" to suppose that it fashions either all or some of the parts and then disappears. If it fashions only some of the parts, then something else fashions the rest; suppose that something else to be the heart -does the heart disappear when it has fashioned the rest of the parts? Aristotle does not really consider here the possibility that it fashions all the parts, then disappears; apparently that would be open to the same criticism as the persistence alternative, for an organizer of that kind would be apparent in semideveloped individuals. Furthermore, whatever fashions all the parts ought to be the same as that which maintains them. Thus whatever it is that fashions the animal persists, and since no such separate entity can be distinguished, we must suppose that it is part of the whole, and present in the semen from the outset, if the semen is that which has the source of movement and development. One might, alternatively, argue that there is some part of the body of the fetus which is present in the semen, and that this "part" constructs and is the source of movement for the rest of the body of the fetus. This suggestion would indeed make the source of movement "internal" at every stage, but Aristotle argues that this position has as many difficulties as the other; the agent must be either internal or external, but it cannot, it seems, be either. This is the dilemma. He argues that it cannot be internal in the manner proposed on the following grounds: either it forms all the parts simultaneously or it forms them successively. By observation we know that the parts are not formed simultaneously, because the heart appears before other parts which are ultimately larger than it. So the parts are formed successively: but is this succession causal, or merely temporal? If the succession is merely temporal, then we must still seek the cause of the generation and the cause of the succession; but if the succession is causal (if the part that is in the fetus makes the heart, and the heart makes the liver, for example)31 then because that which is potentially (dynamei) is generated by that which is in actuality (entelecheia), the heart would have to have the form and shape of the liver. Such an account, he says, would be an absurd fabrication. 30. Note that Aristotle does not entertain the notion that there might be such a thing, unobserved by him. 31. 734a28. Aristotle does not seem to notice one possibility: the part in the semen might be the embryonic heart itself. Although his examination of fertilized chicken eggs might have led him to suggest this possibility, it does not seem supportable by observation of semens.

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It seems to Aristotle, then, that no part of a plant or animal may be present in the semen or seed, whether it makes anything or not; for semen is made by the male parent and is prior to any part. The argument for this position may be restated thus: we are looking for the agent, the source of movement, of the making of the parts and of the whole of the animal; nothing can be maker of itself simpliciter (this is what Aristotle means when he says that the semen is prior to any part). It follows that semen is not, nor does it contain, any part of the animal which is made, if the semen is the source of movement for generation. Thus the dilemma: the agent must be either internal or external, but cannot, according to these arguments, be either. In general, the way to resolve a dilemma is to make at least one distinction, which Aristotle does: "In what sense is it not possible for something to be generated by something external? In one sense it is impossible, in one not." (734b5-7). It is not possible for something to be generated by something external in the sense that it is "separated and not touching," since if it does not touch it cannot have a causal effect. But "external" is subject to some interpretation. There might be something which sets up a movement in a second thing, which in turn effects the movement or change in question. This interpretation, too, seems to bring about the same dilemma, for each mover in the series of movers must be either internal or external to the next member of the series. It might be suggested that semen begins as internal to the male and ends as "internal" to the female, or more properly to the embryo, but then semen must be discoverable in the embryo, which it is not. 3. Toward solution: two analogues and potentiality/actuality

(734b5if). In order to clarify the possible distinction in the senses of internal and external, as applied to moving causes, Aristotle uses an illustrative analogue, that of the self-moving puppets. He also discusses the principle behind the use of this analogue, the distinction between potentiality and actuality: "The parts of these automata have somehow the potentiality in them while at rest; when something external sets the first moving, the next becomes actually."32 According to Aristotle, 32. GA I1.1, 734bll-13,

20

after the texts of Peck and Lulofs.

Aristotle's Generation of Animals if something is to be a source of movement, it must have an active power (potentiality) for that sort of movement or change. An agent which may initiate a series of movements may simply have the active power, and the matter for the change may simply have the appropriate passive power, but anything which is to serve as an intermediary between agent and matter must be both passive and active-passive in relation to the agent, and active in relation to the matter, or in relation to the next member of the series. Such an intermediary is known as a "moved mover." This characteristic of a moved mover may be understood as the same power; for example, the potentiality of some materials for being heated is at the same time the potentiality for transferring that heat to something else. It is in this way that the puppets are understood as passive in relation to the puppet-master, and active in relation to the movement which they initiate. Semen, it is suggested, is similar to the puppets in respect of this kind of potentiality; semen is understood as a moved mover. One may be uneasy with this analogy, however, for semen is not a mechanical device, but a natural product. Pushing the puppet analogy might even reintroduce the preformationist theory in a new and peculiar form. In order to still this uneasiness, I suppose, and to tie the present account to that given in Book I, Aristotle introduces another analogue. He claims that the power of the semen which we are discussing is within the semen in the way in which "housebuilding is in the house" (734b17). Housebuilding is in the house, we may suppose, by virtue of the activity (artistic and intentional) of the builder. Semen is not as mechanical as a puppet, but it is not as human as a builder; the puppet and its parts are moved movers, as semen should be a moved mover; but housebuilding is a kind of unmoved mover, house movement is internal, and initiative of a series of movements. Aristotle no doubt means to suggest that the truth lies somewhere between, but one wonders just how to find that middle ground. The conclusion of the present set of arguments is, as a result, more negative than positive: "It is clear now that there is something which makes, not as an individual, nor present as the first perfected thing."33 Semen is "something" but not as an individual, not as an ousia. That which builds is, strictly speaking, a movement (kinesis), or, in the language of Parts of 33. II.1, 734bl8-19. For the idea of an individual in Aristotle, see Joseph Owens, The Doctrine of Being in the Aristotelian Metaphysics (Toronto, 1957), pp. 334-335; Metaphysics 0 (IX) 7, 1049a22-36.

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Animals (I.1, 641b32), a genesis or "becoming": "Seed (sperma) is a genesis, but the end is an entity (ousia)." Peck translates genesis in this place as "formative process"; a "becoming" or formative process is not thought of as an entity. Aristotle, by referring to the art of housebuilding at 734b, rather than to the builder himself, wishes to abstract or separate the movements from the individual who effects them. Even the puppet is an individual of a sort, but evidently semen is not. It has a form, but only "as a power" and not as "something perfected." 4. Semen and the distinction between energeia and entelecheia (734b21ff). Whatever is generated naturally or artificially is generated by something which is "in activity" (energeia) from something which is such "potentially" (dynamei). Semen is such, and has such a movement and principle, that when the movement stops each of the parts is generated and is ensouled (734b21-25). This is an approach to the positive solution to the problem of the activity of semen. The key word in the understanding of the middle ground which Aristotle attempts to take is energeia, which I here translate "activity," but is often translated elsewhere "actuality." In the passage which follows, and in general, the movement in the semen is described as an energeia rather than as an entelecheia. The latter word does convey a sense of perfectedness, and is reserved for the parent and the offspring, while the semen can only be or have an energeia (734a30, b35, 735al9). The distinction between energeia and entelecheia is precise in the case of the productions of a craft; thus Aristotle calls attention (734b28ff) to the way in which the analogy between art and nature may work out in this regard. Specifically, heat and cold do not make the logos of their product either in art or in nature; whether used in art or nature, it is the "logical movement" which makes the entity. Thus the movement of the tools has the logos, or form, of the art, and this is its energeia, while semen has the logos of the species, and this is its energeia. Aristotle recognizes that using the analogy between art and nature in this way makes his problem arise again, in yet another way: "for art is the source and form of that which is generated, but 'in another'; but the movement of nature is 'in

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Aristotle's Generation of Animals itself,' 'from another' nature which has the form actually" (735a2-5). The translation preserves the opacity of the original in respect of the term "in itself"; one immediately supposes that the "itself" referred to in this case must be the semen, but that is not said either. In the context, Aristotle must show that the movements proper to the eidos are in the semen, but if the semen has these movements, one might suppose that it is a member of the species, for it is matter which has, somehow, the form of this sort of nature. 5. Does semen have soul? (735a5-26) Thus arises the question, "Does semen have soul?" This question takes us to the heart of the matter. In one way, the answer must be affirmative, for semen effects natural production, and only that which has soul can effect natural production. In another way, the answer must be negative, for semen is not an individual entity. Aristotle discusses this question twice in GA II. Book II.1 ends with the assertion that the explanation of generation has been given, leaving only the physical analysis of semen, taken up in I.2. In II.3, however, the question of the relationship of semen and soul is again raised, this time at greater length and depth. The ways in which these passages answer this question differ, since one is predicated on the philosophical analysis and the use of analogies, while the other succeeds a physical investigation of semen. The philosophical argument proceeds thus:

x i ii

The same argument holds for semen as for the other parts of the body: a) there is no soul elsewhere than in that of which it is the soul; b) a part which does not share in soul is a part only "homonymously," like a dead man's eye. It is therefore clear that semen has soul and is soul potentially. The same thing can be potentially in varying degrees, as the sleeping geometer is farther than the waking geometer, and he is farther than the working geometer. Now no part is the cause of this generation; it is rather the first external mover. For nothing generates itself, but when it has been generated, it grows by itself.34

34. II.1, 735a6-14, after Peck. At ii a problem is posed by the word auToU; Louis gives an interpretive translation which gives the correct force to this word: "une meme chose peut etre, en puissance, plus ou moin loin de se reahiser." F. Nuyens, L'Evolution de la psychologie d'Aristote (Paris and Louvain, 1948), pp. 258-259, emphasizes the two negations, marked a) and b) above. "Les deux negations se complMtent 1'une 1'autre.

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It is for this reason that the principle of growth, that which has the nutritive and generative power, is generated first. Aristotle goes on:

x

To be generative of another like itself, this is the function of every animal and plant perfect in nature . . . Though it was generated by something bearing the same name (a synonymon) as a man generates a man, it grows by itself; thus there is something which makes it grow.35

Thus the heart or its analogue is generated first. In an important respect, the problem of semen and soul is evaded in this passage. Aristotle brings himself to say that semen is an organic part of the male parent, and all organic parts share in soul (735a8), namely, the soul of the individual living being of which they are parts. But this is not enough, as can be seen for the ambiguity at i in our translation (Ka't KaL ZuTL [ivx'] Suva'EL). On the one hand, the word dynamis has the effect of taking away something: not actually, but potentially, semen has and is soul. On the other hand, one is X tempted to read the phrase as if it were written "Kai EXEL LaTt soul. of the is the r'nsOvXs 8Uvatits"-both has and power The first interpretation is reinforced by reference to degrees of potentiality. At de Anima 11.5, the same doctrine is applied at length to the account of perception and knowledge. At 417b16, Aristotle speaks of degrees of the potentiality of knowledge, claiming that learning is a change in habits and nature: "The first change in the faculty of perception is generated by the male parent; but when generation is effected the individual already has knowledge, in the sense of perception." The passage in GA II has the effect of inserting the activity of semen into this process. It would seem that semen has the potentiality of perception, but even farther from actuality than the potentiality which the embryo has. Semen must, however, have actually the power of generation, and if it has that power, then it "has and is" soul. We wonder eXIE

Kai

L'ame ne se trouve que dans l'etre vivant, mais non dans un organe particulier de celui-ci. Bien plus, on ne peut meme pas parler d'organe a moins que celui-ci participe a l'&me. L'ame est la force vitale qui, au sein de l'etre vivant, exerce son influence dans tous les organes." If he had said "puissance vitale" his interpretation would be more clearly correct. His interest in this passage is its hylomorphism, easy enough to demonstrate; he does not deal with the problem which hylomorphism causes here for Aristotle's account. 35. 735al8-23, after Louis in text and translation; Lulofs supports this text. Cf. de Anima II.4; Nuyens, Psychologie d'Aristote, p. 259.

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Aristotle's Generation of Animals again just what a dynamis might be. If a dynamis may be ontologically independent, as hot, cold, fluid, and solid seem to be, then perhaps a power of soul may exist without its being the soul of an individual. Aristotle seems to suppose that the soul is a power, or closely interrelated group of powers, which may be the soul of an individual, to be sure, but which may also be transferred by means of semen without being the soul of anything at the moment. Given the kind of statements which Aristotle has made to this point in the GA, we may suppose that the powers of soul in the semen are present as heat, as movements. The male parent imparts movements to the semen, and the semen in turn is said to impart movements to the menstrual fluid. It therefore seems to be the case that semen "transfers" the powers by preserving the form of the movements by which it was itself moved and by imparting them to the menstrual fluid which it finds in the female. Aristotle uses the ambiguity inherent in the word dynamis in his account of semen and soul in II. 1. Semen is, in respect of its dynamis, soul. But the actuality of soul is its exercise, its activity. Semen does not have the sensitive soul in activity; thus it is not an animal. It does, in a sense, have the generative faculty of soul, for its activity is precisely generation.36 It generates an animal, which has the faculties of perception and movement. These latter faculties are thought of as the same power in parent and offspring; this power must be in the semen too, but "potentially." Thus we get the "potential presence of a power" all packed into the word dynamis at 735a9.37 Supposing that we were to accept this theoretical account of the character of semen, there would still remain a specter of the continuing problem which Aristotle faces in his account of generation. This specter is suggested by the pair of terms "homonymous" and "synonymous" at 735a8 and 735a2l, marked "x" in the translation. On the one hand, semen would be only "homonymously" a part if it had no soul; it would be like a dead man's eye. On the other, generation is effected by the "synonymous," as man generates man. Semen is not even homonymously a man, although it is part of a man; it cannot, 36. A problem remains: semen does not grow, so does it really have even the nutritive and generative soul? There are at least two examples of female spermata growing, noted by Aristotle (III.2, 752a24ff, III.4, 755allff), but none of the male sperma, semen, which is at issue here. 37. Cf. Nuyens, Psychologie d'Aristote, p. 259.

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it seems, be said without qualification that semen generates.38 For the time being, it seems that the dilemma posed at 734a (p. 18 above) has been theoretically solved. Generation, it would appear, is effected both by something external and something internal. The male parent is external, but he is the source of the power of soul which is in the semen; the semen ensouls the menstrual fluid largely by fabricating the embryonic heart, which in turn "makes" the rest of the animal. 6. Semen as phenomenon (II.2, 735a30-736a24). In order to show how semen can be the sort of material to be able to carry, to transfer, such a complicated set of movements and powers, Aristotle provides an observational description of semen and attempts to explain its non-generative behavior. This explanation turns upon the hypothesis that semen is a foam of water and pneuma; in support of this notion he calls upon Hesiod's derivation of the name Aphrodite from aphros, or foam. Aristotle is gradually working in his (new) theory of pneuma as the special physical basis of the soul. For the most part, pneuma had no more special connotation of soul in the Greek of his day than does "breath" in English, and Aristotle does not show his hand in this chapter, for he says simply that pneuma is "hot air" (736a2). There is little indication of the role that pneuma will play in the next chapter, where it is even true that the physical characteristics of semen are again investigated (736a30ff). 7. The extended analysis of semen and soul (11.3, 736a24737a34). A seemingly innocent question, "What becomes of the physical part of the semen when it has done its work?" (the problem was already noted at 734a8), introduces one of the most important chapters in the GA. This question arises from Aristotle's position that the male provides the form and the female the matter; semen should provide none of the matter for the embryo (fetation), but it does provide the soul. If it provides none 38. "Homonymous" and "synonymous" are, of course, technical terms developed in the context of Aristotle's logic. It may be noted that these adjectives are as applicable to things as they are to words. Two things are synonymous if they bear the same name and belong to the same class; they are homonymous if they bear the same name but do not belong to the same class, i.e. do not have the same eidos.

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Aristotle's Generation of Animals of the matter, what happens to its physical nature? Before answering this question, Aristotle proceeds to an analysis of the way in which semen has the powers of the soul, including the power of reason (nous). This analysis involves a radical development of the theory of pneuma, which is seen in this chapter as a terrestrial analogue of aither, the matter of the heavenly bodies. Rather than following Aristotle's argument step by step (Moraux does this in his "nous thyrathen" 39) we shall investigate certain key topics: a) what happens to semen? (11.3, 737a8-18); b) energeia in II.3; c) "nous thyrathen"; d) pneuma and the bodily powers of the soul (736b22ff); e) pneuma and aither (736b30-737a8). a) What happens to semen? "It dissolves and evaporates, having a fluid and watery na-

ture" (737al2). The foam "breaksdown" (8taAVeraL) and "tums into pneuma" (rvc6u/aToVTaL). That is, the water evaporates; the pneuma in the semen is already pneuma. It is for this reason that "we should not look for semen leaving the female, nor as a part of the fetations." i) If it all becomes gaseous, even if it were to escape we could not see it. ii) Semen acts on mense as fig juice acts on milk, and we do not look for the fig juice either, "for this also changes (METa3a`AxxE)and is not part of the systematized mass" (a15). Both the fig juice and semen work on their material in virtue of their vital heat,40 but this is not the whole story in the case of semen. There is a sense in which pneuma does not become "part" of the fetation, but there is a sense in which it does too. Pneuma does not become a part of the developing organism in any of the Aristotelian senses of the word morion, for it is neither a homoiomeros nor an anomoiomeros which can be distinguished by anatomical investigation. On the whole, in anatomical investigation Aristotle does not find pneuma in any special location or reservoir, although he surely thinks that there is more pneuma in some parts than in others. But pneuma does become a part of the developing organism in this sense: the pneuma is there, in the organism. It is everywhere in the organism, and is indistinguishable from the compounds into which it enters. The pneuma which the semen 39. Paul Moraux "A propos du voOs Opa&uvchez Aristote," in Autour d'Aristote, Melanges Mansion (Louvain, 1955). 40. GA 11.4, 739b22; cf. III.4, 755al5ff; IV.4, 771b23, 772a22.

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has become cannot be found in the organism because it cannot be distinguished from the organism. It has entered into the compounds which are the homoiomere, giving them their form, maintaining their form, and giving form to the anomoiomere and their activities. Even if we could, in some way, distinguish the pneuma from the other elements of the body, we would still not be certain to have found the pneuma from the father distinguished as a separate nature. i) The female contribution also contains some pneuma, even if insufficient for generation by itself; ii) once the male pneuma has been introduced, and the individual begins to develop, there is a constant production of pneuma which cannot be distinguished from the pneuma which was in the semen or mense. All of the pneuma is now the pneuma of this individual; it is "with" his "nature," symphyton, as the Movement of Animals puts it. b) Energeia in I.3. "Fetations" and "seed" (sperma) have at least the nutritive and generative function of the soul, as was shown in 11.1. As the fetations of animals develop, the sensitive soul appears, for it is the defining characteristic of animals. "'Animal' and 'man,' or'animal' and 'horse,' are not generated simultaneously, for the end is generated last, and the particular is the end of the generation of the individual" (736b2ff). It is thus that the problem concerning nous arises, for mind is the defining characteristic of man: when and how does the individual come to have a mind? In order to get at the problem of the development of the sensitive and motive faculty, and of the development of mind, Aristotle works out some of the consequences of his theory of potentiality and actuality: It is clear that not yet separated (x) spermata and fetations have nutritive soul potentially, but they do not have it in actuality (energeia) before (as separated fetations) they draw in food and exercise the function of this sort of soul; for aUl such fetations seem at first to live the life of a plant.4' Some interpretation is in order. (i) the sense of the word dynamis is unclear. Are both active and passive, or only active, 41. 736b8. On living the life of a plant, see GA II.7, 745b26; IV.6, 774b26; Progression of Animals 4, 705a28; PA IV.10, 686b36; II.3, 650a21; IV.4, 678a10.

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Aristotle's Generation of Animals powers considered? If both, then the semen must have an active power and the menstrual fluid a passive power. The fetation has both, but especially the active. (ii) Saying that something has something potentially is not simply a tricky way of saying that it does not have it. If something has an active power, it is something in virtue of that fact. The soul is a dynamis, or set of powers. This is the point of the analogy of the sleeping geometer: he is a geometer, although he is not actually doing geometry. The faculties of the soul do not simply appear and disappear as the organism acts or desists. Aristotle thus seems to think that the nutritive and generative faculty is in the spermata and fetations, although not yet functioning. (iii) There is a direct correlation between this passage and the definition of the soul in the de Anima: "The soul is the first entelecheia of a natural body having the power of life; such will be whatever is organic" (II.1, 412a27). This passage reads, s8Va,4EL gwn)v EXOVT05) and the GA passage reads, lvx,v. . . 8vva4LL... .Xorra. Just as we read the de Anima passage as affirming the life of that which has soul, so we should read the GA passage as affirming the existence of soul (even if not yet in energeia) in seed and fetations. (iv) It has been suggested42 that Aristotle here tries to resolve a conflict between Diogenes of Apollonia and Empedocles: Empedocles had argued that embryos are alive; Diogenes thought that they did not have life because they did not breathe. At this point Aristotle tries to provide a theoretical basis for the solution of the puzzle, operating in conjunction with his new version of the theory of pneuma as the physiological part of the solution. This theoretical solution must provide for some sort of life, and therefore some sort of soul, in seed and fetations, but Aristotle cannot make the observational errors of Empedocles and Democritus which give spurious independence to seed and fetations. (v) The distinction between dynamis and energeia drawn here may then be the distinction between having (but not exercising) and exercising a power. Two interrelated problems arise from this interpretation: (i) Can it ever be proper, within or without Aristotle's system, to say that something has a power which it cannot exercise now? (ii) Is it not self-contradictory to say something has a soul defined as a first entelecheia) but does not have it in energeia? That is, are not entelecheia and energeia so closely related that the ascription of one necessarily involves the ascription 42. H. A. T. Reiche, Empedocles' Mixture, Eudoxan Astronomy, Aristotle's Connate Pneuma (Amsterdam, 1960), pp. 32-36.

and

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of the other, or else that an energeia is a necessary precondition for the existence of the correlative entelecheia? (i) There is no question that Aristotle envisages powers which are present but need further preconditions for their actualization. The sleeping geometer, and the puppet before being wound up, are cases of entities with potentialities which need prior conditions for their actualizations. In general, given the analysis in terms of potentiality and actuality, one may grant "submerged" potentialities. Matter, for example, has levels of potentialities of the passive sort. As elements, matter has limited immediate potentialities, but immense secondary potentialities; as matter is presented in more and more organized forms, new potentialities come to the surface and become immediately possible. There is no reason I can think of for the same not to be true of active potentialities; they too may be submerged only to appear as their activity works itself out in the presented material. (ii) The supposed contradiction between the presence of an entelecheia and absence of an energeia may be resolved by the specification of the precise entelecheia and energeia involved in the discussion. In the de Anima passage, the entelecheia is the presence of a power which is present by virtue of the presence of an organ. In the GA passage, the energeia is the activity of an organ. Semen is an organ. As for the higher powers, the sentient and rational, Aristotle gives a complicated "rational argument" which purports to exhaust the possible locations and sources of the various powers of soul, in the process of generation:43 I. Possible powers in matter provided by female (mense or egg). A. None in matter, but all generated in it. B. All in matter beforehand. C. Some in matter, some not. II. If A or C is true, then either D. Not by semen (what, is not clear) or E. By semen. III. If generated by semen, when did they come to the semen? (It is suggested that they may come from "outside," though where outside is not specified; we may suppose that the heavenly bodies have something to do with it.) F. All the powers generated by the male come "from outside"; 43. 736bl6ff.

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Aristotle's Generation of Animals G. None of them do; H. Some do, some do not. The combinations are these: B. If all the powers are in the matter, then they are not generated in it, so none of the other possibilities would apply. Aristotle has, of course, argued against this position before, and will do so again at 736b22. FGH apply only to E. A, D: "No powers of the soul in matter, all generated in it but not by semen." This may be the proper understanding of spontaneous generation where the matter is slime or the like; the generation is said to be effected by the sun or by warm weather. C, D: "Some powers in the matter, others generated by a nonspermatic agency." This would, I suppose, be the interpretation of parthenogenesis, for the female spermata (e.g. eggs), even if not fertilized, seem to have some share in life; again, heavenly bodies or the weather effects the rest. AE: No powers of soul in matter, all generated by the male: AEF: all the powers generated in the male from without; AEG: all the powers generated in the male, not from without; AEH: all the powers generated by the male, in whom some are from without, some not. But all the AE possibilities seem unlikely, since Aristotle posits the presence of nutritive and generative soul in the female matter. CE: some power of soul pre-exists in the mense or eggs; the other powers generated by semen. So if nutritive and generative soul in mainly in the matter, CEF: sensitive and rational soul may be generated by semen in which they were generated from without; CEG: same, but not from without; CEH: nutritive in the matter, sensitive and rational generated by semen, in which sensitive is generated not from without and rational generated from without. [Peck thinks that only CEH is operative, but a combination of CD and CE is also possible, since they are not mutually exclusive.] The upshot of the argument seems to be that some of the nutritive and generative power is provided even within the female matter, although some must be provided by the male, in species which generate sexually. Aristotle seems still committed to the view that the male provides the power of the sensitive and motive soul. Unfertilized female spermata (men-

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strual fluid or eggs) develop to a point, then stop short of the development of animal organs which would be sensitive or locomotive; the mnalemust provide something which is "higher." The simplest account which one may dig out of Aristotle's text is that the male semen adds the faculty of sensation and movement to the faculty of nutrition and generation already more or less present in the female spermatic contribution. c) "Nous thyrathen" Aristotle's "most vexed" problem concerns the rational power of the soul, mind (nous). It may be that this problem motivates the exhaustive account of the possibilities of the sources of the faculties, for at 736b5-8 Aristotle has said that his biggest problem is "how and when and whence" mind arises in those that have it.44 This problem arises largely because there is serious question whether mind has any share in physical nature, that is, in matter. Aristotle says in the de Anima that "it is unreasonable to suppose that (mind) is mixed with the body." 45 This theme is developed in de Anima III.4-6; there, mind is opposed to matter in a particularly striking way: not only is mind "separate" and "unmixed," but it is also apathes, which means "un-acted-upon" (being acted upon is the essential characteristic of matter), perfectly active, for its being is activity, and it is immortal. We are reminded of this passage at least twice in GA 11.3 (736b28, 737a8). These passages lead us to suppose that nous cannot be (as aisthesis,

on the contrary, can46)

the name of movements in matter,

at any rate not bodily matter (somatike hyle). If this is so, then there has been no progress toward solving the problem of the origin of mind in the rational being. Although Aristotle uses language47 which seems to indicate that mind has a lo44. This aporia pleiste gives problems to modemn commentators; one such problem is whether Aristotle thinks that he solves the puzzle in this chapter. Nuyens, having analyzed the kinds of statement of aporia, finds that the present instance is the only one called pleiste ("biggest"); he concludes that Aristotle would not introduce a definitive solution in such terms (Psychologie d'Aristote, p. 316). E. Barbotin, in La ThIuorie Aristot61icienne de l'intellect d'apres Theophraste, (Louvain, 1954), pp. 175ff, agrees while Reiche thinks that a solution is offered; although he feels that Aristotle does not find that there is adequate empirical confirmation, nevertheless Reiche believes that Aristotle is satisfied with the conclusion (Empedocles' Mixture, p. 98). 45. III.4, 429a24, suggested at II.1, 413a4-7, and II.2, 413b26. and Aristotle's 5;s," 46. See my "On Dreams 2, 459b24-460a33, Phronesis, 13 (1968), 175-182. 47. For example, "wrapped up in" (IEureptcajup3dEraL), 736alO.

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Aristotle's Generation of Animals cality in semen or fetus, this seems inconsistent with its nonbodily character, for only bodily things can have a place (Physics IV.1-5; Generation and Corruption I.5). If Aristotle were committed to a placeless mind, then his problem would be partly insoluble; one might ask when and how it comes to be for the individual rational animal, but one could not reasonably ask whence, nor could one respond "from outside" (thyrathen). The difficulty may be reduced by the careful handling of the senses of the word choriston, which may mean either "separate" or "separable," and either of those in any of three ways: a) in place, b) in logos, c) by intellection (Metaphysics Iota [XI 1052b17). It seems unlikely that Aristotle means to tell us that mind is "separable in place" (a). The notion of "separable by intellection" seems in the Metaphysics to apply to mathematical objects, which are not (according to Aristotle) separate in reality, but treated as separate for intellectual convenience; this seems unlikely to be applied to mind except in a truistic way. Joseph Owens claims that Metaphysics Eta seems to argue that there are individual separate forms; although Aristotle does not actually say this, he does seem to suggest that the mind is such an eternal separate form.48 Moraux, in his "nous thyrathen," attempts to play down the reference to "mind coming from without" at 736b28. The theory of an adventitious mind is fraught with difficulties, some of which I have hinted at. Yet any interpretation which makes this passage disappear must contend with the Greek commentators, who took this passage as an indication of Aristotle's own position, and even more with the puzzle of Theophrastus: "Nous: how and when it is from outside, and is, as it were, adventitious as well as connate."49 Put in this way, there seems to be a self-contradiction: how can mind be both 48. Metaphysics Eta (VIII) 6, 1045a34-b9 (Owens, Doctrine of Being, chap. 13, pp. 379-399). Owens does not suggest that the discussion of separate forms is applicable to minds, in this chapter, and even suggests that the question is not raised (440, n. 11). On the other hand, he asserts (p. 444) that Aristotle "supposes without question that a form which is not the form of any matter is a mind." Assuming that the supposed separate forms of Metaphysics Eta 6 are minds, are they minds of individual men, or is one of them the mind of the species man, or are they simply disembodied intelligences, while the minds of men are embodied intelligences? I do not think that this question can be answered within the context of Aristotle's own writings, because it was not posed in this way until a later time. 49. Fragment 1 a, in Themistius; translated from the quotation given by Barbotin, p. 188.

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"adventitious," "from without," and "connate"? In spite of the difficulties in Moraux's position, I agree with him that Aristotle does not confidently assert that mind is immaterially adventitious. In spite of all, Aristotle seems to believe, and tries, on the whole, to convince us, that there is no essential difference between the inheritance of mind and the inheritance of the other powers of the soul. This interpretation is, I believe, supported by a casual comment a little later in the chapter: The body of the semen, in and with which some of the psychic principle goes away, some being separable from the body, in those in which there is something divine wrapped up (such is the so-called nous), some being inseparable.50 Here, in the space of a half-sentence or less, Aristotle says that nous "goes away" (from the father) in and with the semen (or body of the semen, which is the same thing); that it is separable (choriston) from body. Not only is it separable, but something divine is "wrapped up in" those individuals which have it. I suppose that here, as in other passages which speak of nous, Aristotle is purposely ambiguous. Certain epistemological considerations lead him to separate or dematerialize the mind, but biological and physiological considerations lead him to avoid dematerializing the mind too much. Perhaps we do not find (try as we might) a solution to the mind-body problem in Aristotle because he intentionally does not give a solution. In fact, there is a way in which the problem of nous is external to the problem of the GA. If nous is immortal, then it is not subject to substantial change; if it is not subject to substantial change, then its generation need not be explained, for it is not generated. It simply manifests itself in the appropriate vehicle. d) Pneuma and the bodily powers of the soul (II.3, 736b22ff) It is not the presence of mind in "some animals" that requires the elaboration of the theory of pneuma, but the generation of the bodily powers of the soul in matter. "Those principles whose energeia is bodily cannot be present without body; for example, "walking without feet" (736b22). This 50. 737a8-1 1; I have followed Peck, Louis, et al., in omitting Tb arippa in line 8, which results in the rendering, "some of the psychic principle" (Peck and Louis: "portion"). The preposition and prefixes in p, 4 auvarcpxeTat are difficult to pack into the English.

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Aristotle's Generation of Animals seems quite clearly to be an Aristotelian (hylomorphist) position, but it seems equally a difficult problem for the explanation of the subrational powers, as Solmsen points out.51 In one respect, Aristotle has already solved part of the problem which this doctrine poses, with his elaboration of the theory of dynamis and the analogy of the sleeping geometer; he has already explained how the power of walking, at several removes, may be present in that which causes the power of walking, in that which becomes the organ for walking. But this power must be in the semen (it is active, and the active powers must be from the male). What is there about semen which has this potential power? After all, "semen is (only) a residue of changing food."52 Yet semen does have in it pneuma; this material must have some special characteristics which allow it to have the powers without being the entity. e) Pneuma and aither (736b30-737a8) Now it seems that the power of all soul shares in a body different from and more divine than the so-called elements; as souls differ in value, so the relative nature also differs. In every sperma there is present that which makes all spermata fertile, the so-called "hot." But this is not fire nor any such power, but the pneuma wrapped up in the spermata and in the "foamy," and the nature in the pneuma, which is the analogue of the element of the stars. Thus fire does not generate any animal, nor is any animal constructed in either fluid or solid under the influence of fire; but the heat of the sun and the heat of animals do generate, not only through the spermata, but also any other residue of their nature likewise has the principle of life. In the preceding chapter Aristotle has simply said that pneuma was "hot air"; now, without giving up that view, he wants to relate pneuma to the divine ait her. Moraux53 argues that it would be a misunderstanding to 51. Solmsen, "The Vital Heat, the Inborn Pneuma, and the Aither," Journal of Hellenic Studies, 77 (1957), 119-123. 52. 736b27. Platt inserted the word "only" into the translation, Solmsen (J. Hellenic Studies, 77), and Moraux (Autour d'Aristote) approve, although no one has it in the text. It is not strictly necessary, although it does emphasize the opposition to preformationism, refuted in 1.17-18. 53. Moraux, "Quinta Essentia," Pauly-Wissowa, Half-volume 47, vol. XXIV, (Stuttgart, 1963), 1171-1263, especially 1196ff. Sections A.3 and B are the most important. He provides an extensive bibliography of discussions of the problem.

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identify pneuma and aither, and in the strongest sense of the word "identify," his position is surely correct. However, Moraux does not provide a clear discussion of the relationship which Aristotle does find between pneuma and aither. One of the most important clues in the present passage is this: "As souls differ in value, so the relative nature also differs" (b32). This statement suggests a proportion or analogia; the suggestion is confirmed when pneuma is called the analogon of aither. Expressed as an "analogy," Aristotle's statement is this: Value of soul A: value of soul B :: value of nature A : value of nature B. If value (timiotes) is expressible as a quantity, or is quasiquantifiable, then if the souls differ in value, then the relative physis (nature) of the souls differs in value. One cannot, or ought not, push the analogy too far: although the physis of animal souls differs, it is all pneuma. But how closely are pneuma and the element of the stars to be related? Peck54 discusses the problem, noting this passage: Air, water, and many solids (are transparent); but neither air nor water is transparent qua water or air, but because the same nature is present in both these as also in the eternal bodies above. Light is the activity (energeia) of this transparent qua transparent (de Anima 11.7, 418b6-10, after Hett). The same physis as is found in heavenly bodies is here supposed to be present in a number of terrestrial materials; the sole physis of the stars is aither. Therefore, there is a sense in which there is aither in some terrestrial objects. In harmony with these passages is GA HII.11, 762al9-27, where Aristotle argues in the course of his explanation of spontaneous generation as follows: Animals and plants are formed in the earth and in water because in earth water is present, and in water pneuma is present, and in all pneuma soul-heat is present, so that in a way all things are full of soul; that is why they quickly take shape once it has been enclosed. Now it is enclosed as the liquids containing corporeal matter become heated, and there is formed as it were a frothy bubble. The object which thus takes shape may be more or less valuable in kind, and the differences depend upon the envelope which encloses the soul principle; the causes which determine 54. Note to 737al and appendices.

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Aristotle's Generation of Animals this are the situations where the process takes place and the physical substance which is enclosed.55 a) It may be that both aither and pneuma are present in water, being different natures, but I doubt that Aristotle makes this distinction. That which is, by inference, called aither in the de Anima is, I feel, called pneuma in this passage. b) The scale of value is again referred to, but with an addition. In the earlier passage (736b32) the value depends directly, it seems, on the relative physis, the quality of the pneuma "which is enclosed" (762a27); now the "place" is said to make a difference.56 We may phrase the theory in modern biological terms. Aristotle seems to suppose that pneuma functions as an organizer, and that the surrounding material is that which is organized.57 This understanding of his account seems confirmed by this passage: The portion of the soul-principle which is enclosed or separated off in the pneuma makes and puts movement into the fetation" (762b16-18). Thus it seems that generation and the character of the result of generation depends upon at least two factors: the character of the pneuma which is either already in, or introduced into, the material for generation; the character of the material which is to be organized by the pneuma. A third factor must be mentioned, regarded as essential at least in plant generation and in a large range of animal generation, including all "spontaneous" generation: the environmental temperature, that is, the activity of the heavenly bodies upon the pneuma and the material. The mention of this third factor re-emphasizes the importance of heat in this analysis; Aristotle finds a qualitative as well as a quantitative difference in various lots of heat. The sun and other heavenly bodies participate in generation because 55. Louis remarks the reference to Thales (all things are full of soul) which occurs similarly in de Anima II.1, 411a7-11. The difficulties of Thales' thesis, which Aristotle now seems to adopt, are alleviated by the doctrine that the activity of the physical powers needs the physical organ, which cannot be present in fire, or air, alone. 56. Peck's translation favors this interpretation where the Greek is ambiguous: T'o arpa Tb 7reptXajuPavo6cvov may mean either "the enclosed body" (i.e. the pneuma), or "the enclosing body" (i.e. the mixture of earth and so on around the pneuma). The latter interpretation would be supported by the words which follow: "in the sea there is plenty of earthly stuff; therefore the nature of the testacea is generated from such a construction." 57. See note 5, above.

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their heat is in fact soul-heat,58 and not radically different in kind, although radically different in value, from the soul-heat of temporal souls. The heavenly bodies thus participate in generation by providing one source of movement and organization for that which is organized. But that which provides the source of movement and generation must, according to the principles already elucidated, have the form. The sun et al. are not actually members of animal species; how then do they have the form? Aristotle seems committed to the sort of distinction made clear later, between formal and eminent reality. The souls of the heavenly bodies must contain all these species eminently, for they do not contain them formally, in the Cartesian senses of these words. 8. The accomplishment of II.1-3. The problem which Aristotle set out to resolve in these chapters was primarily that of the mode of operation of semen in generation. In Book I, semen was supposed to provide the form and source of movement to the female spermatic contribution. How it could do this seemed difficult to explain, and we may be excused if we are still puzzled by the end of II.3. Aristotle sought a material which could serve as a tool when no longer in contact with the male parent, and a material which could at the same time be understood under some teleological rubric. The pneuma, as the special vehicle of vital heat, seems to serve the purpose of the explanation. In the first place, Aristotle has essentially three analogies which converge to an understanding of what "must" be the manner of operation of the semen. These analogies are those of fig juice (or rennet), which comes close in that natural materials are involved in the change; of the puppets, which is as good as Aristotle can do at finding an analogy of a tool acting at a distance; and of the working of a craft, which illustrates the manner in which form and movement are given to matter without becoming a part of the matter. Secondly, pneuma is brought more or less forcibly into line with these analogies. There is no special problem in bringing pneuma into line with the fig juice analogy, because he believes that fig juice coagulates milk by virtue of its pneuma, or the vital heat which the pneuma carries; it is not, in this respect, an analogy but an identity. But the puppet analogy seems forced, for the com58. Metaphysics A (XII) 5, 1071a15; Nicomachean Jaeger, Aristotle, p. 143; Peck GA appendix B #18ff.

38

Ethics VI.7, 1141bl;

Aristotle's Generation of Animals plicated movements of the puppet, and more, are demanded by the results. But those complicated movements are the result of the already quasi-organic structure of the puppet. Semen, and more especially the pneuma in the semen, cannot be understood to have an organic structure in this sense; however it may be that pneuma preserves the dynamic structure (eidos) of the male, Aristotle does not really tell us at all. The analogy of housebuilding or doctoring tells us what must occur, without really telling us how, and it leads Aristotle to suppose that no material of the semen becomes material of the fetation. Since the inspection of semen, and the analogies which occur to Aristotle, seem to be insufficient to carry the day, Aristotle puts a considerable weight on the fateful theory that pneuma is a special material quite unlike anything else one might find in the world, but rather strikingly like the material which one might find out of this world. There is something of the myth-maker about Aristotle here; he seems to bring the gods, or at least the divine, down to earth in order to explain that which he finds otherwise inexplicable. His hylomorphic theory of the soul is up against the wall in the explanation of generation, and he knows it, so he decides to support the notion of a particularly formal material. That it is a rarefied material, his audience would readily accept; that it is to be found everywhere, or nearly so, would also be accepted. But that pneuma may be a kind of divine spiritus would not be too readily accepted, at least not, or especially not, by those who had taken the rest of his physical philosophy seriously. Aristotle knows that he must explain the generation of animals in order to make good his claim to have developed the only philosophy able to account for all kinds of change, while remaining rational and preserving the phenomena. Yet in order to make good his claim he must resort to a quite unverifiable theory. One gets the impression that Aristotle ultimately tries to trick us into thinking that he has solved the problem, for he could as well have admitted his inability to explain the manner in which semen does its work, even while maintaining that his general philosophical principles (for example, the four causes) do in fact work out in animal generation as in all other sorts of change. Yet this criticism may be somewhat harsh. If Aristotle is to gain an understanding of this phenomenon, it must be a theoretical understanding. For only now, with enormous progress in techniques of investigation, are the empirical data accessible. Aristotle knew that there must be something in spermatic secretions which provides the form for development, something

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which would perform the functions which we assign to sperm and egg cells, or more precisely to DNA and related factors. He was mistaken in supposing that his functional equivalent of our DNA was to be found only, or pre-eminently, in the male semen, but he knew that something had to carry the dynamic structure from parent to child. His version of the theory was a considerable advance on the theories of his predecessors. No appreciable further advance was made until the discovery of spermatozoa (1679), and even this discovery was accompanied by a step backward to preformationism. Not until the nineteenth century did epigenesis, the theory suggested by Aristotle, win general scientific approval. C. GENESIS AND ENERGEIA Aristotle uses analogies, the ramifications of the concepts of potentiality and actuality, and the notion of pneuma, to explain animal generation in GA IL.1-3.We tend to suppose that pneuma is the central concept in those chapters, partly because it appears so surprisingly. The concept of energeia is, however, still important in II.1-3, as we have shown above. Elsewhere in the GA the central explanatory concepts are energeia and the related four causes, and dynamis. This is important to note, for it is the use of these concepts which most closely relate the GA to theoretical works such as Physics, Metaphysics, and de Anima. We begin this part with a discussion of an especially important passage, GA II.4, 740b13-741a3, which gives an account of generation relying upon the nuances in the sense of energeia and upon the analogy of art and nature, rather than upon the concept of pneuma. The particular emphasis here is the relationship between activity and the matter of which it is the activity. A finer appreciation of the character of the concept of energeia may be gained by an investigation of some of the concepts related to it; we will summarize some of our findings concerning Aristotle's concept of matter and his concept of a source of movement. We may then show how energeia, form and end, develop out of the idea of a dynamic moving matter. Of the appearances of the notion of energeia in the GA which we will note in this part, one of the most important is its use in the explanation of resemblance and lack of resemblance of offspring to parents, in Book IV. It is an especially difficult problem for Aristotle, as shall be shown; it is a phase in his explanation of generation which requires a

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Aristotle's Generation of Animals further development of the concept of energeia. Some general remarks concerning the character of the teleology of the GA conclude this essay. 1. Summary of the explanation of animal generation (II.4, 740bl 3ff). (1) The differentiation of the parts is generated . . .59 because the residue of the female is potentially such as the animal is in nature, and the parts are in it potentially, but not actually (in energeia), for this reason (cause) each of the parts is generated; and because the active and the passive, when they touch, in the way in which the one is active and the other passive (by "way" I mean how, where, and when), immediately the one acts and the other is acted upon. The female provides the matter and the male the source of movement. (2) (a) Just as that which is generated by art is generated by means of tools, it would be truer to say by the movement of tools, and this is the activity (energeia) of art, "art" being "the shape of that which is generated in another," so also the power of the nutritive soul, (b) as it even later effects growth in animals and plants, using as tools hot and cold (for its movement is in these, and each thing is generated according to a certain logos), so also from the beginning it constructs that which is generated naturally; (c) for as it is the same matter by which it grows and from which it is constructed at the beginning, so also the making power is the same;60 now if this is the nutritive soul, then it is also the generator; and this is the nature of each thing, present in all plants and animals. (3) But the other parts of the soul are present in some living things, and not present in others. The passage is structured thus: (1) A general account of generation, including both reproduction and embryological development, is proposed in terms familiar in Aristotle. (2) Two analogies and one other argument develop the account; analogy (a) extends over analogy (b), and argument (c) supports analogy (b) without being a part of analogy (a). (3) Concluding remarks tie the passage together. (1) Aristotle begins with the matter provided by the female, 59. Omitted here is a reference to the theory of "like to like," cursorily refuted in this place. Cf. de Anima I.2, 404b18, 405b15; 5, 409bl9ff.

60. Secl.

rz et dpXi s jue4gdv be

aJT,7

eJTiV

with Peck. Lulofs retains.

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explaining its role by reference to the distinction between dynamis, in the sense of potential, and energeia, in the sense of actual. Physis and energeia seem identified. Having distinguished matter and end, Aristotle proceeds to the source of movement, which he presents in terms of the distinction between active and passive powers.61 The process of generation depends upon the contact of the (right) active and passive powers. (2) The role of analogy (a) may be stated thus: art : movement of tools (energeia of art) : product of art :: nature : movement of hot and cold in semen and in the body = power of the nutritive/generative soul (seems = energeia of nature) : natural product. Art is defined as the shape of that which is generated in another, which implies the definition of nature, the shape of that which is generated in itself. This definition is comfortable in the case of later growth [as in the first part of analogy (b) and at (3)], but he is not altogether clear about what the "itself" is that the shape of nature is "in" in the first movement of generation. There is, I find, a continuing uneasiness on Aristotle's part that the account of the first generation is too like the case of the productions of art, and a certain relief once the nutritive soul is installed in the heart. This is, I suppose, the motivation of analogy (b) and argument (c); he must make perfectly clear to himself and to his auditors that it must be the same power, movement, and activity in both cases, that the nutritive/generative soul has a continued existence between parent and offspring, although one cannot clearly say whose soul it is during this period. The problem is not actually solvable by simple empirical investigation. One has to have at hand some criterion for what will count as "soul" in order to discover it (or not) in the phenomena. By implication, but not straightforwardly, this passage says that the power of the nutritive soul is the energeia of nature. From analogy (b) it is clear that the semen generates by virtue of the nutritive soul; if it also provides sentient soul, as Aristotle surely believes, it must do so by means of the nutritive/generative soul. One may put it this way: the nutritive/ generative soul of animals is such as to generate sentient 61. Cf. Movement of Animals 8, 702al2-21, where animal movement is explained in a similar way, including the "immediacy" of action by active and passive powers. See I. Diiring, Aristoteles (Heidelberg, 1966), pp. 337345.

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Aristotle's Generation of Animals beings. The parts or functions or powers are not very strictly delimited from each other. The form of argument (c) is: if matter 1 is the same in character as matter 2, then mover 1 is the same in character as mover 2; if mover 2 is nutritive soul, then mover 1 is nutritive soul; but matter 1 is the same in character as matter 2 (cf. 740b3). Thus it is the same power which makes the heart (or its analogue) as constructs the rest of the body once the heart (or analogue) is established. 2. Matter in generation. Given the role of the notion of "matter" in this account, and its similar role in the other related accounts of generation, it will be worthwhile to examine Aristotle's concept of matter more generally than we have done thus far. Aristotle's matter is not the atoms of the atomist, nor the matter of anyone who finds material reduction to be an adequate explanation. Indeed, he does not have just one sort of matter at all; his four elements are not even roughly comparable to the modern 100-odd elements. Aristotle's matters form a hierarchy; in the biological books (cf. PA II.1, 747b13) the lowest level is occupied by the simple powers. These powers are qualitative, and their presence or absence is discovered more or less empirically.62 They are also "causative," for as active and passive they enter into changes and constructions. The most truly active power (the hot), as active, seems to have a share in finality, for it is (at least insofar as it does something good) a cause for the noble and good. In the world as it is, these powers are found in somewhat organized materials; in the biological books Aristotle almost always is interested in the materials which one finds in living bodies. As materials, these are homoiomere, the parts of the body of which the (gross) constituents are identical, or bear the same name. This does not mean that homoiomere are unanalyzable; they are, after all, compounds of the elements and powers. Blood, for example, is a rather complicated homoiomeros.63 These homoiomere either compose the anomoio62. Only roughly, since much of Aristotle's account of the heat of various parts of the body is fantasy, or a pTiori reasoning at best. Cf. G. E. R. Lloyd, "Right and Left in Greek Philosophy," Journal of Hellenic Studies, 82 (1962), 56ff; also On the Heavens (de Caelo) IV. 63. Aristotle's homoiomeries must not be confused with those attributed to Anaxagoras. Cf. A. L. Peck, "Anaxagoras, Predication as a Problem in Physics," Classical Quarterly, 25 (1931), 27-37, 112-120, esp. p. 34.

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mere (arms, legs, head, etc.), or are food for the parts (e.g. blood), or are "residues." Menstrual fluid, semen, and milk are residues and homoiomere. They are the result of concoction, and semen is more concocted than menstrual fluid. Concoction changes a material more or less radically;64 in these cases the concoction is toward greater perfection, toward more closely resembling in heat and movement the character of the heat and movement of the parent animal. Through the concoction, pneuma is either generated or gains the character of the animal in which it finds itself. I hesitate to say that it becomes organized, for it has no organs, but one might say that it becomes more sophisticated, or at any rate it becomes more useful for generating new individuals of the same species. Of the generative materials, the male semen, Aristotle repeats again and again, provides the source of movement (only). We have found reason to suppose that the semen becomes some of the pneuma of the new individual, but Aristotle does not see this as providing material for generation. Two sorts of consideration may lead to this position. The first is that semen is said to have little or no earthy material in its makeup.65 The second sort of consideration is presented in Generation and Corruption: When a whole changes, nothing perceptible persisting as its substrate, e.g., blood from the whole semen or air from water or water from a lot of air, this is already a genesis of the one, a destruction of the other; especially if the change occurs between perceptible and imperceptible (either to touch or to all the senses), as when water is generated from or is destroyed into air; for air is more or less imperceptible (Generation and Corruption I.4, 319bl5ff, my translation). In this passage, semen is said to be literally destroyed in the generation of blood, and there is no perceptible substrate or material which persists through the change. It is interesting and important to note that the process is immediately compared to the change between water and air, which, I have already argued on the basis of GA II.3 itself, is the essential answer to the problem of the disappearance of the semen. In Aristotle's view, if there is a change in the matter, then matter is not contributed, since it is not the same matter. 64. PA 11.3, 650a5. Concoction and metabole occur through the power of the hot. 65. 11.2, 735b37, cf. 736a6-8.

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Aristotle's Generation of Animals According to Aristotle's generally stated position, only the menstrual fluid is the matter from which the new individual is generated; the female provides all the matter for the initial fetation, and for the first period of growth and development. In comparison with the male, what the female lacks in concoction of its material is made up for in readiness for construction, like the materials of the artist or builder. The female contribution may be passive, but it is conditionally necessary. It is worked up to the point that it is almost a new individual, lacking only something of a push, some formation by the power of soul. It is proximate matter. This is even more clear in the case of the oviparous or larvaproducing animals; indeed, in the latter instance, there seems to be relatively little need for a male animal at all, for they sometimes produce parthenogenetically (as the bees do-GA III.10, 759a8ff), or are even produced out of the lively earth or sea by the power of the heavenly bodies (GA III.11, cf. II.6, 743a35). 3. The source of movement. The proximate source of movement in generation is normally the male semen; this is in turn derived from the male parent. The sun, and to a lesser extent the other heavenly bodies, seem to have a role in imparting movement and form to the proper matter for generation. The male parent got his proper movements from the semen of his father, and so on, more or less indefinitely. One might mention that these proper movements are preserved in the individual by his eating his proper foods, which contain a measure of the natural heat by which they grew; this heat, in the pneuma apparently, takes on the proper degree and form so as to become co-natural with the individual. The sun and moon also have a role in the well-being of the individual.66 4. Energeia, and movement, form, and end. An examination of the occurrences of the word energeia will help us see how movement, form, and end are closely interrelated, yet distinguished in the GA. According to Bonitz and to my own examination of the text, there are just two occurrences in Book I, but several in II and III. The first appearance contrasts dynamis and energeia in the familiar potentiality/ actuality way: what each part of the developed animal is in energeia, the sperma is in dynamis (I.19, 726b17). Such pas66. IV.10, 777bl7ff, cf. Physics II.2, 194b13, Metaphysics A 5, 1071al5.

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sages by themselves (they are frequent enough in II) might suggest that the energeia or entelecheia67 exerts some occult finalistic causation on the development from seed. The second occurrence (1.22, 730b21) should show otherwise. Here nature in generation is compared to the carpenter; as the carpenter uses his tools, so nature uses semen "as a tool which has the movement in energeia." The implication here is that the natural tool has a movement of itself which counts as an activity, and indeed the right activity. The tools of the carpenter have only potential movement in and of themselves. The first appearance of the word energeia in II is at 1, 734b13; here it is used in connection with the analogy of the puppet, which somehow has a dynamis when at rest, and an energeia when set in motion. We infer that semen has both the dynamis and the energeia. Aristotle goes on to assert that a source of movement must be something which "is actually,"08 and that it acts upon something which "is such potentially (dynamei)." It was at this point that the problem about semen and the energeia arose, for as activity, semen must have the energeia, but as actuality there is some doubt, since this seems to imply the possession of the eidos actually. One wants to say (735a5) that only parent and child have the eidos in the full "actuality" sense of the word energeia. The difficulty, seen from the direction of the use of the word energeia, may be expressed as a tension between the rather simple sense, "'activity,"and the metaphysically portentous sense of "reality." This, or something very like this, must be the reason why the unexercised movements of the semen are no longer called unqualifiedly energeiai in II.3; these movements are now called powers (dynameis), in the active rather than the passive sense of that word. They seem to be energeiai only as they work upon the matter and as the individual begins to develop (736blO). Everything in the physical world, Aristotle believes, must exist as a dynamis before it can exist as an energeia (736bl6); however, if it is only a dynamis, and not an energeia, then it cannot be, properly speaking, a source of movement at all-it needs a prior energeia. Only mind seems to escape this particular difficulty (736b29), since its energeia seems to be independent of physical activities. So semen and fetations are said to have soul as a power (dynamis) but not as an 67. The word occurs rarely; cf. II.1, 734a30, 734b35. 68. energeia ontos, 734b22; entelecheia is used comparably at a5, b35, and Metaphysics 0 (IX) 8.

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Aristotle's Generation of Animals energeia (737al8). Aristotle has not satisfied himself on the question of the relationship of semen and energeia. The problem is not so difficult in the case of the female contribution (737a24). It is easy enough to fall back on the passive sense of dynamis, and to say that the matter has all the parts potentially, but none in energeia. In II.4 Aristotle traces some of the implications of his course of reasoning. The heart, he believes, is the first part to have the energeia, to be actually (740a4), and this is reasonable enough when the problem is seen from the direction of the menstrual fluid.69 He still has the problem of semen. Recalling the analogy of the semen and the tools of art (740b26), he says that the movement of the tools of art is an energeia. But while this implies that the movements of the tools of nature are energeiai too, he forebears and says simply that the movements in the semen are "the power of the nutritive soul." He goes on to say (II.5, 741a12) that they are the cause of the sensitive soul in the developing animal. Sitting directly on the fence, he says that nothing can be the part of an animal unless there is in it the sensitive soul, "either energeia or in dynamis, either qualified or unqualified." By 11.6 Aristotle has gone back to the language of Book I. Describing the solidification of bones and sinews by the agency of heat, he argues that there must be the right material and the right mover: for that which is potentially will not be (brought into being) by a mover which does not have the energeia, nor will that which has the energeia make it from just anything; just so the carpenter cannot make a chest out of anything but wood, and without him there will be no chest from the wood. The heat is present in the spermatic residue, and has the right amount and character of movement and energeia as is proportionate to each of the parts (743a23-29, after Peck). This seems to be the original position. Semen, or the heat in the semen, has the "movement and energeia" without worry about its status as an entity. It is considered as an organ of the father, but a very special organ, which has all the powers but can actually exercise only one of them: the power to pass the rest along to the proper matter. 69. As in II.4, 740b21; II.5, 741bl5; II.6, 742al3, 744a8.

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5. The explanation of resemblance between parents and offspring (GA IV). The term energeia appears again in the explanation of resemblance and lack of resemblance of offspring to parents.70 Aristotle begins Book IV with an explanation of the generation of offspring of different sexes. Treated briefly at the beginning of II, Aristotle's explanation is presented in the context of a discussion of earlier writers on the subject.7' He begins his positive account with a reprise of the definitions of male and female, this time emphasizing the difference in ability to concoct seminal residue. The male can concoct semen, but the female gets only as far as menstrual fluid. The semen in turn concocts the menstrual fluid, and always "tries" to make it into a male; sometimes, however, it fails, due to the recalcitrance of the material.72 The next best thing is the sexual opposite, a female. The same general principles are applied to the degrees of lack of resemblance of offspring to parents (767a35ff). Some offspring resemble their grandparents, or more remote ancestors; some do not have the form (idea) of a man, but of a monster (teras, 767b6). This is a kind of scale of deviation of nature; although females are a conditionally necessary deviation, they are also naturally necessary for the species (767b9). Monsters, however, are necessitated by the character of the material, for they do not, in themselves, serve a purpose. The movement or power in the semen constructs the menstrual fluid according to the logos which it has; this logos depends not only on the immediate parent but also on the ancestors, the species, and the genus. All of these "are at work in the act of generation,"73 but the individual parent most of all, "for this is the ousia" (767b34). That which is generated has a character like the movements in the semen, but it too is an individual, a tode ti, "and this too is the ousia." Here, at 768al2ff, the distinction between energeia and dynamis is recalled in order to explain two different kinds of departure from similarity to the male parent. As I understand the passage, Aristotle supposes that resemblance to the female 70. IV.3, 768a13, b4; cf. 769b2. 71. He mentions by name Anaxagoras, Empedocles, Democritus, and Leophanes, and refers to "other physiologers" (763b32). Peck adds references to Aeschylus, and Hippocrates' "On Diet," "On the Sacred Disease," "Airs Waters Places"; Louis adds Euripedes and Parmenides.

72. 766a19: pLflKpaTfl. 73. Peck's translation of Ev, at 767b33.

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Aristotle's Generation of Animals parent (or ancestors on the female side) is due to "departing from type and changing over"74 to the opposite. Resemblance to ancestors rather than immediate parents is due to "relapsing" (lyesthai). "Some of the movements are present in energeia, some in dynamis; in energeia, those of the generator and of the universals (e.g. 'man' and 'animal'), in dynamis those of the female and of the ancestors." Two questions arise for us: where are the "movements present in energeia and dynamis," and what are the senses of the words energeia and dynamis in this passage? Peck gives his answer to the first question by inserting the words "the semen" at 768a12, and "the seminal substance" at 768b4 and 7. The second sort of insertion must be closer to the truth than the first, although both have something to be said for them. Aristotle cannot possibly be taken to be saying that the semen is in energeia generative of an individual like the father and in dynamis generative of an individual like the mother with all her individual peculiarities. The father cannot have, even in dynamis, these peculiarities. VVhathe can have "in dynamis" as opposed to "in energeia" are the faculties of the female, or (in general) characteristic variations of the species which he does not show in energeia. This is possible, in Aristotle's system, if the operative dynamis is rational (meta logou). In Metaphysics O(IX) 2, 1046a36-b28, he argues that a rational dynamis can have opposite effects, and he gives us as examples the powers of art, especially the medical art (1046b8). We have seen that Aristotle argues constantly in the GA that the power of generation is rational, analogously with the powers of the arts. Just as the doctor, having the power to heal, passes over to the opposite and kills when he is defeated by the recalcitrance of the material, so semen, having the power to generate a male, when it is defeated by the recalcitrance of the material, passes over to the opposite and generates a female. Or, perhaps, "dark eyes" pass over to "light eyes." These would be cases of going over to the opposite, as I understand Aristotle's account. Aristotle makes clear that the movements (whether energia or dynamis) are also present in the female contribution to generation,75 so that seminal substance more closely designates the 74. Peck's translation of 14haTaT-Oat Kai ,ierafa3cXXEtv75. 768a25. It does not seem necessary to oMit Ka' t1rt 7-O)vepp4vwVKaL with Aubert and Wimmer, Aristotelis de Generatiome E7rt TWv ojXEp Animalium (Leipzig, 1860), and with Peck, or Kal &wZrCox appipwivwith Lulofs, but it does not matter for our argument.

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location of the movements; I should prefer the plural (seminal substances), supposing that Aristotle understands arcpliaut after at the places noted above. It is of course equally true that v all of these movements are especially "present in" the first process of systematization of the original fetations. Energeia and dynamis seem to be understood in both the ways which we have distinguished, in this part of Book IV. Particular movements seem to be both energeia and dynamis in that the energeia has the dynamis of bringing about its opposite (that is, femaleness), and in that an energeia has or is the dynamis for bringing about the generation of the developing individual. A further nuance in the distinction between energeia and dynamis may be found in its use in explaining resemblance to more remote ancestors. Aristotle seems to understand that movements proper to the immediate parents are all, in some sense, present in energeia in the spermatic materials, while the movements proper to the ancestors as individuals are present as dynameis but not energeiai. Somehow the various senses of dynamis are combined here again, thus: they are back-up powers, which come into action (energeia) when the primary (parent) powers fail to do the job. They are potentially generative, as the second and third lines of soldiers are potential first-line soldiers. These lines, or powers, come into action, says Aristotle, "because the active is also acted upon by the passive, as the cutter is blunted by the cut, and the heater is cooled by the heated." 76 Thus GA IV.3 illustrates, in the stress of a difficult explanation, how Aristotle sees the interaction of mover and moved, how both mover and moved have their formal character (but more so the mover), and how mover, form, and end in the sense of activity for an end are intertwined in a genetic explanation. Conclusions: Teleology in Aristotle's Explanation of Generation There is little indication that the teleology of the GA has anything to do with the notion of a consciousness of purpose. At most there are hints that minds, and the intelligent stars, are the best things that exist; self-consciousness belongs to them, to be sure, but it is not by virtue of that self-consciousness that the process of generation may be understood teleologically. Semen acts purposively, but not consciously. 76. 768bl6-18;

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Cf. Generation and Corruption 324a33ff.

Aristotle's Generation of Animals Aristotle does not make much use of the notion of "obstacle avoidance" in his explication of the finality of generation. He does say that "everything strives to share in the eternal and divine insofar as it is able," but he does not attempt to show that this might be accomplished in a variety of ways. There is an element, in the account of the resemblance of offspring to parents, of the notion of reserve troops to combat the recalcitrant matter; in more general terms, the existence of many kinds of natural beings is also a sort of cosmic obstacle avoidance. Being is as much as possible. Professor Cherniss criticizes Aristotle's teleological account of generation thus: . . . the fortuitous birth of monsters is "contrary to nature" not in the sense that it does not proceed from natural causes, but only because the form fails to master the matter completely; the whole doctrine rests upon a circular argument, for when the variations are frequent they can no longer be called monsters.78 The argument is circular, however, only if regularity is the only argument for teleology, or if nature has only one sense, the regular. However, nature does have several senses (Met. A, 4, Phys. 11.1), and regularity is only one, not the most important, argument for teleology. Specifically, in the case of generation the steady repetition of generation of new individuals of a species is not what shows it to be a teleological process, but rather the way in which the end, being, is achieved. To be sure, Aristotle believes that monsters do not reproduce themselves, but if they did, the result would be something inferior to the species which originated it. The end in generation is rather understood as "perpetual being," a cyclical return of genesis and decay for individuals, but the permanent existence of the kind, the preservation of the eidos. This is the end of the development of the individual and its parts. Since this perpetuity of the species has a cyclical character, there is indeed a repetition of processes. It is for this reason that the repetition of processes may be taken as prima facie evidence of the purposiveness of the process, but it is only prima facie evidence; one must ask further, "Does it serve the end of more being, more soul, and so on?" Aristotle 77. GA II.1; de Anima I.4, 415a28ff. 78. Aristotle's Criticism of Presocratic Philosophy (Baltimore, 252.

1935), p.

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believes that the species which do exist do serve being in this way. If we ask how final causation operates in the individual case, we are led inevitably to see the close relationship between the notions of form and end in Aristotle's account. Teleological explanations of a non-Aristotelian sort are sometimes offered in the case of generation and development: some have supposed that there is a special "thing" in developing embryos which "guides" its development; others have supposed that the final state of the mature individual exerts an influence upon the development, from the future.79 Both sorts of statement might find some support in Aristotle's text; the first might find it in his theory of pneuma; the second might find it in his often repeated assertion that the developed individual is the end or telos. We must be careful, however, with the idea of causation. If Aristotle gives the "end," it is in answer to the particular question, "What is this for?" or "in order for what?" The end does not, per se, exert a movement on the material, surely not backward in time. Pneuma, on the other hand, is able to structure the material because it already has, Aristotle supposes, the dynamic structure of the individual, although it does not have the (earthy) material. The soul, the ousia in the sense of form of the individual living being, reproduces itself for its own sake. This is because the soul is somehow both individual (and mortal) and universal (and immortal), at least in that it continues itself through spermata to offspring, perpetually. This is its nature, form, and end. 79. These characterizations more closely resemble those provided by critics of teleology than those given by its defenders, but they do seem common notions.

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Descartes' Physiological Method: Position, Principles, Examples* THOMAS S. HALL Department of Biology Washington University St. Louis, Missouri

When Descartes wrote as a biologist-especially in his Treatise of Man (1632)1 and his Description of the Human Body questions was he trying to answer? (circa 1648) f2-what How much-and what-did he accept as (a) already factually established, but (b) still needing to be explained? What axioms *Part of a wider study of Descartes' biomedical works supported by Washington University and by National Science Foundation Grants GS-967 and GS-1985. The author appreciates the help received from Robert Penella, Harvard University, a research assistant whose appointment was made possible under the indicated grants. jAbbreviations used in the notes: AT Charles Adam and Paul Tannery, Oeuvres de Descartes (Paris: Cerf, 1897-1910 [republ., 1956-7, and 1964-7]), cited by volume and page. K C. G. Kuhn, Medicorum graecorum quae exstant (Leipzig: Knobloch, 1821-1833), cited by volume and page. 1. Written in French in 1632. First published posthumously in a Latin transl. by F. Schuyl, De Homine Figuris et Latinitate Donatus (Leyden: apud P. Leffen & F. Moyardum, 1662). Original publ. later, L'Homme de Rene Descartes [et un Traitte de la formation du foetus du mesme auteur, see below, note 2] avec les Remarques de Louys de la Forge, . . . (Paris: Angot, 1664), AT 11:119-202. Note on the terms biology, biological, biologist. Objections are sometimes raised to the anachronistic application of these terms to events or persons that antedated the introduction of the terms themselves. But this objection seems narrow. From Greek times, science has investigated the conditions and varied manifestations of life in general, and from this point of view it Descartes-as biologists, and seems permissible to think of Aristotle-and of their endeavors as biological. 2. "La Description du corps humain" (alternate title "De la formation du foetus"), first publ. jointly with "L'Homme de Rene Descartes" (see above, n. 1), AT 11:223-290. Journal of the History of Biology, Vol. 3, No. 1 (Spring 1970), pp. 53-79.

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and assumptions chiefly governed his explanatory procedures? What were, in effect, his interpretive methods and goals? It is not easy to find, in the secondary literature on Descartes, satisfactory answers to these questions. His biology has been on the whole rather sparingly studied by scholars. Sebba's bibliography (1964),3 which covers the period 1800 to 1960, lists only a handful of titles on this subject. The two best of these deal critically, one of them -xtremely well-with Descartes' physiological theories (Georges-Berthier, 1914, 192021) and the other-usefully but less well-with his biomedical ideas (Dreyfus-Le Foyer, 1937).4 More recently, A. C. Crombie has analyzed the epistemological posture of Descartes as well as his contributions to physiological optics (for example, Descartes was the first to insist that the lens changes shape according to the distance of the object). Finally, L. Chauvois has monographed Cartesian physiology (mostly its weaknesses) as presented in the Fifth Part of the Discourse on Method and K. E. Rothschuh has provided important new insights on the historical setting and sources of Man and Description of the Body in his just published German translations of those works.5 The present paper will differ from those mentioned in focusing sharply on three physiological topics selected to illustrate (a) 3. Gregor Sebba, Bibliographia Cartesiana (The Hague: Nijhoff, 1964). 4. A. Georges-Berthier, "Le Mecanisme Cartesien, et la physiologie au 170 siecle," Isis 2 (1914), 37-89; 3 (1920), 21-58. H. Dreyfus-Le Foyer, "Les conceptions m6dicales de Descartes," Revue de m6taphysique et de morale, 44 (1937), 237-286. See also P. Mesnard, "L'Esprit de la physiologie cartesienne," Archives de philosophie, 13 (1937), 181-220. The longer monograph of B. de Saint-Germain, Descartes conside&r comme physioloqiste et comme medecin (Paris: Masson, 1869), is reportorial and not helpful from an interpretive point of view. There are useful materials on Descartes' biology in J. Roger, Les Sciences de la vie dans la pensee frangaise du XVIII siecle (Paris: Colin, 1963). See also the commentaries on the "Fifth Part" of the Discourse on Method by E. Gilson, R. D., Discours . . . texte et commentaire (Paris, Librairie Philosophique, 1939), pp. 293348, and K. E. Rothschuh on D.'s biological theories in his Physiologie vom 16. bis 19. Jahrhundert (Freiburg: Albert, 1968), pp. 111-115. 5. A. C. Crombie, "Descartes," Scientific American, 201 (1959), 160-173; also "Some aspects of D.'s attitude to hypothesis and experiment," Collection des travaux de l'Acad6mie d'Histoire des Sciences (Florence: Bruschi, 1960), pp. 192-201; and "The mechanistic hypothesis and the scientific study of vision, etc." in S. Bradbury and G. L'E. Turner (eds.) Historical Aspects of Microscopy (Cambridge, Eng.: W. Heffer for the Royal Microscopical Society, 1967), esp. pp. 66-112. L. Chauvois, D.: Sa methode et ses erreurs en physiologie (Paris: tditions du Cedre, 1966). K. E. Rothschuh, tUber den Menschen . . . (Heidelberg: Lambert Schneider, 1969). See, also, Rothschuh's "R. D. und die Theorie der Lebenserscheinungen," Sudhoif's Archiv 50 (1966) 25-47.

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Descartes' Physiological Method the kinds of sources used by Descartes and (b) his detailed explanatory method. But, first, a suggestion concerning the position of Descartes in biomedical history. Does the relative neglect of his biology imply a general disregard for its merits? Georges-Berthier has sampled three centuries of opinion on this subject and has found that estimates vary. One can read that as a biologist Descartes "blazed new trails on which, however, he then went astray" (La Mettrie, 1745).6 And, in another century, that what he built was a "physiology of fancy, almost entirely imagined" (Bernard, 1872).7 But one may also read that it was Descartes who "founded biology, by first explaining life in a scientific, naturalistic way" (Lemoine, 1862).8 And that he "laid the foundations of modern physiology-just as he did of modern physics" (Fouillee, 1893).9 A negative judgment, acceptable even though not wholly documented, is put forward by Dreyfus-Le Foyer-namely, that Descartes' biological essays are marked by "inadequate rigor in regard to verification, inordinate rigor in regard to explanation," and that "C'est pour lui [Descartes], l'essentiel n'est pas de constater juste mais de 'rendre compte.'" 10 Georges-Berthier concluded that the biological effort of Descartes was scientifically unsuccessful (its premises were not new, its conclusions not accepted) but philosophically sound (it sought a common method for science as a whole, biology included). Whatever position we adopt on these questions, it seems worthwhile, for three reasons, to examine Descartes' analytical method. First, more consciously and clearly than any contemporary thinker, he articulated the crucial biological question of the day. The point at issue was the nature of the latent cause, or causes, of the patent phenomena of life. Were these causes essentially psychic (as almost all earlier biologists had believed) or, rather, physical (as Descartes quite strongly affirmed)? Some historical notes on this question are contained below, in section one. Second, without succeeding admirably himself (because his 6. J. 0. de LaMettrie, "Histoire naturelle de l'ame," first publ., The Hague, 1745, Oeuvres philosophiques (Amsterdam, 1753), 1, 24; see also the translation by C. G. Bussey et al., of extracts only, published with Man a Machine . . . (Chicago: Open Court, 1912), p. 158. 7. Claude Bernard, Le!gons de pathologie expe'rimentale . . . (Paris: Bailliere, 1872), p. 481. 8. [Jacques] Albert [Felix] Lemoine, L'Ame et le corps: etudes de philosophie morale et naturelle (Paris: Didier, 1862), p. 206. 9. A. J. E. Fouillee, Descartes (Paris: Hachette, 1893), p. 65. 10. Dreyfus-Le Foyer, n. 5 above, p. 261.

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method was too deductive), Descartes clearly showed what the future goal of physiology must be, namely, the construction of conceptual micromodels to "explain" life as it presents itself to the senses. This form of reductive analysis was not new to biology, but it had scarcely been undertaken earlier on nonpsychistic assumptions (however, for certain exceptions to this statement, see below).1' Third, by limiting soul-functions to mind-functions, and by insisting rigorously on the distinction between mind (res cogitans) and body (res extensa),12 Descartes gave a special cast to the mind-body problem and largely laid down the lines along which physiological psychology, and psychology in general, were thereafter developed and debated. This aspect of his influence will be treated in this paper only in that one of our three examples will be of a neurophysiological nature. I. NONPSYCHISTIC BIOLOGY Historians have often noted but not always sufficiently stressed one of the central facts of the conceptual revolution that overtook biology during the seventeenth century-namely, the effective (though by no means immediate or total) overthrow of putative psychic causes of physiological function. The most persistently influential Greek thinkers (Plato, Aristotle, Galen) had attributed life-as-action (usually bios, zoe) to a variously depicted causal life-soul (psyche). This idea, transmitted to Western science by the Arabs, had been elaborated in the Schools and reaffirmed (and altered) by sixteenth-century 11. The most serious approach to a kind of nonpsychistic biology in earlier Western thought had been that of Epicurus, who considered all phenomena of life, including cognition, to result from the proper configuration of immanently inanimate atoms. Yet even Epicurus was an animist in that he supposed that four sorts of small rapidly moving atoms composed the soul, larger and slower atoms the body. However, Epicurus looked on the organism as a diphase system comprising two interlocking being able somatic and the other psychic-neither networks of atoms-one to function adequately in the absence of the other. Thus soul-materialized, a crucial role in Epicurus's physiological scheme. See, to be sure-plays e.g., Lucretius, De natura rerum, bk. 2, lines 944-961, and bk. 3, lines 548-557. 12. See esp. Rene Descartes, Meditationes de prima philosophia . (Paris: Soly, 1642), the Third Meditation. And his Principia philosophiae, first publ. in Latin (Amsterdam: Elzevir, 1644) and then in a translation by "un de ses Amis" (Picot), Les Principes de la philosophie (Paris: Le Gras, 1647), pt. 1, sects. 8, 53 (AT 8:7, 25 [and 9:28, 48]).

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Descartes' Physiological Method authors, including Fernel (1542, 1554-55),13 Pare (1561),14 and Piccolhomini (in a version adapted to church doctrine, 1586),'15 to mention but three. We usually and rightly think of the mechanization of biology as an extension of the partly antecedent but still continuing Imechanization of the world picture" 16 in general. But it is important to keep before us what "mechanization" entailed. In cosmology it had involved, among other things, an incomplete and irregular, but generally progressive, substitution of physical for psychic (and often transcendental) causes of celestial motion. In biology-in the seventeenth century-a similar development began, namely, a substitution of physical for psychic (and often transcendental) causes of vital motion. Thus the biological revolution partly took the form of a cogent and ultimately decisive assault on the Greek (and Medieval and Renaissance) idea of a causal, physiological soul. To whom should we chiefly attribute the soulless biology that now began-with many false starts and backslidings-to gather momentum? This question will be considered by the author in a separate paper, but, on partial evidence, a tentative judgment may be offered here. It is reasonable to think not of one but of three arguments about the cause of vital functions as developing during the seventeenth and early eighteenth centuries. In one of these, with Descartes (from 1637) its principal but not only inceptor,'7 the very existence of the life-soul was questioned. In another, culminating with Stahl (from 1684),18 the life13. Jean Fernel, De naturali parte medicinae (Paris: apud S. Colinaeum, 1542); rev. ed. in Medicina (Paris, 1554), trans. C. de Saint-Germain, Les VII Livres de la Physiologie . . . (Paris: J. Guignard, 1655), of which bk. 5 deals especially with the physiological soul and its several faculties. 14. See Ambroise Par6, Anatomie universelle (Paris: Le Royer, 1561), p. cxliv ff; also, "Livre de la generation de l'homme, recueilly des anciens et modernes," Oeuvres (Paris: Buon, 1575), pp. 802-850. 15. Archangelo Piccolhomini, Anatomicae praelectiones (Rome: Bonfadini, 1586), pp. 11-14. 16. The phrase is adopted from the title of Dijksterhuis' indispensable book on the subject (London: Oxford University Press, 1961, 1964). 17. The automatism question had occurred in various forms in Scholastic thought, and Gomez Pereira had suggested a mechanical conceptual model of man in his Antoniana Margarita (Medina del Campo: de Millis, 1554). See, on the unoriginality of Descartes' bioautomatism, Georges-Berthier, n. 4 above, 1914, pp. 80-85. 18. See, e.g., G. E. Stahl, "Medicinae dogmatico-systemicae partis theoreticae sectio I quam constituit physiologia," Theoria medica vera (Halle: Orphanotrophei, 1707, 1708), passim but esp. p. 260. For an earlier

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soul was affirmed but its mode of intervention was argued. And in a third, the life-soul was acknowledged-but neglected: it was not denied but neither was it used in detailed explanation. The first of these three debates-over what we may term not always pay much attenphilosophical mechanicism-did tion to body-functions, and, when it did, derived its explanations of them from its answers to larger, axiomatic questions. Is man, are animals, soulfull-or, are they soulless? If soulfull, how is soul allied to the body? Is its role both physiological exclusively the latter? These and related and cognitive-or problems continued to be debated for more than a century by Descartes' defenders, developers, and detractors.19' 20 They were to become central issues of the Enlightenment at least as far as psychology was its concern. The second-explicitly psychistic-tradition, as embodied in Stahl, was partly a counter-reaction to the mid-seventeenthcentury drift away from the life-soul idea. But Stahl's was not a reversion to the conventional (Galenic) idea of different soul-faculties for different physiological functions. He saw the soul as governing, rationally, every detailed operation of the body, and as doing this either consciously and deliberately (his term for this sort of soul function was ratiocinatio) or through unconscious but nevertheless rational intervention (ratio) at what we should think of as the molecular level.2' Stahl was not the first to think animistically in other than strictly Greek terms. Something similar to his two-level interpretation of soul-function (ratio and ratiocinatio) had appeared slightly earlier, for example, in the Tractatus de Homine of Honoratus Faber (1677),22 whose ideas, however, were otherwise still Galenic. Earlier still, there had been van Helmont (d. 1644) with his concept of mind linked with soul, both mind and soul holding sway in the pyloric end of the stomach soul governs the body, according to van Helmont, -whence statement, "De sanguificatione in corpore semel formato," first publ. Jena, 1684, trans. T. Blondin, in Oeuvres medico-philosophiques de C. E. Stahl (Paris: Baillihre, 1859), 6, 556-562. 19. On medical Cartesianism, see Georges-Berthier, n. 4 above, 1920, pp. 23, 29. 20. See, on the philosophic consequences of Cartesian physiology, A. Vartanian, Diderot and Descartes (Princeton, N.J.: Princeton University Press, 1953), esp. ch. 4; and L. C. Rosenfield, From Beast-Machine to ManMachine (New York: Oxford University Press, 1940). 21. See esp. G. E. Stahl, Propempticon inaugurale de differentia rationis et ratiocinationis (Halle, 1701). 22. H. Faber, "de Homine," Tractatus duo . . . (Nuremberg: sumpt. Endteri, 1677).

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Descartes' Physiological Method through the intermediation of a hierarchy of directive "archei." 23 The third argumentative tradition engaged many of the experimentally-and in some cases quantitatively-oriented biologists on whom we usually think of the real progress of physiology as depending. They occupied a variety of conceptual positions between the outright mechanicism of Descartes and the equally explicit animism of Stahl. Some-among them Gassendi (before 1655)24 and Thomas Willis (1672)25-not only acknowledged but "materialized" the life-soul; they gave it a corpuscular constitution. Others were less definite; they admitted the life-soul's existence, but used it rarely in their explicative procedures; this was true, for example, of Harvey (1651),26 Hooke (1665),27 Mayow (1674),28 and Borelli (be23. J. B. van Helmont, "Sedes animae," and "Jus duumviratus," short treatises first publ. posth. in Ortus medicinae (Amsterdam: Elzevir, 1848). 24. Gassendi's position was a blend of Epicurean, neo-Platonic, Aristotelian, and ecclesiastic elements, involving a corporeal, mortal (in these respects Epicurean) nutrient-sentient soul that animals share with men, and a separate incorporeal rational soul (a Platonic and, incidentally, Cartesian conception) which is immortal (as denied by Epicurus but demanded by ecclesiastic Aristotelianism). See P. Gassendi, 'Liber tertius: De anima,' "Physicae: sectio tertia," Syntagma Philosophicum, first publ. posth. in Opera omnia . . . (Leyden: Anisson & Devenet, 1658); republ. in facs. (Stuttgart-Bad Cannstart: Frommann, 1964), II, 250-259. 25. Willis adopted a five-element chemistry (spirit, sulphur, salt, water, earth) and saw the corporeal soul-particles as based on the first two of these elements. The soul has three parts, vital (equated with vital spirits), animal (equated with animal spirits), and genital (an abstract of the other two); see T. Willis, De anima brutorum . . . first publ. London, 1672, trans. S. Pordage, "Two Discourses Concerning the Soul of Brutes," (= Treatise XI in) Dr. Willis' Practice of Physick .. . (London: Bassett and Crooke, 1684), pp. 4, 6-7, 39. 26. Harvey makes blood "the generative part, the fountain of life, the first to live, the last to die, and the primary seat of the soul." See William Harvey, Exercitationes de Generatione Animalium, first publ. London, 1651, trans. R. Willis, The Works of William Harvey, M.D. (London: Sydenham Society, 1847), p. 377. 27. Hooke mentions (apparently only once) "an anima or for-ma infor-mans that does contrive all the Structures and Mechanismes of the constituting body, to make them subservient to the great Work or Function they are to perform." Micrographia (London: Martyn & Allestry, 1665), p. 95.

28. Mayow disagreed with Willis who (see above, n. 25) "corpuscularized" the soul; Mayow saw the "nitroaerial" corpuscles as vehicles for an immaterial soul associated with the soul of the cosmos. See John Mayow, Tractatus quinque medico-physici ... (Oxford: Sheldonian Theater, 1674); trans. A. C. Brown and L. Dobbin, Medico-physical Works . . . (Edinburgh: Alembic Club, 1907), p. 259. For the influence of Descartes on Mayow, see W. Bohm, "John Mayow und Descartes," SudhofFs ATchiv fur Geschichte der Medizin und der NatuTwissenschaften, 46 (1962), 45-68.

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fore 1680).29 Finally, many made no open issue of the life-soul problem, passing it over for the most part in silence; this group included Steno (1669),30 Redi (1688),31 Keill (1698),32' and Baglivi (1700, 1703)33 as well as later "iatromechanists" ranging in their methods from the highly speculative Boerhaave (1708)34 to Stephen Hales, who was capable of cautious, quantitative experimentation (from 1727). The story of the decline of psychistic biology must be sought in the evolution and mutual accommodation of the two mechanistic biologies just mentioned-one philosophical (which undermined the life-soul idea by open opposition), the other scientific (which undermined it by progressive inattention). The influence of Descartes on the former-the philosophical-tradition is apparent enough. His scientific influence is more difficult to assess. 29. According to Borelli, the soul "as principle and as efficient cause of animal movements" was that "through which the animate live (animantia per animam vivant)," but he rarely mentioned the soul in his explanations of function; see G. Borelli, De Motu Animalium, first publ. posth. Rome, 1680-81; 2nd ed. (Leyden: vander Aa, 1865), pt. 1, ch. 1, pp. 1-4. 30. Steno praises the endeavor of Descartes, but disagrees with his scientific results; see N. Steno, "Discours sur l'anatomie du cerveau," first publ. Paris: Ninville, 1669, Opera Philosophica (Copenhagen: Tryde for Carlsberg Foundation, 1910), 2, 7-12; in a rare allusion to the individual soul, he dismisses it from his interpretive scheme, "De solido intra solidum naturaliter contento," first publ. Florence, 1669, Opera, 2, 188-189. 31. See, e.g., F. Redi, "Esperienze intorno alla generazione degl'insetti," first publ. Florence, 1688, Opere (Milan: Soc. Tipogr. de' classici Italiana, 1809-11), 3, 13ff. 32. See J. Keill, Anatomy of the Humane Body, abridged, first publ. London, 1698; many subsequent editions. 33. Without singling out the physiological life-soul in particular, Baglivi attacks ancient assumptions and urges mechanistic and micromechanistic analytical procedures. He mentions favorably, but does not develop, Descartes' solutions of the mind-body problem. See the introductory chapters in G. Baglivi, Specimen quatuor liborum de fibra motrice et morbosa (London and Basel: Konig, 1703). 34. Georges-Berthier (above, n. 4) says Boerhaave got his physics from Newton rather than Descartes; and indeed Boerhaave was sometimes critical of Descartes (see J. Roger, n. 5 above, p. 150). But Boerhaave's biophysics was more Cartesian than Newtonian; like Descartes, he builtelaborate largely deductively, nonexperimentally, and non-numerically-an conceptual micromodel of the patent functions of the body. Boerhaave was Cartesian, likewise, in his view of man as comprising body plus mind. Boerhaave owed much, to be sure, to Baglivi and especially to Harvey (Boerhaave made the body an "hygraulic machine"). See H. Boerhaave, Institutiones medicae . . ., many editions from 1708; especially that of von Haller, Praelectiones academicae . . . (Amsterdam, 1739-42), trans. anon. Dr. Boerhaave's Academical Lectures (London: Innys, 1742-47), 1, 65. On Boerhaave's micromechanics, see T. S. Hall, Ideas of Life and Matter (Chicago, Ill.: University of Chicago Press, 1969 ), ch. 26.

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Descartes' Physiological Method The new physiology was to be-by contrast with the old-experimental, quantitative, reductive, and nonpsychistic. All of this, Descartes quite clearly proclaimed. But was the new physiology really new? What Descartes proclaimed was, to some extent, already in the air. For example, even Vesalius (1543) had adopted, on the life-soul as on so many subjects, an agnostic position. And some of the post-Vesalian anatomists, notably Columbus (1559) and du Laurens (1600), had made little use of soul as an explicative device. Again, Sanctorius' influential Medicina Statica (1614) had proceeded mathematically, experimentally, and nonpsychistically in a spirit more modern than anything Descartes himself was later to produce. Thus, Descartes was partly focusing and crystallizing a trend that was already present, if somewhat diffuse. Moreover, his own effort at crystallization, his proposal that the life-soul be given up entirely, was, as just seen, neither promptly nor universally adopted. But the latter point is not entirely to Descartes' discredit. It may even suggest that he was ahead of his time. The fact is that gradually and unevenly-but irrevocably-the life-soul was destined to disappear (with unimportant exceptions) from the main line of physiological inquiry. Separately, the present author is making a detailed study of late seventeenth-century attitudes toward Descartes' physiological theories. Pending the outcome of that study, it may be suggested that his causal role in biomedical history was illuminative and accelerative rather than inceptive or decisive. He helped the new biology move forward by pinpointing the goals toward which it was already groping. II. EXPLANATORYPRINCIPLES AND PROCEDURES Certain broad features of Descartes' biology are familiar to readers of his Discourse on Method (1637),35 which includes a partial paraphrase of the slightly earlier (but only posthumously published) Treatise of Man. In the Discourse, Descartes sharply separates life (which men and animals have in common) from soul (only present sensu stricto in man). Life is an ensemble of functions that have their kinetic origin in heat-specifically a certain "fire without light" that burns, in men and animals, in the heart. 35. DiscouTs de le methode pour bien conduiTe la Taison, & chercher la verite dans les sciences, first publ. anon. with La Dioptrique, les meteores, et la geometrie (Leyden: Maire, 1637), often republished and translated. AT 6:1-78.

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The feu-sans-lumiere of the heart resembles corpuscularly the sort of lightless fire-or heat-that occurs in various kinds of fermentation.36 In Descartes' Treatise of Light, designed for simultaneous publication with the Treatise of Man (and, like it, suppressed), we hear that all invisible (as well as all visible) heat is reducible to the rapid movement of particles not indivisible (atomic) particles37-of a certain -though fiery matter that is the first of three elements acknowledged by Descartes.38 The other two elements are: a second, airy substance (matiere de ciel) whose particles are somewhat coarser; and a third, or earthy, element whose particles are coarser still. The matter of which the elements are composed is the same for all three, the differences residing in the shapes and sizes of the particles into which this matter is subdivided. The first element composes the sun and fixed stars; the second, the interstellar heavens; the third, the tangible contents of the earth, the planets, and the comets. In tangible bodies, including man's, the interstices between the earthy particles of the third element are occupied by airy particles of the second whose own interstices in turn are completely filled by the fiery

particlesof the first.39 Readers of the Treatise of Man and of the Discourse are by God, excluespecially made aware that the soul-given, sively to man40-lacks the lower faculties (those permitting 36. AT 6:46. For Descartes on this, see also AT 1:521-534; 4:573; 8:256 (9:250-251); 11:23, 228, 333, 538, 599, 631-632. 37. Descartes placed no lower limit of divisibility on his constitutive corpuscles ("particules," "petites parties"). For references to his explicit objections to Democritean atomism, see E. Gilson, Index ScolasticoCart6sien, first publ. Paris, 1912 (New York: Franklin, 1913), p. 31. 38. Specifically, fire entails continuous direct agitation of third-element particles by first-element particles without intermediation of second-element ones. See also Descartes on the same subject in his Principles, n. 12 above, AT 8:218, 249-250 (and 9: 215-217). 39. AT 11:23-31. In the Principles (1644), Descartes no longer calls the second element airy, because familiar, atmospheric air comprises primarily, in his view, particles of the third or earthy element. Descartes presents his doctrine of matter commencing at pt. 3, sect. 46, AT 8:100 (and 9:124). The characterization of the atmosphere as composed of detached delicate, feather-like particles of the third element also appears in the Principles, AT 8:23 (and 9:225-226). 40. On the automatism (which meant above all, for Descartes, soullessness) of animals see esp. the Discourse (AT 6:57-60), and letters to Mersenne (AT 3:121), the Marquis de Newcastle (AT 4:573), and Henry More (AT 5:276-279). It is, however, not quite true that Descartes always eliminated the soul from animals. He sometimes acknowledged at least a one place (letter to Buitendijck, AT 4:64) material equivalent of soul-in equating it with blood whose subtlest part separates off in the brain as

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Descartes' Physiological Method generation, nutrition, and unconscious motion) with which the Greeks and many Medieval and early Renaissance thinkers had endowed it.41 The soul is concerned, according to Descartes, with conscious perception, voluntary motion, and the intellective activities of memory, imagination, and reason.42 There is no need to elaborate these widely known aspects of Descartes' biological-and psychobiological-program. Our object, rather, will be: to show how he used his nonpsychistic, triadic, particulist physics in a reductive explanation of familiar biological function. For, such explanation was the central core of his entire physiological effort. That such was his goal becomes clear as soon as we read what he wrote (and we shall do this in a moment) about such cardinal physiological problems as (1) assimilation, (2) the initiation of embryonic differentiation, and (3) the receptor action of sensors. If we pay close attention to Descartes, we find that he used a kind of strategy of inquiry which, far from being new with him, had been extensively developed in Greek biomedical science. This classic procedure (which began with the preSocratics and culminated, in antiquity, with Galen) proved fruitful-and flexible-enough to be used in all subsequent periods. (Indeed, from a certain point of view it is the strategy still followed by physiologists today). Its cardinal assumption is that the goal of physiological inquiry is to discover the latent equivalents of patent biological function. It is true of Descartes' pursuit of this goal-and this point is crucial-that neither the patent phenomena he interpreted nor the latent equivalents he posited were fully original with him; most of the explanations he offered were only partly his own. Indeed, the principal point we wish to make about Descartes' physiological method is that the explanations he developed were corpuscularized, nonpsychistic versions of psychistic explanations put forth earlier by others (namely, by the major Greek biological writers, by Scholastic authors whom Descartes is known to have studied,43 and by a group of Renaissance animal spirit, and elsewhere arguing that being corporeal, the dog's soul cannot be separated from the body and saved (Letter to Voetius, AT 8;167-168). Thus soul in animals is res extensa rather than, as in man, res cogitans. 41. AT 6:46, and esp. Man, AT 11:202, and Description of the Body, AT 11:224-225; see also Descartes to Plempius, AT 1:523, and to Regius, AT 3:369-370 and 371-375. 42. See, esp., Descartes Passions of the Soul, first publ. Les Passions de l'ame (Paris: Le Gras, 1649), pt. 1, arts. 17-20, AT 11:142-144. 43. Possible Medieval sources of Descartes' world-system are suggested by E. Gilson in his Index, n. 37 above.

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anatomists whom he rarely mentions in his published works but with whose ideas he was clearly acquainted).44 Descartes created a problem for historians by generally omitting any reference to his sources. This omission was in line with his goal of building biology anew, by reasoning logically from certain axioms that seemed inescapably clear. He wished to extend to biology the logic that had served him so well in his mathematical investigations. But, despite this aim of disengagement, we sense, in the examples that follow (and in almost everything he wrote about biology), a thorough immersion in already existing ideas, ideas within the context of which, and not outside them, his own opinions were developed. We obtain, in consequence, a paradoxical impression: his explanations seem new on the one hand, yet strangely familiar on the other. The paradox is less surprising when we realize that what Descartes had to offer were not explanations of fact. They were explanations, rather, of other peoples' explanations (often dismembered and reassembled with various additions and deletions). III. EXAMPLES The present author will shortly publish English translations of the Treatise of Man and the Description of the Body, with suggestions concerning the origins of the ideas that Descartes borrowed and inserted into his own interpretive machinery. The following illustrations of his method could be multiplied many times by sampling the texts of the treatises more or less at random. Example 1: Assimilation of nutriment to the body solids Descartes' interpretation of assimilation is, in effect, a "cartesianized"-that is, corpuscular and antipsychistic-amalgamation of two already established interpretive traditions. The first of these was a classic concept concerning the central nature of nutrition. According to this idea, a prime distinction of living systems is their continuous and balanced involvement in material displacement and replacement. Elsewhere, we have considered this idea-of life as opposed transformation-as it appears in pre-Socratic and Hippocratic theories; in Plato, who speaks in the Timaeus of the body's emptying (anachoresis) and filling (plerosis); in Aristotle and Galen; in several Arab authors; in Arnald of Villanova, Paracelsus, and Francis Bacon 44. For biomedical sources of Descartes, see Georges-Berthier, n. 4 above, 1914, pp. 43-44.

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Descartes'PhysiologicalMethod (who speaks of "depradation" and "refection'); as well as in a sequence of post-Cartesian theorists up to, and into, the twentieth century. All of these thinkers endeavored to lay bare the latent equivalentsof patent intake and output.45 The second tradition which Descartes incorporated in his scheme envisioned the body-solids as composed of subvisible fibers. Galen, in a reformulation of even earlier ideas about fibers, had given muscle a fibrous microstructure, supposing that within the muscle the terminal subdivisions of nerves combine with the terminal subdivisions of ligaments to form fibers that emerge from the farther end of the muscle as tendons.46 With the reaffirmation of Galenic doctrine in Europe, variations on the fiber-theme were proposed by many theorists, including such immediately pre-Cartesian authors as Fernel (1542),47 Vesalius (1543), Jacques Dubois,48 and Jean Riolan (1610) who extended Fernel's ideas to make fibers the basis of the "whole architecture" of the body.49 With respect to assimilation, Descartes thus envisioned his task as one of describing, in the language of his own corpuscular physics, how the body's constitutive fibers are continuously displaced and replaced. He saw the fibers as being constantly added to by the arterioles (at the tips of which they arise), and constantly eroded (at their free outer ends by friction or evaporation). Descartes had developed his own re-explanation of Harvey's explanation of the circulation,50 but the idea of a 45. Thomas S. Hall, "Life as Opposed Transformation," J. Hist. Med. Allied Sci. 20 (1965), 262-275. 46. De placitis Hippocratis et Platonis, bk. 1, ch. 9, K 5:204. Galen gave certain viscera a triple muscle-coating of circumferential, longitudinal, and oblique fibers, to account for their various functional capacities. 47. J. Fernel, Medicina, see above, n. 13, bk. 7, ch. 10. 48. Jacobius Sylvius (Jacques Dubois), Introduction sur l'anatomique partie de la phisiologie d'Hippocras & Calien, trans. J. Guillemin (Paris: Hulpeau, 1555), pp. 43ff. 49. According to A. Berg, "Die Lehre von der Faser als Form- und Funkder Organismus," Virchow's Archiv fur pathologische tions-Element Anatomie und Physiologie, 309 (1942), pp. 394ff. This paper details important aspects of the history of fiber-theory. 50. Blood is volatilized by the heat of the heart, and the resulting expansion induces diastole: Man, AT 11, from 123; Description of the Body, AT 11, from 228; Discourse, AT 6, 48-49; letters to Plempius, AT 1:521534, and Beverwijck, AT 4:3-6. Harvey thought the innate heat of the blood caused it to swell in the auricles, causing them first to dilate and then contract in response, driving the blood into the ventricles where a similar cycle of dilation and contraction occurs. The swelling of the blood is reminiscent of, but is not in fact, fermentation, in Harvey's opinion. See W. Harvey, "A second disquisition to John Riolan . first publ. Cambridge, 1649, Works, n. 23 above, pp. 132, 140-141.

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closed capillary connection between the arteries and veins had not been introduced at this time;51 hence, there was nothing to prevent the tips of the arterioles from giving rise to fibers. Listen to Descartes himself on the subject: For, at the moment when the arteries are inflated [by the pulse],52 the blood particles they contain will here and there strike the roots of certain fibers which-emanating from the the ends of the branchlets of the arteries-compose bones, flesh, membranes, nerves, and brain, and the rest of the solid parts according to the different ways in which they are joined or interlaced. They [the escaping particles] thus have force enough to push [the fibers] before them slightly, and so to replace them. Then, at the moment when the arteries are disinflated, each such particle stops where it is and is united, by that fact alone, to the particles [of the fiber] it touches, in accordance with what was said heretofore. Now if it is the body of a child that our machine represents, its matter will be so tender and its pores so easily stretched, that the part[icle]s of the blood which enter thus into the composition of its solid members will generally be a little coarser than those whose places they take, or it will even happen that two or three together will replace a single one, which will be the cause of its growth. However, the matter of its members will harden little by little so that after a few years its pores will no longer be able to stretch so much; and so, ceasing to grow, it will represent the body of an older man.53 In unpublished notes inspired by his own experience in the dissecting room, Descartes distinguished between appositive and immutative [intussusceptive] accretion (a dichotomy not original with him);54 the picture just drawn is his own reduc51. Harvey thought that the blood percolated through pores or channels in the tissues; see esp. his Exercitatio de motu cordis et sanguinis in animalibus (Frankfurt: sumpt. Fitzeri, 1628), ch. 7. 52. Descartes did not have the idea of a pulse wave but of a simultaneous enlargement of all arteries synchronized with the forced diastole of the heart. AT 11:125. 53. Man, AT 11:126-127. 54. For Descartes on this, see Anatomica quaedam ex Mt? Cartesii (a manuscript from the hand of Leibniz), first publ. in Oeuvres inedites de Descartes (Paris, 1859-60), AT 11:596-598. See also Galen, De naturalibus facultatibus K 2:82. Also, for a possible Scholastic source of Descartes on this concept, Gilson, Index, n. 37 above, art. 508, p. 333.

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Descartes'PhysiologicalMethod tive interpretation of immutation (as far as the body-solids are concerned). Incidentally, the idea of maturation, of aging, as a process of gradual hardening and drying was also preCartesian. Aristotle had said that "the matter of which bodies are composed among the living consists of hot and cold, dry and moist. But as they grow old they must dry up.""55 Galen said that "that which all men commonly call old age is the dry and the cold constitution of the body." 65 During the early Renaissance, the idea was common that the body's innate or "radical humour," being-unlike the other parts -irreplaceable, gradually dries up. Thus Pare (1575): "Now in old age men are cold and dry . . . [because of] the consumption of the radical or substantific humour proceeding from the multitude of years." 57 For Fernel, a body engendered of blood and semen must begin by being hot and wet. Weighing whether maturation is primarily a cooling or a drying process, he decided that both are involved.58 In Descartes' later Description of the Human Body (written 1648), he altered his model of fiber formation somewhat and made the fibers emerge from pores along the arterial walls instead of at the tips. He specified how they are eroded at their free outer ends. He also, in the later treatise, stipulated that particles of the first and second elements flow alongside the fibers, encouraging the continual outward movement of each from its arterial base.59 To sum up, what Descartes advances in connection with assimilation is an eclectic, reductive restatement of classical ideas, adapted to fit his own cosmological and physical doctrine. Example 2: Initiation of differentiative development We hear about generation, from Descartes, in some of his letters as well as in disconnected posthumous fragments and especially in the Description of the Human Body. Studies of generation had dealt, traditionally, with certain recurrent questions. What is the constitution of the seed-stuff, or germ? Where and how does it arise in the bodies of the parents? Do both father and mother contribute something to the offspring? 55. Aristotle, De longitudine et brevitate vitae, trans. W. S. Hett, On Length and Shortness of Life (Loeb Classical Library Series, Cambridge, Mass.: Harvard University Press, and London: Heinemann, 1935), 466a20ff. 56. Galen, De sanitate tuenda, trans. R. M. Green, Galen's Hygiene (Springfield, Ill.: Thomas, 1951), bk. 5, ch. 9. 57. A. Pare, Oeuvres, n. 14 above, bk. 1, ch. 9. 58. Jean Fernel, Physiologie, n. 13 above, bk. 3, ch. 10. 59. AT 11:245-250.

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If so, is the contribution of both sexes the same? What triggers the beginning of development? Through what subsequent events and in what sort of sequence does the organism acquire and mature each organ? This way of resolving the general problem into subproblems had been charted by Greek biomedical theorists and variously elaborated in Medieval and early Renaissance science. Descartes addressed himself to some-but only some-of the classic subproblems into which the general problem of generation had been resolved by earlier thinkers. Though influenced, here as elsewhere, by a mixture of past and prevailing ideas, he perhaps came nearer on this than on other biological subjects to a theory distinctly his own. Assume three elements, he argues, differing only in the shapes and sizes and motions of the particles they comprise. And assume these particles to be subject to orderly varieties of mechanical interaction. How account, on the basis of these assumptions, for the sequential appearance-commencing with an undifferentiated initiative substance-of: first, the future left ventricle of the heart; next, the future aorta with its primary branches the carotids and spermatics; then, related to the foregoing, the rudiments of the brain and genitalia; then, in relation to the brain, the sensory nerves and sense organs; also, at about the same time, certain major arteries (and their branches) and veins (and their branches); and, finally, the fibrous micro-units that constitute the solid organs? Note, in the following example, how Descartes sees the process as beginning and how he gives it a typically Cartesian, corpuscular interpretation. I assert nothing definite touching the shape and arrangement of the particles of the seed. Suffice it to say that the seed of plants, being solid and hard, may have its parts arranged and situated in a definite way which could not be altered without their being rendered ineffective. But it is not the same with the seed of animals, which, being very fluid and ordinarily produced by the coming together of the sexes, seems to be only a mixture compounded of two liquors which, serving each as a ferment to the other, are so heated that some of the particles, acquiring the same agitation that fire has, move apart and press against others, and by this means gradually arrange the latter in the way required to form the members [of the body].60 The foregoing introduction to the subject of generation is 60. AT 11:253.

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Descartes' Physiological Method partly an echo of pre-Cartesian opinion. The Greeks had transmitted three speculations, or streams of speculation, about the seminal substance. Either (1) (2) (3)

two similar-or equally important-seminal substances are involved, one supplied by each parent; or a single substance is needed, and this is supplied by the father; or two different substances are supplied: semen by the father and blood (or blood and female semen) by the mother.61

The two-semina theory appeared, pre-Platonically, in the writings of Democritus (probably)62 and in the Hippocratic treatise On Regimen.63 The idea of a single-seed stuff was taken over from Alcmaeon by Plato, whose "panspermia" (also "marrow," myelos) is depicted as descending from the brain, by way of the spinal canal, to the urethra for transfer to the "plowed soil" of the womb.64 Plato gives this idea a rather cryptic formulation; it had a number of Medieval and Renaissance revivals.65 61. For a rather different classification of ancient ideas on the seed-stuff, see Erna Lesky, 'Der enkephalomyogene Samenlehre,' "Die Zeugungs-und Vererbungslehren der Antike und ihr Nachwirken," Abhandlungen der geistes- und sozialwissenschaftlichen Klasse, Akademie der Wissenschaften und der Literatur in Mainz (Wiesbaden, 1950), pp. 1233-1254. 62. As reported by Aristotle, De generatione animalium, 721b6-722a2; see also Hermann Diels, Die Fragmente der Vorsokratiker, 7th ed., edited with additions by W. Kranz (= a photographic repr. of the 5th ed.), Berlin: Weidmann, 1954, 68 A 41 and B 32. 63. "On Regimen," Hippocrates, trans. W. H. S. Jones (Loeb Classical Library Series, Cambridge, Mass.: Harvard University Press, and London: Heinemann, 1931), vol. 4, bk. 1; the theory is contained, also, in the two treatises (which Littr6 combines into one) "On Generation" and "On the Nature of the Child," Oeuvres completes de Hippocrates, ed. Littr6 (Paris: Bailli6re, 1851), 7, 470-543. 64. Plato, Timaeus, 90E - 91D. A similar idea appears in the Hippocratic treatise "On Generation," see above, n. 63, pp. 472-473, but, there, it is in both sexes that seminal substances descend from the head, via the spine, to the genitalia. The metaphor of the womb as a field was a commonplace thereafter until ca. 1700. 65. One of the famous coition-figures of Leonardo shows two channels in the penis, one connected with the spinal marrow, the other with the testes. For a rather late pre-Cartesian adaptation of this theory, see Jacobus Sylvius (Jacques Dubois), Livre de la generation de 1'homme recueilly des antiques & plusseurs autheurs de medecine & philosophie, trans. G. Chrestian (Paris: Morel, 1559), p. 25 and esp. his "Livre de la nature et utilit6 des moys des femmes," bound with the foregoing, pp. 113-116. (Note: these two works combined were published earlier as De mensibus mulierum et hominis generatione . . . Jacobi Sylvii . . . commentarius (Paris, 1555; Basel, 1556.)

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The third idea, of semen and blood, was elaborated by Aristotle, who linked generation with nutrition. Both parents' bodies, he said, "concoct" the nutriment they absorb in order to ready it for assimilation to the tissues, a process entailing the actualization of the food's unexpressed morphological potential. An unassimilated residue of concocted nutriment passes to the genitalia for further concoction into semen in the male and catamenial blood in the female. The latter is less highly elaborated-possessing only vegetative potentialities-than the former, which possesses sensitive potentialities as well. Semen acts on the catamenia, at coition, to commence an actualization of its morphogenetic potentialities by a kind of "setting" or curdling effect. The process is abetted by an indispensable but inadequately explicated pneuma brought in with the semen.66 Galen gives two principal, and a number of peripheral, accounts of the origin of the offspring. One of these is a Galenized adaptation of Aristotle's idea; it depicts the seminal pneuma as vehicle for an alterative faculty which changes the blend of elements in the catamenia so as to convert it into tissues.67 The other account eliminates the catamenial blood as a seminal substance, substituting intravascular blood from the mother. The semen, with its pneuma, is coagulated by contact with the womb and forms a capsule. Vascular (not catamenial) blood of the mother, along with pneuma and heat, penetrates the capsular membrane in multiple streams, which come together inside to form the umbilical vein. Some (mostly fleshy) organs derive from the blood; other (mostly more solid and membranous) organs, from semen.68 The foregoing and other Greek ideas reappeared, variously modified and combined, in Medieval and Renaissance physiological theory. The two-semina scheme was variously adapted by Paracelsus,69 Fernel (see below), Jacobus Sylvius (Jacques Dubois),70 du Laurens,71 Realdo Colombo,72 Caspar Bartho66. De partibus animalium, 647b4-7, 650a4-15, 678al-20; De juventute, 468alO, 469a27 to b20. De generatione animalium, 727b30, 729a22-b19, 732alO, 738b20ff. 67. Galen, De naturalibus facultatibus, n. 54 above, bk. 2, ch. 3. 68. Galen, De semine, bk. 1, chs. 9 and 10, K 4:545-552. 69. Paracelsus, "Das Buch von der Gebarung der empfindlichen Dinge in der Vernunft (Tractatus secundus)" Paracelsus samtliche Werke, eds. K. Sudhoff and W. Matthiessen (Munich: Barth, and Munich and Berlin: Oldenbourg, 1922-33), 1, 257-265. 70. Sylvius' theory appears to be a synthesis of Galenic with Hippocratic beliefs, especially as the latter were developed in the two treatises

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Descartes' Physiological Method and many others. Note, for example, Fernel's adaptation which served with others as grist for the mill of Descartes: Fernel posited similar male and female seminal fluids, made of highly elaborated blood supplemented by three pneumata (vital, natural, and animal) plus a set of other pneumata that rush to join the seminal fluid at the moment of orgasm. These pneumata carry corresponding faculties of the soul. The activation of the seminal me'lange is due to a power peculiar to the womb which creates a capsule that is relatively warm and subtle within, cold and earthy without. Within this capsule a special faculty arises to guide the steps of morphogenesis, commencing with bladders representing the future liver, and brain, and heart.74 Fernel's theory is an amalgamation of Galenic facultative pneumatology with the pre-Platonic "similar semina" doctrine. Descartes retains the sinilar semina but substitutes micromechanisms for the faculties and pneumata:

lin1,73

And for this [reciprocal fermentative activation] the two [male and female seminal] liquors need not be very different. For, as we see that old dough can make new dough rise, and that the foam that beer throws up suffices as a ferment for other beer, so it is easy to believe that the seminal liquids of the two sexes, being mingled, serve as ferments to each other. Now I believe that the first thing that happens in the mixture of seminal fluid, and that makes every drop of it stop resembling every other drop, is that heat is excited there which, acting as in effervescent new wines, or in hay when stored before dry, makes some of the particles gather near a particular part of the containing space; and these particles, expanding there, press against others that surround them; which starts to form the heart.75 mentioned in n. 65 above. Sylvius seems to envision (a) apparently equivalent male and female semina (Galen, too, acknowledged a female semen but assigned it an auxiliary rather than a participative role, De semine, K 4:536-538) and also (b) the catamenia; (a) and (b) give rise, as in Galen, to seminal and sanguinary tissues respectively, see above n. 68. 71. A. du Laurens, Toutes les oeuvres de M' A. du Laurens . . , trans. T. Gelee (Rouen: R. du Petit Val, 1621), pp. 240-242. 72. Realdo Colombo, De re anatomica libri XV (Venice: Bevilacqua, 1559), p. 246. 73. C. Bartholin, Anatomicae institutiones (Strassburg: Scher, 1626), pp. 125-126. 74. J. Fernel, Physiologie, n. 13 above, bk. 7. 75. AT 11:253-254. Descartes' point that the male and female semina need not be very different from each other harks back to an earlier idea he

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Descartes follows, here, his usual procedure of recrystallizing mixed traditional ideas along the lines of Cartesian particle theory. There was nothing new in linking the act of conception to the manifestation of ebullient heat. The Hippocratic treatise "On the Nature of the Child" had argued that a heating of newly mixed male and female semina in the womb produces pneuma in a process comparable to burning green wood or foliage (a smudge)3.78 Aristotle, discussing the fate of the semen, depicted it as dissipated through vaporization,77 and Galen, writing critically, later asked whether Aristotle meant this process to resemble the effervescence of wines. Galen agreed with the idea of a vapor produced at about the time of conception but not with an Aristotelian anathymiasis of the semen as a whole; both the pneuma and the semen persist, Galen said, and are used in building the brain and other parts.78 Pre-Cartesian Renaissance theorists (ca. 1542-1632) had mostly attributed the initiative heat in the semen to the influence of its intra-uterine surroundings. The womb arouses the dormant developmental faculties of the semen, according to this view; and it also provides a milieu for "fomentation." Fernel, for example, compared the effect of the uterus on semen to that of the stomach on food.79 Externally effected fomentation-rather than spontaneous fermentation-was acknowledged by du Laurens,80 Bartholin,8' Crooke,82 and other pre-Cartesian authors. Descartes' view differed from theirs. It was in line with his physiological method to liken the heat of conception to a fermentation or leaven. Such heat-producing chem"fire-without-light"-were a subject ical actions-generating had that the lungs and liver form first, and that spirits from the former and blood from the latter then meet in a heat-producing, combative interaction to form the heart. AT 11:508-511 and 599. He has changed his mind. 76. Oeuvres de Hippocrates (ed. Littre), n. 63 above, vol. 7, p. 487. 77. Aristotle, De generatione animalium, 737al0-15. 78. Galen, De semine, bk. 1, ch. 8, K 4:540. 79. J. Fernel, Physiologie, n. 13 above, p. 733. 80. A. Laurentius, Toutes les oeuvres, n. 71 above, p. 248. 81. C. Bartholin, Anatomicae institutiones, n. 73 above, pp. 125-126. 82. Thus: "The wombe rowzeth and raiseth upp the sleepy and lurking power of the seeds, and that which was before but potentiall, it bringeth into act ... The generative faculty which before lay steeped, drowsie, and as it were intercepted in the seede, being now raised up by [the] heat and inbred propriety of the wombe breaketh out into acte, as raked Cinders into a luculent flame." He goes on to attribute to pneuma ("wherewith the frothy seed swelleth") the role of a builder or painter, acting in response to the soul (Mikrokosmographia . . ., 2nd ed. [London: Sparke, 1631], pp. 262-264).

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Descartes' Physiological Method he never wearied of discussing, usually in terms of his own corpuscular theory of matter. An important chapter in the scientific system of Descartes is his handling of the physics of motion, that of individual particles as well as of aggregates thereof. All motion in his theory was, originally, God-given. And, though motion may be transferred from particle to particle or from aggregate to aggregate, the total amount of it in the universe as a whole, he said, never varies. A particle or aggregate, once moving, tends, unless resisted or deflected, to continue moving without any change of direction.83 But, since the universe is a plenum, the movement of particles or aggregates to any locus entails a displacement of the particles or aggregates already there. A natural destination for those thus displaced is the former locus of those that displaced them. The resultant movement is circular in pattern, and the cosmos, as depicted by Descartes, contains many examples of cycles of displacement-and-replacement.84 Note how Descartes utilizes his theory of motion in continuing his analysis of embryonic development. Next, since particles thus expanded [by fermentative heating in the heart-region] tend to continue to move in a straight line; and since the heart-beginning to take form-resists them, they move off a certain distance and make their way toward the place where the base of the brain will later be formed, and in so doing they displace certain other particles which circle back to replace them in the heart. There, after a brief period needed to bring them together, they expand and move out along the same path as the preceding [toward the future brain region]. And this causes some of those that went there before and which happen still to be there to come again to the heart-along with others that come in from other places to take the place of those that, all this while, have been leaving. And those [that thus arrive in the heart], being promptly expanded, leave in their turn. And it is this [heat-induced] expansion, occurring over and over, that constitutes the heartbeat, or pulse. Outflow from the heart is thus circularly balanced by inflow so that arteries and veins are generally formed in pairs. Such 83. Principles, pt. 2, sects. 36-43 (AT 8:61-67 and [9:83-88]). 84. Such cycles had an antecedent in the biophysics of Plato discussing respiration, said that exhaled fire and air particles environmental fire and air particles which, in his opinion, enter the pores of the body to replace those being exhaled. The cycle then itself, fire and air moving out through the pores and displacing fire and air which make room by being inhaled (Timaeus, 79A-E).

who, in displace surfacereverses ambient

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flows are envisioned as liquid streams moving in a less liquid matrix (morphogenetic currents had been stipulated by Galen;85 after a rather uneven career they were still being invoked in 1809 by Lamarck).86 With considerable ingenuity, and fidelity to his physics, Descartes goes on to detail how, later, the currents become surrounded by membranes (blood-vessel walls). And how, in the case of arteries, the pulsing of the membranes permits the extrusion of particles-and the consequent formation of fibers that constitute the body-solids. Among such solids are the walls of the heart, whose derivation from particles extravasated by the coronary artery is fully detailed. Descartes continues to construct the body with a kind of gratuitous precision that is likely to repel the reader who forgets the author's intention. What he is building, so he assures us, is a hypothetical model-mostly a micromodel-not of man but of a mechanism that simulates man. To this model-a kind of conceptual robot-we shall return after listening briefly to Descartes on the subject of sensory reception. Example 3: The sense of smell In the microneuroanatomy of Descartes, the functional peripheral units are hollow nerve-tubules, each containing several longitudinal fibrils surrounded by animal spirits.87 The fibrils, if peripherally disturbed, act (comparably to a bellpull)88 to initiate, in the brain, a reflexive outflow of spirits. Flowing back through the same-or out through other-nerve-tubules, the spirits act, in a very special way, to trigger muscular contraction.89 We shall not concern ourselves here with the role of the pineal gland, man which Descartes notoriously saw as intermediating-in 85. See above, n. 78. 86. J. B. Lamarck, Philosophie zoologique . . ., first publ. Paris, 1809, 2nd ed. (Paris: Bailliere, 1830), 1, 409. 87. Discourse AT 6:54; Corps humain AT 11: from 129. Spirits were for Descartes just as inanimate and corpuscular as other things. They are "all bodies consisting of terrestrial particles that [a] are bathed in subtle matter and [b] are more agitated [by their direct contact with particles of the first element] than those of air but less so than those of flame" (Letter to Adolphus Vorstius, AT 3:687). For several score further references to animal spirits (psychic pneuma) in Descartes, see Gilson's Index, p. 99. 88. The arriving spirits do not pump up the muscle; they operate certain valves that regulate the flow of spirits, already present, from the flexor to the extensor or vice versa. See Man, AT 11:133-137 (here he discusses the reciprocal action of muscle antagonists on which see also AT 11:336) and 142 (here he uses the bell-pull analogy, on which see also AT 11:337). 89. Considerable detail on the micromechanism of reflexes is given at Man, AT 11: from 170, and at Passions, AT 11:338-342; also, letter to Mersenne, AT 3:123.

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Descartes' Physiological Method alone-between the intracerebral flow of spirits on the one hand and the activities of the consciously motive-and-perceptive soul.90 What we wish rather to stress is that whatever aspect of Descartes' neurobiology we examine (we have chosen olfaction for convenience), we find the same corpuscularizing analysis that we discovered in his treatment of assimilation and the initiation of development. We find, too, that what he explained were not empirical data but earlier writers' opinions (disassembled and reassembled with additions and deletions). Thus, his theory of sensation in general was a reductively reinterpreted synthesis of already existing ideas-mostly sixteenth- and early seventeenth-century revisions of Platonic, Aristotelian, and Galenic theories of perception. Galen had regarded the sensory nerves of the head as protrusions of the brain-substance, permitting an extension of the sensitive faculties of the soul to the organs of special sense. The nerves contain psychic pneuma which acts as a substrate for the faculty extended by the nerve. Galen had considered smell to be the only sense mediated entirely inside the brain. He reasoned that odoriferous matters pass first through holes in the ethmoid bone and then through the presumably permeable floor of the brain itself, within which the soul's olfactory faculty resides. The same apertures in bone and brain permit an inflow of air (for conversion to animal spirits) as well as an outflow of excremental excesses (if these are superabundant; otherwise, they drain postnasally via the palate).'il During the sixteenth century, the status of the mamillary processes (our olfactory tracts with terminal bulbs) was debated. Should they or should they not be thought of as nerves? Vesalius (1543) was noncommittal on this subject.92 Realdo Colombo (1559) considered the terminal thickenings of the mamillary processes (our olfactory bulbs) to be the proper organs of smell.93 Piccolhomini (1585) agreed, and to him this seemed to project smell to a locus outside the brain (though not outside the cranial cavity); he called the olfac90. For which see: Man, AT 11: from 175; Passions, AT 11:351-352; and two letters, D. to Meysonnier, AT 3:18-21 and especially D. to Mersenne, AT 3:262-265. Descartes' chief reason for choosing the pineal was that he wanted a single organ "inside" the brain ventricles where impressions from paired organs (eyes, ears) could form a single image; more generally the soul was, for him, unitary. 91. Galen, De usu partium, bk. 8, cbs. 6, 7, K 3:647-656. 92. A Vesalius, De humani corporis fabrica (Basel: ex off. Oporini, 1543), pp. 322-323, 643. 93. R. Colombo, n. 72 above, pp. 193-194.

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tory tracts nervi odorati because they seemed to connect the bulbs with the brain.94 Caspar Bauhin (from 1597) took a similar position, pointing especially emphatically to the existence of olfactory nerves (where we see olfactory tracts).95 In this sequence of ideas we witness a tendency to place the olfactory receptor farther and farther from the brain ventricle, where it had been located by Galen. Descartes carried the same tendency one step farther. The sense of smell, as well [he has just been speaking of taste], depends on several fibrils that extend from the base of the brain toward the nose beneath those two little hollowed-out parts that anatomists have likened to nipples [olfactory tracts, termed processus mammilares by Renaissance anatomists]. And these fibers are in no way different from nerves that serve for touch and for taste, except that [a] they do not extend outside the cavity of the head that contains the whole of the brain and [b] they can be moved by smaller earthy part[iclels than can the nerves of the tongue both because they are slightly finer and because they are touched more directly by the objects that move them. For you should know that when this machine [this hypothetical mechanical analog of a real man] breathes, the subtlest air part[iclels that enter it through its nose, passing through the pores of the bone denominated spongy [ethmoid] penetrate if not all the way into the brain cavity [as stipulated by Galen] at least as far as the space between the two membranes that envelop the brain [the subdural space]. From this space, particles may simultaneously leave through as, recipthe palate [again, as stipulated by Galen]-just rocally, when air leaves the chest, its particles can enter this [subdurall space by way of the palate and leave by way of the nose.96 [You should] also [know] that on entering this [subdural] space they encounter the ends of the [aforementioned] fibrils which are quite bare, or covered with so extremely delicate a membrane that little force is needed to move them. The foregoing paragraphs well illustrate the paradoxical imby novelty combined with familiarity-created pression-of 94. A. Piccolhomini, n. 15 above, p. 292. 95. See, e.g., C. Bauhin, Theatrum anatomicum (Frankfort: Becker, 1605), pp. 643-644. 96. For Descartes elsewhere on the respiratory current see his Excerpta Anatomica, n. 54 above, AT 11:599-600.

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Descartes' Physiological Method Descartes' explanatory method: the paradox stems, in this case, from his fusion of a rather new view of biodynamics (based on his own corpuscular physics) with ancient errors of Galen about the flow of matters into and out of the brai and skull. The correct idea that olfactory fibrils actually traverse the cribiform plate was put forward, with certain errors, *by Conrad Schneider (1654) and Thomas Willis (1664).97 You should also know that these pores [in the ethmoid bone] are so arranged, and so narrow, that they prevent access to these fibrils of particles coarser than those which, in speaking earlier on this subject, I designated Odors-except, perhaps, for certain ones that constitute eaux de vie because their shape renders them especially penetrant. Finally, you should know that among the extremely small earthy particles that are always found in greater abundance in air than in other composite bodies, only those which are [a] a little coarser or [b] a little finer than the others-or which because of their shape are more or less easily moved -will be able to occasion in the soul the different sensations of odors. Similarly, only those in which these excesses are vcry moderate and mutually tempered will cause agreeable sensations, for those which act only ordinarily will not be able to be sensed at all; and those that act with too much or too little force cannot but be unpleasant.98 CONCLUSION The three cases just cited are merely examples of Descartes' analytical method, but they typify rather well his approach to physiology in general. Whatever the immediate explanandum -heart action, respiration, reciprocal innervation, muscular antagonism, secretion, digestion, absorption, blood-formation, is discovnervous action, bio-optics, bio-acoustics-Descartes ered to follow a fairly predictable practice-namely, a reductive (corpuscular, nonpsychistic) interpretation partly of empirical fact but primarily of earlier Renaissance revisions of Greek physiological doctrine. His sources are often only semirecognizable because of the reconstruction to which he submits them in preparing them for "cartesianization." As for the physics to which he assimilates his biological data, that too 97. C. Schneider, Liber de osse cTibriforme . . . (Wittenberg: Mevius & Schumacher, 1655), p. 169, and T. Willis, Cerebri anatome, first publ. London, 1644, trans. in S. Pordage, Dr. Willis's Practice of Physick, see n. 25 above, p. 112. 98. Man, AT 11: 148-149.

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is partly his own and partly an altered conceptualization of earlier elementary-particle theory.99 To what extent is Descartes' procedure-that, namely, of re-explaining not empirical data so much as earlier explanations-the procedure of theorybuilders in general? We leave this question for separate and more extended exploration. From the point of view of scaling, it would be correct to think not of one but of three mechanical sciences as arising during the sixteenth and seventeenth centuries: a celestial or megamechanics treating such very large things as the earth and the heavenly bodies; an intermediate mesomechanics having to do with usable machinery, automata,100 and so on, and with their biological analogs, namely, plants and animals and their visible parts; and finally a micromechanics concerned with subvisible things, ranging downward in size from those which would presently become visible through the microscope all the way to elementary particles.'01 Descartes reasoned as a mechanist on all of these levels. In his biology he drew a number of comparisons between the body-parts and various sorts of visible machinery, water-works, clocks, and the like. He made no sharp distinction between meso- and micromechanics, but if we take the lower limit of (unaided) visibility as the line of division, the mesomechanical allusions in his works are, though trenchant, relatively rare: his biology is mostly microrather than mesomechanical. A question finally remains as to the epistemological status of the "Man" whom (or which) Descartes portrays. With what in mind does he picture not man himself-so he assures usbut, rather, a hypothetical analog of man? A clue is contained in the Treatise of Light, where we read that the "World" that Descartes would portray is not the one that actually exists. It is merely a possible world, one that God could have created 99. The Greek, Medieval, and Renaissance sources of Cartesian physics have been the subject of much historical study. See Marie Boas, "The Establishment of the Mechanical Philosophy," Osiris, 10 (1952), 412-541; and J. R. Partington, "The Origin of the Atomic Theory," Annals of Science, 4 (1939), 245. 100. See, on automata and mechanicism, D. J. de S. Price, "Automata and the Origins of Mechanism and the Mechanistic Philosophy," Technology and Culture, 5 (1964), 9-42. 101. Thus, Robert Boyle in 1674: "the mechanical affections of matter are to be found and the laws of motion take place not only in great masses, and in middle sized lumps, but in the smallest fragments." See his "Of the Excellency and Grounds of the Mechanical Hypothesis," The Works of the Honourable Robert Boyle in Six Volumes .. . (London: J. & F. Rivington, 1772), 4:71.

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Descartes'PhysiologicalMethod had he wanted to construct a mechanical analog of the world he created in fact. Did this seemingly ambiguous presentation -of man and the world-stem from Descartes' willingness to guard himself, or his system, against ecclesiastical censure? Historians have supposed that it did; and we know that the example of Galileo partly caused Descartes to postpone publication of his own Treatises of Light and of Man. Another interpretation of Descartes' tentativeness has often been suggested: He was notoriously aware of the limitations of sensation, but he was also aware of the limitations of reason. He saw himself not as stating the truth but as developing a model -a metaphor-that somehow squared with truth on the one hand and with sensory experience on the other. In the Principles, he expresses the wish that "what I shall write be taken as only an hypothesis which may be very far from the truth"; and he continues that "even though it be such [only an hypothesis] I shall think I have done much if all the things which shall be deduced from it are entirely conformant to experience; because if that be the case, it will be no less useful to life than if it were true, because one will be able to use it just as well in arranging natural causes to produce desired effects." 102 102. Principles, pt. 3, sect. 44 (AT 8:99 [and 9:123]).

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Harvey and Fludd: The IrrationalFactor in the Rational Science of the Seventeenth Century ALLEN G. DEBUS Department of History University of Chicago

Only a few years ago Charles Singer, referring to William Harvey's sources in the De Motu Locali Animalium, remarked: "He mentions Laurentius (once), Vesalius (once) and-save the mark-the Rosicrucian Robert Fludd (once). The last is important for the history of human error, but can have no place in the history of science." 1 Singer's statement is understandable if we insist upon evaluating the science of earlier periods from our post-Newtonian vantage point. If we do this, the "mystical-alchemist" Fludd surely seems far removed from the "experimentalist" Harvey. When the relevant texts are viewed in historical context, however, the relationship between these men becomes far less difficult to understand.2 Not only do we find that they were colleagues and friends; we also find that they were deeply 1. Book review of William Harvey's De Motu Locali Animalium, 1627, ed., trans., and introduced by Gwenneth Whitteridge, M.A., D. Phil., F.S.A. (Cambridge: University Press for the Royal College of Physicians, 1959) in Brit. Med. J. (1960), 1202. 2. "Instead of selecting data that 'make sense' to the acolyte of modern science, the historian should therefore try to make sense of the philosophical, mystical or religious 'side-steps' of otherwise 'sound' scientific workers that are usually excused by the spirit or rather of the past-'side-steps' backwardness of the period. It is these that present a challenge to the historian: to uncover the internal reason and justification for their presence in the mind of the savant and their organic coherence with his scientific ideas. In other words it is for the historian to reverse the method of scientific selection and to re-state the thoughts of his hero in their original scientific and the non-scientific-will setting. The two sets of thought-the then emerge not as simply juxtaposed or as having been conceived inspite of each other, but as an organic whole in which they support and confirm each other. There is no other way to lay the savant open to our understanding." Walter Pagel, William Harvey's Biological Ideas: Selected Aspects and Historical Background (Basel/New York: S. Karger, 1967), p. 82. Journal of the History of Biology, Vol. 3, No. 1 (Spring 1970), pp. 81-105.

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concerned at an early stage of their work-and at the same time-with the problem of the blood flow. For both "circulation" was seen to be the answer.3 And if on the one hand-much to the distaste of Singer-we see that Harvey was willing to cite Fludd, on the other hand we find Fludd convinced that the work of Harvey essentially represented a confirmation of his own mystical scheme of the circulation. Even more important, we find that some of their contemporaries agreed that the work of these two men should be discussed together. Both in the first notice of Harvey's book by continental authors (Mersenne and Gassendi, 1629), and also in the debate between John Webster and Seth Ward (1654), the relative merits of the works of the two English physicians were compared and evaluated. It is not the purpose of the present paper to show that Fludd's system of the circulation was "scientific" in a modern sense-it was not. Rather, the purpose is to give additional information on the genesis of Fludd's views, and further, through a study of the disputes of 1629-1633 and 1654 to show that what he had to say was taken so seriously by recognized scholars that it had to be discussed in relation to the work of Harvey. THE GENESIS OF FLUDD'S VIEWS ON THE BLOOD It should not be necessary here to review the scientific background of Harvey's discovery of the circulation. This is done admirably by Pagel in William Harvey's Biological Ideas. Suffice it to say that before Harvey there had been a long history of circular symbolism comparing macrocosmic and microcosmic events. This is an analogy which is clearly stated in the De motu cordis.4 Similarly, one finds in the earlier literature 3. Walter Pagel, Harvey's Biological Ideas, p. 337. Fludd's early acceptance of Harvey's discovery was first noted by Pagel in his "Religious Motives in the Medical Biology of the XVIIth Century," Bull. Hist. Med., 3 (1935), 277-278. The present author has discussed this problem in "Robert Fludd and the Circulation of the Blood," J. Hist. Med., 16 (1961), 374-393, and in The English Paracelsians (London: Oldbourne Press, 1965), pp. 114-118. 4. The relevant passages are discussed in detail by Pagel, pp. 82ff. and passim. In addition to Fludd, John Donne was expressing an interest in the blood flow in the early seventeenth century. Thus, he speaks of this topic in his Second Anniversarie (1611); and in a sermon delivered April 8, 1621, he refers to the capacities of bodily organs, including the ventricles of the heart. See F. N. L. Poynter, "John Donne and William Harvey," J. Hist. Med. 15 (1960), 233-246, and Geoffrey Keynes, Kt., The Life of William Harvey (Oxford: Clarendon Press, 1966), pp. 121f.

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Harveyand Fludd references to a circulation of the blood in the concept of a chemical "distillation-circulation" of the pelican. These views were not expressed in folios gathering dust in forgotten libraries, but rather were part of the current literature in Harvey's formative years. Nor need one "force" the evidence to show Harvey's connection with Fludd. The former was only four years younger than the mystical alchemist, and he was admitted as a Fellow of the Royal College of Physicians just two years prior to Fludd. Both men were interested in anatomy, and Fludd refers not only to his own dissections, but also to having watched Harvey perform anatomical dissections of the heart in search of the elusive interventricular pores of the septum. At the same time, Harvey cites Fludd more than once in his writings, and the two men shared a common interest in pharmacological problems, particularly the compounding of chemical medicines.5 More important perhaps is the fact that Fludd calls Harvey his friend, and that Fludd was instrumental in having Harvey's work printed by his own continental publisher. The special significance of Fludd derives from the fact that his Pulsus (completed in 1629) contains the first approval of Harvey's doctrine in print and that he himself had discussed the circulation of the blood in the traditional "spiritual" and "chemical" contexts in his Anatomiae Amphitheatrum, which had been completed in 1621 and published two years later.6 As Pagel points out, it was sometime in the early 1620's that the concept of the circulation took firm shape in Harvey's mind.7 Even from the limited number of documents still in existence from the period prior to 1620 we may assume that neither author had yet elaborated a circulatory system. In 1616 Harvey began his Lumleian lectures at the Royal College of Praelectiones Anatomiae Universalis. In his Physicians-the lecture notes the section on the heart does not refer to any circulation of the blood, and although most of the extant 5. This evidence is presented by Pagel in Harvey's Biological Ideas, pp. 113-119. 6. Robert Fludd, Pulsus seu nova arcana pulsuum historia, e sacro fonte radicaliter extracta, nec non medicorum ethnicorum dictis & authoritate comprobata (Frankfort?, n.d.), p. 11. A large folding plate, generally missing from this work, is dated 1630. On the Pulsus reference see Pagel, "Religious Motives," p. 277f. The circulation scheme in the Anatomiae Amphitheatrum was first noted by the present author in the article cited above in fn. 3. 7. Pagel, Harvey's Biological Ideas, pp. 119, 213ff.

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manuscript have been dated 1616-1618, it was only at some later date-and in an empty space on the back of a sheetthat Harvey in a short note compared the heart with a water bellows and referred to the total circulation of the blood.8 With Fludd we find an interest in the blood and its motion occurring at precisely the same time (1616/1617). The relevant passage is found in his defense of the Rosicrucians. Today it is difficult to understand the excitement caused by the publication of the Fama Fraternitatis and the Confessio in 1614 and 1615. These works decried the atrophied learning of the Schools and made a plea for a new learning.9 Contemporary scholars, their author complained, stiUl pore over the works of Porphyry, Aristotle, and Galen when instead they should be seeking a more perfect knowledge of the "Son Jesus Christ and Nature." All truly learned men agree, he asserted, that the basis of natural philosophy is medicine and that this is a godly art. No other Philosophy we have, then that which is the head & sum, the foundation and contents of all faculties, sciences and arts, the which (if we behold our age) containeth much of Theology and medicine, but little of the wisdom of Lawyers . . ." 10 8. See ibid., p. 213, and note 16 on the dating of this note. the early Rosicrucian texts will be discussion-of 9. A listing-and found in F. Leigh Gardner, A Catalogue Raisonn6 of Works on the Occult Sciences, vol. 1: Rosicrucian Books, intro. by Dr. William Wynn Westcott, 2nd ed. (privately printed, 1923). Here, on pp. 4-6, 21 (items 23-29, 144), will be found a discussion and description of the first editions of the Fama Fraternitatis and the Confessio (1614, 1615). From this it would appear that seven editions of the Fama were published in the years 1614-1617 in German and High Dutch. An English translation of the Fama and the Confessio is available in The Fame and Confession of the Fraternity of R:C: Commonly, of the Rosie Cross. With a Praeface annexed thereto, and a short Declaration of their Physicall Work, by Eugenius Philalethes (Thomas Vaughan), London: J.M. for Giles Calvert, 1652. All references will be made to this edition. No satisfactorily unbiased modern history of the Rosicrucians exists. The most extensive discussion in English will be found in A. E. Waite, The Real History of the Rosicrucians, founded on their own manifestoes and on facts and documents collected from the writings of Initiated Brethren (London, 1887). An early discussion of their work will Kirchen- und Ketzerbe found in Gottfried Arnold, Unparteyische Historie vom Anfang des Neuen Testaments biss auff das Jahr Christi 1688, 4 parts in 2 vols. (Frankfurt on Main: Thomas Fritsch, 1700), I (pt. 2), 613-628. Frances A. Yates touches on the Rosicrucian movement and early modern science in her "The Hermetic Tradition in Renaissance Science," Art, Science, and History in the Renaissance, ed. Charles S. Singleton (Baltimore: John Hopkins Press, 1968), pp. 255-274. 10. Fame and Confession, pp. 1-2, 36.

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Harvey and Fludd The founder of the Rosicrucian Order, allegedly one Christian Rosenkreuz, had learned great secrets of medicine and mathematics in Africa, where he had translated the incomparable Book M into Latin from Arabic. These truths, according to the author of the Fama, were available to the Brothers of the Order, but if one searched with open eyes, he would find in the Germany of that day many learned Magicians, Physicians, and Philosophers. Such a one likewise hath Theophrastus been in Vocation and Callings, although he was none of our Fraternity, yet nevertheless hath he diligently read over the Book M: whereby his sharp ingenium was exalted; but this man was also hindered in his course by the multitude of the learned and wise-seeming men, that he was never able peaceably to confer with others of his knowledge and understanding he had of Nature. And therefore in his writing he rather mocked these busie bodies, and doth not shew them altogether what he was: yet nevertheless there is found with him well grounded the aforenamed Harmonia, which without doubt he had imparted to the Learned, if he had not found them rather worthy of subtil vexation; then to be instructed in greater Arts and Sciences; he then with a free and careless life lost his time, and left unto the World their foolish pleasures."' Beyond the special books of Christian Rosenkreuz and the Brethren themselves, only the work of Paracelsus is mentioned as being concealed within the hidden vault of the Rosicrucians. This was a neo-Paracelsian and an alchemical movement. At the same time there was a missionary zeal associated with these documents. How much might be accomplished if the truly learned scholars of Europe united for the benefit of mankind? Yet the truly learned must be found elsewhere than at the universities. They must declare themselves and join the Brotherhood in this great reformation. Accordingly, one reads a plea in the Fama Fraternitatis addressed to all the learned scholars of Europe to examine their art "and to declare their minde, either Communicatio consilio, or singulatim by Print." Both the Fama and the Confessio were to be published simultaneously in five languages so that no one could excuse himself while the and say that he had not seen the message-and Brothers refused at this time to give out their names or an11. Ibid., p. 10.

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nounce their meetings, they were willing to assure those who answered their call that their words would not go unnoticed.12 The response to this appeal must have been beyond the wildest dreams of the promoters. In the course of less than ten years several hundred books and tracts appeared, debating the merits of this secret group. The most incredible part of this phenomenon is that neither then nor now has there been any proof that such a group existed. Although most who took part in this debate are not well known today, there were some participants of considerable renown. One of these was the German mystical alchemist Michael Maier (1568-1622), a man who was in England shortly before Fludd published his first book, perhaps to prepare a Latin translation of Thomas Norton's Ordinall of Alchimy. Deeply involved in the Rosicrucian controversy, Maier was to publish his commentary on the laws of the Rosicrucians, the Themis Aurea in 1618.13 Far less complimentary to the proposed new order was the respected and renowned iatrochemist, Andreas Libavius (1540-1616), who had for years attacked the mystical alchemists and Paracelsians. Among his last publications are the De Philosophia harmonica magica Fraternitatis de Rosea Cruce (1615) and the Analysis Confessionis Fraternitatis de Rosea Cruce (1615), in which he strongly censured the Order. Whether Fludd was inspired to write through contact with Maier in England,14 or simply through his reading of the surely appealed to him-is Fama and the Confessio-which unknown. We do know, however, that it was his strong feelings about this controversy that first moved him to publish at the age of forty-two. His Compendiaria Fraternitatis de Rosea Cruce (1616), a short pamphlet of twenty-three pages, was quickly revised into a two hundred-page book, the Tractatus Apologeticus, which was printed the following year.'5 Here 12. Ibid., p. 31. Gardner's bibliography of Rosicrucian works (see above, n. 9) lists seven editions, 1614-1617, but in only two languages. 13. Michael Maier, Themis Aurea, hoc est de Legibus, Fraternitatis Rosea Crucis (Frankfort: L. Jennis, 1618). An English translation dedicated to Elias Ashmole was printed by N. Brooke in 1656. Brooke published Ashmole's Theatrum Chemicum Britannicum (1652) and his The Way to Bliss (1658). 14. There is much doubt concerning Maier's activities in England. His relationship with Fludd has been discussed by C. H. Josten in "Truth's Golden Harrow. An Unpublished Alchemical Treatise of Robert Fludd in the Bodleian Library," Ambix, 3 (1949), 91-150, esp. pp. 99-101. 15. Robert Fludd, Compendiaria Fraternitatem de Rosea Cruce suspicionis et infamiae maculis aspersam veritatis quasi Fluctibus abluens et abstergens: auctore R. de Fluctibus, M.D. London (Leyden: G. Basson,

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Harvey and Fludd will be found Fludd's defense of the Rosicrucians against what he felt was an unwarranted attack by Libavius. Here also are his proposals for a new and reformed science, his appeal to the members to make themselves known to him-and here will be found evidence of his deep concern with the problem of the blood flow. Robert Fludd's concept of a newly reformed philosophy of nature is far removed from the concepts of the later mechanical philosophers. Fludd's search for the mystical secrets of the Pythagoreans and the hidden lore of other secret groups who professed a knowledge of the truths of antiquity led him to deep studies in the writings of kabbalistic, hermetic, and alchemical authors. His views on the macrocosm and the microcosm led him to a firm belief in a connecting link between the divine Creator and man. This essential link is the divine spirit-indispensable for our life, health, and spiritual well being. The source of this spirit is the sun, which is midway between earth and man in the heavens.16 As neither plants nor animals can exist without the sun and its light, we may be certain that it imparts something vital to us.17 1616); Tractatus Apologeticus Integritatem Societatis De Rosea Cruce defendens. In qua probatur contra D. Libavii et aliorum ejusdem farinae calumnias quod admirabilia nobis a FTaternitate R. C. oblata, sine improba Magiae impostura, aut Diaboli, praestigiis et illusionibus praestari possint. (Lugduni Batavorum: G. Basson, 1617). At the same time Fludd prepared in Libros tres distributus: quorum the Tractatus Theologo-Philosophicus (i), De Vita, (ii), De Morte, (iii), Resurrectione . . . collecta, Fratribusque d Cruce Rosea dictis dedicata, a Rudolfo Otreb Brittano (Oppenheim: J. Theo. de Bry, 1617). 16. A. G. Debus, "The Sun in the Universe of Robert Fludd," Le Soleil a la Renaissance-sciences et mythes, Colloque International tenu en Avril 1963 (Brussels, 1965), pp. 259-278. 17. A. G. Debus, "The Paracelsian Aerial Niter," Isis, 55 (1964), 43-61, and The English Paracelsians, pp. 114-115. C. H. Josten has discussed the important "wheat experiment" connected with Fludd's speculations on the aerial spirit in his "Robert Fludd's 'Philosophicall Key' and his Alchemical Experiment on Wheat," Ambix, 11 (1963), 1-23. The aerial spirit features prominently in all the Fluddean writings. Again in 1617 he writes, "Quare voluit Creator, ex su& bonitate, ut hi radii dispersi simul colligerentur, qub operationes eorum essent fortiores, ad fortiorem & vivaciorem creaturam producendam. Praeterea spiritus sancti tabernaculum erat Sole, quo efficaciores forent ejus operationes ad perficienda mandata divina (Otreb [Fludd], Tractatus Theologo-Philosophicus, p. 18). The medieval origins of this belief in the significance of wheat seem to arise from the "black grain" which represented the heart. "It is in this corn of the heart that both worlds are joined together, that of the angel and that of Satan. From each point a new circle does develop which turns as circle and stops still as centre" (Pagel, Harvey's Biological Ideas, p. 112).

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This is the aetherial spirit of the Lord, which is in reality nothing else but the aerial saltpeter and the traditional universal spirit of the alchemists. Fludd as a physician had a special interest in the study of man, the microcosm. However, his mystical views of the Creation and the relation of the greater and lesser worlds had convinced him that nothing of value could be learned of man without a true understanding of the macrocosm. If we are diligent in our search we shall be able to correctly interpret the book of nature and perceive the true intentions of the ancient sages in their obscure teachings. In this search we must maintain a definite plan with key questions forever standing foremost in our minds. We must consider the Act of Creation -and especially the part played therein by the divine light of the Lord (Spirit). We must seek out the relation of this light to life and motion for we know that, whenever there is a deficiency, illness or death will follow. In our study of the macrocosm we must concern ourselves with all aspects of this divine spirit, paying special attention to the views on the invisible fire of Zoroaster and Heraclitus as well as the atomic theory of Democritus.18 When we turn to the microcosm we must focus our attention primarily on the assimilation of this spirit in the body. We know that all animated faculties require air, which contains the aetherial spirit, and its divine light hidden within. The Stoics seemed to understand this, Fludd believes, because they defined the soul as the substance of fire converted through air into water.'9 Even all generation is ultimately from air, which has been thickened into the form of sperm, itself an aerial spirit.20 These are all topics of importance to us, he contends, but we must single out for special study the problem of how this celestial balsam or quintessence nourishes our bodies.2' We will find that this occurs in a twofold fashion 18. Fludd, Tractatus Apologeticus, pp. 187-189. 19. Ibid., pp. 188-190. 20. The concept of all life deriving from air is ultimately Aristoteliansee De gen. anim., III, 11:762a. Here animals and plants are described as coming into being in earth and in liquid because there is water in earth and air in water. There is vital heat in all air. This is discussed in relation to Servetus in Pagel, Harvey's Biological Ideas, p. 148f. 21. Fludd, Tractatus Apologeticus, p. 190. Quomodo quodlibet animal, ab hac quinta essentia invisibiliter, duplici ratione nutriatur: Videlicet vel quatenus delitescit in aere. Unde fit ut attractus aer I pulmonibus per inspirationem ad cor, omni attractione, occultam quintae essentiae portionem secum rapiat, quae dispergitur & dispensatur per arterias ad nutriendos spiritus vitales in ijs contentos & congregatos. Aut quatenus

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Harvey and Fludd and that it involves the respiratory process. It seems clear that the spirit in the air is attracted through inspiration to the lungs and is then carried to the heart. The precious occult portion of the quintessence, separated from the rest, is then dispersed and dispensed as the vital spirit through the arterial system. The grosser part, on the other hand, is mixed in the veins with the blood for its completed form, so that from it the more solid parts are nourished and conserved. Surely it is necessary that some of the celestial fire be united with the thickened part of the elements for the conservation of the whole frame. One may even from the orifices of the arteries derive the greater portion of that supplemental chain. "The ocular proof (I say) is that neither animal, nor vegetable, nor mineral could live, last, or exist for one minute of an hour without this lucid fire." 22 The section ends with a ringing defense of the interdependence and need of air and spirit for life. One could hardly argue that this statement is a sophisticated anatomical or physiological study of the vascular system. It is brief and contains little original thought. It is important, however, because of its timing, for it was printed precisely at that time when Harvey was giving his Lumleian lectures. And as Harvey does not yet (1616-1618) discuss the circulation of the blood, so too we find that Fludd (1616-1617) gives no indication of a circulatory system such as he was to elaborate in 1621. On the other hand, here-in his search for recognition by the secret society of the Rosicrucians-he points to a program centered around a study of the universal spirit and its assimilation in the body as a prime aim in a newly reconstructed science. And it is in the next few years that we find Fludd elaborating this theme. In his unpublished Philosophicall Key (c. 1618-1621) he again discussed the blood colligitur & condensatur occult6 in corpore composito ipsius cibi, seu nutrimento ita ut haec lux caelestis congregata, faciat ignem spissum caelestem, visibilem facultates nutritivas & naturales, nutrientem, cujus etiem pars spiritualior per diapedisin ad cordis thalamos penetrat & transportatur atque passim per arterias dispergitur. Crassior vero ejus pars cum subtili elementorum, in venis cum sanguine, ad completam ejusdem constitutione permiscetur, ut ab eo partes solidores nutriantur & conserventur. Atque necesse est ut aliquid hujus ignis coelestis cum elementorum spisso uniatur, ad conservationem ligamenti naturalis totius compositionis; licet, etiam ab arteriarum orificijs, major illius vinculi supplementi portio derivetur. Testis (inquam) ocularis fui quod nec animal, nec vegetabile, nec minerale per unicam horae minutam sine hoc igne lucido vivere, durare, aut existere potuisset. 22. Ibid., p. 190.

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flow. The scheme is far from Harveyan, but at one point he states: Then in his midle spheare shalt thou erect a pavillion called the hart, Which lik the sonne in the greater World, shall send forth the essentiall beames circularly from his centre that therby they may animat and vivify euery member of this so Well erected a Microcosme.23 Whether or not Harvey and Fludd discussed the problem together, they surely were becoming convinced of the importance of the study of the blood at the same time. Circulation their views here was to be the answer for both-although were to differ markedly. Again, we find a certain silarity between the two men regarding their view of the blood. Pagel points out that Harvey modified his adherence to the Aristotelian primacy of the heart in favor of the blood.24 It is true that Fludd was to state that as God placed his tabernacle of the great world in the sun, he had placed his tabernacle in man in the heart.25 Yet, as early as 1617, we find that Fludd's primary concern is not with an anatomical study of the heart, but rather with the blood which has the all-important function of carrying the spirit of the Lord to all parts of the body. Again, in his study of The Generation of Animals, Pagel has pointed out that Harvey held that blood incorporates in itself the two vital spirits of the ancients: Innate Heat and Radical Moisture. It derives from the immediate germ of the embryo and all its parts. This germinative liquid, though "simple" and formless in itself, is capable of assuming all forms-potentially. It is protean, it is Prime Matter. Being generative implies that it is also nutritive-for all generation is a form of nutrition and vice versa. The body "is what it eats"-no physician nor any philosopher ever denied this. The primordial fluid, then, is a nutritious dew, a ros primigenius.26 Although blood and semen are not the same for 23. Robert Fludd, A Philosophicall Key. Or Ocular demonstration, opening and decyphering a great deale of the hidden mysteries of Nature, partly by an experimental conclusion, as also by an intellectual speculation . . . Trinity College (Cambridge) MS 0.2.4.6, fol 24 verso. I am currently in the process of preparing a critical edition of this text. 24. Pagel, Harvey's Biological Ideas, p. 251. 25. Robert Fludd, Anatomiae amphitheatrum effigie triplici more et conditione varia (Frankfort, 1623), p. 266. At this point Fludd proceeds to set forth his mystical concept of the circulation. 26. Pagel, Harvey's Biological Ideas, p. 257.

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Harveyand Fludd Harvey, they may both be considered in terms of a nutritive faculty. Harvey had a vast treasury of similar speculations to drawn on, both from contemporary and ancient authors, as Pagel clearly shows, but it remains interesting that Fludd in 1617 suggested that the aerial spirit in thickened form becomes the sperm, and that this suggestion may be found on the same page in which he discussed the blood flow in terms of the distribution of the spirit in the body. One may conclude that both Harvey and Fludd were deeply concerned with the same problems in the middle years of the second decade of the century. And if the mystical speculations of the one were to lead to an alchemical-symbolical concept of the circulation in the space of a few years, the sober observational and experimental lecture notes of Harvey already foreshadow the future De motu cordis. THE PLACE OF THE DE MOTU CORDIS IN THE FLUDDGASSENDI DEBATE In addition to his Rosicrucian works, Robert Fludd published the first of a series of folio volumes on the "Fluddean" system of the world in 1617. The initial volume, the Utriusque Cosmi Maioris scilicet et Minoris Metaphysica, Physica, atque Technica Historia, appeared at Oppenheim in 1617. The views expressed in this and in succeeding volumes brought him to the attention of continental authors of note. An appendix to the Harmonices Mundi of Kepler (1619) attacked the Utriusque Cosmi in detail. Two years later Fludd replied to this critique in his Veritatis Proscenium. In 1622 Kepler answered in his Prodromus, which included a reprint of his Mysterium Cosmographicum of 1596. The Apologia against Fludd was replied to in turn by the English physician in his Monochordum Mundi of 1622. The details of Fludd's debate with Gassendi are no less complex. Gassendi's friend, Father Marin Mersenne, had been clearly disturbed by the growing popularity of the mystical alchemists in the 1620s. He had specifically singled out Fludd for attack in his Quaestiones Celeberrimae in Genesim (1623), where he had accused Fludd of dealing with magic,27 and two years later he had offered another attack on the alchemists in his La Verite' des sciences. Fludd's reply to Mersenne appeared in the Sophiae cum Moria Certamen (1629), to which 27. Robert Lenoble, Mersenne ou la naissance Vrin, 1943), pp. 134-153.

du mdcanisme

(Paris:

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was appended a defense of the claims of the Rosicrucians, the Summum Bonum-a work by one "Joachim Frizius" which has been attributed to Fludd. Even prior to Fludd's reply, Mersenne had appealed to his friend Pierre Gassendi to aid him in this attack on the alchemists. From a letter we know that before December 2, 1628, Mersenne had sent a group of Fludd's works to Gassendi, and that along with these he had included a copy of the newly published De Motu Cordis of William Harvey.28 Gassendi turned immediately to his task, and his refutation of Fludd's work, the Epistolica exercitatio in qua principia philosophiae Roberti Fluddi, medici, reteguntur, et ad recentes illius libros adversus R.P.F. Mersennum ... respondetur (Paris, 1630) was completed in manuscript form on February 4, 1629.29 In Gassendi's critique of Fludd's views there appears the first significant printed abstract of Harvey's concept of the circulation-here placed in contrast to the mystical symbolical system of Fludd. In sending the works of the two authors together, it had evidently been Mersenne's hope that Gassendi would set forth just such a comparison. Fludd's mystical and chemical schemes of the circulation had been outlined in his Anatomiae Amphitheatrum (1623). Here the circulation of the aetherial spirit in the microcosm (through the arterial system) was compared with the macrocosmic circulation of the sun about the earth.30 One would hardly expect this mystical scheme to have appealed to the experimentalist Gassendi. We find, however, that although he preferred Harvey's work to that of Fludd, he rejected both schemes. Before the Epistolica had appeared in print, Gassendi wrote to his friend Peiresc (August 28, 1629) to give him an account of his views on the circulation.3' From this letter it be28. P. Marin Mersenne, Correspondance du P. Marin Mersenne, ed. Cornelis De Waard and Rene Pintard, 10 vols. (Paris, 1932-1967), II, 181. 29. Pierre Gassendi, Opera omnia, 6 vols. (Florence, 1727). This includes the Epistolica as the Examen philosophiae Roberti Fluddi medici in III, 195-245. For the date of completion see p. 243. 30. Fludd, Anatomiae Amphitheatrum, p. 266. See also my discussion in J. Hist. Med., 16, 376-382. 31. Nicolas-Claude Fabri de Peiresc, Lettres de Peiresc publi6es par Philippe Tamizey de Larroque . . . Tome quatrieme. Lettres de Peiresc d Borrilly, d Bouchard et & Gassendi. Lettres de Gassendi 2 Peiresc 16261637 (Paris, 1893), p. 208. "Le livre dont Mr. Valois vous a parl6, Mr. du Puy en a un exemplaire pour vous envoyer. Je l'avois desja veu avant que partir pour l'Allemagne et en avois dit mon sentiment en ma lettre a P. Mersenne qui enfin se verra peut estre bientost inprimee. Son opinion de la continuelle circulation du sang par les arteres et veines est fort vraysemblable et establie mais ce que je trouve k dire en son fait est qu'il s'imagine que le sang ne sauroit passer du ventricle droit du coeur au

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Harvey and Fludd comes apparent that his forthcoming refutation of Fluddinsofar as the circulation of the blood is concerned-would be directed not at Fludd, but rather at Harvey, with whom Gassendi disagreed because of his own firm belief in the interventricular pores of the septum. Gassendi's promise to Peiresc was fulfilled in the published Epistolica, in which he summarized Fludd's views on the blood -centering his discussion on the question of where the arterial blood originated. Citing the Anatomiae Amphitheatrum, Gassendi stated that Fludd believed that the subtle blood which is in the arteries is not seized from the veins either in that manner in which it is commonly taught, that is, by an inflow in a branch of the vena cava into the right ventricle of the heart out of which the blood, having crossed over into the left side rushes forth then into the arteries,-or in that manner in which the extremes can be fashioned together, that is, the venous capillaries with the extreme arteries, and then out of the veins the blood is instilled into the arteries. For he teaches that this (the arterial) blood is simply from that duct by the inspiration of the aetherial Spirit . . . and he does not admit anything from the venous blood "for this reason," he says, "if the heavy and elementary blood, destined for procreating the solid members, mixes itself with the spirits of life, confusion would follow." 32 But, continues Gassendi, "since the common method of explanation did not please Fludd, he should have listened to Harvey, his countreyman." 33 Harvey's account was based on gauche par le (septum), 1A oi il me souvient que le sieur Payen nous a fait voir autrefois qu'il y a non seulement des pores, mais des canaux tres ouverts . . . qui rendent cette doctrine evident. Vous verrez quelque jour ce que j'en ay dit." 32. Pierre Gassendi, Epistolica exercitatio in qua principia philosophiae Roberti Fluddi, medici, Teteguntur, et ad recentes illius libros adversus R. P. F. Marinum Mersennum . . . respondetur (Paris, 1630), pp. 128-129 ". . . sanguinem illum subtilem, qui est in arteriis, non hauriri ex venis, seu eo modo, quo vulgo docetur; scilicet ramo venae cavae influente in dextrum ventriculum cordis, ex quo sanguis trajectus in laevum, in arterias deinde prorumpat; seu eo modo, quo fingi posset coire extremas, seu capillares venas cum extremis arteriis; sicque ex venis sanguinem in arterias instillari. Docet proinde hunc sanguinem esse dumtaxat ex ducto illo per inspirationem aethereo spiritu; . . . "sanguis grossus elementaris, & ad membra solida procreanda destinatus se permisceret cum spiritu vitae, atque sequeretur in opere humano confusio." 33. Ibid., p. 132. Denique nisi Fluddo communis explicandi ratio placeret, audiendus forte fuerat Harveus ejus conterraneus.

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experiment rather than analogies of the macrocosm and the microcosm. Further, his insistence on the connection of the veins and the arteries through capillaries required a basic identity between the two types of blood. The Galenical system, as understood by Gassendi, required this because of the percolation of the venous blood through the pores of the septum. Gassendi felt then that the real problem was to decide between the Galenical and the Harveyan positions on this question, and he went on to summarize Harvey's work in addressing Mersenne: If you have not seen it [Harvey's book], the argument is that the blood out of the trunk of the vena cava goes into the right ventricle of the heart through a branch, and then is throwvn together through the venous artery into the left ventricle, out of which, after having been sent into the arteries, at length it is returned into the veins through insensible openings of the arteries and junctions with the veins; and in the same way again out of the veins through the lungs and the heart into the arteries in a perpetual circuit.34

However, although Gassendi esteemed the work of the "learned anatomist," he had witnessed dissections at Aix at which an industrious surgeon by the name of Payanus had demonstrated the existence of the interventricular pores. "Therefore there definitely are these passages . . . and since they are really there, they ought not to be useless, and there is a purpose in readiness, the percolation of the blood out of the right vessel into the left; and I might argue that the arterial blood is derived in this manner."35 Thus, in a detailed criticism of Fludd's philosophy, Fludd's own views on the blood flow are summarily brushed aside and Gassendi gives rather a refutation of Harvey's thesis which more nearly coincides with his own concept of the relation of the venous and arterial blood. 34. Ibid., pp. 132ff. "Nisi videris, argumentus est, quod sanguis ex venae cavae trunco per ramum transversum in dextrum usque ventriculum cordis, & inde in pulmones per arteriosam venam respersus colligatur, atque coniiciatur per venosam arteriam in sinistrum ventriculum, ex quo in arterias immisus, per insensibilia tandem arteriarum, veniarumque juncta oscula regrediatur in venas; similque rursum modo ex venis per pulmones, & cor in arterias, circuitione perpetua." 35. Ibid., pp. 133-136. "Hi igitur meatus sunt . . . ut cum illi revera sint, nec frustra esse debeant, & aliunde causa sit in promptu, ipsa sanguinis e dextro sinu in laevum percolatio; sanguinem illum arterialem hac derivari arguerem."

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Harveyand Fludd Yet, although this was a pointed attack on Harvey, it was not Harvey who answered. Prior to the appearance of Gassendi's Epistolica, Fludd had completed his Pulsus (on October 19, 1629), in which he affirmed that his view of the microcosmic circulation of the blood "exacte" seems to confirm the feeling and opinion of the learned William Harvey, a most skilful doctor of medicine, most clear in the art of anatomy, and yet highly versed in the mysterious profundities of Philosophy, a man who is my esteemed compatriot and the most faithful of the college: about which he instructed the world advisedly and prudently in his little book Exercitatio De cordis sanguinis in animalibus motu, and he declared remarkably well with reasons produced from the treasure of Philosophy as well as with demonstrations for the eye.36 It is likely that the Pulsus was not published until the following year,37 but from this quotation it is evident that, for Fludd, an attack on Harvey could be taken as an attack on his own views. By 1631 Fludd had written his reply to Gassendi's criticism, a detailed point-by-point discussion which appeared as the Clavis philosophiae et alchymiae Fluddanae (Frankfort, 1633). Here, although he maintained his belief that the arterial blood is not formed from the venous blood,38 he specifically turned to Gassendi's attack on Harvey. From the standpoint of modern science Fludd never appeared to better advantage. Surely, he 36. Fludd, Pulsus, p. 11. "Hoc exact6 illam viri grauissimi Gulielmi Haruei, Medicinae Doctoris pertissimi, arte anatomica, quam clarissimi, necnon in profundi philosophiae mysteriis versatissimi, compatriota mihi charissimi, & collegae fidelissimi, sententiam atq; opinionem confirmare videtur; qua, idq; consulte & prudenter satis mundum in libello quodam suo, cui titulus est: Exercitatio anatomica, de cordis sanguinisq; in animalibus motu; instruit, insigniterq; cum rationibus a Philosophiae arca depromptis, tum multiformi demonstratione oculari declarat motum ipsius sanguinis esse circularem." The date of the completion of the Pulsus is given on p. 93. 37. On the dating of the Pulsus see above, n. 6. We know that Fludd had read Gassendi's Epistolica by 1631. In Doctor Fludds answer unto M. Foster or, the squeesing of Parson Fosters sponge, ordained by him for the wiping away of the weapon-salve (London, 1631) Fludd compares Gassendi's scholarly refutation of his works with Foster's "ill mannered" and "impudent" reply, and he mentions in several places his own reply to Gassendi (the Clavis philosophiae et alchymiae Fluddanae). 38. Robert Fludd, Clavis philosophiae et alchymiae Fluddanae, sive Roberti Fluddi Armigeri, ut medicinae doctoris, ad epistolicam Petri Gassendi theologi exercitatem responsum (Frankfort, 1633), p. 33.

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writes, Gassendi must be wrong about the interventricular pores. Either Payanus had forced the opening in the septum with his probe, or perhaps in a few abnormal individuals such openings occur naturally; . . . but from this single case it should not be considered to be universal. For thus in one man such a single vein may be come upon, while in another there may be no spleen and with many others there may be strange and unexpected deviations from the ordinary human composition . . . Certainly one or even a second example does not prove the does one swallow prove the advent of argument-neither summer. In contrast to Gassendi's one example, Fludd had personally observed Harvey search repeatedly for these pores, but, he asserted, "not in any one out of the many cadavers that he examined did he find such a septum; and neither I nor any others who with most acute and almost lynx-like eyes saw this when we examined the septum of the heart." Thus, the mystical alchemist informs the observationalist Gassendi that "we know and speak as experts" when we "assert with confidence that the septum of the heart is not ordained by nature to that purpose called for by Gassendi." 39 39. Ibid., pp. 33ff. "Inprimis sciendum quod autor hic noster, ad suam caussam, ad relationem experimenti cuiusdam statuminandum Aquis-Sextiis peracti recurrat, vbi (ait) se Anatomica inquisitione, mediante spathula seu ferro quodam obtuso, porosos in septo cordis meatus inuenisse, quos mille in septo ianuas, licet occultas vocat; vnde concludit cum Medicis & chirurgis, qui dissectioni tali adfuerunt, esse transmissionem sanguinis A dextro sinu in laevum licet insensibilem. Idque aduersusme . . . Memini ego Chirurgum quendam iactitasse, se probo aut spathula, irregulari quadam dilatione, admirabilem illum plexum retiformem ab implicatione venae & arteriae factum, qui descendit a vasis seminariis praeparantibus ad testiculos, penetrasse; quod quidem propter contexturae subtilitatem impossibile est, sine meatuum occultorum violatione; nisi raro in aliquo particulari hoc contingat . . . Praeterea ista inquisitio a pluribus collegarum meorum, & praecipue a D. Harueo Anatomico expertissimo, saepius instituta est magna cum diligentia, quatenus ipse ad suam in sanguinis circulationis causam, cum fatigatione huius rei experimentum fecit; sed ne in vnico quidem, ex pluribus cadaueribus inuenit ipse tale quidpiam; nec ego, nec alii, qui oculis acutiissimis & quasi lynceis cordis septum sumus scrutati. Quare concludendum est, quod, licet in vno cadauere vel alio (si ita esset sine spathula violatione) res talis AquisSextiis comparuerit, hoc tamen particulare non inferat vniversale. Sic enim in vno homine, vnus inuenitur Ren tantum; in alio nullus Lien, atque per multa alia ab ordinaria compositione humana aliena atque incosueta. At anne ob eam caussam concludendum, nimirum ob aliquos partium defectus vel augmenta in vno honine, necessario omnes debere ita se

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Harvey and Fludd Although he had spoken of an exact confirmation of his views, it is somewhat questionable whether Fludd could have accepted the Harveyan thesis of the circulation in its entirety. While he could speak in terms of the circulation of the spirit of the blood, he certainly was opposed to a mixing of the venous and the arterial systems, which he normally speaks of as distinct and separate. This is a question I have discussed earlier,40 and it is a technical point that need not concern us here. What is important for the present problem is that (a) Harvey's work was taken up for discussion by the Mersenne circle in 1628-and in a Fluddean context; (b) Fludd himself felt that Harvey's work confirmed his own mystical views of the circulation; and (c) when Gassendi attacked him, Fludd answered not only the arguments directly aimed at him, but also those on the constitution of the septum which had been directed primarily at his friend and colleague, Harvey. This was surely the first significant controversy over the circulation of the blood and there can be no doubt that in the mind of Mersenne the work of the two English physicians could be considered together-and surely Fludd would have heartily concurred in this. It is with Gassendi that we see a recognition of the basic difference between the works of the two men. Here, however, the issue is clouded by Gassendi's own misjudgment and Fludd's remarkable reply to him. FLUDD AND HARVEYIN THE PURITAN ERA-THE WEBSTER-WARDDEBATE (1654) Although Robert Fludd could complain with some reason that his work was esteemed more on the continent than in his own country,41 he was not completely ignored by English authors. There was a strikingly increased interest in "occult" thought of all kinds in England in the middle years of the habere? Vnum cert6 vel alterum exemplum, non probat argumentum, nec hirundo vnica probat aestatis aduentum. Quare experimento a Gassendo prolato, partim haud fidem habebimus, & partim si pro concesso illi detur, dicimus quod vnum particulare non demonstret generale. At in generali agnoscimus, & experti loquimur, istam Gassendi assertionem esse falsissimam: ac proinde asseuerare haud erubescimus, quod septum cordis, non ad illud officium A Gassendo nominatum a Natura ordinetur. Nam si hoc esset pro statuto, sequeretur, quod nulla esset inter sanguinem Arterialem & Venalem differentia. 40. Debus, J. Hist. Med., 16, 388-391. 41. And he did so repeatedly; as an example, see his Answer unto M. Foster, p. 24.

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century. Of special note was the new concern with Paracelsism and Helmontian thought interpreted as the proper key to nature.42 In an inflammatory work we find Noah Biggscalled a "Psittacum Helmontii" by an opponent-demanding chemistry as the proper model science for students at the universities (1651), while John Hall, addressing An Humble Motion to the PARLIAMENT of ENGLAND Concerning the ADVANCEMENT of Learning: and Reformation of the Universities (1649), asked: "Where have we any thing to do with Chimistry, which hath snatcht the keyes of Nature from the other Sects of Philosophy, by her multiplied experiences?" 43 During the Puritan era, more Paracelsian and mystical chemical works were translated than in the entire century before 1650. At the same time, and with good reason, there was an increased interest in the Rosicrucian movement. John Heydon gave his books such titles as A New Method of Rosie Crucian Physicke (1658) and The Rosie Crucian Infallible Axiomata (1660), while Eugenius Philalethes (Thomas Vaughan) prepared a lengthy introduction to a translation of The Fame and Confession of the Fraternity of R:C: (1652). The connection was quite clear for George Hakewill, who praised the "Chimiques, Hermetiques, or Paracelsians (& a branch of them as I conceive is the order Roseae Crucis)."44 And John Wilkins, referring to the perpetually lighted lamp "in the sepulchre of Francis Rosicrosse, as is more largely expressed in the confession of that fraternity," 4.5 could confidently cite Robert Fludd as an authority on clocks, automata, and perpetual motion.46 42. The social aspects of this problem have been investigated by P. M. Rattansi in his "Paracelsus and the Puritan Revolution," Ambix, 11 (1963), 24-32, and "The Helmontian-Galenist Controversy in Restoration England," Ambix, 12 (1964), 1- 23. 43. Noah Biggs, Chymiatrophilos, Mataeotechnia Medicinae Praxews. The Vanity of the Craft of Physick . . . With an humble Motion for the Reformation of the Universities, And the whole Landscap of Physick, and discovering the Terra Incognita of Chymistrie (London, 1651), sig. b.1 recto. Biggs was attacked by William Johnson in his preface to Leonard Phioravant's Three Exact Pieces (London, 1652), p. 1; J(ohn) H(all), An Humble Motion To The PARLIAMENT of ENGLAND Concerning The ADVANCEMENT of Learning: and Reformation of the Universities (London: John Walker, 1649), p. 27. 44. George Hakewill, An Apologie or Declaration of the Power and Providence of God in the Government of the WoTId, 3rd ed. (Oxford: William Turner, 1635), p. 276. 45. John Wilkins, Mathematicall Magick. Or the Wonders That may be performed by Mechanical Geometry (London: M. F. for Sa. Gellibrand, 1648), pp. 236-237. Other editions appeared in 1648, 1680, and 1691. I am indebted to Dr. Frances A. Yates for this reference. 46. Ibid., pp. 163, 264.

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Harveyand Fludd Indeed, this was the time when the only one of the major works of Robert Fludd, the Philosophia Moysaica (1638), appeared in English translation (1659). The new interest in the chemical philosophy, which was combined with a plea for a reformed educational system, came into direct conflict with the mechanical philosophy. Here, too, there was an opportunity for comparisons of Harvey and Fludd. This may be clearly seen in the debate between John Webster and Seth Ward in 1654. John Webster (1610-1682) is a typical English Helmontian of the mid-seventeenth century. He turned to the works of Bacon and the Paracelsians in the search for a replacement for the outdated and "heathenish" learning taught at the universities.47 Fludd especially seemed of interest to him because his whole philosophy was allegedly based on Christian principles. Attracted early to the study of nature and religion, Webster studied chemistry under John Hunyades (c. 1632) and was ordained a minister shortly after that date. With his Puritan sympathies he served as a surgeon and chaplain with the Parliamentary Army during the Civil War. By 1648 his reaction against the Established Church had forced him to become a nonconformist. Although most of his writings are on religious topics, he was later to prepare an important survey of current views on metals, his Metallographia (1671). John Webster's strong views on the reform of the universities had led him into controversy in the early part of the 1650s, a fact which prompted him to write his Academiarum Examen in 1654. This work, "offered to the judgements of all those that love the proficiencie of Arts and Sciences and the advancement of Learning,"' is in reality a call for reform in the same terms advocated by Robert Fludd in his Tractatus Apologeticus of 1617. Reacting against the sterile-and to his mind atheistic-writings of Aristotle taught at the universities, Webster spoke instead of the "mysterious and divinely-inspired Teutonick [Jacob Boehmel, and . . . the highly illuminated fraternity of the Rosie Crosse." 48 As the Paracelsians teach, 47. See the article on John Webster by Bertha Porter in the Dictionary of National Biography. The work of Hunyades has been discussed by F. Sherwood Taylor and C. H. Josten in "Johannes Banfi Hunyades 15761650," Ambix, 5 (1953), 44-52, where it is suggested that Hunyades arrived in London sometime between 1623 and 1633, and "Johannes Banfi Hunyades. A Supplementary Note," Ambix, 5 (1956), 115. of 48. John Webster, Academiarum Examen, or the Examination Academies. Wherein is discussed and examined the Matter, Method and

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true Christian knowledge of Nature will be taught best by ocular demonstrations learned by putting "hands to the coals and furnace." In this way, Webster asserted, we shall learn the importance of the three principles, while we must also seek out the secrets of natural magic and "Cabalistick Science." In general if we are to properly reform our knowledge on Christian principles we must surely seek to build up tables true of axioms as Bacon has suggested, but we must-as Christians-seek a knowledge of nature. that is grounded upon sensible, rational, experimental, and Scripture principles: and such a compleat piece in the most particulars of all human learning (though many vainly and falsely imagine there is no such perfect piece to be found) is the elaborate writings of that profoundly learned man Dr. Fludd, than which for all the particulars before mentioned (notwithstanding the ignorance and envy of all opposers) the world never had a more rare, experimental and perfect piece. In addition, the new philosophers are told to avoid Aristotle and to turn to the works of Ficino, Plato, Gilbert and Hermes Trismegistus-as interpreted by the Paracelsians. As might be expected, experimental chemistry is to be the new key to nature, and the medicine of Paracelsus and van Helmont is to replace that of Galen.49 It is in his discussion of medicine that Webster turns to recent advances in anatomy, which indicate that this part seems to be growing, and arising towards a Zenith of perfection; especially since our never-sufficiently honoured Countreyman Doctor Harvey discovered that wonderful secret of the bloods circulary motion: yet for all this there comes small advantage by it in practise, and application, for the more certain, safe, and easie curing of diseases: for though it bring great satisfaction to speculative Customes of Academick and Scholastick Learning, and the insufficiency thereof discovered and laid open: As also some Expedients promoting of all kind of Science. Offered to the judgements of all those that love the of Arts and Sciences and the advancement of Learning proftciencie (London: Giles Calvert, 1654), p. 26. A more extensive discussion of the Webster-Ward exchange will be found in my Science and Education in the Seventeenth Century. The Webster-Ward Debate (London: Macdonald and Co., in press). 49. Ibid., pp. 71, 75, 104ff, 105.

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Harvey and Fludd understanding, and help to cleer many intricate doubts, yet doth it little to remove dolor, danger, or death.50 Rather, Webster continues, in the long run we shall find that this vulgar anatomy-and here he would clearly include the work of Harvey-is defective as to that vive and Mystical Anatomy that discovers the true Schematism or signature of that invisible Archeus or spiritus mechanicus, that is the true opifex, and dispositor of all the salutary, and morbifick lineaments, both in the seminal guttula, the tender Embrio, and the formed Creature, of which Paracelsus, Helmont, and our 50. Ibid., p. 74. This is all reminiscent of Viscount Conway's advice to his daughter-in-law, Anne Conway, to avoid William Harvey as her personal physician. A learned man and "a most exelent Anatomist" he may be, he declared, "but to have a Physitian abound in phantasie is a very perilous thing, occations in diseases are very often suddaine, therefore one ought to have a Physitian that should be governed only by his judgment, as one puts the best man to the Helme in a storme." (Keynes, Life of William Harvey, p. 393). Dr. Lester S. King has drawn to my attention several other similar statements from the same period. In the Usefulness of Experimental Philosophy (published 1663, but Part I written about 1650) Robert Boyle stated that the new anatomical advances do not really help the progress of medicine and, he added, if science was to serve medicine then recourse must be made to chemistry (The Works of the Honourable Robert Boyle, 6 vols. [London: J. and F. Rivington, L. Davis, W. Johnston, 1772], II, 145, 163). In his Medela Medicina. A Plea for the free profession and a renovation of the art of physick; out of the noblest . . . writers . . . (London, 1665) Marchamont Nedham argued that no science had made such marked advances as medicine in the previous twenty years (p. 215). To this John Twysden replied that the new discoveries had not aided medicine at all (Medicina Veterum vindicata or an answer to a book entituled Medela Medicinae . . . [London, 1666], pp. 197-199). A few years later Henry Stubbe wrote that ". . . it was thought at first that this Circulation of blood would overthrow all the usual Methods of Physick, and introduce new and beneficial discoveries in that part of Medicine which is Therapeutick. But Harvey denieth that it varieth the Medicine of the Ancients; and Slegelius asserts the same opinion, avowing it to be rather an happy illustration, then a subversion of the former praxis, though it alter the Theory much. In fine, those little advantages and Diorismes, which we derive from that Invention merit not our notice; nay, any man shall with more assurance bleed in many diseases in sundry manners and different places, upon diverse indications upon the old observations and rules, then on the new hypotheses, wherein as to the use of parts, and nature of humors, there is as little of clearness and certainty; as there is efficacy in that practise, which is regulated most thereby. Henry Stubbe, A Specimen of Some Animadversions upon a Book, Entituled Plus Ultra, or Modern Improvements of Useful Knowledge . . . (London, 1670), p. 114.

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learned Countreyman Dr. Fludd, have written most excellently.51 In short, Harvey's work is splendid and a glory to the nation, but Robert Fludd's works represent a true foundation for a new Christian interpretation of the universe. His mystical anatomy of the blood gives us a far more profound understanding of the subject than the more superficial anatomical work of Harvey. John Webster's book was scathingly attacked by Seth Ward (1617-1689).52 Educated at Cambridge, Ward had early showed an aptitude in mathematics and astronomy. Although he had been chosen mathematical lecturer at Cambridge in 1643, Ward was deprived of his Fellowship in 1644 because of his staunch adherence to the Established Church. After a period of wandering, he moved to Oxford (1647) where he was appointed Savilian Professor of Astronomy. This was a period when Oxford was rapidly becoming a center for the new science. In addition to Ward, Robert Boyle, Thomas Willis, Jonathan Goddard, John Wallis, and John Wilkins were at Oxford in the early fifties. As a group, these men formed the nucleus of the "Philosophical Society of Oxford," and this, in turn, was a forerunner of the Royal Society of London. It is little wonder with these scholars about him that Seth Ward took exception to John Webster's attack on the universities. As a result, he wrote the Vindiciae Academiarum (1654), a tract which contained an introduction by his friend, John Wilkins. Perhaps forgetting that at Oxford the lectures in astronomy had fallen into neglect before his appointment in 1649, Ward emphasized the high level of scientific work at the universities -and how improper and inconsistent Webster's suggestions really were. The prelude to the attack may be seen in John Wilkins' introduction. Speaking of Webster, he states: The man doth give me the freest prospect of his depth and braine, in that canting Discourse about the language of nature, wherein he doth assent unto the highly illuminated fraternity of the Rosycrucians. In his large enconiums upon Jacob Behem, in that reverence which he professes to iudiciall 51. Webster, Academiarum Examen, p. 74. 52. On Ward's life see the Dictionary of National Biography. Influenced by Kepler (through Bullialdus), Ward is perhaps best known for having taught that the planets move in elliptical paths.

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Harvey and Fludd Astrologie, which may sufficiently convince what a kind of credulous fanatick Reformer he is like to prove.53 Ward-in a point-by-point rebuttal of Webster's work-devotes much space to the state of mathematics at the universities, which he admits could be bettered, but hardly in the fashion proposed by his adversary. He questions the wisdom of completely discarding the Galenic medicine, and he argues that Webster is unfair in his accusation that chemistry is unknown at the universities-surely the group at Oxford included men who were active in this field.54 And if Webster pointed to the writings of Francis Bacon as basic for a reform of nature, Seth Ward could hardly agree more-but, how does Webster follow up this suggestion? He says, indeed, that "the second Remedy is, That some Physicall Learning may be brought into the Schooles, that is grounded upon sensible, Rationall, Experimentall, and Scripture Principles, and such an Author is Dr. Fludd; then which for all the particulars, the World never had a more perfect piece." Surely this is too much! "How little trust there is in villainous manl" Although a moment before he had recommended Francis Bacon "for the way of strict and accurate induction," now he is fallen into the mysticall way of the Cabala, and numbers formall: there are not two waies in the whole World more opposite, than those of L. Verulam and D. Fludd, the one founded upon experiment, the other upon mystical Ideal reasons; even now he was for him, now he is for this, and all this in the twinkling of an eye, 0 the celerity of the change and motion of the Wind. And if he turns in Philosophy to Plato, Democritus, Epicurus, Philolaus, and Gilbert, why should there be any need in this, for "if De Fluctibus be so perfect, what need we go any

farther?"55 Ward's reaction to Webster's statements on anatomy are predictable. Here he clearly defended Harvey and damned Fludd and the Rosicrucians. Yes, he agrees, 53. Seth Ward, Vindiciae Academiarum Some briefe containing, Animadversions upon Mr. Websters Book, Stiled The Examination of Academies. Together with an Appendix concerning what M. Hobbs, and M. Dell have published on this Argument (Oxford: Leonard Lichfield for Thomas Robinson, 1654), p. 5. From the preface signed N. S. (John WiLkins). 54. Ibid., p. 47. 55. Ibid., p. 46.

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The practice of Physick hath been bottomed upon experience and observation. And that this is the reason, that the discoveries of the Circulation of the blood, of the venae lacteae, both Mesentericall and Thoracicall, of the vas breve, and severall new ductus, vasa lymphatica &c. have not made an alteration in the practice of Physick, answerable to the advantage they have given to the Theory; and the security and confirmation they have brought to the former waies of practise. But Webster prefers the mystical anatomy of Paracelsus, van Helmont and Fludd: As for his Postulatum of discovering the signatures of the Invisible Archeus by Anatomy, it is one of his Rosycrucian Rodomantados; would he have us by dissection surprize the anima mundi, & shew him the impressions of a thing invisible? Yet the Schematismes of nature in matters of sensible bulke, have been observed amongst us, and collections made of them in our inquiries, and when the microscope shall be brought to the highest (whether it is apace arriving) we shall be able either to give the seminall figures of things, which regulates them in their production and growth, or evince them to lye in quantities insensible, and so to be in truth invisible. No matter what the Rosicrucians may tell John Webster, the state of medicine and surgery is highly advanced, and the study of the subject is greatly affecting its practice. Although there are many specific cases of cures Ward might mention, he must surely point out the Royal College of Physicians in London which is the glory of this Nation, and indeed of Europe, for their Learning and felicity, in the cures of desperate Ulcers and diseases, even of the Cancer, and those he (ignorantly) mentions, which have been diverse times performed, by D. Harvey and others.56 In this tirade Ward conveniently ignored the fact that Fludd had been an honored member of the "glorious" college for nearly thirty years. The Webster-Ward debate may in fact be seen as part of the larger conflict of the Paracelsian-observationalists on the one hand and the mechanical philosophers on the other. The spe56. Ibid., pp. 35-36.

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Harvey and Fludd cific occasion made it possible to weigh the value of the work of Harvey and Fludd once again. If Webster had found the Fluddean approach more valid, Seth Ward was willing to relegate the folios of the mystical alchemist to permanent oblivion. CONCLUSION It is tempting to separate Harvey and Fludd-the experimental biologist and the mystical alchemist-and Charles Singer did just that in 1960 no less decisively than had Seth Ward three centuries earlier. Yet Pagel has suggested that the influence of Fludd may have sensitized Harvey to the notion of circularity. This thesis is not weakened, but given added weight, when we examine additional evidence in relation to the intellectual currents of that period rather than our own. The earliest writings of Fludd in defense of the Rosicrucians appeared precisely at the time when his friend began the Lumleian lectures. Fludd's early tracts give a distinct indication of his concern with the blood flow and the assimilation of the universal spirit in the body. His answer to this problem, a scheme of the mystical circulation, appeared in 1623-the same year that Harvey was reaching his own conclusions. We note a further similarity between the emphasis they both place on the role of the blood over that of the heart-and also in their views on the origin of the semen. And if today we separate the mystical-alchemical symbolism of the one author from the observational-experimental approach of the other, we find that this was not mandatory in the seventeenth century. This is clear in the two scientific debates referred to in this paper. Fludd thought that Harvey's work confirmed his own, while Mersenne and Gassendi felt that Harvey and Fludd could be rightly discussed together. John Webster went so far as to assert that Fludd's contribution was more significant than that of his colleague. There is surely no question about Harvey's place in the history of the biological sciences, but his roots were deep enough in Ancient and Renaissance thought for him to be claimed as a fellow traveler by Paracelsists and Hermetic philosophers in his own day.57 57. Another case where a professed alchemist praises Harvey and Fludd in the same breath is where Elias Ashmole speaks of the glories of the Royal College of Physicians. Theatrum Chemicum Britannicum . . . collected and annotated by Elias Ashmole, A Reprint of the London Edition 1652 with a new Introduction by Allen G. Debus (New York: Johnson Reprint Corporation, 1967), p. 460.

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Towards a Synthesis: Population Concepts in Russian EvolutionaryThought, 1925-1935 MARK B. ADAMS Department of the History of Science Harvard University

Between 1925 and 1935, the Russian School of population geneticists, consisting mainly of students and colleagues of Sergei S. Chetverikov (1880-1959)1, made many important contributions to the development of the modern evolutionary viewpoint. One such contribution which highlights some of the salient similarities and differences between the Russian investigators and their Western counterparts is the Russian attempts to formulate a genetic explanation for the effects of isolation and population size on evolution. I use the word "attempts" because there were in fact three: one by Sergei Chetverikov in 19262 and two more in 1931 by two of Chetverikov's students, D. D. Romashov 1. Only within the last ten years has Chetverikov's work come to be generally recognized as a landmark in the history of population genetics and evolutionary thinking. See, for example, Theodosius Dobzhansky, "Evolution of Genes and Genes in Evolution," Cold Springs Harbor Symposia on Quantitative Biology, XXIV (1959), pp. 15-30; I. M. Lerner, "Introductory Note" to S. S. Chetverikov, "On Certain Aspects of the Evolutionary Process From the Standpoint of Modern Genetics," translated by Malina Barker, Proceedings of the American Philosophical Society, vol. 105, no. 2, April, 1961, pp. 167-195; L. C. Dunn, A Short History of Genetics, McGraw-Hill, 1965; Theodosius Dobzhansky, "Sergei Sergeevich Tshetverikov, 1880-1959," Genetics, vol. 55, no. 1, January, 1967, pp. 1-3; Mark Adams, "The Founding of Population Genetics: Contributions of the Chetverikov School, 1924-1934," Journal of the History of Biology, vol. 1, no. 1, Spring 1968; pp. 23-39. For an example of Chetverikov's recognition in the Soviet Union, see B. L. Astaurov, "Two Landmarks in the Development of Genetical Concepts" (in Russian), Biulleten' Moskovshogo Obshchestva Ispytatelei Prirody, LXX (1965), 25-32; see also Genetika, 1965, no. 1, pp. 204-205. For additional information on Chetverikov, see his "Autobiographie" in Nova Acta Leopoldina, n.s., no. 143, pp. 308-310 (1959). 2. Sergei S. Chetverikov, "0 nekotorykh momentakh evoliutsionnogo protsessa s tochki zreniia sovremennoi genetiki," Zhurnal Eksperimental' Journal of the History of Biology, Vol. 3, No. 1 (Spring 1970), pp. 107-129.

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and N. P. Dubinin.3 The theories of Romashov and Dubinin had much in common with that of Sewall Wright on genetic drift, published in 1930 and 1931.4 So far as I can tell, however, the theories of Wright, Romashov, and Dubinin were of independent derivation, although in 1932 Dubinin and Romashov integrated their views in a joint theoretical paper.5 In order to put the work of the Russian School in perspective, we should understand its relation to the development of the so-called "synthetic theory of evolution." When Julian Huxley used the word "synthesis" in Evolution: The Modern Synthesis (1942), and when G. G. Simpson used the word "synthetic" in The Meaning of Evolution (1949), they were highlighting the fact that the new view of evolution had emerged from the results of many biological disciplines and was providing for them a common theoretical framework." We might characterize the emergence of this synthesis in the following way. Three separate modes of evolutionary explanation led into the 1920's that were distinct in their approaches: the genetic, the naturalist, and the biometric, or statistical.7 The work of R. A. Fisher and J. B. S. Haldane, noi Biologii, 2 (1926), pp. 3-54. The Russian original is reprinted in Biulleten' Moskovskogo Obshchestva Ispytatelei PriTody (BuUetin of the Moscow Society of Naturalists) Biological Section, LXX (1965), no. 4, pp. 33-74. For an English translation, see that done by Malina Barker, edited by I. M. Lerner, "On Certain Aspects." In general quotations of Chetverikov in the text are taken from the Barker translation, occasionally amended by the author. 3. D. D. Romashov, "Ob usloviiakh 'ravnovesiia' v populiatsii," Zhurnal Eksperimental'noi Biologii, 7, (1931), No. 4, pp. 442-454; N. P. Dubinin, protsessy i ikh znachenie dlia mekhanizma "Genetiko-avtomaticheskie organicheskoi evoliutsii," ibid., nos. 5-6, pp. 52-95. 4. Sewall Wright, "Evolution in Mendelian Populations," Genetics, 16: 97-159 (1931); Sewall Wright, "The Roles of Mutation, Inbreeding, Crossbreeding and Selection in Evolution," Proceedings of the Sixth International Congress of Genetics, 1932. 5. N. P. Dubinin and D. D. Romashov, "Geneticheskoe stroenie vida i ego evoliutsiia: 1. Genetiko-avtomaticheskie protsessy i problema ekogenotipov," Biologicheskii Zhurnal, 1 (1932), nos. 5-6, pp. 52-95. 6. Julian Huxley, Evolution, The Modern Synthesis, George Allen & Unwin Ltd., London, 1942 (see especially chapter 1, "The Theory of Natural Selection"); G. G. Simpson, The Meaning of Evolution (New Haven: Yale University Press, 1949), p. 277. 7. More frequently a distinction is made between the "naturalists" and the "experimentalists" in the first few decades of the twentieth century. According to this distinction, the biometricians are considered as "naturalists." It is true that the battlelines in the debate on the validity of Darwin's theory of evolution by natural selection tended to divide biologists into these

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Russian Population Concepts and to a certain extent that of Sewall Wright, achieved a kind of synthesis between two of these three modes: genetics and biometry.8 The primary thrust of their work during the decade 1925-1935 was the demonstration that selection, viewed as a statistical process, was compatible with particulate genetics and was in fact reinforced by it. In formulating this demonstration, Wright, Fisher, and Haldane all expressed themselves primarily in mathematical terms, using a mode of theoretical expression developed by two generations of biometricians. The formulation of a more total view of the evolutionary process which incorporated the results of genetics, biometrics, and natural history was only the product of the subsequent decade (1935-1945). Its creation was signaled notably by such works as those of Theodosius Dobzhansky (1937), Julian Huxley (1942), Ernst Mayr (1942), G. G. Simpson (1944), and G. L. Stebbins (1950), which treated problems of natural variation, speciation, two groups. [For example, see Th. Dobzhansky, Genetics and the Origin of Species (Ist ed.), (Columbia University Press, 1937); N. W. TimofeefRessovsky, "Mutations and Geographic Variation" in The New Systematics, ed. Julian Huxley (Oxford, 1940); and Julian Huxley, Evolution: The Modern Synthesis (New York: Harpers, 1942). Probably the most complete discussion of the nature and causes of the naturalist-experimentalist split appears in the relevant chapter of Dr. Garland Allen's unpublished thesis on T. H. Morgan, Harvard University, 1967.1 Such a distinction, in my view, can obscure the very real difference in philosophy, methodology, and theoretical approach which separated the more orthodox natural historians from the statistical biometricians. The biomedical camp, in which I would include such figures as Karl Pearson and W. F. R. Weldon, tended to treat natural selection as a statistical process acting on continuous variation, and their investigations fall as much in the discipline of statistics as of biology. Although this theoretical approach gained great currency at the turn of the century, there was a concurrent tradition of evolutionary explanation which had developed among Continental Darwinians, exemplified by Moritz Wagner's theory of speciation by "isolation." This type of theoretical biology tended to rely much more heavily on studies of the processes occurring in natural populations. Most turn-of-the-century taxonomists, systematists, "ecologists," and zoologists did not rely on statistical models and can be seen as members of the naturalist tradition. 8. This is especially true of R. A. Fisher; see for example, The Genetical Theory of Natural Selection (London: Oxford, 1930). J. B. S. Haldane's papers in the mid-1920's are heavily mathematical, but by 1932 he was already attempting to relate his mathematical work in evolutionary theory to natural populations-with some credit to Russian evolutionists: see The Causes of Evolution (London, 1932). For Wright's main theoretical papers, see Wright, "Evolution" and "The Roles." Of the three, Wright is closest to the Russians in the attention he gives to evolution in natural populations.

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paleontology, ecology, biogeography, and allied disciplines in genetic terms systematically for the first time.9 It is interesting to note that in his important 1926 paper, the Russian biologist Sergei Chetverikov also used the word "synthetic" (sinteticheskii) in roughly the same sense as Simpson was to use it some two decades later. As we shall see, Chetverikov was looking forward to the coming development of a general theoretical framework which would explain all the results of systematic biology, and he looked upon his own work as a conscious first attempt at such a total synthesis. Thus, in contrast with comparable Western work of the same period, some Russian evolutionists during the decade 1925-1935 strove for a general synthesis whose scope would encompass all evolutionary phenomena. In all fairness, it must be admitted that their work lacked the mathematical sophistication of their Western counterparts. Nonetheless, the Russians did consciously seek to bridge the genetic, naturalist, and biometric traditions, and to relate their conclusions to the broadest possible body of literature. A glance at Chetverikov's 1926 bibliography tells the story. Of 67 entries, roughly a third deal with biometry (20), another third genetics (23), and the remainder natural history (24). Significantly, 14 of these entries -more than one fifth of the total-are concerned with the theory of isolation.10 Chetverikov felt it necessary to deal at length with the evolutionary effects of isolation because of a dual conviction which underlay the thinking of Russian population geneticists during this period. First, they believed that Mendelian genetics was of general applicability, that it explained the mechanism of inheritance and hence would have to be at the core of evolutionary explanation. Second, they believed that the generalizations of natural history were valid and hence had a genetic explanation. In particular, this meant that they believed that natural adaptation occurred and that its cause was natural selection. But it also meant that they believed that species were real, that divergent speciation occurred, and that its cause was isolation. 9. Th. Dobzhansky, Genetics and the Origin of Species (New York. Columbia University Press, 1937); J. Huxley, Evolution; E. Mayr, Systematics and the Origin of Species (Columbia University Press, 1942); G. G. Simpson, Tempo and Mode in Evolution (Columbia University Press, 1944) and G. L. Stebbins, Variation and Evolution in Plants (Columbia University Press, 1950). 10. See the bibliography of S. S. Chetverikov (1926).

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Russian Population Concepts These considerations become clear when we examine the discussion of the genetic effects of isolation in Chetverikov's 1926 paper. The paper is constructed around a distinction between "two processes . . . strictly distinct in their causes as well as in the consequences resulting from them . . . one is the process of differentiation, of splitting-up, leading in the end to speciation-its basis is isolation; the other leads to adaptation and its cause is . . . natural selection." " Chetverikov was by no means the only one to suggest that adaptation and speciation have different causes;12 but he was one of the first to attempt a genetic explanation for both. Providing a genetic explanation for speciation was important to Chetverikov because he accepted the reality of the species. This view had lost favor both among some orthodox Darwinians (following Darwin's views on the arbitrariness of species distinctions) and among Mendelians, who had seen interspecific differences as only quantitatively, not qualitatively, different from intraspecific genetic variation. In contrast to these views, in which species were generally defined in terms of their morphological differences, such outstanding naturalists as Karl Jordan and E. B. Poulton defined species in terms of their reproductive isolation.'3 Chetverikov clearly sided with this latter group: Potentially, all the individuals of a single species are able to cross freely, without encountering any hindrance either in the process of fertilization itself or in the viability or fertility of the offspring.14 He maintained that such a species definition "corresponds most closely to our genetic and systematic ideas." 15 11. Chetverikov, "On Certain Aspects," Barker translation, pp. 188-189. 12. See, for example, the almost simultaneous work of Albert Eide Parr, Adaptiogenese und Phylogenese, 1926. 13. For an exposition of Karl Jordan's species concept, see Ernst Mayr, "Karl Jordan's contributions to current concepts in systematics and evolution," Trans. Roy. Entomol. Soc. London 107 (1955), 45-66. For Poulton's species concept, see E. B. Poulton, "What is a species?" Proc. Entomol. Soc. (London) (1903), lxxvi-cxvi. Chetverikov refers to an article by Karl Jordan in the bibliography to his 1926 paper: "Der Gegensatz zwischen geographische und nichtgeographische Variation," Z. wiss. Zool. 83 (1905). Elsewhere in his 1926 paper, Chetverikov makes clear his debt to German authors for his species ideas: "The natural state of a species pre-supposes precisely a state of a "freely crossing community"-"Paarungsgemeinschaft" of the German authors." 14. Chetverikov, "On Certain Features.. .", 174. 15. Chetverikov discusses his species concept, pp. 174-175 of the Barker translation. Just how far Chetverikov's species concept is from the

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Furthermore, Chetverikov felt compelled to explain divergent speciation without any recourse to selection. He knew the literature stemming from J. T. Gulick's nineteenth-century work on Hawaiian land snails and found it to be a convincing demonstration of speciation in the absence of selection.'6 Chetverikov believed that the process of mutation is continually occurring in natural populations, and that due to the Hardy-Weinberg law, new mutants would be maintained at a constant frequency in freely-crossing populations.'7 Chetverikov morphological species concept is made clear in the following excerpt from these pages: "Undoubtedly, an accumulation of a greater or lesser number of morphological or physiological differences always precedes the appearance of reproductive isolation, but the very fact of existence of these differences, in whatever form they should appear, is not a sufficient basis for the formation of a new species. And from our genetics experiments on the most diverse animal and plant organisms, we now know that it is possible to create two groups of such organisms, which will differ from each other by a perfectly concrete complex (theoretically speaking, as large as one wishes) of morphological characteristics, which are not connected by intermediate forms; that is, having a so-called morphological hiatus, but at the same time belonging genetically to the same species." 16. Rev. J. T. Gulick, "Diversity of Evolution Under One Set of External Conditions," J. Linnean Soc. London (Zool.), 11 (1872); "Divergent Evolution Through Cumulative Segregation," ibid. 20 (1887), 189-274. In addition to the above two works of Gulick, Chetverikov's bibliography lists works by A. Garret, Mayor, and H. Crampton. Gulick's 1872 paper had documented the existence of some 175 species of Achatinella land snails represented by some 800 varieties in the forest region that covers one mountain range of Oahu, each mountain valley and ridge having its own indigenous species. Gulick felt that this speciation could not have been the result of selection, "lst. Because in different valleys, on the same side of the mountain, where food, climate, and enemies are the same, there is still a difference in the species. 2nd. Because we find no greater difference in the species when we pass from the more rainy to the drier side, than when we compare the forms from valleys on the same side of the mountain, separated by an equal distance" (p. 498). Chetverikov's acceptance of Gulick's results and conclusions forced him to give an explanation of how isolation could produce speciation and divergence in the absence of selection. 17. G. H. Hardy, "Mendelian Proportions in a Mixed Population," Science, 28 (1908), 49-50. Wilhelm Weinberg, Ver. vateri. Naturk. Wurttemberg, 64: 369-382. Chetverikov, like many evolutionists of the period, was unaware of the work of Weinberg and only made reference in this connection to Hardy. Hardy's contribution took the form of a letter to the editor of Science countering the view of Udny Yule that "if brachydactyly is dominant 'in the course of time one would expect in the absence of counteracting factors, to get three brachydactylous persons to one normal,'" Hardy demonstrated that, supposing "that the numbers are fairly large, so that the mating may be regarded as random, that the sexes are evenly distributed among the three varieties, and that all are equally fertile," each Mendelian character would be maintained at a constant frequency in the

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Russian Population Concepts argued mathematically that continuous natural mutation, preserved by free-crossing, would produce divergence even without selection. Let us consider his argument. Chetverikov assumed that a new mutation has entered a population as a homozygote and that free-crossing causes both alleles to enter immediately into heterozygous combinations.18 What is the likelihood, under random mating, that the mutant will become expressed homozygously? If the population size is 'N+ 1', Chetverikov reasoned, the probability of the two mutants coming together in any given generation is equal to '1/N'. For a large population, this probability will be small. But if mutation occurs continually, there will be a second pair of alleles, and a third pair, and so forth. The probability that some one of these mutants (but not any particular one) will be expressed homozygously then equals the sum of the separate probabilities minus the chance of their joint occurrence. Chetverikov derived a general equation for the probability of at least one of 'm' mutants being expressed homozygously: 19Pm= 1- [(N-1)/N]m. In Chetverikov's view, this general equation expresses a fundamental relationship between the size of a population and the manifestation of its genotypic variability. Here we are dealing with two opposing tendences: on the one hand, the more numerous the population, the greater are the chances for the origin of new mutations within it. Thus the frequency of origin of variability is directly proportional to the size of the freely-crossing population. On the other hand, the less numerous the population population. Thus, he concluded that "there is not the slightest foundation for the idea that a dominant character should show a tendency to spread over a whole population, or that a recessive should tend to die out." Chetverikov applied this rule to recessive mutations occurring in natural populations, which led him to conclude that a new mutation "will not be destroyed, will not be dissolved in the mass of normal individuals. It will exist in the heterozygous state remaining hidden from the eye . . . generation after generation." 18. Chetverikov writes: "Let us begin the analysis with a case of the not infrequent appearance in nature of a recessive homozygous mutation (aa)" (Barker trans., p. 176). Here he does not discuss the much more likely possibility that mutations will occur singly in the heterozygous condition. He could have suggested, but did not, that the entry of a new allele into a population might occur by the migration of a member of another population, which could well introduce a new allele (aa) in homozygous condition. 19. The Barker translation, due no doubt to a misprint, gives this equation incorrectly as P =1 - (N-l)m/N. Both Dobzhansky ("Evolution of Genes") and the 1965 Russian reprint give the equation in the correct form.

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(that is, the smaller the value of N) the greater is the probability of manifestation in it in homozygous form of mutations absorbed by it earlier. If we imagine that the total number of individuals of a given species, N, is subdivided into a series of isolated colonies, then the frequency of origin of new mutations within the limits of the entire species will not suffer, but the probability of reappearance of each such mutation will be once more considerably increased, depending on the reduced size (n) of the colony, within which it originally arose . . . isolation entirely automatically leads to a differentiation within a species, to the fact that the colonies of one species, isolated from each other, begin, with time, to manifest differences in individual characters . . . And so isolation, under the conditions of a process of continuous accumulation of mutations becomes, by itself, a cause of intraspecific (and consequently eventually also of interspecific) differentiation.20 Indeed, on the basis of his calculation, Chetverikov establishes a 'law": "all other conditions being equal, the degree of differentiation within a species is directly proportional to the degree of isolation of its separate parts." 21 What are we to think of Chetverikov's argument? His assumption that recessive mutants enter as a pair and remain two seems arbitrary and inappropriate, since it was known some time before 1926 that mutation occurs at a given locus at a statistical rate. Further, his notion that recessives will not be exposed to any selection until homozygously expressed is curiously incompatible with his own clear arguments demonstrating the role of the genetic background, to be found in the same essay a few pages later. Finally, his implicit assumption that the occasional homozygotic expression of mutants in individuals Will cause divergence between populations is clearly unfounded, for Chetverikov does not provide a mechanism, in the absence of selection, whereby such mutations will spread throughout a small population to become one of its characteristics. There is some internal evidence that Chetverikov was not perfectly satisfied with his explanation.22 It almost seems as though he knew what had to be explained, and knew the 20. Chetverikov, "On Certain Aspects," pp. 178-179. 21. Ibid., p. 180. 22. For example, he writes: "Naturally, it should not be thought that the factors named exhaust the full essence of the phenomenon of differentiation in the realization of which a series of still other processes not considered here also participates." (Ibid., p. 188.)

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Russian Population Concepts form the explanation would have to take, but was somehow unable to bring it off. Why then would he treat the problem of the genetic effects of isolation? Chetverikov made clear in his introduction that "the basic aspects of genetics . . . must penetrate the depths of the various departments of our science," and that what he himself was attempting to create was "an orderly elaborated system of evolutionary knowledge based on contemporary genetic foundation."23 Hence, Chetverikov cannot avoid treating the problem of isolation and its effect on speciation genetically. He realizes that his own explanation is not complete and forecasts in his conclusion a key prerequisite for a "synthetic theory": . . . it is still too early to speak of a synthetic formulation of the evolutionary process. Only after we have disentangled the basic principles . . . underlying the evolution of organisms . . . as well as the phenomena of speciation, only then will we finally be able to attempt a reconstruction of the definitive structure of evolution.24 Later he reiterates: "But only after we have unravelled the basic principles, those of speciation as well as of the whole evolutionary process, shall we be able to take stock."25 Thus Chetverikov set the tone for Russian thinking on evolution, and gave priority to the problem which Dubinin and Romashov were to elucidate more satisfactorily some five years later. Two groups of workers at the Institute of Experimental Biology were engaged in detailed analyses of the genetic structure of wild populations in the five years following the publication of Chetverikov's 1926 essay.26 One group, headed by Aleksandr Serebrovskii, had begun even before 1926 a series of studies of the genetics of populations of domesticated fowl in the Caucasus. The goal of these studies had been to determine the genetic differences between various semi-isolated populations of fowl and thereby to trace the historical and 23. Ibid., p. 169. 24. Ibid., p. 193. 25. Ibid. 26. For a description of the work of the Institute in general and of Chetverikov and Serebrovskii in particular, see N. K. Kol'tsov, "On the Works of the Institute of Experimental Biology in Moscow" (Russian), Uspekhi Eksperimental'noi Biologii, 8 (1929), no. 1, 15-25.

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geographical origins of these populations.27 N. P. Dubinin had participated in these studies in 1929 and 1930, and their specific impact on his work will be noted later. A second group, working under Chetverikov, had been investigating the genetic structure of wild populations of Drosophila. D. D. Romashov participated in these investigations, and it will be worth our while to trace in more detail the development and results of this work in order to understand its relevance to Romashov's later theoretical views. Three separate lines of thought had led Chetverikov to conclude that natural populations should contain a large amount of cryptic variability. First, he believed that mutation occurred continually in nature and that most of the resultant mutants would be recessive. Second, he believed that "free-crossing," following the equilibrium formulated by G. H. Hardy,28 would maintain these mutants in a natural population. Finally, on the basis of the accompanying table by H. T. J. Norton (see Table 1.),29 he believed that selection would eliminate dominant mutations much more quickly than recessives and hence would favor the build-up of recessive (vs. dominant) mutations in the wild. Since these recessive mutations would only be phenotypically expressed in the homozygous condition, their heterozygous presence would be masked by phenotypic uniformity. In order to test this reasoning, Chetverikov and his students at the genetical laboratory of the Institute of Experimental Biology undertook in the summer of 1925 an "analysis of the constitution of the wild species of the genus Drosophila." 30 This study was in fact the first genetic analysis of a natural popula27. A. S. Serebrovskii, "A genetical analysis of a population of domesticated fowl of Dagestan," Zhurnal Eksperimental'noi Biologii, 3 (1927), 62-146; A. S. Serebrovskii, N. P. Dubinin, and R. 1. Serebrovskii, "The genogeography of domesticated fowl in Kabardy and Balkariia," cited as "in press" in Dubinin's 1931 work, p. 476. Serebrovskii's important and influential role in the development of population genetics in Russia is worthy of a more complete study, on which the author is now working. 28. See n. 17 above. 29. Norton's table first appeared in R. C. Punnett, Mimicry in Butterflies, Cambridge, 1915. Dunn, in his Short History, suggests that Chetverikov was the first to see the implications for evolution of Norton's table. The table reproduced here is taken from the Barker translation, p. 182. The table as it originally appeared in Chetverikov's work in Russian is given on p. 55 of the Russian reprint. ." lists the students who participated 30. Astaurov, "Two Landmarks.. in "this first work in Moscow": B. L. Astaurov, E. I. Balkashina, N. K. Beliaev, S. M. Gershenson, I. F. Rokitskii and D. D. Romashov (p. 26). The quotation is from Chetverikov, Barker translation, p. 178.

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Table 1. Percentage of total population formed by old variety

99.9 98.0 90.7 69.0 44.4 25 11.1 2.8 .03 .008 .000 a

Percentage of total population formed by the hybrids

.09 1.96 9.0 27.7 44.4 50 44.4 27.7 9.0 1.96 .09

Percentage of total population formed by the new

NORTON'S TABLEa

Number of generations taken to pass from one in the percentage of different individu -. . .i. A. A. Where the new varety iS dominant 100 100 100 100 10

variety

50-

75

90

99

.000 .008 .03 2.8 11.1 25 44.4 69.0 90.7 98.0 99.9

4 2 2 2 2 4 10 36 170 3840

10 5 4 4 4 8 17 68 333 7653

28 15 14 12 12 18 40 166 827 19,111

300 165 153 121 119 171 393 1,632 8,243 191,002

From Lerner translation of Chetverikov, 1926, Proc. Amer. Phil. Soc. 105 (1961), 183.

50

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MARK B. ADAMS

tion, and it marks the inception of experimental population genetics. Chetverikov's analysis of the progeny of 239 female Drosophila melanogaster revealed no fewer than 32 different hereditary characters which had been masked heterozygously. Chetverikov's study had immediate impact in the circle of his colleagues and students, and its success led many of them to perform further studies of the genetics of Drosophila populations. Even before the first results were published, N. V. Timofeev-Resovskii, one of Chetverikov's former students31 had completed and published a very similar analysis of 78 wild female Drosophila melanogaster from Berlin.32 Members of the Chetverikov group extended their analysis to other Drosophila species from around Moscow (D. phalerata, D. transversa, D. vebrissina, and D. obscura) and studied a population of Drosophila melanogaster from Gelendzhik, near the Caucasian coast of the Black Sea. D. D. Romashov had never been one of the group of seven who had performed the early work in Moscow, and he made a study in 1929-1931 of three populations of Drosophila funebris (from Moscow and Kiev) which, along with the other studies, was to have a significant influence on his thinking.33 One might suppose that these experimental results would have been regarded as a brilliant proof of the validity of Chetverikov's reasoning, for they confirmed the presence of an enormous amount of cryptic variability in natural populations which he had predicted. But these experiments also produced other, unexpected results which could not be explained by the effects of mutation and selection alone.34 In both the Berlin and the Gelendzhik populations of Drosophila melanogaster, "very many mutations were represented . . .not by one but by a significantly large number of heterozygous individuals."35 Furthermore, the majority of such muta31. Both N. V. and H. A. Timofeev-Resovskii (the joint authors of the 1927 study) had been students of Chetverikov in Moscow. They left Russia in 1925, establishing themselves at Buch, just north of Berlin, but they maintained close contact with Russian workers while in Germany. Since the war, Timofeev-Resovskii has been working in Russia and has published there since 1956. 32. Timofeef-Ressovsky, H. A. and Timofeef-Ressovsky, N. W., "Genetische Analyse einer freileben Drosophila melanogaster Population," Roux Archiv Ent. 109, 70-109. 33. The most complete reporting of these results appears to be in Dubinin and Romashov (1932), "Geneticheskoe stroenie." 34. Romashov, "Ob usloviiakh" (1931), pp. 448-450. 35. Ibid., p. 448. Translations from the works of Dubinin, Romashov, and Russian authors other than Chetverikov were done by the author unless otherwise noted.

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Russian Population Concepts tions were "manifestly 'harmful' deviations from the norm ... or were identical to mutations in these Drosophila species studied earlier which decrease viability, lower fertility, etc." 36 Lethal recessives had not been looked for in these initial studies, since it was assumed that they would not be present in significant concentrations due to the strong selection against them. Indeed, the technique of brother x sister mating, used by both Chetverikov and Timofeev-Resovskii, would not even reveal the presence of autosomal recessive lethals.37 But Timofeev-Resovskii's study did unexpectedly uncover sex-linked lethals in the progeny of 3 of the 78 females analyzed.38 A second significant outcome of these investigations was the discovery of apparently nonadaptive differentiation of different populations. Romashov summarizes this finding as follows: . . . both studied populations were sharply delineated from one another by their genetic structures: the majority of mutations characteristic for the Berlin population were not found in Gelendzhik, and conversely, characteristic Gelendzhik mutations were absent from the Berlin population . . . Completely similar results were obtained in the analysis of three mutually isolated colonies of Drosophila funebris from Moscow, Petrovskii-Razumovskii (outside of Moscow), and Kiev.39 How are these two results to be explained? Clearly, selection cannot be responsible for high frequencies of harmful or lethal alleles; and, according to Romashov's reasoning, mutation could be responsible for these frequencies only if mutation rates were to differ from place to place-a possibility Romashov rejects.40 36. Ibid., p. 449. The omitted portion lists examples of such mutations: in Dros. melanogaster, cluboid-2, pedes taxi; in Dros. funebris, balloon, alae curvatae. Romashov reasoned that the presence in natural populations of such mutants in high frequencies could not be due to natural selection, which would tend to eliminate such genes rather than spread them and increase their frequencies. 37. This is because flies homozygous for this gene would die very early in development and would simply not appear in the progeny to be counted. 38. Sex-linked lethals would be detectable by the technique used because they would cause differential survival of males and females in the progeny, whereas on the basis of Mendel's laws equal numbers of both sexes should appear. 39. Romashov, "Ob usloviiakh," p. 448. 40. According to Romashov (ibid., p. 449) such an assumption "contradicts numerous observations of the mutational process in Drosophila melanogaster, which in the most diverse places where this species has been studied has proceeded with great constancy in relation to the relative frequencies of the origins of various mutations."

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The inadequacy of selection and mutation in explaining the build-up of harmful mutants and some genetic differences between populations suggested a very significant conclusion: some factor other than selection and mutation must be having a major influence on the frequencies of mutant genes. The point of departure for Dubinin and Romashov's modified view of the evolutionary process was an observation made by Aleksandr Serebrovskii, who along with Chetverikov had been in charge of Drosophila genetics at the Institute of Experimental Biology since 1922. In his 1930 report on the "genofund" of domesticated fowl in Daghestan,4' Serebrovskii noted that the Hardy-Weinberg equilibrium would not necessarily maintain new mutations at a constant frequency as Chetverikov had supposed. He reasoned as follows. Newly arising mutations would presumably arise heterozygously. Since only half of a diploid genotype can be carried by a haploid gamete, there would be a 50 percent chance that the new mutation would not have the same frequency after reproductionsince there would be a 1/4 chance that the mutant would be doubled, and a 1/4 chance that it be lost altogether.42 Dubinin and Romashov realized the great evolutionary implications of Serebrovskii's observation. If Chetverikov's expectation about the maintenance of a new mutation at a constant frequency was unfounded, what effect would diploid reproduction have, in the long run, on a new mutation's frequency? In order to answer this question, Dubinin and Romashov ran a series of experiments simulating what seemed to them the germane features of diploid breeding.43 In an urn, they placed 100 markers, numbered 1-100, representing a population of 50 diploid individuals. From the urn they drew 25 pairs of markers, replacing the first of each and 41. A. S. Serebrovskii, "Problems and methods of genogeography," (Russian), Trudy Vsesoiuznogo S'ezda po Genetike, II (1930). 42. If one imagines a mutation occurring at a statistical rate in a population of infinite size, then of course such random effects would cancel out. The importance of Serebrovskii's observation lies in his recognition of the ways in which theoretical, mathematical results must be modified to deal with events occurring in actual populations. 43. Dubinin and Romashov, "Geneticheskoe stroenie" (1932) pp. 57-61. Clearly Dubinin and Romashov were aware in 1931 of the importance of population size, but in their works of that year there is no reference to any such experiments. Whether runniing the series of urn drawings was suggested by Wright's work of 1931, which is listed as a reference in the 1932 work, is unclear. In any case, it is indicative of their experimental bias that they chose to run an "urn model" experiment rather than to derive general mathematical statements.

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Russian Population Concepts adding another with the same number, and discarding the second. Such a drawing represented one generation, resulting in a loss of 1/4 of the alleles and the duplication of another 1/4. They continued the "drawings" until the urn contained markers with only one number, whereupon they noted how many drawings (generations) had been required to make the urn population "homozygous." Dubinin and Romashov considered three findings resulting from their urn experiments of special relevance to evolution in natural populations. First, although the number of required generations varied, all the populations went to complete homozygosity. This result led them to conclude that "in the end every limited population, without the participation of selection and the mutational process, will go to complete homozygosity of one or another chance allele." 44 Second, since different markers became homozygous in different populations, automatic genetic processes lead to the differentiation of the genetic contents of populations through the loss of alleles and through the reproduction of several genes to significant concentrations.45 Finally, the experiments had shown that the speed with which a population went to homozygosity was inversely correlated with population size. The evolutionary implications of diploid sexual reproduction provided Dubinin and Romashov with one mechanism capable of altering gene frequencies without selection or mutation. They found a second in the phenomenon of "population waves," or, to use Chetverikov's term, "waves of life" (volny zhizni). 46 Chetverikov had published an article while still a student at Moscow University (1905) which had called attention to the evolutionary significance of radical variations in the size of insect populations over time. It can be said without any exaggeration that the fauna is not permanent for a minute. With each day, practically with each moment, its equilibrium is disturbed, some species undergoing "high tide," others "ebb," and at the same time it is perfectly right to emphasize that there is no species which doesn't from time to time undergo these ebbs and 44. Ibid., p. 60. 45. Ibid., pp. 60-61. 46. S. S. Chetverikov, "Volny Zhizni" (waves of life), Dnevnik zootdeleniia, III, no. 6 (1905). Chetverikov studied at Moscow University from 1900 through 1906.

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flows. Hence anyone who has more or less carefully studied some fauna of a single location knows that no two years are the same: that which last year was rare or even absent is met this year in abundance, and conversely that which last year struck the eye with each step now demands careful search.47 Dubinin and Romashov cited extensively from more recent literature documenting and charting data on variations in insect numbers, irregular "depressions and rebirths" as well as periodic seasonal, annual and long-term fluctuations.48 What effect would such variations have on the genetic structure of populations? One effect Dubinin and Romashov noted may well have been suggested by Dubinin's experience with Serebrovskii in the summers of 1929 and 1930 in a study of chickens in a region of the Caucasus. There Dubinin came upon a case which directed his attention to the influence of radical changes in population size on the genetic structure of populations. In the 1930 expedition in Eastern Bashkiriia we managed to discover a whole region . . . where there were but a few fowl, owing to the crop failure of 1929. There was enough interference because of this factor that the structure of the future population of chickens, arising from a small group of remaining individuals, appears distinctly recon47. Ibid., cited in Dubinin and Romashov, "Geneticheskoe stroenie" (1932), P. 70. N. V. Timofeev-Resovskii credits Chetverikov with being the first to point out the evolutionary significance of "population waves," which he sees as one of the major factors (along with selection) which cuts down genetic variability in populations. See N. V. Timofeef-Ressovsky, "Mutations and geographical variation," in Julian Huxley, ed., New Systematics (London: Oxford University Press, 1940), pp. 73-136, esp. p. 114. 48. In discussing periodical fluctuations in population numbers, Dubinin and Romashov cite works by M. J. Bertoni (1919), K. Chapman (1931), I. I. De-Gryse (1929), C. S. Elton (1924), H. Gasow (1925), C. G. Hewitt (1921), N. A. Kholodkovskii, W. B. Johnson and L. Lloyd, T. W. Kirkpatrick (1923), N. M. Kulagin (1930), K. Lampert (1900), M. A. Menzbir (1930), I. K. Parker (1930), A. N. Promptov (1933), H. L. Seamens (1923), W. L. Thompson (1928), E. T. Seton (1912), and B. P. Uvarov (1931). The number of Russians in this group, taken together with the fact that virtually all of these studies are on insect populations, suggests the importance of studying the development of Russian entomology around the turn of the century. How large a group of Russian entomologists were there? What species concept did they employ? What were their evolutionary views? It should be recalled that Chetverikov himself was an entomologist, as were most of the group around him in the middle 1920's, including D. D. Romashov.

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Russian Population Concepts structed, and the given population is losing all the basic elements of similarity with other populations historically connected with it.49 Dubinin believed that these effects could also be explained by a consideration of the implications of diploid breeding. It should be recalled that the assumption was made that each parental pair left exactly two offspring in calculating the chance of 1/4 of the loss of a heterozygous mutant. But if a parental pair left twice the offspring, the chances of the loss of an allele heterozygous in one parent is quartered. In short, the more offspring left by a given parental pair, the greater the likelihood that any mutations they contain will be maintained and spread. Likewise, if an organism contributes no progeny, the probability that the mutations it contains will be maintained is nil. Hence the decimation of a population would lead to the elimination of a great number of alleles, and its regrowth would greatly increase the frequency of those genes possessed by the survivors-whether or not they had contributed to survival. The recognition of the importance of the nature of diploid breeding and population waves made possible a theoretical framework within which the earlier experimental results made sense. The high concentrations of neutral and harmful mutants, and the nonadaptive differentiation of different populations of the same species, could now be viewed as predictable consequences of random factors operating in small natural populations. In addition to removing the consternation caused by experimental results, such a theoretical framework made possible a new view of the evolutionary process and of the role played by isolation. Chetverikov had understood adaptation to be the result of selection acting on new mutations, eliminating relatively unfit mutations and spreading fit ones to eventual homozygosity. But because of the implications of Norton's table, he realized that selection "proceeds extremely slowly" in increasing the frequency of rare recessives. Hence he regarded the frequency of each such allele as "distinctive' (svoeobraznyi) and the relative concentrations of alleles as the result of "a continuous history of many years regulated by selection." 50 Serebrovskii had concurred that genetic structure of populations reflected a "thousand-year 49. Dubinin, "Genetiko-avtomaticheskie 50. Chetverikov, "On certain aspects."

protsessy" (1931), p. 476.

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history," and had seen genetic analyses of natural populations as capable of supplementing the "results of archaeology." 51 By contrast, the recognition of the evolutionary importance of automatic genetic processes acting in small populations gave Dubinin and Romashov reason for believing that evolution could occur much more rapidly. For if diploid reproduction and population waves resulted in the chance elimination of many newly arising mutations, they also were capable of greatly increasing the frequencies of certain chance mutations to a level where selection could act more quickly and effectively.52 Thus, according to Norton's table, even a selection intensity of 50 per cent requires some 1,920 generations to raise the frequency of a recessive allele from 0.09 per cent to only 1.96 per cent, whereas in a sufficiently small population the automatic genetic processes could achieve the same result in only a few generations. Most mutations made frequent by chance would be less fit than the common allele, in which case negative selection would result in their gradual elimination. But for the few mutations which were useful, selection could then act to spread them to near homozygosity in a considerably shorter time than would have been possible under the action of selection alone.53 Dubinin and Romashov were thus able to come to terms with the problem Chetverikov had posed earlier: the genetic effects of isolation. Since the effectiveness of automatic genetic processes varied with population size, they reasoned, the isolation of a population from the rest of the species, either spatially or temporally, should have three results. First, some alleles present in the species would be lost to the isolated population. Second, at the moment of isolation, the frequency of a given allele present may greatly increase; and since under random mating, the chance of an allele entering a homozygous combination increases with the allele's frequency, the phenotypic appearance of recessive alleles would hence also increase. (This was the one effect of isolation noted by Chetverikov.) Finally, and most significantly, random fixation would proceed 51. A. Serebrovskii, N. Dubinin, and R. Serebrovskii, "The genoprotsessy" geography"; quoted in Dubinin, "Genetiko-avtomaticheskie (1931), p. 476. 52. The efficiency of selection corresponds to the rate at which the allele selected for displaces the other. In the case of two alleles, 'A' dominant and Ca'recessive, Aq = sq2(1-q)/(1-sq2), where 's' is the selection coefficient of 'aa' and 'q' is the frequency of 'a'. It can be shown by simple calculation that selection proceeds most efficiently when q=2/3 (assuming that the heterozygote 'Aa' is selectively indistinguishable from 'AA'). 53. Dubinin, "Genetiko-avtomaticheskie protsessy" (1931), p. 472.

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Russian Population Concepts more rapidly after isolation due to the decreased size of the population. To Dubinin and Romashov, then, isolated populations are the primary arenas of evolutionary change. Such populations provide the most favorable setting for mutant alleles to reach significant frequencies, permitting selection to preserve and spread useful mutations more effectively. Hence, isolated populations would be expected to become partially differentiated from one another in their possession of both adaptive and non-adaptive traits. Dubinin and Romashov recognized several causes of natural "isolation": population size, geographic separation, or the formation of colonies in new habitats by a few individuals from the larger populations.54 The fact that such isolation takes place in nature led Dubinin and Romashov to formulate the notion of the ecogenotype: Real species are made up of a large number of populations, more or less independent of one another. The genetic contents of these populations, different for each, we propose to call ecogenotypes, emphasizing by this the interdependence and role of a whole complex of biological conditions of species life for the fate of genes and of the genetic structure of populations.55 In 1931, Dubinin and 14 collaborators undertook a large-scale study of the ecogenotypes of Drosophila melanogaster from nine localities in the Caucasus and one in Central Russia. This study was completed in 1932, and its published version in 1934 demonstrates that the theory of automatic genetic processes played a key role in the design of the experiment.56 54. Dubinin and Romashov, "Geneticheskoe stroenie" (1932), pp. 80-81. They are thus apparently the first to discuss what Ernst Mayr later termed the "founder principle." See Mayr, Animal Species and Evolution, Harvard University Press, 1963, p. 211; and Mayr, Systematics and the Origin of the Species, pp. 32, 237. It is also interesting to note the similarity between the ideas of Dubinin and Romashov and those expressed by Ernst Mayr in "Change of Genetic Environment and Evolution," in Evolution as a Process, ed. by Julian Huxley, A. C. Hardy, and E. B. Ford, 1954 (Collier ed. 1963). Mayr writes on p. 206: "Recessives will have a much greater chance to become homozygous in the reduced population and thus become more exposed to selection." And on pp. 209-210: "The genetic reorganization of peripherally isolated populations, on the other hand, does permit evolutionary changes that are many times more rapid than the changes within populations that are part of a continuous system. Here then is a mechanism which would permit the rapid emergence of macroevolutionary novelties without any conflict with the observed facts of genetics." 55. Dubinin and Romashov, "Geneticheskoe stroenie" (1932), p. 89. 56. See, for example, the English summary given in N. P. Dubinin and

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Dubinin and collaborators found some 61 identifiable mutants. As expected, the concentrations and nature of the uncovered mutants varied with the geographical source and within one source from year to year. Dubinin wished in particular to evaluate the frequency of autosomal lethals, which he expected would appear rather frequently due to random fixation and population waves. When new techniques of genetic analysis were used,57 the frequency of lethal mutations in the 10 natural populations was found to range between 0 and 21.4 per cent. In particular, it was found that 10-20 per cent of the total number of second chromosomes analyzed carried recessive lethals. This outcome was viewed by Dubinin as a striking proof of the correctness of his theoretical reasoning; to others it was completely unexpected. Theodosius Dobzhansky called it "a novel result-and a very startling one." 58However, follow-up experiments done by a number of investigators corroborated Dubinin's findings.59 Dubinin and co-workers had completed another major analysis by 1936.6f0 By 1939, Iu. M. Olenov and collaborators had shown that concentrations of lethals grow from August to October, demonstrating the predicted dependence of such concentrations on population cycles.61 Other studies by Dubinin resulted in a major analysis of lethals in natural populations.62 14 co-authors "Eksperimental'noi analiz ekogenotipov D. melanogaster," parts 1 and 2, Biologicheskii Zhurnal, 3 (1934), nos. 5-6, pp. 52-95. One example of the use of the theory of "genetico-automatic" processes in the design of the experiment is reflected in the thoroughness of the genetic analysis, which was designed to detect autosomal lethals. Another is the number and distribution of populations studied. 57. For a discussion and explanation of these techniques by Dubinin himself, see his book, Problems of Radiation Genetics (Oliver & Boyd, 1964), a translation by G. H. Beale of N. P. Dubinin, Problemy radiatsionnoi genetiki (Moscow, 1961). In the translation, pp. 158-161. 58. Th. Dobzhansky, "Concepts and Problems of Population Genetics," Cold Springs Harbor Symposia in Quantitative Biology, Vol. XX (1955), p. 4. 59. For example, C. Gordon (1955), "The frequency of heterozygosis in free-living populations of Drosophila melanogaster and Drosophila subobscura," J. Genet., 33 (1936), pp. 25-60. Sturtevant also did a follow-up study, as he mentions in his A History of Genetics (New York, 1965), p. 110. 60. N. P. Dubinin, M. A. Heptner, Z. A. Demidova, and L. I. Djachkova, "The genetical structure of the population and its dynamics in wild Drosophila melangaster," Biologicheskii Zhurnal, 5 (1936), pp. 939-976. 61. Iu. M. Olenov, I. S. Kharmak, K. F. Galkovskaia, and G. D. Muretov, "Factors responsible for the genetic composition of wild Drosophila melanogaster populations," Doklady Akademii Nauk SSSR, 24 (1939), pp. 466-470. 62. N. P. Dubinin, "On lethal mutations in natural populations," Genetics, 31 (1946), pp. 21-38.

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Russian Population Concepts Chetverikov's failure to effect the synthetic theory he foresaw is reflected in his own inconsistency in dealing with the evolutionary importance of population size. In his 1926 treatment of the way in which selection acts to produce adaptation, Chetverikov made the usual assumption of infinite population size. However, as we have seen above, in his treatment of the way in which isolation contributes to speciation, Chetverikov concluded that population size is a crucial parameter responsible, even in the absence of selection, for genetic divergence. Even so, Chetverikov saw chance as having only a limited role in a population of any size: chance mutations, and their expression via random mating, serve only to allow the allele to be expressed phenotypically and hence to be exposed to the action of selection. Dubinin and Romashov went much further in the direction Chetverikov had indicated. To label as "simultaneous discovery" the concurrent realization of the importance of chance factors and population size by Wright in America and Dubinin and Romashov in Russia might be to overestimate the similarity of the two viewpoints and to underestimate the influence on their views of earlier workers, notably Hagedoorn. Clearly, both recognized the importance of sampling errors for the evolutionary process in natural populations. But Wright treated selection and drift as quite independent phenomena, and felt that while small population size favors drift, it has no relation to selection, whereas Dubinin and Romashov argued that small population size increases the efficiency of selection in spreading adaptive mutations. They thus anticipated a position taken by Ernst Mayr in 1954,63 and in this respect their work has a distinctly modern flavor. There is no doubt that the Russian realization is independent from Wright's, for it is the product of a separate line of reasoning and a distinct biological tradition. The hallmark of the Russian approach was its emphasis on the natural historian's viewpoint-the continuous concern for natural populations in the wild and the continuous harkening back to them for demonstration, example, and theoretical inspiration. This concern may well have been a source of theoretical 63. Emst Mayr, "Change of Genetic Environment and Evolution," in Evolution As a PTocess, ed. Julian Huxley, et al., Collier, 1963, pp. 188-213. Mayr's 1963 work, Animal Species and Evolution, includes a brief passage on the history of "genetic drift," p. 204. Both Wright (1930) and Dubinin and Romashov (1932) mention the 1921 work of A. L. and A. G. Hagedoorn, The Relative Value of the Factors Causing Evolution, The Hague, 1921.

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shortcomings. In contrasting the development of the Russian thinking with that of such Western workers such as Sewall Wright, one is struck by the enormous difference in mathematical sophistication. By means of an abstract statistical analysis, Wright could deal with population size as a mathematical parameter; in contrast, Dubinin and Romashov resorted to a probability "experiment" with markers in an urn, whose results are not generalized but only suggestive. There are indications that Dubinin and Romashov were aware of their mathematical shortcomings, and by 1934 they had enlisted the aid of the brilliant probability theorist, A. N. Kolmogorov, who had been one of Dubinin's fellow students at Moscow University in the 1920's. But, so far as I can tell, the article they proposed to write with him was never published.64 But if the work lacked mathematical sophistication and fruitful-theorizing, aplomb, it nonetheless was sound-and and it laid sure foundations for later work. It was natural that the first outgrowth of Dubinin and Romashov's theory was not its mathematical development but its experimental testing by the study of Soviet ecogenotypes of Drosophila melanogaster beginning in 1931, and published in 1934 and 1936. Eventually Wright joined with Theodosius Dobzhansky, who had realized sooner than most the significance of Dubinin's work, in an evaluation of the effect of drift in a natural population published in 1941.65 By then, however, there was already an impressive literature on the subject-almost all of it in Russian.66 Dubinin and Romashov's theoretical views seem modern to us for yet another reason: they consciously attempt to 64. Mention of the work with Kolmogorov appears in an article on the work of the Institute of Experimental Biology, "Raboty Instituta Eksperimental'noi Biologii Narkomzdrava," Biologicheskii Zhurnal, 3 (1934), No. 1, p. 223. The proposed work was to be called "Uchenie o ravnovesii populiatsii i protsess izogametatsii," (A study of population equilibrium and the process of isogametization). 65. Th. Dobzhansky and S. Wright, "Genetics of natural populations. V. Relations between mutation rate and accumulation of lethals in populations of Drosophila pseudoobscura," Genetics, 26 (1941), 23-51. 66. For much of this literature, see the bibliographies to Dubinin, "On lethal mutations . . ." (1946), and also to the article by C. Gordon, H. Spruway, and P. A. R. Street, J. Genet., 38 (1939). The references listed at the end of the 1939 piece, twenty-five in all, include most genetic analyses 18 in number. Signifiof wild populations done prior to that time-some cantly, 13 of these had been done by members of the Russian School: Chetverikov or his students, N. V. Timofeev-Resovskii, or Dubinin and colleagues.

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Russian Population Concepts integrate information from many biological disciplines in formulating evolutionary explanations of concrete cases, borrowing heavily from genetics, ecology, biogeography, and entomology. Like Chetverikov, they believed genetics had to be at the core of evolutionary explanation but also realized that genetics alone could not generate evolutionary explanation; that evolutionary principles could not be deduced from genetic ones. They shared with Chetverikov the naturalist's desire to come to terms with isolation and its role in producing genetic divergence between populations. The evolutionary thinking of Chetverikov, Dubinin, and Romashov achieved the beginnings of an integration of experimental genetic approaches with a concern for natural populations in the wild. One of Chetverikov's main goals had been to give the theory of isolation a genetic foundation. The success of Dubinin and Romashov's theory in achieving this goal made possible a view of evolution which was more "synthetic" in Chetverikov's sense. ACKNOWLEDGMENTS I wish to express my gratitude to Theodosius Dobzhansky, I. M. Lerner, Ernst Mayr, and Everett Mendelsohn, who read the manuscript and made several helpful suggestions.

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Historical Aspects of F. W. Putnam's Systematic Studies on Fishes' RALPH W. DEXTER Department of Biological Sciences Kent State University, Kent, Ohio

Frederic Ward Putnam (1839-1915) was a student of Louis Agassiz specializing on fishes, especially the blind fishes and the darters. In the former group he described one new species (Chologaster agassizii Putnam, 1872) and in the latter group he described two genera (Cottogaster, now included with Percina, and Microperca, now included with Etheostoma), and two new species, M. punctulatum and M. kennicotti, both of which are still valid. The first species is now known as Etheostoma microperca Jordan and Gilbert (Putnam's name was preoccupied in Etheostoma) and the latter is known as E. kennicotti (Putnam). Putnam is given brief mention by Hubbs as one of the notable ichthyologists of the third quarter of the last century.2 Putnam served as Curator of Fishes for the Boston Society of Natural History from 1860 to 1868, and Curator of Vertebrates for the Essex Institute at Salem, Massachusetts, from 1864 to 1867. He continued in the latter capacity at the Peabody Academy of Science, an outgrowth of the Essex Institute, from 1867 to 1874. Putnam was also Director of the Museum at the Peabody Academy, founded by George Peabody in 1866, and was part-time Curator of Ichthyology at Harvard's Museum of Comparative Zoology between 1876 and 1878. His first publications, as a young naturalist age 16, were reports on fishes 1. Paper read at the 19th annual meeting of the Society of Systematic Zoology, New York City, 27 December 1967. Quotations are taken from the F. W. Putnam papers in the Archives of Harvard University, with permission of the Archives and the Putnam family. 2. C. L. Hubbs, "History of Ichthyology in the United States after 1850," Copeia, 1964, pp. 42-60. Journal of the History of Biology, Vol. 3, No. 1 (Spring 1970), pp. 131-135.

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he collected from Salem Harbor. During his career as a zoologist he published thirty-three brief articles and notes of a general nature and nine papers of a systematic nature on fishes. For nine years (1860-1868) he published an "Annual Report of the Curator of Ichthyology" in the Proceedings of the Boston Society of Natural History. He also prepared the Annual Report of the Commissioners on Inland Fisheries for the State of Massachusetts during the years 1882 to 1888. His first systematic paper was "List of the Fishes of Essex County, Massachusetts" published in 1856 when he was a firstyear student of Agassiz.3 He then developed a special interest in the Etheostomidae (darters). He wrote to his fiancee on 25 September 1859, "These are found only in North America. Heretofore there were only some 20 species known and described, and I have made out about 80 or 90 species." Later that year he wrote to her again, "I am getting to be quite widely known among scientific men. Prof. Hind of Canada has sent me a fish which he wishes me to describe for his report on the Exploring Expedition of the Upper Part of British Provinces-it will be quite a feather in my cap." Putnam recorded in his notebook for 1859: "I have resolved to spend my life in the study of ichthyology and to do all in my power to elevate the study and put it upon its true basis." The next year he began a manuscript entitled "Notes on Etheostomoids." In the introduction to this work he stated, "The following catalog is preliminary to a monograph of the family of Etheostomidae," but the monograph was never completed. Agassiz placed Putnam in charge of the fishes collected by the U. S. Exploring Expedition. At the same time they planned a major work to be published jointly. Putnam wrote in his diary on 8 January 1862, 'We spoke over a work on American Ichthyology to be undertaken between us if we could get a publisher to take it-this is my great hope and aim." But this projected plan, too, was never completed. Early in his career Putnam tangled with Theodore Gill on taxonomic matters, and a bitter controversy was continued for many years. Putnam soon became friends with E. D. Cope, but his first impression of Cope was that he seemed to be "a fine fellow, only given too much to species making." Cope encouraged Putnam, writing to him on 26 February 1862, "I am very glad to hear that you are at work on the Catastomidae. I have often attempted to study them, but have found them very 3. F. W. Putnam, "List of Fishes of Essex County, Mass., 1855-56," Proc. Essex Institute, 1 (1856), 144, 148, 201-231.

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F. W. Putnam's Studies on Fishes imperfectly illustrated. When will your Etheostomids be out? I want to see these about as much as any work on North American Fishes." Following the split between Agassiz and his students, known as the "Salem Secession,"4 Samuel Scudder, former fellow student and lifelong friend of Putnam, wrote to him on 11 July 1867, I promised Prof. Agassiz to communicate to you an offer of his, and to request you to respond to it.-He offers to put all your material on the Etheostomoids into your hands, and in addition, all which has accumulated in the Museum since you left, and give you all the illustrations you desire if you will put the manuscript in shape for publication in the Illustrated Catalogue.-I hope you will accept his proposition. -It seems to me to savor of a more generous spirit and I have tried to judge it from the point of view of advantage to scientific truth apart from personal considerations.-but I shall not judge you harshly if you decline, but only think privately that you have made a mistake. This broke the ice between Agassiz and Putnam. Putnam agreed to the proposal, and on 9 May 1868 Agassiz wrote him: "I accept your proposition to work up fully the Etheostomoids. I shall have the necessary drawings made as you proceed with the fishes.-Please let me know when you are likely to be able to begin." After the death of Agassiz (1873), his son Alexander wanted Putnam to complete the work on fishes obtained by the U. S. Exploring Expedition. Alpheus Hyatt, another close friend and former fellow student, wrote Putnam, "Agassiz said before us all, that if anything happened to him he should wish you to have the continuance of his work on the Wilkes Expedition fishes." Putnam was both willing and anxious to undertake these tasks, but in a year's time he was appointed Curator of the Peabody Museum of American Archaeology and Ethnology at Harvard University and turned abruptly from ichthyology to American archaeology. Unfortunately, he never found time to complete the projects on the Etheostomoids or the fishes of the Exploring Expedition, although upon request of the Agassiz family he did prepare for publication several of Agassiz's unpublished papers on fishes. 4. R. W. Dexter, "The 'Salem Secession' of Agassiz Zoologists," Essex Institute Hist. Coll., 101 (1965), 27-39.

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In 1871 Putnam published a "Note on E. D. Cope's Classification of Fishes." Putnam was "not prepared to accept all the changes that were proposed," but regarded Cope's work as the nearest approach to a correct classification of the fishes that had ever been made. The next year he published his study on "Blind Fishes of Mammoth Cave and Their Allies" and a "Synopsis of the Family Heteropygii," including his description of Chologaster agassizii as a new species.5 The former paper brought forth criticism from Sir Richard Owen, who did not agree with the interpretation of optic lobes in blind fishes as given by Putnam.6 He also published that year his "Summary of the Etheostomoids,"7 which included about 40 species and was to be his major work on the taxonomy of fishes, since his projected monograph was never completed. After the publication of his last two papers on systematic ichthyology in 1874,8 Putnam's attention was given entirely to his new interest in American archaeology. In the summers of 1873 and 1874 Putnam taught systematic ichthyology at the Anderson School of Natural History, the marine station on Penikese Island founded by Louis Agassiz. One of Putnam's students was David Starr Jordan. Jordan wrote to Putnam in January 1874, "I will send you a list of our fishes [of Indiana] sometime if you will do me the favor to correct the synonyms and bring the names up to the present time." Shortly before the second and last summer session on Penikese in 1874, Jordan wrote again: "I have besides a large number of Etheostomoids and small Cyprinoids which I will bring with me to the island [Penikese] and work them up with your assistance." Jordan expressed his appreciation of the help given to him by Putnam in a letter of 11 January 1875: "-You are almost the only ichthyologist who doesn't prefer to keep all he knows to himself lest somebody forestalls him and makes a genus first." 9 With Putnam's aid, Jordan followed up the work of Rafin5. F. W. Putnam, "Note on E. D. Cope's Classification of Fishes," Amer. Natur., 5 (1871), 593; "Blind Fishes of Mammoth Cave and Their Almes," ibid., 6 (1872), 6-30; "Synopsis of the Family Heteropygii," 4th Ann. Rep. Peabody Acad. Sci., 1872, pp. 15-23. 6. R. W. Dexter, "Sir Richard Owen's Interpretation of Optic Lobes in Blind Fishes," Amer. Natur., 100 (1966), 271-272. 7. F. W. Putnam, "The Etheostomoids," ibid., 6 (1872), 109-115. 8. F. W. Putnam, "Remarks on the Li paridae," Proc. Boston Soc. Nat. Hist., 16 (1874), 114; "Notes on the Myxinidae," ibid., pp. 127-135, 156160. 9. See R. W. Dexter, "Can One Predict Success in Science?" Science, 139 (1963), 670.

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F. W. Putnam's Studies on Fishes esque on Ohio River fishes. On 24 December 1875, Jordan wrote to Putnam: Would it be possible for love or money for me to borrow your copy of Rafinesque's Ichthyologia Ohioensis long enough to copy the descriptions of those species not yet identified or only doubtfully so. -I am so located now that I can go and seine all those streams where Rafinesque fished and I am morally certain that some species very abundant here -not traced to him-could not have escaped him." Jordan acknowledged receipt of the volume and commented (10 January 1876): "I find the old man's descriptions are much better than he has been given credit for." 10 Dr. John Newberry, Chief of the Ohio Geological Survey, wanted Putnam to prepare a volume on "Fishes of Ohio." This task was turned over to Jordan on the recommendation of Spencer F. Baird with the blessings of Putnam. Newberry wrote Putnam (4 November 1876). I had been so desirous that you should prepare the report that, as you know, I waited two years for your affairs to come into such shape that you could do it. Now that you find it impossible, I am driven to do the next best thing and at the recommendation of Prof. Baird have made an arrangement with Dr. Jordan to write the report. I do not know Dr. Jordan personally, nor have I any satisfactory means of judging for myself in regard to his merits, but Baird speaks in the highest terms of him, and thinks that he will do the work well. Summary: As a student and collaborator of Louis Agassiz on the study of fishes, F. W. Putnam gave promise of becoming a leading ichthyologist with special interest in taxonomy generally and the Etheostomidae in particular. While he was noted briefly in these fields, contributed a number of minor papers, and aided in the posthumous publications of some of Agassiz's work on fishes, he neither reached his original goal nor completed his major projected works. For in 1874 he switched careers and was appointed Curator of the Peabody Museum of American Archaeology and Ethnology at Harvard University, and is remembered today primarily as a founder of American archaeology rather than as a systematic ichthyologist. 10. R. W. Dexter, "Jordan on Rafinesque," System. Zool., 5 (1956), 46-47.

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VertebratePaleontology,an EarlyNineteenthCenturyTransatlanticScience PATSY A. GERSTNER Howard Dittrick Museum of Historical Medicine Cleveland, Ohio

Questions have often been asked concerning the attitude of the European scientist toward his counterpart in America during the first half of the nineteenth century, particularly whether or not there was any respect for American work or any use made of it in Europe. The answer is not yet available for all the sciences, but whatever the relations of American and European students of the more established sciences might have been, it is clear that in the burgeoning field of vertebrate paleontology American descriptive studies were respected and used during the first half of the century. An attitude of skepticism toward American studies of vertebrate fossils, however, did prevail in Europe during the first quarter of the century.' Although Americans commonly made every effort to learn of European activities concerning fossils the interest was not reciprocal. Until about 1830 Americans were, for the most part, looked upon as fossil collectors with dubious qualifications for making descriptive studies or interpreting the results. After 1830, however, the European attitude changed. This change was based in part on the adoption in America of the theoretical framework established by Georges Cuvier, the framework within which most paleontology was then being done in Europe, so that there existed in the second quarter of the nineteenth century a similar basis for paleontology in America and in Europe. More specifically the change was based on a rapid development in the ability to use the methods of Cuvier 1. This was not without exception. Professor John C. Greene has given some examples of British interest in work done in America before 1820 in The Death of Adam; Evolution and Its Impact on Western Thought (Ames, Iowa: Iowa State University Press, 1959), chap. 4. Journal of the History of Biology, Vol. 3, No. 1 (Spring 1970), pp. 137-148.

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in studying fossils. Cuvier's work was based largely on the reconstruction of fossil animals which was made possible by an extensive knowledge of comparative anatomy. His methods had little impact in America until about 1820. At that time the study of comparative anatomy was growing in some places, and several people, mostly physicians, began to appear with more than casual interest in that study. For those who happened also to be interested in fossils, an access to the works of Cuvier was provided through this knowledge of comparative anatomy.2 The change brought about by these men was apparent soon after 1830 in a new feeling toward American studies among Europeans. A distinct indication that a new status was achieved is to be found in a statement by Roderick Murchison, who, in reflecting on a similar controversy in America and in Europe about the zoological position of a fossil elephant, noted that American naturalists had not been backward in studying fossils and that it was "not therefore, a little remarkable that the same difference of opinion as to the generic and specific identity of the animals that prevailed across the Atlantic, is presented in the memoirs which have recently been read before us [the Geological Society of London]."3 Similar abilities were requisite to an appreciation of American studies of fossils, and since similarity is well attested in the "difference of opinion" to which Murchison referred, the circumstances leading to his statement are of interest. In 1830 John D. Godman, physician and naturalist of Philadelphia, described before a meeting of the American Philosophical Society a mastodontoid animal preserved in the Society's collection.4 The bones were of the common American 2. For a more complete statement of Cuvier's influence on American paleontology and on the interests in comparative anatomy which characsee my unpublished dissertation terized some physician-paleontologists, entitled "The 'Philadelphia School' of Paleontology: 1820-1845" (Case Institute of Technology, 1967). 3. R. I. Murchison, "Anniversary Address of the President [1843]," Proc. Geol. Soc. Lond., 4 (1842-1845), 145. Though Murchison's statement was made in 1843, it reflects events of the preceding several years and is thus appropriate as an indication of a changing status in the 1830's. 4. John D. Godman, "Description of a New Genus and New Species of Extinct Mammiferous Quadruped," Trans. Amer. Phil. Soc. 3 (1830), 478485. Godman's paper was probably read at the January 1 meeting. The proceedings for that date were not recorded, but at the January 15 meeting, which was the second meeting of the year, Godman's paper was approved for publication by a committee appointed to consider it. The specimen described by Godman came from a site in Orange County, New York, not far from the famous 1801 mastodon excavation of Charles Wilson Peale.

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Vertebrate Paleontology Pleistocene mastodon, the Mastodon giganteum of Cuvier. However, this was an anomalous specimen having small tusks in the lower jaw as well as the characteristic large pair in the upper jaw (giving a total of four tusks instead of the usual two), and Godman, choosing to emphasize the presence of the two small tusks and a slightly elongated jaw over characteristics in common with Mastodon giganteum, assigned the specimen to a new genus and christened it Tetracaulodon mastodontoideum .5 Immediate reaction to Godman's interpretation came from Richard Harlan in Philadelphia and William Cooper in New York. Both noted in particular that, except for the additional tusks, all characteristics of the specimenr were in keeping with a normal variation of Mastodon giganteum. Therefore it seemed unreasonable to assume that the lower tusks did not also represent a normal variation, in this case a variation that could be accounted for as either a juvenile or sex-linked characteristic.6 Dr. Isaac Hays, a friend of Godman who shared his enthusiasm for fossil studies, and George Featherstonhaugh, a naturalist of diverse interests, became outspoken defenders of the views of Godman and Harlan respectively, and the true nature of Tetracaulodon became a point of bitter debate among the fossil devotees of America.7 5. The ancestors of the American Pleistocene mastodon had well developed lower as well as upper tusks. As the animal evolved toward its American Pleistocene form, the lower tusks underwent a reduction in size but were never completely eliminated from the population. Henry Fairfield Osborn noted that although the lower tusks are wholly wanting in the majority of American Pleistocene mastodons their presence cannot be termed rare. The lower tusks are found in some jaws embedded in the bone, never erupted, while in others one or two did erupt (Proboscidea. A Monograph of the Discovery, Evolution, Migration and Extinction of the Mastodonts and Elephants of the World [New York: American Museum of Natural History Press, 1936], 1, 174). 6. For Harlan's opinions see the Proc. Amer. Phil. Soc. 22, April 16, 1830 and his "Critical Notices of Various Organic Remains hitherto Discovered in North America," Trans. Geol. Soc. Pa., 1 (1834), 52. Cooper's opinion was published in his "Notices of Big-Bone Lick," Mthly. Amer. J. Geol. Nat. Sci. 1, (1831), 163, fn. 7. For Hays's comments and information on his position, see his "Description of the Inferior Maxillary Bones of Mastodons in the Cabinet of the American Philosophical Society, with Remarks on the Genus Tetracaulodon (Godman), etc.," Trans. Amer. Phil. Soc., 4 (1834), 317-339; the report of the monthly meeting in J. Franklin Inst., 8 (July, 1830), 6-8; and the review of "The Monthly American Journal of Geology and Natural Science," J. Franklin Inst., 8 (August, 1831), 143-144. For Featherstonhaugh's views see particularly his "To Readers and Correspondents," Mthly. Amer. J. Geol. Nat. Sci., 1, 9 and 140-144.

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Partly because the differences of opinion were published in jourmals available to Europeans and partly because of the intensity of the arguments, the controversial nature of the specimen was well known in Europe from the early 1830's.8 Europeans did not really study the problem, however, until Albert Koch, renowned collector of old bones, arrived in England in 1841 with several specimens of Tetracaulodon. Members of the Geological Society of London who examined these specimens found themselves in the same predicament the Americans had experienced several years earlier. Opinions expressed by Robert E. Grant, Professor of Comparative Anatomy and Zoology at University College, London, and Alexander Nasmyth, painter and architect by profession but widely known for his scientific interests, were identical to the position of Godman and Hays and relied on emphasizing one or two characteristics over the whole.9 On the other hand, Richard Owen held a view like that of Harlan and Cooper which emphasized the whole over the part, and in an event reminiscent of the American scene Owen and Grant engaged in angry discussion about Tetracaulodon before the London Geological Society.10 Each 8. Godman's and Hays's papers were probably available through European distribution of the Trans. Amer. Phil. Soc. In addition, Godman's paper appeared in the Annales des Sciences Naturelles, 20 (1830), 292-301. Harlan's longest published statement on the Tetracaulodon tusks as juvenile characteristics appeared in the form of a synopsis of a letter he sent to the Bulletin des Sciences Naturelles et de Geologie, 22 (1830), 320. His views were also available in his "Critical Notices of Various Organic Remains hitherto Discovered in North America" which was published both in Trans. Geol. Soc. Pa. and in the Edinburgh New Phil. J., 17 (1834), 342363, and 18 (1834), 28-40. Cooper's view and Featherstonhaugh's were available in the Mthly. Amer. J. Geol. Nat. Sci., which was probably distributed in Europe (see the "Prospectus" published in vol. I, 1831, of the journal, 1-4). In addition to these opportunities to know Tetracaulodon, casts of the specimen were available at the Jardin des Plantes and the Geological Society of London through Harlan's efforts (see his "Critical Notices," Trans. Geol. Soc. Pa., 50-51, and the Proc. Geol. Soc. Lond., 3 [February 27, 1833]). On February 5, 1834, still more casts reached the Geological Society of London, this time from the American Philosophical Society (see Trans. Geol. Soc. Lond., 4 [February 5, 18341). 9. Robert E. Grant, "On the Structure and History of the Mastodontoid Animals of North America," Proc. Geol. Soc. Lond., 3 (1838-1842), 770771; Alexander Nasmyth, "On the Minute Structure of the Tusks of the Extinct Mastodontoid Animals," Proc. Geol. Soc. Lond., 3 (1838-1842), 775-780. Grant based his position in part on the form of the molar teeth and Nasmyth on the cellular structure of the tusks. The basic problem of defining generic characteristics adequately was the same, however. The opinions of Grant and Nasmyth are also discussed by Murchison in his "Anniversary Address of the President" for 1843. 10. Richard Owen, "Report of the Missourium Now Exhibiting at the

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Vertebrate Paleontology man was clearly well versed in what Americans had said about Tetracaulodon; there was no difficulty or question about the description or the methods of the Americans. In fact, Owen credited Cooper with the first essentially correct statement about the animal's zoological position."1 The only real problem on both sides of the Atlantic concerned the fundamental nature of a valid generic characteristic. In neither England nor America could this problem be immediately settled to the satisfaction of the majority, but the similar responses of both groups were strikingly obvious to everyone and resulted in Murchison's acknowledgement of the equality of American and European paleontological work. With varying degrees of success, each party to the Tetracaulodon debate in America was responsible for the development of paleontology to the point that made it acceptable in Europe, but by far the most important was Richard Harlan. Harlan was the most skillful of the American paleontologists in the decade of the 1830's, and it is in his relations with Europe that the new attitude toward American work is most often apparent. Because he did work more extensively with fossils than any of his immediate contemporaries, but also because he was able to visit Europe and meet those most active in the field in France and England, he was in a position Egyptian Hall, with an Inquiry into the Claims of the Tetracaulodon to Generic Distinction," Proc. Geol. Soc. Lond., 3 (1838-1842), 689-695. The debate between Owen and Grant is referred to by Gideon Mantell in his journal on June 15, 1842 (The Journal of Gideon Mantell, Surgeon and Geologist, Covering the Years 1818-1852, Cecil E. Curwen, ed. [London: Oxford University Press, 1940], p. 159). Owen combined the idea that the tusks were juvenile anomalies with the idea that they were sexual characteristics. He believed that a pair of small lower tusks was present in all the young animals belonging to the Mastodon giganteum, that a single tusk was preserved in the adult male, and that none remained in the adult female. Young specimens with two tusks had been described by Godman and by Hays. In the collection Koch brought to England, several clearly adult specimens showed evidence of the eruption of only a single tusk (either one or both tusks can erupt as noted in fn. 5). Since there were certainly many mastodon skulls known without lower tusks, Owen reasoned that the presence of a tusk in an adult mastodon indicated a male. He cited analogous situations in the animal kingdom, particularly the narwhal, where the young of both sexes have two rudimental tusks, only one of which develops beyond the rudimental stage in the adult male. Owen and others who believed the lower tusks were sexual characteristics were partly correct. Osborn (Proboscidea, p. 175) noted that when lower tusks do occur they occur in the male only. This does not, however, mean that males without lower tusks did not exist. 11. Owen, "Report of the Missourium," 690; Cooper, "Notices of Big-Bone Lick."

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both to encourage the development of what may be termed a transatlantic science and to participate in that science. Harlan (1796-1843), a graduate of the medical department of the University of Pennsylvania and a resident of Philadelphia, had shown an intense interest in comparative anatomy from the time of his graduation in 1818.12 Within the following few years he became completely enamored of Georges Cuvier the man and Georges Cuvier the paleontologist, holding him as a personal hero and a scientific guide. Harlan's career as a paleontologist began in 1823 and lasted until his death in 1843.13 As a member of both the American Philosophical Society and the Academy of Natural Sciences of Philadelphia, he was in constant contact with much of the paleontological work in America. From this vantage point, and as his own work progressed, he came to realize more than any of his contemporaries that paleontology was by its very nature an international study. He was acutely aware of the need for adequate communication of ideas to Europe. Prompted by this realization and by requests from European scholars he met while visiting Europe in 1833, he prepared an extensive article in 1834 summarizing developments in paleontology in America entitled "Critical Notices of Various Organic Remains hitherto Discovered in North America." The article was meant to serve a twofold purpose, demonstrating that Americans had been increasingly active in the area of fossil study and presenting their work in a convenient form for use by Europeans.14 This synopsis was published almost simultaneously in Europe, and its availability was quickly made known to members of the Geological Society of London by George Greenough.'5 In 1835 Harlan took another step in bringing 12. In 1821 Harlan was appointed lecturer on comparative anatomy at the Philadelphia Museum Company, an outgrowth of Peale's Museum, and served in this capacity for a number of years. He was reported at one time to possess the finest collection of specimens for comparative anatomy in Philadelphia which further underlines his interests along this line ("A Sketch of the History of the Museum," The Philadelphia Museum, or Register of Natural History and the Arts, 1 [18241, 3). 13. Harlan's first published piece on fossils appeared in 1823: "Observations on Fossil Elephant Teeth, of North America," J. Acad. Nat. Sci. Philadelphia, 3 (1823), 65-67. 14. This article appeared in volume I of Trans. Geol. Soc. Pa. (1834), 46-112. Harlan's purpose for compiling the information is stated on p. 46. 15. In Europe the "Critical Notices" was published in Edinburgh New Phil. J., 17 (1834), 342-363 and 18 (1834), 28-40. The American edition included both vertebrate and invertebrate paleontology, but the invertebrates were omitted from the Edinburgh edition. Greenough's announcement is found in his "An Address Delivered at the Anniversary Meeting of the Geological Society of London on the 20th of February, 1835," Proc. Geol. Soc. Lond., 2 (1833-1838), 159.

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Vertebrate Paleontology American information to Europe by seeing that a volume containing some twenty of his own papers on fossils, called the Medical and Physical Researches, was available in Europe.', The appearance of information on American paleontology was not without precedent in Europe as several French and British journals frequently published American papers, in abstract, after 1830, and publications such as the Monthly American Journal of Geology and Natural Science, which contained fossil reports, were circulated in Europe.17 Harlan's efforts reflect the existing situation while at the same time they represent an important extension of it. Harlan's most important efforts in encouraging the development of paleontology as a joint European-American science took the form of two trips to Europe, one in 1833 and one in 1839. During the first trip he received the requests for information that led to his "Critical Notices" and generally stimulated interest in what Americans were doing. During the second trip a cooperative study with Richard Owen aided in the solution of a problem that was of major interest to European paleontologists. Harlan's first trip, in 1833, included an extended tour of the British Isles and the continent. During his travels, and especially during longer stops in Paris and London, he met many of Europe's leading paleontologists. In France he exchanged information with Henri Ducrotay de Blainville and Alexandre Brongniart; in England he joined in the Third Annual Meeting of the British Association for the Advancement of Science held at Cambridge in June of 1833.1' Here he had 16. Philadelphia: Lydia R. Bailey, printer, 1835. This volume also contains papers on general biology and physiology. In an effort to secure wide distribution of this work Harlan wrote to his close friend, John James Audubon, who was then in England, and asked him to announce the availability of the work to the Linnaean Society, the Geological Society of London, the Zoological Society, the British Museum, and, because it also contained papers of medical interest, to several medical and surgical Historical organizations (January 3, 1836, Gratz MSS, Pennsylvania Society). He also sent several copies to a London distributor to make sure they were available to any interested parties. 17. Journals publishing abstracts or entire papers included the BibliothWque Universelle des Sciences, the Bulletin des Sciences Naturelles et de G6ologie, the Philosophical Magazine, and the EdinbuTgh New Philosophical Journal. See also fn. 8 for information on the availability of American ideas in Europe. 18. His activities are chronicled in a letter to J. J. Audubon on January 9, 1834 (Misc. MSS, New York Public Library). Harlan wrote the date on this letter as 1833. It should clearly read "1834" since the letter was written after his return to the United States.

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the opportunity to meet the most influential proponents of fossil studies in England. Harlan did more than simply attend the meeting; he participated in it. He served on the Committee of Geology and Geography and read a paper on some saurian reptiles found in America.19 Gideon Mantell, an attendant at the meeting, reported that Harlan's paper and comments on American paleontology led to at least one extensive discussion.20 One of the specimens described by Harlan was a large vertebra which he assigned to a new genus called Basilosaurus. Subsequently proven to be the vertebra of a whale, Harlan identified the bone as that of a reptile. Several years later, on the basis of more material and therefore a more extensive description, Basilosaurus became significant in the EuropeanAmerican interplay of science. It became an important part of an argument over the class of animals to which a few fossils from Secondary deposits, specifically the Middle Jurassic Stonesfield slates in England, belonged. These fossils were first discovered about 1814 and were correctly identified as mammals by William Buckland and Georges Cuvier. It was generally believed, however, mainly because of Cuvier's concept of a succession of worlds and his extensive investigations of the fauna that characterized each world, that mammals had not existed until the Tertiary. Cuvier suggested that the Stonesfield deposits had been misinterpreted and were, in fact, Tertiary. However, studies made by William H. Fitton in 1828 offered convincing proof that the deposits were Secondary. Buckland and some other followers of Cuvier were able to accept the seeming contradiction between fact and theory and continued to believe the fossils were mammals; others could not do so and reasoned that the fossils must be those of reptiles.2' The strongest supporter of the latter interpretation 19. "On Some New Species of Fossil Saurians Found in America," Report of the Third Meeting of the British Association for the Advancement of Science, Held at Cambridge in 1833 (London: John Murray, 1834), 440. The paper dealt with two fossils, Ichthyosaurus missouriensis and Basilosaurus. The former was subsequently described in "Notice of the Discovery of the Remains of the Ichthyosaurus in Missouri, N. A.," Trans. Amer. Phil. Soc., 4 (1834), 405-408. The Basilosaurus was described in "Notice of the Fossil Bones Found in the Tertiary Formation of the State of Louisiana," Trans. Amer. Phil. Soc., 4 (1834), 397-403. This paper on Basilosaurus had been read before a meeting of the American Philosophical Society on October 19, 1832, some eight months before it was read to the British meeting. 20. The Journal of Gideon Mantell, p. 117, and letter from Mantell to Benjamin Silliman, October 3, 1833 (Silliman-Mantell MSS, Yale University). The nature of the discussion is not made clear. 21. A concise history of the early arguments about the Stonesfield mam-

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Vertebrate Paleontology was Henri Ducrotay de Blainville, and the concise and forceful presentation of his reasons to the Academy of Sciences in Paris in 1838 started a controversy in France and England that was to last for several months.2 From its beginning in 1838 this controversy involved Richard Harlan and Basilosaurus. Blainville believed that the characteristics of the Stonesfield fossils in general were like those of reptiles, but the teeth presented a particular obstacle to this interpretation.23 The teeth were of two kinds, some having double roots. This was generally considered a mammalian characteristic. In an effort to counter this general belief Blainville turned to Harlan. Blainville had undoubtedly been aware of Harlan's Basilosaurus since 1833 when Harlan visited him in Paris, but it mals may be found in George Gaylord Simpson, A Catalogue of the Mesozoic Mammalia in the Geological Department of the British Museum (London: The British Museum, 1928), p. 3-5. Henri Ducrotay de Blainville, as a participant in the controversy, gives an interesting and short history of ideas about the fossils, including mention of Cuvier's doubts of the age of the Stonesfield deposits, in his "Doutes sur le pretendu Didelphe Fossile de Stonefield, ou a quelle Classe, a quelle Famille, a quel Genre doit-on rapporter l'animal auquel ont appertenu les Ossements Fossiles, ai Stonefield, d6sign6s sous les Noms de Didelphis Prevostii, et Didelphis Bucklandii, par les Pal6ontologistes," (Comptes Rendus, 7 [1838], 402-418), an English translation of which appeared in The Magazine of Natural History, 2 (1838), 639-654. However, neither Simpson nor Blainville takes into account the opinion of Charles Lyell, which undoubtedly brought further emphasis to the reptilian interpretation. Lyell argued that the presence of even one mammal in Secondary deposits was completely fatal to the doctrine of a geological succession of worlds and as completely favorable to his uniformitarian concept (Principles of Geology, 2nd ed. [London: John Murray, 18321, 1, 173). The fossils from the Stonesfield slates that figured in the controversy belong to two distinct orders of primitive mammals, the Pantotheria and the Triconodonta (Simpson, pp. 67 and 107). These are among the most primitive of the mammals and among the earliest to evolve from a reptilian ancestry. The zoological characteristics and affinities of such primitive groups are often confusing even today, and it is not surprising to find that much of the controversy over the nature of these fossils in the 1830's sounds wholly zoological rather than geological. However, the relative roles of the zoological understanding of the characteristics of the Stonesfield fossils versus the theoretical complications arising from geology have not, so far as I am aware, been determined. 22. This controversy may be traced in the Comptes Rendus de 1' Academie des Sciences for 1838 and in the Proceedings and Transactions of the Geological Society of London for 1838 and 1839. The duration of the arguments is specifically referred to by William Whewell in his "Address to the Geological Society, Delivered at the Anniversary on the 15th of February, 1839," Proc. Geol. Soc. Lond., 3 (1838-1842), 86. 23. For Blainville's detailed opinions see his "Doutes sur le pretendu Didelphe Fossile" and his "Nouveaux Doutes sur le pretendu Didelphe de Stonefield," Comptes Rendus, 7 (1838), 727-736.

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was the more extensive article, published by Harlan in 1835 and based on the discovery of more bones, that provided Blainville with the help he wanted.24 This article contained a good description of the teeth and jaws of Basilosaurus as well as vertebrae, ribs, and long bones. Blainville agreed with Harlan's conclusion, which was based on a consideration of all parts of the fossil, that Basilosaurus was a reptile. Harlan had noted that Basilosaurus had teeth with double roots, and Blainville used this information to uphold his contention that the Stonesfield fossils were reptiles in spite of some generally accepted ideas about tooth structure: M. le D. Harlan of Philadelphia described and figured many years since, some gigantic fossil bones, which he referred to a new genus of the class Reptilia, named by him Basilosaurus. Now a portion of the jaw of this animal displays implanted teeth of two kinds, the first being simple, among which there are even some resembling canines larger than others; the second compressed, triangular, and provided with two roots, fixed in the jaws, and projecting beyond their edges; and as at the slightest glance we cannot refuse to acknowledge their great analogy to what is described and drawn of the Stonesfield animal ... we can neither hesitate to admit that if this Basilosaurus be a reptile, which fact appears placed beyond doubt by the form of the vertebrae, humerus, etc., it is more than probable that it is an animal of the same kind as that found at Stonesfield.25 When, late in 1838, the Stonesfield fossils became a point of discussion among members of the Geological Society of London, Richard Owen entered the developing controversy in opposition to Blainville and decidedly in favor of the mammalian classification of the Stonesfield fossils.26 While the controversy was developing in Paris and London, 24. "Description of the Remains of the 'Basilosaurus', a Large Fossil Marine Animal Recently Discovered in the Horizontal Limestone of Alabama," Trans. Geol. Soc. Pa., 1 (1835), 348-357 and the Medical and Physical Researches, pp. 349-358. Blainville refers to this article as it appears in the latter. 25. Blainville, "Doutes sur le pr6tendu Didelphe Fossile" (English translation), 653-654. 26. Owen, "Observations on the Fossils Representing the Thylacotherium prevostii (Valenciennes), with Reference to the Doubt of Its Mammalian and Marsupial Nature Recently Promulgated; and on the Phascolotherium bucklandii," Trans. Geol. Soc. Lond., 6 (1841-1842), 47-66, or Proc. Geol. Soc. Lond., 3 (1838-1842), 5-9 and 17-21.

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Vertebrate Paleontology Harlan in America was planning a second trip to Europe. Aware of the involvement of his Basilosaurus in the Stonesfield problem, Harlan arrived in London early in January of 1839 with the teeth and bones of the specimen. Owen was well aware of the importance of the Basilosaurus teeth in supporting the arguments for the reptilian nature of the Stonesfield fossils, but up to that time had declined any comment, not because he felt there was any reason to mistrust Harlan's identification of Basilosaurus as a reptile, but because the teeth of that animal had never been examined microscopically.27 However, Harlan's arrival in England and the permission which he "liberally granted" Owen to have sections of the Basilosaurus teeth prepared for microscopic examination enabled Owen to arrive "at such conclusions, as to the nature of the Basilosaurus, as can be safely founded on the dental characters ascertainable by this mode of investigation."28 The microscopic structure of the teeth proved to be unlike that of the saurians in general and like the teeth of cetaceans.29 Owen went on to suggest further similarities in the vertebrae, ribs, and so on, of the whales and Basilosaurus, proving quite adequately that Basilosaurus was a mammal and therefore undermining a major argument for the reptilian character of the Stonesfield fossils. Owen had not reached his conclusion without consultation with Harlan. Owen made it clear to members of the Geological Society of London, and through their publication to the learned community in general, that he and Harlan had arrived at the conclusion together after a joint examination of the specimen.30 In this cooperative effort paleontology as a transatlantic pursuit emerged full-grown. In the years after 1830 Europeans respected American paleontology, and they accepted both descriptive and interpretive American fossil studies. That the descriptions provided by 27. The use of the microscope for fossil studies had been pioneered by Owen only a few years before, but had proved such a valuable adjunct to his work that he quickly came to rely on it. 28. "Observations on Basilosaurus of Dr. Harlan (Zeuglodon cetoides, Owen), Trans. Geol. Soc. Lond., 6 (1841-1842), 69-70. Though sometimes called the Zeuglodon today, the animal properly retains its original name, Basilosaurus. 29. Ibid., 72-73. At first Owen thought the teeth were those of a herbivorous whale but later decided on the basis of the vertebrae that the specimen was a carnivorous whale (Odontography: or, a Treatise on the Comparative Anatomy of the Teeth; Their Physiological Relations, Mode of Development, and Microscopic Structure, in the Vertebrate Animals [London: Hippolyte Bailliere, 1840-1845], p. 364). 30. Owen, "Observations on the Basilosaurus of Dr. Harlan," 75.

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Americans were generally accepted is clear in European statements about Basilosaurus and Tetracaulodon. This is implied also in the foreign requests for information which Harlan fulfilled in part with the "Critical Notices." But these were not isolated incidents. Owen used Harlan's description of an edentate, or sloth-like animal, in describing and defining the genus Mylodon of North and South America. In fact, Owen's description of the North American Mylodon was based on the description of that animal as published by Harlan.31 Yet it is apparent that European interest in American work was in no way limited to an appreciation of descriptive talent. Blainville expressed a belief in American interpretation when he accepted Harlan's view of Basilosaurus. In a similar vein, Murchison's appraisal of the Tetracaulodon controversy did the same. Owen's acknowledgement of Cooper's primary role in the proper identification and interpretation of Tetracaulodon, his hesitancy to challenge Harlan on Basilosaurus, and the cooperation between Harlan and Owen speak, perhaps, most clearly. American interpretations were sometimes wrong, but this is of little consequence. The most important prerequisite for the existence of a transatlantic science was a genuine and mutual respect for each other among the paleontologists on both sides of the Atlantic. Certainly, Americans had all the regard for Europeans that was necessary. In the events of the several years after 1830, Europeans clearly returned the feeling. 31. In the Medical and Physical Researches, "Description of the Jaws, Teeth, and Clavicle of the Megalonyx laqueatus," 334-336. For Owen's use of Harlan's study see his The Zoology of the Voyage of H. M. S. Beagle, under the Command of Captain Fitzroy during the Years 1832-1836, Pt. I: Fossil Mammalia (London: Smith, Elder and Co., 1840), pp. 63-68. Owen assigned the specific name haTlani to the North American Mylodon and referred to Harlan as "that indefatigable Naturalist who has contributed to natural sciences so much valuable information respecting the zoology, both recent and fossil, of the North American continent" (ibid., p. 68). Owen and Harlan never came to an agreement as to the genus involved, Harlan believing the animal to be Megalonyx laqueatus, but their differences were based on reasonable grounds. For their varying opinions see: Harlan, "Notice of Two New Fossil Mammals from Brunswick Canal, Ga., with Observations on Some of the Fossil Quadrupeds of the United States," Amer. J. Sci. 43 (1842), 141-144; "Description of the Bones of a New Fossil Animal of the Order Edentata," ibid., 44 (1843), 69-80; "Remarks on Mr. Owen's Letter to the Editors on Dr. Harlan's New Fossil Mammalia," ibid., 45 (1843), 208-211; Owen, "Letter from Richard Owen, Esq. F. R. S., F. G. S., etc. on Dr. Harlan's Notice of New Fossil Mammalia-Published in this Journal, Vol. XLIII, p. 141," ibid., 44 (1843), 341-345. Unfortunately, Harlan ill-advisedly accused Owen of using his description of Megalonyx laqueatus as the basis for the genus Mylodon.

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The Historyof the Naming of the LoblollyBay EDMUND BERKELEY Central Virginia Community College Lynchburg,

Virginia

Many of our students have long been convinced that taxonomists lie awake at night thinking of horrendous names for otherwise attractive plants. They may feel vindicated to learn that several very competent botanists required fifteen years to complete the naming of the Loblolly Bay, that handsome tree of the Southern Coastal Plain. The attempt to honor friends by naming plants for them, so prevalent in the eighteenth century was both natural and admirable, but it frequently provides a source of considerable confusion for present-day researchers. A name proposed by an American botanist, and referred to by him in a number of letters, was often later rejected and the name applied to a different plant. Some of the original correspondence may have been lost, while some survived. It is not difficult to be misled. In the case of the Loblolly Bay, the correspondence has fortunately been largely preserved. In December 1755, Dr. Alexander Garden (1730-1791) of Charles Town, South Carolina, sent a large shipment of seeds to John Ellis (ca. 1710-1776) of London. Students of this period of the history of biology will be familiar with both of these gentlemen. For the benefit of others, Alexander Garden was a young Scot who had received his education at Marischal College in Aberdeen and at the University of Edinburgh. He had spent several years as a surgeon's mate in the British Navy before settling in South Carolina in 1753. Ellis eventually named the Cape Jessamine Gardenia, but that is another long story, as is Garden's effort to name a plant Ellisia. Ellis was a native of Ireland, but spent most of his life in London. He was for a time a merchant, but later became King's Agent for West Florida and for Dominica. An able naturalist, he was a very active member of the Royal Society of London. His Journal of the History of Biology, Vol. 3, No. 1 (Spring 1970), pp. 149-154.

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major contribution was his establishment of corals as being animals rather than plants, a contention which even Linnaeus was reluctant to support. He had a large number of correspondents in various parts of the world who sent him a great variety of biological specimens, and to whom he sent many. Prominent among them were Alexander Garden and Carolus Linnaeus.' On Garden's "List of Seeds Shipped Aboard of Capt. Ball's Vessel for Mr. Ellis," this December day, number 2 is "Gordonia or Loblolly Bay caled by Catesby Aleca Floridana." Mark Catesby, in his beautifully illustrated Natural History of Carolina, had pictured the painted finch poised on a petal of the Loblolly Bay. Neither Garden nor Ellis seem to have been concerned with what Catesby had called the plant, but then Garden took a dim view of all of Catesby's work. Garden's shipment to Ellis also included young plants. Number 4 on this list was "Gordonia or Loblolly Bay, 8 young plants three years old & will flower this year." A few days later he sent more seeds to Ellis on board the Charming Martha. Number 18 on this list was again "Gordonia Loblolly Bay. This is a new genus as p[er] characters." 2 In a long letter to Ellis in March 1755, Garden included a "short account of the various genera whose characters I send you by this opportunity." He commented on the first four of these, and then added "The fifth, No. 5, is the Gordonia, which I call so in honour of my old master, Dr. James Gordon, at Aberdeen, a very ingenious and skilful physician and botanist, who first initiated me into these studies, and tinctured my mind very early with a relish for them. It is the Loblolly Bay which Linnaeus classes among the Hyperica, just with as much propriety as I might join Oak and Ambrosia, nay, indeed, there is not nigh such an affinity, as you will readily see, from comparing the characters of this and those of the Hypericum. I think you may take Catesby's cut of this to join with the characters." Garden yielded to no one in his admiration for Linnaeus, but he never hesitated to disagree with him when the occasion called for it, as this one clearly did.3 1. John Ellis, An Essay Towards the Natural History of the Corallines and other Marine Productions (London: 1755); Sir James Edward Smith, A Selection of the Correspondence of Linnaeus and other Naturalists (London: Longman, Hurst, Rees, Orme, and Brown, 1821), I, 79-81, 360-362. 2. Ellis Manuscripts, unnumbered, Linnean Society of London; George Frederick Frick and Raymond Phineas Stearns, Mark Catesby, the Colonial Audubon (Urbana: University of Illinois Press, 1961), p. 61. 3. 22 March 1756, Smith, Correspondence of Linnaeus, I, 378-379.

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The Naming of the Loblolly Bay Since the Loblolly Bay today bears the name Gordonia, one might reasonably assume that it bears the name given it by Garden at this time and that he would be credited with having named it. Such is not the case. Its name is Gordonia Lasianthus (L.), Ellis, and it was not named for Doctor James Gordon whom Garden had served as an apprentice at Aberdeen. It was not given this name until 1770, and during the fourteen years which intervened there was occasional mention of it in correspondence. In a long letter to Ellis on May 6, 1757, Garden wrote: "There is one thing which I entirely forgot to mention to you, and that is, that you need not call the Lobloly Bay, Gordonia. I shall leave the denomination to you. You know all those who I think merit the compliment among my own acquaintance besides yourself, Dr. Hales and Dr. Huxham. After these I shall give you the trouble of denominating the others as you please. You may, if you think proper, call the Loblolly Bay Huxhamia, or what you chuse." Why he had changed his mind about honoring Doctor Gordon is not clear. He was no more successful in conferring honor upon Stephen Hales or John Huxham, with whom he also corresponded.4 Two years later Garden seems to have changed from indifference to definite objection to naming this plant Gordonia. He wrote to EJlis on February 17, 1759: "I have sent you the only specimen which I have of the Loblolly Bay. This must not be called the Gordonia. Name it yourself, or let Linnaeus name it." Two days later he wrote again about various matters and again mentioned the Loblolly Bay: "this must not be called Gordonia. Name it yourself, or send it to Linnaeus to baptise." No explanation for this definite objection is apparent.5 In spite of all of Garden's efforts, Ellis was having no luck in getting the Bay to grow for him. In March 1759, he wrote to Garden asking him to send seeds of various kinds covered with melted tallow, in connection with studies he was making on how best to preserve them in transit. He included in this list "Loblolly Bay, or Gordonia, in the husks," and he commented that "the Loblolly Bay is a very rare plant with us, and has been lately sent to Mr. Collinson, in young plants, by Mr. Lamborn [Lamboll] of your town. These are likely to do well. Not any of the seeds which you sent have ever yet 4. John Kunkel Small, Manual of the Southeastern Flora (New York: published by author, 1933), p. 877; Smith, Correspondence of Linnaeus, I, 407-408. 5. Ibid, p. 436; ibid, 19 February 1759, p. 437.

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appeared, for they are so thin that they perish immediately; but I am in hopes the tallow will preserve them in their capsules." The eight three-year old plants which Garden had sent in 1756 had been lost when the ship on which they were sent failed to reach England. This was a very frequent occurrence and a source of great frustration to Garden and to his friends abroad. He could not let their mutual friend, Peter Collinson, steal a march on Ellis, so he hurriedly sent him seeds of the Bay covered not with tallow, but with myrtle wax. These did survive, and Ellis later reported that they had germinated. He also desired that the Loblolly Bay should be named Gordonia in honor of James Gordon, the famous nursery-gardener of Mile End, London, to whom both Ellis and Garden were indebted for various favors.6 Early in 1761 Garden wrote to Ellis: "Let the Loblolly be, as you desire, a Gordonia; if it be agreeable to you it makes me happy." Another year went by, and he again commented in a letter to Ellis: "I shall be happy to see the Loblolly Bay called Gordonia, in honour of our friend Mr. Gordon. It certainly is a new genus." One might suppose that after all of this Ellis would have confirmed its name with Linnaeus, but on November 19, 1764, the issue was raised again. Garden wrote to Ellis: "In one of your former letters to me, you desired that I would name the Loblolly Bay Gordonia, after Mr. Gordon at Mile End. I answered that I had no objection. However, if it is not done, you may, if you think proper, omit it, because I intend to put this and some others, into a little Fasciculus Stirpium Carolinensium Rariorum, or what the French call a Bouquet, which I may send into the world when the hurry of the day is a little over with me, which time I hope is approaching and almost at hand." Garden was thinking about retirement somewhat prematurely, at the age of thirtyfourl Unfortunately, he never published his little Fasciculus, and most of his rare Carolina plants have been credited to others. Two years later he again inquired of Ellis: "What have you called the Loblolly Bay? Please to name it." 7 None of Gordon's prodding seemed to arouse Ellis on the subject of Gordonia, although they corresponded about a variety of other matters, and Garden patiently inquired again on May 12, 1770: "What did you with the Loblolly Bay, which 6. June 7. ibid,

Ibid, 25 March 1759, p. 440, and postscript to letter, p. 442: ibid, 13 1760, p. 494. Ibid., "About January, 1761," p. 504; ibid., 26 February 1762, p. 517; p. 525; ibid, 2 June 1767, p. 555.

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The Naming of the Loblolly Bay you once intended to make a Gordonia? I think you should send it to Linnaeus, or if you choose the characters to be re-examined and drawn, I will endeavour to do them in the best manner I can, and send them to you." At long last his needling produced action. On September 25 of that year Ellis wrote to Linnaeus: "The Hypericum Lasianthus (Anglice, Loblolly Bay) is a new genus; the petals join in a tube at the base; besides the five ciliated subrotund foliola of the Calyx, there are constantly four bracteal leaves, like a calyx, under it, but not regularly placed, one of them being always lower than the rest. It is now going into flower; and I shall have an exact drawing made of it, with a description, which I shall send you, proposing, with my friend Dr. Garden, to have it called Gordonia, after our friend James Gordon.8 Ellis presented the characters of Gordonia to the Royal Society of London in November, and the following month sent them also to Linnaeus as promised, with the following comment: I am sorry I cannot oblige you in changing the name of Gordonia to Lasianthus as it has been presented to the Royal Society, and my worthy friend James Gordon has accepted this compliment from me: but I shall retain the trivial name of Lasianthus. For my part I do not know a greater proficient in the knowledge of plants, particularly their culture, nor so warm a friend of yours; for he always toasts your health, as the King of Botany, by the name of My Lord Linnaeus, and that before he drinks the King's health. His son has your books in his hands of tener than the Bible, and is now assisting a person here in translating your Genera into English.9 In January 1771 Ellis informed Garden that the Loblolly Bay was at last officially Gordonia. There is occasional mention of it in their later correspondence. Three points are noteworthy in connection with the official name. First, when Garden sent the seeds and plants in January 1756, he declared 8. Ibid, p. 577; ibid, 1770, pp. 250-251. 9. Ellis' account of the Gordonia was published as "A Copy of a Letter from John Ellis, Esq., F. R. S. to Dr. Linnaeus, F. R. S. &c. with the Figure and CharacteTs of that elegant American Evergreentree, called by the Gardeners, the Loblolly-Bay, taken from Blossoms blown near London, and shewing that it is not an Hibiscus, as Mr. Miller calls it; nor an Hypericum as Dr. Linnaeus supposes it; but an intire new Genus, to which Mr. Ellis gives the Name of Gordonia," in the Philosophical Transactions, 60 (1770), 518-530; 28 December 1770, Smith, Correspondence of Linnaeus, 1, 254.

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it to be a new genus and he sent the characters or description of it. Second, he later referred to Ellis having requested that "I would name the Loblolly Bay Gordonia." Third, and last, is that Ellis, writing to Linnaeus, said: "proposing, with my friend Dr. Garden [italics mine] to have it called Gordonia." Yet the name of the plant, like many others sent by Garden, has been solely credited to Ellis.10 10. Ibid, January 1771, pp. 582-583.

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The Historyof Embryologyas Intellectual History FREDERICK B. CHURCHILL Department of History and Philosophy of Science. Indiana University, Bloomington

Jacques Roger, Les Sciences de la vie dans la pense'e frantaise du XVIIIe siecle, la ge'neration des animaux de Descartes a l'Encyclopedie (Paris: Armand Collin, 1963). Howard B. Adelmann, Marcello Malpighi and the Evolution of Embryology, 5 vols. (Ithaca, N.Y.: Cornell University Press, 1966). Elizabeth Gasking, Investigations into Generation, 1651-1828 (Baltimore, Md.: Johns Hopkins Press, 1966). Jane M. Oppenheimer, Essays in the History of Embryology and Biology (Cambridge, Mass.: The M.I.T. Press, 1967). The Interpretation of Animal Form, Essays by Jeifries Wyman, Carl Gegenbauer, E. Ray Lankester, Henri Lacaze Duthiers, Wilhelm His and H. Newell Martin, 1868-1888, translations and introduction by William Coleman (New York and London: Johnson Reprint Corporation, 1967). Forty years ago, in the introductory chapter of his monumental work on Chemical Embryology, Joseph Needham paused for a moment to reflect upon the nature of the embryologist: It can hardly be a coincidence that so many among the great embryologists of the past were men of strongly philosophic minds. It would be absurd to support this opinion by citing Aristotle, but it holds less obviously true of William Harvey, whose book on generation is full of thoughts about causation, and in the cases of Ernst von Baer, Ernst Haeckel, Wilhelm Roux, Hans Driesch, d'Arcy Thompson, and J. W. Jenkinson, there is no doubt about it. It is not Journal of the History of Biology, Vol. 3, No. 1 (Spring 1970), pp. 155-181.

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really surprising, for of all the strange things in biology surely the most striking of all is the transmutation inside the developing egg, when in three weeks the white and the yolk give place to the animal with its tissues and organs, its batteries of enzymes and its delicately regulated endocrine system. This coming-to-be can hardly have failed to lead, in the minds of those most intimately acquainted with it, to thoughts of a metaphysical character.1 Today's embryologist would emphasize the genetic code and polypeptide synthesis in describing "the transmutation inside the developing egg," but he would unquestionably concur with Needham's characterization and, in fact, would add Needham himself to the list of "philosophic minds." This same philosophical bent has undoubtedly turned the embryologist to writing in the history of his field, and it is no accident that the four standard English surveys of the history of embryology were written by embryologists, or at least practicing zoologists.2 It is surprising, however, that professional historians of science, those renegade quasi-scientists who delight in the dance of ideas behind the scenery of specific scientific achievement, have been so slow to recognize the exciting philosphical implications and ramifications of the study of animal development. They have been particularly slow, in fact, considering that the general surveys are, one and all, limited in chronological scope and now seem handicapped with their historical perspective of a generation past. It is highly important, then, that in the last few years a number of new studies have appeared which focus on the history of embryology. It is even refreshing when the respective authors and editors hail from a range of disciplines, from professional embryologists to historians of science, to an intellectual historian. They all should be able to offer valuable perspectives to a field which is potentially boundless in scope. For this reviewer, outstanding among these works is Jacques Roger's Les Sciences de la vie dans la pensee frangaise du 1. Joseph Needham, Chemical Embryology (Cambridge, Eng.: Cambridge University Press, 1931), I, 7-8. 2. Francis Joseph Cole, Early Theories of Sexual Differentiation (Oxford: Clarendon Press, 1930); Arthur William Meyer, The Rise of Embryology (Stanford, Calif.: Stanford University Press, 1939); Joseph Needham, A History of Embryology (Cambridge, Eng.: Cambridge University Press, 1934); Edward Stuart Russell, Form and Function, A Contribution to the History of Animal Morphology (London: John Murray, 1916).

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Essay Review siecle.3 Despite the fact that this work is now nearly six years old, I feel it is absolutely essential to discuss it because it fashions a standard of incredible breadth against which all subsequent histories of the biological sciences must be measured. In fact, it must become one of the seminal works in the interpretation of eighteenth-century science. If further justification is necessary for leading first with this ace, it is because it has been neglected in the English literature, and two of the authors discussed later would certainly have been better prepared if they had taken notice of Roger's monumental efforts. At first glance, Roger's title appears somewhat deceiving: by implication it promises a survey of eighteenth-century French biology; yet the work gives much less. Moreover, in no way does the title warn the reader that he will plunge to the bottom of Aristotelian and Galenic theories of generation and Renaissance compromises and rise to the heights of seventeenthand eighteenth-century philosophy and theology before he gets to the 779th, or final page-in this respect, the work gives far more. The subtitle, La Ge'neration des animaux de Descartes a l'Encyclopedie, helps somewhat; and should the reader study the detailed table of contents (here the French publishers really excel), he should be prepared for the assault. After reflection, the title gains in meaning. The study of generation does, in fact, encompass the sciences of Life. It is the study of the origin of form, the meaning of vitality, the distinction between the inert and the animate, and the manner and moment of the appearance of the soul, as well as the more obvious questions, such as the possibility of spontaneous generation, the essence of sexuality, the mechanisms of regeneration, the significance of monstrous births and hybrids, the definition of species, and the coming-to-be of diversity. This is surely enough grist for a much larger list of "philosophic minds" than Needham has given. The text itself is divided into three major divisions: (1) "The End of the Renaissance (1600-1670)," (2) "The Philosophy of the Savants (1670-1745)," and (3) "The Science of the philosophes (1745-1770)." While the first of these sections serves to create a setting for the author's real objective -the eighteenth century, it also forms in its own right an excellent 150-page portrait of the medical and biological concerns of the seventeenth century.

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3. This work was the author's These principale de Doctorate 6s-lettres and won the Grand Prix Gobert de l'Acade'mie frangaise.

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During the waning of the Renaissance, the currents and countercurrents of biological thought were particularly tumultuous. The Aristotelian world picture was collapsing, and with it disappeared a successful solution of generation founded within the framework of the four Aristotelian causes. It was a time, too, when the medical profession, which had been revitalized a hundred years earlier by the humanist recovery of Galenic texts and subsequent advances in human anatomy, settled back once more into an inflexible educational system -indifferent to research and dominated by extraneous social demands. In an extremely effective chapter, "Le Combat contre les ombres," Roger explores a wide spectrum of scholastic and Galenic opinions about the contribution of each parent, the development of the embryo, and the matter of species, sex, and trait determination. The scholastics emphasized the role of the soul as the formal cause and on the importance of the male parent, whereas the Galenists sought refuge behind a battery of faculties and intermediary causes and gave the female parent equal determinative status with the male. Roger believes that scholastics were at their best when arguing on a logical level and that the physicians excelled at the factual and anatomical base. Roger insists, however, that neither school, nor any of the bewildering compromise positions, resorted to either of two extreme alternatives in explaining the most fundamental question, the origin of animal form. Neither side placed all the burden of the creative process on the sole action of the material or on a divine or spiritual act.4 Roger also examines other views that emerged in the seventeenth century, expressed by such men as van Helmont, Feyens, and Sennert, who insisted upon a spiritualized cosmos and upon the complete passivity of matter. The outstanding contribution of William Harvey to the field of generation rates a special study unto itself. Harvey excelled as an experimenter both in his De Motu Cordis and in his Generatione Animalium, yet Roger, unlike the surveys mentioned at the outset, does not waste effort trying to promote Harvey's modernity. Instead, he describes the curious blend of scholastic, Galenic, and spiritualist doctrines which determined Harvey's perspective. Finally, an examination of the revival of Hippocratic or atomistic solutions, refined by the eclectic hands of Gassendi, and the outright mechanistic solutions offered by Descartes conclude the array of studies of Part One. 4. Jacques Roger, Les Sciences de la vie, pp. 92-93.

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Essay Review By the time he reaches the description of Descartes' tourbillon vital the reader should be impressed by the over-all strategy of Roger's presentation. The author is not concerned about offering a collection of individual essays in a chronological sequence. He has chosen and blended each study to illustrate a much grander story than one would suspect from the isolated parts. From Aristotle's formal cause to Descartes' tourbillon vital, or from Harvey's punctam saliens to the atomist's mixing of parental matter, the primary and omnipresent question which the student of generation had to face was an explanation of the origin of animal form. This is the theme which Roger hammers home in a way which no earlier survey has done; it is a theme which binds together the innumerable embryological texts and gives them an importance far beyond the technical innovations or failures which superficially appear to be the primary story in the history of generation. The question of the origin of animal form confronts not just the embryologist as technician, but forces him to have at minimum certain implicit commitments about the cosmos. This would be as true today as in any of the yesteryears described by Roger. The origin of animal form as a fundamental problem, moreover, permits those who crusade primarily for certain explicit cosmic commitments to feel competent to issue judgments about the biological technicalities of generation. This happens, unfortunately, even in the twentieth century. In emphasizing so strongly the origin of animal form as the focus of other problems, Roger has drawn forth one of the most important issues in the history of science. The first part of his book fashions not only a background or portrait, but a preview of how the other parts of the volume treat the eighteenth century. Part Two surveys the period between 1670 and 1745-a time when the mechanical philosophy became the established framework for all the sciences. In France the Cartesian tradition of mechanism ruled supreme-a priori in its basic assertions and dogmatic about the exclusiveness of matter and motion as the principles of change. The Cartesian world picture also placed God and science in a particular relationship -God, the creator and legislator of natural laws; the scientist, the student of these laws, who paid little heed to the purpose of their creation. The biological sciences lagged somewhat behind the physical ones in entering this new framework, but by the beginning of the eighteenth century the Academie des Sciences encour-

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aged biological problems at its forum and the Paris Medical Faculty showed some flexibility toward new ideas. Anatomy profited from the mechanistic philosophy and the peculiar analogies which it engendered. At the same time, moreover, microscopy became the recognized frontier of biology, and it was this area of research which added a new dimension to the debates about generation. This added perspective, of course, included many new anatomical discoveries, outstanding among which were the presumed mammalian egg of de Graaf and the animalcules in the male semen. Equally important perhaps was a change in the aspirations of the scientist. "These animal machines [insects] are more complex, more admirable and made perhaps with greater skill than those which interest us the most and of which we have the greatest idea," declared Reaumur; and, echoed Malebranche, "One insect is more in touch with the divine wisdom than the whole of Greek and Roman history."5 These revelations held catastrophic implications for the eventual understanding of the origin of animal form, as the disillusioned Cartesian, Fontenelle, most eloquently stated: You say that animals as well as watches are machines? But place a machine-sire and a machine-bitch, one next to the other, and there may result from this a third, small machine; on the other hand two watches may be set next to one another their whole life without ever making a third watch.6 As men such as Fontenelle and Reaumur became increasingly aware of the intricate structures of the Lilliputian world, they found it increasingly hard to conceive of its generation in mechanistic terms-the biologist simply was left to gaze and to marvel. This particular trend in biology, Roger argues, reinforced and in turn was reinforced by a general movement toward deism in science. No longer did God appear as the legislator of simple mechanical laws but as the architect of an unbelievably complex universe. Where Descartes had felt confident about understanding the consequence of God's legislation, Reaumur, Fontenelle, Vallisneri, and others began to question the scientist's ability to comprehend the phenomena fully. A skepticism swept into fashion, and with it came one of the 5. Ibid., p. 237-238. 6. Ibid., p. 346.

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Essay Review most notorious explanations in all of the history of embryology -emboitment. This extreme form of preformationism is now so commonly ridiculed that its original appeal becomes lost. When viewed in juxtaposition with the triumphs and failures of the mechanical philosophy, the microscopic advancements and the rise of deism, Roger presents it as a persuasive doctrine. With it the problem of the orgin of all animal forms was relegated to a single creative act at the hands of God; the biologist literally renounced his intentions to solve any generative process more basic than the accretion of material-a simple task for the mechanics of the age. Part Three, "The Science of the Philosophes (1745-1770)," forms the obvious climax toward which the earlier two sections progressed. Since it comprises nearly a half of the whole text, there is an imbalance which reflects Roger's own deeper interest with the second half of the eighteenth century. The asymmetry is further accentuated, for Roger concentrates on the works of a few men rather than the common ideas of many; thus the reader will find a 60-page chapter devoted to Buffon and a 100-page chapter to Diderot. Despite the author's acknowledgment of this change of emphasis,7 this reviewer finds it a drawback in the structure of the work. How can the historian really compare two halves of a century when they have been treated so differently? In a fascinating opening chapter, entitled "Precursors and Sharp-shooters" (Francs-tireurs), Roger stresses the importance of the Newtonian and Leibnizian doctrines which by the 1740's had entered the scientific fray on the continent. In that Newtonian science considered attraction at a distance an acceptable explanation and in that the Leibnizian monads added the dimension of desire and perception to matter, both of these intellectual currents made it possible for embryologists to re-examine the generative capacity of matter. Nature did not have to be confined to the explanatory limitations of matter in motion, and all four savants whom Roger next introduces exploit these new possibilities. Maupertuis, as one of the earliest Newtonian physicists in France, turned his mathematical abilities toward analyzing polydactylism, a hereditary trait which could not be explained away in terms of emboitment. Maupertuis offered first the laws of attraction and then a material memory as possible explanations for this aberrant animal form. On the other hand, Turberville Needham, the 7. Ibid., p. 458.

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outstanding English microscopist known for his supposed demonstrations of spontaneous generation, developed a complex system of active and reactive forces combined with a sensible principle to account for all forms of generations. As a cleric, Needham had no intention of pre-empting the power of God, but as a serious biologist he brought nature back into consideration as a creative, organizing component. Le Mettrie and de Maillet argued that the laws of nature were fully capable in their own operation of explaining the origin of form. It is not surprising to find Buffon holding the center of attention in Roger's next chapter, for without question he was one of the pivotal naturalists of the eighteenth century. Roger himself is a Buffon scholar, having edited a critical edition of Buffon's Les Epoques de la nature and coedited a 60-page bibliography on Buffon.8 Buffon's theory of generation indicated the same traits as those examined in the previous chapter. By declaring a distinction between organic and inorganic molecules and by invoking the action of an internal mold and a penetrating force, Buffon returned the generative process to nature. The Newtonian and Leibnizian roots of his concepts are clear, and Buffon's microscopic examination of male semen (made in company with Needham) became his empirical starting point. The reader finds in Roger's study of Buffon, however, much more than an elaboration of just another theory of generation promulgated by the "science nouvelle." On the question of generation (and, in fact, throughout his L'Histoire naturelle) Buffon played as much the role of a philosopher as of a biologist. This inclination permits Roger to expand upon a second major theme which has occupied him since the outset and to which I have not yet alluded. The construction of any theory of generationupon a indeed, of all theories in the natural sciences-rests theory of knowledge. Such foundations manifested themselves in a number of subtle ways during the collapse of the Aristotelian world picture, notably in the form of a general skepticism about the constancy of nature's laws. The Cartesian system momentarily reestablished a confidence in the ability of a priori reasoning, but, according to Roger, biologists not only resorted to emboitment in 8. Les Epoques de la Nature. Edition critiques avec le manuscript, une introduction et des notes, ed. Jacques Roger, in Memoires du MusOum national d'histoire naturelle, n.s., ser. C (Sciences de la Terre), vol. X (Paris, 1962); "Bibliographie de Buffon," ed. E. Genet-Varcin and Jacques Roger, in Oeuvres philosophiques, ed. Jean Piveteau (Paris: Presses Universitaires de France, 1954) pp. 513-575.

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Essay Review order to enhance the specter of God's hand, but because of a skepticism about their ability to fathom the form problem. The preformationists not only relegated the generative act to a remote period at the beginning of life, but they banished it beyond the pale of scientific inquiry-it was no longer a fit subject for study. The next generation of investigators, including Maupertuis, Needham, Le Mettrie and Buffon, having employed Newtonian and Leibnizian weapons to broach once again the origin-of-form question, accorded to man the possibility of understanding generation insofar as he understood nature. Buffon himself was swayed by the English empiricist attitudes about human knowledge; by maintaining a relativistic rather than absolutist position toward scientific achievements, he appeared to keep within a skeptical tradition. His skepticism, however, was not a cry of despair. The world for Buffon was ordered, and much of it was accessible to man's reason: The theory of generation which Buffon proposes corresponds exactly to his theory of knowledge and to his concept of man. It allows the same ambiguities and raises the same problems, which still concern the relation of God and his creation. Anarchy and chance in the production of living beings which pure mechanism is not able to eliminate, the radical weakness of the human mind, the animality and the diversity of the human species are the joint ideas which Buffon rejects completely. Buffon believes in the order of nature, in the intelligence, the unity and superiority of man. But neither the order in nature nor human reason need find a guarantee in God. They exist from fact, not from right. We are inclined to believe that at this date such an attitude implies other things than scientific prudence and that Buffon in 1749 was practically an atheist.9 By repeatedly interjecting such epistemological questions into his study, Roger performs a task which few historians of science take on. He adds an essential dimension for a fuller history of science, since the history of scientific ideas depends in part upon the history of scientific methods and rationale. The chapter on Diderot is not only a full-scale study of the changing perspectives toward matters of life and generation of the great encyclopedist, but it reflects the changing attitudes which the biologists have experienced as well. The study of Diderot is executed with such detail, examining four different 9. Ibid., p. 558.

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stages of Diderot's career, that one wonders whether Roger's entire work did not grow out of this chapter. Diderot, prior to his Lettre sur les Aveugles (1749), reflected the sentimental deistic view typified in Reaumur-an awe in God's creation and a belief in preexistence. By 1749 Diderot recognized that preformation was simply a denial of an explanation and began to attack any recourse to a final cause. By 1753 Diderot began to utilize Buffon's opinions about generation, but it was not long before he went way beyond Buffon's provisional distinction between organic and inorganic molecules. Perhaps the most interesting pressure molding Diderot's intellectual development came from some lingering strains of medical spiritualism. Theophile de Bordeu, a professor at the medical school at Montpellier and a contributor to the Encyclopedie, had been malcontent with a deistic recourse as an explanation for animal form. His own philosophical roots lay with a medical tradition which drew its nourishment from Van Helmont, Glisson, and Stahl. It was undoubtedly through their influence that Bordeu placed such great stock in the explanatory potentialities of an animal "sensibility." In short, Bordeu-in fact, the Montpellier tradition-resorted to a vital principle, which Diderot found as a timely aid to the understanding of generation. After the completion of his Reve d'Alembert (1769) Roger finds Diderot resorting to a curious extreme. On the one hand, Diderot envisioned an Epicurean flux of unstable material forms, yet on the other he confided in a universal and pervasive sensibility to organize matter. What is important to remember while reading this chapter is that Roger describes the response of a well-informed philosopher to biological questions, not the formation of new scientific ideas. It is significant to find an important biological issue contributing so fundamentally to Diderot's development. The final chapter describes the resistance to the "science nouvelle." In the case of the Abbe de Lignac, this remained on a polemical level, but with von Haller, Bonnet, and Spallanzani there issued a very real scientific debate and a reformulation of preformationism in adjustment to the recent biological findings. It is unfortunate that Roger does not devote as much space to elaborating their positions as he did with Buffon's. Since none of the above trio was a Frenchman, this shortcoming perhaps reflects the limitations of a national history. Roger, however, is certainly not parochial, and he is able to make some valuable generalizations about these biologists which in fact support the

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Essay Review thesis he has constructed about eighteenth-century preformationism. These savants were far superior to their opponents of the "science nouvelle" in experimental and observational abilities; their empirical investigations, on the other hand, Roger argues, appear designed to confirm their convictions in preformation. As a group they discounted the possibility of epigenesis and spontaneous generation. They were mechanists but also devoted Christians; so neither a vitalistic nor Epicurean solution was tenable. No choice remained but to make God responsible for the impression of form onto matter. None of these later preformationists held to the extreme doctrine of preexistent miniatures nested within the loins of the first parents, but they all thought in terms of divinely created germs which bore the adult organization in some rudimentary and material way. In his portrayal of the eighteenth century, Roger describes a drift away from the tenets of preformation and the intellectual baggage which this seemingly innocuous biological theory entailed. At the scientific level this drift became obvious with the increasing popularity of secondary formative causes; on the philosophical level it revealed itself in the increasing awareness of the epistemological implications of scientific knowledge, and it was no accident that the philosophically inclined biologist found himself attracted to the "science nouvelle." As preformation came under attack, its deistic appeal suffered as well. Maupertuis and Needham accepted the existence of a final cause, but a belief in epigenesis and the creativeness of nature enticed Diderot, and finally d'Holbach, down the road to materialism. Roger has demonstrated how biological, philosophical, and theological arguments were inextricably bound to one another; his greatest triumph has been to show how history of science is very much a part of the history of ideas. The sheer bulk of Professor Adelmann's Marcello Malpighi and the Evolution of Embryology is likely to stagger the most inquisitive historian. Five large volumes compressing over 2000 exquisitely printed pages, this is a work which was created for the ages and which will obviously confront and even defy generations of scholars. It is a work which was severely criticized, with some justification, when it first appeared. The extent of its detail, the wearisome digressions both in the text and in the footnotes, the lengthy and occasionally inopportune quotations from easily accessible sources, led to bewilderment and then to frustration on the part of the hurried reader. But it is too important a work to be dismissed because of a lack of

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patience: It is a labor of love, by an embryologist about an embryologist, and if understood as such it can become an extremely useful historical tool. The first and largest of the volumes comprises the biography of Malpighi. Arranged in a chronological panorama and highlighting the major controversies and publications of Malpighi's career, this biography has been fashioned out of the author's close scrutiny of both the published record and the extensive Malpighi manuscripts. The biography is readable in spite of excessive details; it will remain invaluable because of them. A tabular account of Malpighi's life appended to the end would have helped the reader immensely to keep the whole forest in view and would have made it easier for future historians to locate desired particulars. I find the only really inappropriate chapter to be the discussion of the Studium of Bologna. To be sure, it is important to take account of the academic setting which created the extraordinarily intransigent Galenic opposition to the new sciences represented by Malpighi, and to do this effectively the reader must be made aware of the conservative nature of the European universities. Yet this hundred-page history of the Studium, completely derivative in nature, defeats the admirable objective. Adelmann's careful narrations about and extensive paraphrases and translations from Malpighi's microscopical works are perhaps the most valuable components of this first volume. From Malpighi's epistles on the structure of the lungs and his studies of the viscera to his incomparable monographs on the silkworm, the development of the chick, and the anatomy of plants, the reader is given a virtual history of anatomy up to and including Malpighi's contributions. The most interesting feature of all of these studies, however, is not the extraordinary structural details which Malpighi was able to capture through the microscope, but the mechanical rendering of their functions. Malpighi was truly a protege of Giovanni Alphonso Borelli and the Academia del Cimento at Pisa. For a time master and disciple complemented one another's talents as they both pursued a mechanistic reconstruction of the organism. Borelli, primarily a mathematician, introduced Malpighi to the Galilean and Democritean modes of thought, encouraged him in his dissections, and broadcast his discoveries. In return, the younger Malpighi, primarily a physician, helped Borelli through the complexities and anomalies of anatomy. Adelmann follows the interaction of these two men very closely; he reproduces a great deal of the correspondence between them which adds significantly to the

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Essay Review understanding of Malpighi's works, and he identifies Malpighi, along with other disciples of Borelli, such as Lorenzo Bellini and Carlo Fracassati, as a coterie of "neoterics" who endeavored to promote the new sciences and mechanical philosophy in Italy. Yet Adelmann fails to present a coherent picture of Malpighi and the new sciences; the reader must piece it together himself from Malpighi's works, from Borelli's comments, from Malpighi's voluminous correspondence with Henry Oldenburg, and finally from the acrimonious broadsides of his Galenic rivals-all of which can be found in their rightful chronological position in the volume. Since the reader continually stumbles across reminders and examples of Malpighi's mechanistic proclivities, it is worth reproducing several excerpts to indicate how the historian can piece together this fuller picture. These particular examples come from a discussion of Malpighi's research on glandular bodies: It is therefore probable that the juice separated in the surrounding follicle is conveyed through the tubular structures, as through excretory vessels, into the cavity. From their ends a network extends, the unequal spaces of which represent so many stigmata [the orifices of simple glands], so to speak, by which the expelled mucuous juice is first received and then led off into the cavity . . . Thus far it seems that Nature proceeds by a single, simple method: she attaches to an excretory vessel one or sometimes more membranous follicles or acini, and by means of them separates a peculiar humor from the vessels and, when it has been collected, expels it. I have elsewhere pointed out that Nature employs the same device in constructing the viscera too, when I announced that the liver, brain [!], and kidneys are glands; and to these we may add the mammary glands, the testes, and other similar parts . Malpighi next attempts to demonstrate that the pericardium, pleura, peritoneum, and tunica vaginalis testis are all glands. He believed this because when he squeezed them he found drops of fluid issuing from tiny orifices. The modern reader will be surprised by Malpighi's interpretation of the structure of the stomach, which, he regarded, in effect, as one large spread-out gland; he believed that the tubules, or fistulaethat is, the gastric glands themselves-are the excretory ducts of the single huge follicle, or locule, represented by the sinewy membrane (that is to say, the submucosa), and that the proventricular glands of the fowl are so many little stomachs,

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which are all "rounded up into a ball because of the narrow quarters." 10 Surely embodied here are interpretations of structure formed within the context of a iatromechanical, even iatrochemical, science. Here we find the success of the mechanical philosophy as it challenged the humoral physiology of attractions and faculties; yet at the same time we recognize the overextension of a particular mechanical analogy. Adelmann does not pursue such obvious implications, but the virtue of his text is that other historians can follow through if they choose. Another fascinating story which emerges from the text of this biography relates to the ferocity of the Galenic physicians as they attacked the new sciences in general and Malpighi in particular. Roger commented upon their reaction in his own account, and now Adelmann confirms it with some carefully detailed examples. Malpighi spent most of his career in Bologna, which appears to have been a veritable hive of medical conservatism. At times these attacks involved petty jealousies and personal ambitions, as seems to have been the case with Malpighi's former student, Paolo Mini. At other times, however, there were some basic issues at stake which are worth further exploration if the historian is to understand more fully the scientific revolution of the seventeenth century. Malpighi's life-long rival, Giovanni Sbaraglia, brought one issue brazenly to the bar with the comment: "The brain, for example. The more it is subjected to the anatomist's knife, the more obscure its use becomes. So many labors of the most distinguished men, so many observations on the cortex, the medulla, and the combinations of nerves have removed none of the obscurity, and so the physician cannot educe any certain remedies from the anatomy of the brain." 11 Here is the sign of a very deep gulf which, in fact, has divided the medical profession throughout history-the extent to which scientific research adds to a patient's well-being. This question alone is worth a volume, so it is a pity that Adelmann does not attempt to draw together an over-all evaluation of the Galenic opposition, for he could have focused further research on many such issues of general historical interest. These are left, however, scattered like broken ears amongst the sheaves. When the reader turns to the second volume he arrives at the real objective of Adelmann's efforts, the explication of 10. Howard Adelmann, Marcello Malpighi, I, 453-455. 11. Ibid., I, 561.

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Essay Review Malpighi's role in the evolution of embryology. In 1672 Malpighi communicated two essays on the development of the chick to the Royal Society of London, both of which appeared forthwith in the Philosophical Transactions. Adelmann has reproduced them together with their ten magnificent plates. He has added a translation from the Latin and provided an extremely detailed footnote commentary to run with the text. Only an experienced embryologist could have clarified Malpighi's descriptions in such a thorough fashion. Any historian who wants to understand what Malpighi saw and how his observations of specific anatomical details compared with those of other historical vintages will have to study Adelmann's comments with great care. The historian will be disappointed, however, to discover how little Malpighi discussed any general theory of development. This, after all, was the object of Roger's scrutiny as he built up a convincing case that the origin of form was an issue which touched very deep philosophical and theological chords. Adelmann comments on Malpighi's neutral presentation, too, and devotes much of the second volume to extrapolating Malpighi's implicit theory of development. Where Roger focused attention on the philosophical and theological impact of the origin-of-form question, Adelmann concentrates on a more traditional level. Was Malpighi a preformationist, as is so commonly assumed, or was he an epigenesist? Since this dichotomy is one of the most familiar yet most obscure problems in all of the history of embryology, it is worth examining Adelmann's analysis of the distinction. Epigenesis is the easiest side of the dichotomy to grasp. Harvey coined the term, and Adelmann implies that in its strictest sense epigenesis entails "a de novo production of heterogeneity out of homogeneity." 12 He adds that this "neoproduction" is usually construed as a sequential development of parts (Harvey's epigenesis) but might instead consist of a simultaneous development of parts (Harvey's metamorphosis). This definition is broad enough to accommodate the Aristotelian "potentially existent" form in the homogeneous mensis, Harvey's homogeneous colliquament of the chick egg and homogeneous material of the completed insect "pupa," and possibly Malpighi's colliquament in animal eggs and plant seeds. The catch phrase with Adelmann's definition is the modifier de novo, which is redundant if "heterogeneity out of 12. Ibid., II, 872.

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homogeneity" is to be taken literally; moreover, it appears to me to add confusion, since one can imagine a "de novo production" when a heterogeneous mass changes into a second heterogeneous mass. Preformationism is a much more difficult concept to define. Adelmann sees it logically arising "from the sound assumption that without the introduction of a deus ex machina the origin of an organized individual from chaos or from utter [?] homogeneity is inconceivable." 13 That Adelmann considers preformationism as the only alternative in Malpighi's day to epigenesis is likely but not certain, particularly since his argument that the modern explanation "incorporates the residual truths in both the old theories," 14 indicates that a compromise alternative now exists. That Adelmann then adds a "deus ex machina" in such a way as to make it appear as a necessary corollary to his earlier definition of epigenesis leads to unfortunate confusion. Certainly one would not want to relegate Buffon's moule inte'rieur or Needham's vegetative force to the status of ad hoc devices brought in solely for development, despite the fact that both these savants are customarily considered adherents of epigenesis. Having introduced some equivocations about this traditional dichotomy, Adelmann then distinguishes two extreme positions in preformation: (1) a particulate form in which the generative material of both parents is heterogeneous and comprised of particles derived from the parental parts; and (2) emboitment, or "preformation in miniature." Adelmann nevertheless adds that "the premise common to all forms of [preformation] is that there is no production or generation de novo during de-

velopment."15 Should the reader collect all these notions under two rubrics, he would find that epigenesis implied a homogeneous germinal material producing de novo diversification under the influence of a deus ex machina, and that preformation implied the opposite characteristics. Since Malpighi nowhere subscribed to either extremes of preformation, since he clearly ascribed to a neo-production in chick eggs, and since he called upon a "plastic spirit" to organize the homogeneous colliquament,l6 Adelmann concludes that "Malpighi's provisional theory is thus rather a theory of epigenesis." 17 This is a revisionist 13. 14. 15. 16. 17.

Ibid., Ibid., Ibid., Ibid., Ibid.,

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875. 884-885. 875. 866. 885.

Essay Review interpretation, but Adelmann gains an advantage by showing that the statements used in the past to support Malpighi's supposed preformationist leanings refer not to prefecundated eggs but to the fertilized though pre-incubated eggs after the neo-production could already have taken place. My objection to Adelmann's presentation is not to his conclusions about Malpighi, but to his incomplete analysis of the preformation-epigenesis dichotomy. As this analysis now stands, either there must exist a third clearly distinct alternative or Adelmann's definitions must be revised. After all, given the generally accepted embryological sense of de novo, there certainly can be a "neo-production" starting from a heterogeneous and highly organized substrate. Moreover, aside from revising the exclusive character of the definitions of this awkward dichotomy, it takes little ingenuity to recognize that a thorough analysis must tangle with those intractable questions about the meaning of "novelty," "emergence," "comingto-be," and "form." This in turn means that the historian of embryology must master the current and technical philosophical literature on these matters, to say nothing of the equivalent literature written during the epoch subjected to his historical scrutiny. As the reader works his way through the historical surveys which lead up to Malpighi's contributions, he begins to sense another very marked difference between the approach of Adelmann and that of Roger. Where Roger emphasized the philosophical pressures which influenced the development of scientific theories, Adelmann concentrates on the empirical side of science. This is his obvious concern when he cites Hippocrates' entreaty for systematic observations on the developing chick and then laments that "no one took the advice of the Hippocratic author for over two millenniums.""8 When he gets deep into the Renaissance, Adelmann judges the value of embryological work in terms of the extent to which each author contributed new and reliable observations about development. Thus he can conclude a very brief discussion about competing theories of sex with the following revealing comment: We should gain little from continuing this analysis of the unprofitable discussion of worn-out issues. The important fact is that none of these men followed the Hippocratic injunction to investigate systematically at close intervals 18. Ibid., 747-748.

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the development of the chick, and none had investigated methodically the whole progress of development in any other animal.19 Such a criticism of sixteenth- and seventeenth-century embryology is unquestionably valid, and Adelmann is in a stronger position than most historians to pass such a judgment. But to summarily reject the subject is to leave a fragmented picture. Compare Adelmann's brief dismissal with the brilliant article by Robert Herrlinger and Edith Feiner entitled "Why did Vesalius not Discover the Fallopian Tubes?"20 In a scant six pages these authors persuasively demonstrate that aparticular anatomical discovery depended upon a functional interpretation which rested on a time-honored philosophical point of view -in this case the analogy between reproductive organs and the superior status of the male sex. Herrlinger and Feiner have shown not only how much can be added to our understanding of factual discoveries, but also that with careful scrutiny we certainly can gain from the analysis of "worn-out-issues." More than missed opportunities, however, are at stake when Adelmann emphasizes the factual revelations in Malpighi's embryological texts. There is no question that these two texts are paragons of empirical accomplishment. With his singleminded concentration on the description of development, Malpighi, in fact, appears to have followed Hippocrates' injunction. Yet one cannot help read Malpighi's text without noticing some of his recurring metaphors: "rivulets of colliquament," "restraining zones," "embankments," vascular networks of different-sized "meshes," "mingled nutritive and fermentative juices," and so on. Though such metaphors, even collectively, do not add up to any clear mechanical model of development, knowing Malpighi's earlier use of such models (e.g., the agitative function of the capillary network in the lungs), his relation with Borelli, and his commitment to the mechanical philosophy, the reader surely might wonder whether there were not some philosophical presumption shaping these seemingly neutral observations. This suspicion appears all the more probable when the reader turns to the Opera posthuma where, again referring to chick development, Malpighi describes Nature's plan as delineating "pens" of solid material in which "pores"or "glandular sieves" "separate(s) . . . fluid from fluid." 21 19. Ibid., 755. 20. Medical History, 8: 335-41, 1964. 21. Adelmann, II, 866-867.

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Essay Review It may turn out that the mechanical philosophy contributed not one whit to Malpighi's two great embryological dissertations, but this we will only ascertain after an explicit study. Adelmann unfortunately holds a concept of science which precludes such historical considerations. "The progress of science," he reports elsewhere, "consists essentially in the accumulation of new data arrived at by progressively more accurate observation going hand in hand with the rejection of erroneous earlier interpretations of the data and an attempt at revaluation." 22 It seems to me that the most remarkable contribution by Adelmann is contained in the last three volumes. Here the reader will find twenty-eight "Excursuses," each of which traces the observations and reflections of the outstanding embryologists prior to the twentieth century as they pass judgment on particular anatomical features. For example, there are 200 pages surveying the literature on the embryonic heart from Aristotle through Harvey to Wolff, von Baer, and the elder His. Adelmann has supplied extensive translations from Latin, French, and German, and has included his own running commentary on the structures revealed in these texts. All of these "Excursuses" focus on references in Malpighi's two dissertations, but the author's intention appears to be twofold: first to make available the classical literature covering these particular subjects to the humanistically inclined embryologist and the historian of science; second, to put each of these texts (but particularly Malpighi's) into the perspective of the whole development of descriptive embryology. The author's metaphor, used to elucidate his intentions, reveals once again his concept of the development of science. "But to return to the excursuses," he writes in his Note to the Reader, "in the longer ones the gradual clarification that came with time may be followed step by step, and the effect is as if the microscope of time were at first out of focus, or its lenses defective, sometimes badly so, and as if with improved lenses and continual adjustment the details were brought into ever-clearer resolution," 2"3-again, the lineal ascent of science with the emphasis on factual clarification. But let us pause for a moment, we intellectual historians, before we assert too surely our chosen way! Adelmann, with these "Excursuses" ranging from observations on the structures in the unincubated egg, to the embryonic organs, to extra-embryonic membranes, has provided another kind of 22. Ibid., I, 499. 23. Ibid., xxiv.

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synthesis. Here are three volumes of source material on specialized subjects, complete with original texts, translations, and exegeses. Without the guidance of a trained embryologist, classicist, and archivist the intellectual historian would be hard put to make his way through a single one of these texts. In sum, they give an invaluable picture of observational techniques and advances from one historical period to the next. Only after mastering these details will we be able to write a fuller history of embryological concepts; with these "Excursuses" Adelmann has made this ultimate task feasible. I only wish that he had added "Excursuses" on the egg itself, the sperm, and the adult reproductive organs. The bibliography is extensive, but one wishes that it had not been limited simply to "Literature cited"; it would also have been more useful if Adelmann had divided it into specialized subjects. The index is even more extensive and more helpful. Considering the expense and labor involved in putting this formative work together, it is unfortunate that plates were not added to supplement the "Excursuses." With Elizabeth Gasking's Investigations into Generation, 16511828 the reader turns to a work of a very different caliber, produced for a general rather than professional audience. Essentially a series of essay-chapters written by a professional historian and philosopher of science, this short volume consists of thumbnail sketches of the major embryological theories from Harvey to von Baer. The book thus courses the same ground covered by Roger and Adelmann, yet the similarity ends abruptly. Gasking's essays are brief; they appear much in the style of lectures delivered before a college class, and perhaps for this reason are loosely argued and inadequately documented. Gasking has looked at the main classics in the field, but she leans heavily on excerpted translations or delves into the originals only to extract the most relevant items. Thus she ignores the strongly vitalistic side of Harvey, which permeates his De Generatione Animalium and which deserves comment if one is to argue as strongly as she does that Harvey represents "The Breakaway" from Aristotelian embryology. It is significant that she does not cite Walter Pagel's "William Harvey and the Purpose of Circulation," 4 which suggests the depth of Harvey's vitalistic leanings. Gasking very rightly 24. Isis, 42:22-39 (1951). Pagel's more extended analysis of Harvey's theory of generation found in William Harvey's Biological Ideas, Selected Aspects and Historical Background (Basel/New York: S. Karger, 1967) was, of course, not available to her.

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Essay Review points out Buffon's interest in scientific methods, but she leaves a confused description of Buffon's concepts of the moule interieur and organic molecules-a subject which Roger so expertly handled. Adding further confusion, she describes "structural units Iwhich] produce more units like themselves," a reference probably to entities quite different from the organic molecules. And how possibly can she leave wholly unexplicated the orphaned statement, "[Buffon] came as near to inferring the cell theory as was possible on the mere facts of regeneration"? 25 In her chapter on Wolff, Gasking makes some useful suggestions about Wolff's influence on the later Naturphilosophen, but the reader should be disturbed by the seepage of historical accuracy: Wolff did not "dedicate" his Theoria Generationis to von Haller as alleged's nor did he "translate" it into German.27 In this last assertion, Gasking probably meant Wolff's Theorie von der Generation, in zwei Abhandlungen erklart und beweisen (1764), which was an entirely altered and greatly extended volume.28 Gasking's chapter on von Baer appears to depend completely upon Meyer's introduction and translation of von Baer's "Commentar"29 and upon Hall's excerpts from T. H. Huxley's translation of the "fifth Scholion" from the first volume of von Baer's Entwickelungsgeschichte der Thiere.30 Because of her third-hand selection, Gasking misses the opportunity to employ von Baer's devastating criticism of preformation (translated, incidentally, by Aldemann) which would have restrained her from remarking at one point, "The dispute between the preformationists and epigenesists was to be resolved as soon as the cell theory was established." 31 A reading of E. S. Russell would have suggested that she might say something about von Baer's germ-layer theory and its connection with that hoary dispute. A familiarity with Jane M. Oppenheimer's essay on von Baer's mechanistic insights32 would have helped her 25. Elizabeth Gasking, Investigations, p. 89. 26. Ibid., p. 101. 27. Ibid., p. 98. 28. A facsimile edition of both works edited by Robert Herrlinger has recently been published (Hildesheim: Georg Olms, 1966). The useful "Introduction" contains valuable references to earlier secondary literature on Wolff, almost none of which was used by Gasking. 29. Arthur William Meyer, Human Generation, Conclusions of Burdach, Dollinger and von Baer (Stanford, Calif.: Stanford University Press, 1956). 30. Thomas S. Hall, A Source Book in Animal Biology (New York and London: Hafner Publishing Co., 1964). 31. Gasking, Investigations, p. 106. 32. "K. E. von Baer's beginning insights into causal-analytical relationships during development," Developmental Biology, 7:11-21 (1963).

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tone down Meyer's strong interpretation of von Baer's naturphilosophische leanings. But enough of this random sampling of the pudding! Gasking does present a unifying message for her fourteen chapters, and this is the most challenging aspect of her book. She starts out by explaining that the historian of science, if he is to understand the prevalence of certain types of explanations at certain periods, must take into account not only the "experimental facts," but the "general biological outlook" and the "general views . . . of science." 33 She herself considers all three factors, but concentrates on the "biological outlook," and it is significant in this respect that she frosts the opening and closing chapters with quotations from Thomas S. Kuhn's The Structure of Scientific Revolutions. In fact, Gasking's work appears to be a conscious attempt to understand the eighteenth-century theories of generation in terms of Kuhn's thesis of a prevailing paradigm. She wisely avoids that troublesome term, yet the apparatus is there: A theory gains in plausibility if it accords with the current world picture; but it cannot even be considered unless it is in accordance with recognized principles within its own field. Preformation was completely in accord with the central biological assumption of the time . . . This central assumption might be described as the law that the animals function because they have organs, for it assumed that all animals have organs, and that their essential vital activities were always associated with the appropriate organs.34 Here is an exciting notion, wrapped and ribboned for the intellectual historian. Gasking retums to it on numerous occasions and in different forms.35 It is this "central assumption" which promotes the eighteenth-century theories of preformation, and only when a new central assumption, the cell theory, replaces it, can further progress in embryological theories be forthcoming.36 How magnificently this fits Kuhn's message; yet Gasking unfortunately fails to establish her point. Aside from a somewhat distorted quotation from Aristotle and two sentences from the philosopher Malebranche,37 she brings forth no concrete evidence that vitality implied the possession of organs to the eighteenth-century biologist. She may indeed 33. 34. 35. 36. 37.

Gasking, Investigations, p. 14. Ibid., pp. 43-44. Ibid., pp. 82, 111-112, 119, 157. Ibid., p. 173. Ibid., pp. 44 46.

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Essay Review be right, but it is curious that neither Roger nor Adelmann mentions this pervasive assumption. It is a shame that this explicit test of Kuhn's thesis is not adequately documented. Jane M. Oppenheimer's Essays in the History of Embryology and Biology stands in marked contrast to the previous volume. Here is a collection of scholarly essays written by a professional embryologist over a span of twenty-five years. Although the last four essays deal with topics of the seventeenth and eighteenth centuries, the majority concentrate on the post1800 period, which not only has been rarely surveyed but has been poorly understood. All of these essays exhibit a thoughtful analysis of the primary sources, and a "Postscript" catches the reader up with the most recent secondary literature. It is a pleasure to have so many of Oppenheimer's essays together, because the reader has an opportunity to follow her development through the years as she has established herself as the most important historian of modern embryology. The oldest essay in the collection, "The Non-Specificity of the Germ-Layers,"38 written in 1940, is a chronological survey of the germ-layer doctrine and of its opponents. Her primary objective is to shed light for a modem evaluation and so she concludes: "As a result of all the experiments that have been enumerated . . . the doctrine of the absolute specificity of the germ-layers as enunciated in the last century must be abandoned." 39 In an essay written seven years later, "John Hunter, Sir Thomas Browne and the Experimental Method," Oppenheimer sets herself a very important historical question: Why did Thomas Browne in the seventeenth century differ so widely in his attitude toward experimentation from John Hunter in the late eighteenth century? She carefully examines Browne's view of science, but leaves the question begging, simply suggesting that the intervening years brought a change in intellectual climate and the appreciation of "an animal as a mechanism."40 By 1955, in an historical survey of the entire history of embryology, she weighs the interplay of the empirical and speculative aspects of science,41 and declares that "history has learned to traffic as profitably in 38. It is unfortunate that the publishers did not include her earliest foray into the history of science, "Historical Introduction to the Study of Teleostean Development," OSiTis 2: 124-148, 1936. 39. Oppenheimer, Essays, p. 286. 40. Ibid., p. 337. 41. "Analysis of Development: Problems, Concepts and Their History."

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ideas as in events." 42 In a pair of essays in 1959, she has become very much a part of that traffic as she wrestles with a number of worthwhile problems. "A scientist can proceed to his work only if he has in his mind certain assumptions," 4 she explains, as she sets about comparing some of the basic working assumptions in embryology of the pre- and post-Darwinian eras.44 In the second, and very important essay,45 written in 1959, Oppenheimer traces some of von Baer's conclusions about development as they were incorporated and distorted by Darwin. Her essays in the 1960's reveal this same interest in the impact of basic ideas on scientific events. Thus she relates Harrison's interest in bisymmetry to his perfection of tissue culture techniques,46 and elucidates the mechanically, mathematically, and cytologically oriented quests of Roux, Driesch, and Boveri, respectively.47 It is important to emphasize, however, that, while perfecting the art, Oppenheimer does not force her intellectual history. Her generalizations come naturally from the documents she has examined with care and professional acumen. One final work deserves mention in this Essay Review. Although The Interpretation of Animal Form represents a collection of essays written between 1868 and 1888, theirs is a timely resurrection for the history of embryology. The editor, William Coleman, a historian of science by training, has selected six essays (translating two of them) to illustrate the variety of intellectual pressures which, during the post-Darwinian period, besieged the investigator of the origin of form. Coleman has included a useful introduction which explains the intellectual setting of each selection and refers to a number of useful bibliographical items. The volume in size and scope supplements an earlier selection of readings edited by Benjamin H. Willier and 42. "Embryological Concepts in the Twentieth Century," ibid., p. 48. 43. Ibid., p. 207. 44. "Embryology and Evolution: Nineteenth Century Hopes and Twentieth Century Realities." 45. "An Embryological Enigma in the Origin of Species." 46. "Ross Harrison's Contributions to Experimental Embryology." 47. "Questions Posed by Classical Descriptive and Experimental Embryology." Two of Oppenheimer's later essays, which were not incorporated in this collection, are worth noting: "The Growth and Development of Developmental Biology," in Major Problems in Developmental Biology (New York: Academic Press, Inc., 1967, pp. 1-27), and "Some Historical Relationships Between Teratology and Experimental Embryology," in Bull Hist. Med. 42:145-159 (1968).

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Essay Review Jane M. Oppenheimer48 which contains classical papers in experimental embryology written between 1888 and 1939. The editors of both these books have been guided by the conviction that a reader should examine a scientific paper in its entirety in order to understand the framework of the author's endeavors. Four of the six essays chosen by Coleman actually champion a particular investigative approach rather than offer an explicit explanation of form. Thus Carl Gegenbauer argued that comparative morphology "stands . . . in contrast but not in opposition to physiology, which solves other problems by other means." 49 Henri Lacaze Duthiers defended zoology as an experimental and explanatory science against the criticisms of Claude Bernard; Wilhiam His insisted on a mechanical analysis of development, while H. Newell Martin elaborated upon the sort of laboratory training a fin de siecle morphologist should receive. This is a curious selection, for the reader will assume from the title of the volume that he will find essays actually interpreting animal form. In fact, only the remaining two essays, one by Jefferies Wyman and the other by E. Ray Lankester, do this by showing the impact of Naturphilosophie and "recapitulation" theories on embryology. By choosing four essays that emphasize the methods and rationale of research, Coleman has pointed the historian of science in the very direction pursued by Roger. It is not enough for the historian to track down specific theories or metaphysical digressions; he must draw forth from his historical quarry a confession about his feelings toward the scientific estate. To observe, to experiment, to employ sister disciplines as implements of discovery, or to insist on one approach and to exclude another, tells much more than the empirical contents of a given piece of research. Such a confession tells what the scientist believes an explanation entails; it tells what the scientist conceives to be the boundaries of scientific knowledge. These essays and the many others like them are essential for the intellectual historian. Let us step back for a moment and make some general observations about the state of the history of embryology. It should first of all be clear that the current historical efforts are largely devoted to the seventeenth and eighteenth centuries and that there remains a relatively uncharted sea thereafter. Oppenheimer and Coleman, however, have brought in enough 48. Foundations of Experimental Embryology (Englewood Prentice-Hall, Inc., 1964). 49. William Coleman, Interpretation, p. 50.

Cliffs, N.J.;

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sightings to show what rewards the nineteenth and early twentieth centuries might offer. It should also be clear that the history of embryology contains some fundamental problems, the most basic of which is emphasized by Roger and implied by Coleman-namely, the origin of animal form. In the postDarwinian period this quest was given a new solution; yet, why was it that embryologists, such as Haeckel, Roux, Driesch, Delage, and even T. H. Morgan, were unappreciative of the role of natural selection in the "form problem"? There are other questions to be posed as well. The epigenesis-preformation dichotomy is mentioned by Roger, Adelmann, and Gasking, but none of them points out that the dilemma re-emerges at the end of the nineteenth century. What are the common and what are the unique features in these separate eras of debate?50 Roger explicitly, Adelmann with reference to the Galenics, Oppenheimer in a number of essays, and Coleman in his choice of material, all recognize the value of studying the justifications for particular scientific attitudes, but where does this lead us? It is obvious that specific approaches will set the scene for certain observations and certain consequent theories. More is needed, however. Not until we can show the actual emergence of a given type of theory from a given epistemology will we make the connection between philosophy and the history of ideas. Roger alone, in his analysis of Buffon and Diderot, approaches this goal. It need hardly be added that we must generate detailed studies on more of the major embryologists. To mention von Haller, von Baer, Lankester, Roux, and Spemann would be simply to single out the most obvious figures. A final word about the philosophic contents of embryology so extolled by Joseph Needham: I have used his characterization as an inducement for the intellectual historian and have measured the books under review principally by their involvement with the history of ideas. Thus I have seen Roger as setting the standard, Oppenheimer and Coleman as approaching the mark in lesser but important ways, Adelmann as missing the opportunities which he has amply fashioned, and Gasking as forcing scientific concepts without the substantive 50. Edward Stewart Russell, The Interpretation of Development and Heredity, A Study in Biological Method (Oxford: Clarendon Press, 1930), Ludwig von Bertalanffy, Modern Theories of Development, An Introduction to Theoretical Biology, trans. J. H. Woodger (New York, 1933; Harper, 1962), and J. H. Woodger, Biological Principles, A Critical Study (London: Kegan Paul, Trench, Truber & Co., Ltd., 1929) are the standard English sources for this later period.

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Essay Review backing. Intellectual history (and this is certainly a cloak of many hues) has captivated the imagination of the historian of science for the last fifteen years, so I make no apology for my bias. Should this, however, be the only objective for the history of embryology? To answer with an unguarded reflection, it is worth noting that only Adelmann and Oppenheimer, the two professional embryologists, have made any effort to say something about the techniques and institutions for embryological research.

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The J. H. B. Bookshelf JUDITH P. SWAZEY

Chambers, Robert. Vestiges of the Natural History of Creation. Introduction by Sir Gavin de Beer. New York: Humanities Press, 1969; 390 pp.; $7.50. A welcome reprint of the first edition (1844) of Chamber's highly controversial work, which should be read both as an important precursor of Darwinian evolution and, more broadly, as a fascinating document in the history of Victorian ideas. Chamber's book attracted both the curious and the hostile, for it contradicted current understanding of the Bible and the accepted tenets of natural theology. The anonymously published Vestiges went through twelve editions, and it was not until edition 12 (1884), thirteen years after Chamber's death, that his identity was revealed. Cutright, Paul R. Lewis and Clark: Pioneering Naturalists. Urbana, Ill.: University of Illinois Press, 1969; xiii + 506 pp., illus.; $12.50. Based upon field studies along the route of Lewis and Clark's 1804 expedition and years of research on all facets of their epic joumney, Cutright presents a detailed account of the exploration and its significance. The author writes from a naturalist's perspective, and surpasses by far previous accounts of the expedition's scientific and technical aspects and the impact of Lewis's and Clark's discoveries upon contemporary science and scientific institutions. Darwin, Charles. The Life and Letters of Charles Darwin. Edited by Francis Darwin. Sources of Science, No. 102. New York: Johnson Reprint Corporation, 1969; 3 vols., 395, 393, 418 pp.; $50.00. The Life and Letters remain one of the basic and fascinating sources for students of the Darwinian story. Fleming, Donald, and B. Bailyn, eds., The Intellectual Migration. Europe and America, 1930-1960. Cambridge, Mass.: Harvard University Press, 1969; 748 pp., ilus.; $12.95. A superb collection of 14 essays and memoirs, combining the best that can be offered by trained historians and by

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those who write their history from direct experience, depicting the impact of migration from Hitler's Europe on many strands of American intellectual life. Donald Fleming's often provocative history of the origins of the double helix concept, "Emigre Physicists and the Biological Revolution," should be read with interest by anyone concerned with the development of molecular biology. French, Roger K. Robert Whytt, The Soul and Medicine. London, England: The Wellcome Institute of the History of Medicine, 1969; 182 pp.; ?2.5S. Though he has long been cited as a significant figure in the history of neurophysiology and neurology, this monograph is the first detailed study of the eighteenth-century Scottish physician, Robert Whytt. In this most welcome study, Dr. French analyzes Whytt's important studies of reflex action, hydrocephalus, and nervous disease. He also considers at length the historical development, unifying role in Whytt's work, and the place in eighteenth-century physiology of the idea of the co-extensive soul. Fullmer, June Z. Sir Humphrey Davy's Published Works. Cambridge, Mass.: Harvard University Press, 1969; 112 pp.; $6.50. An annotated bibliography of writings by Davy, published during his lifetime and posthumously, including many papers not found in the 9-volume Collected Works. Professor Fullmer's cataloguing of translations, critical reviews, and reports of Davy's papers are a valuable inclusion for those interested in tracing the diffusion of scientific developments. Harrison, Ross Granville. Organization and Development of the Embryo. Edited by Sally Wilens. New Haven, Conn.: Yale University Press, 1969; xxiv + 290 pp., illus.; $15.00. R. G. Harrison's fundamental contributions to developmental biology, including the method of in vitro tissue culture, are a major component of the history of experimental embryology since the 1890's. Included in this handsomely prepared volume are the five papers on which Dr. Harrison based his 1949 Silliman Lectures at Yale, lectures whose publication were preempted by Harrison's death. His longtime associate, Miss Wilens, has skillfully interpolated excerpts from his Silliman Lecture notes with the papers, revised the bibliography prepared by Dr. Harrison, and added a complete bibliography of his published works. Herber, E. C., editor. Correspondence between Spencer Foulerton Baird and Louis Agassiz-Two Pioneer American Naturalists. Washington, D. C.: Smithsonian Institution Press,

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The J. H. B. Bookshelf 1963; 237 pp., illus.; $5.00. All the known correspondence between two of the leading naturalists and museum builders of the nineteenth century, arranged in chronological order and covering a wide range of scientific, business, and personal subjects. History of Science, vol. 7, 1968. Edited by A. C. Crombie and M. A. Hoskin. Cambridge, England: W. Heffer and Sons, Ltd., 1969; 148 pp.; 42s. History of Science is an annual review offering analyses of recent publications, discussions of prospective research topics, essay reviews, and notices of doctoral dissertations. The present volume includes a study by J. Schiller on "Physiology's Study for Independence in the First Half of the Nineteenth Century," and two essay reviews on the history of medicine. Lopez Pifiero, J. M. Medicina Historia Sociedad. Barcelona: Ariel, 1969; 343 pp., paperbound. An anthology of excerpts from medical classics, dating from Hammurabi's Code to the nineteenth century, prepared by the Professor of the History of Medicine at the University of Valencia. Medawar, Peter B. Induction and Intuition in Scientific Thought. Philadelphia, Pa.: Memoirs of the American Philosophical Society, vol. 75, 1969; ix + 62 pp.; $2.00. Medawar, Director of England's National Institute for Medical Research and the 1960 Nobel laureate in medicine, combines his outstanding scientific ability and experience with a gift for philosophical reflection in these three essays on scientific methodology. Holding that "what passes for scientific methodology is a misrepresentation of what scientists do or ought to do," Sir Peter explains what he feels is wrong with the traditional system of inductive reasoning. He then presents as an alternative scheme of how scientists think the "hypothetico-deductive" process, a blend of imaginative insight and critical appraisal. Olby, Robert C. Origins of Mendelism. New York: Schocken Books, 2d printing, 1967; 204 pp., illus.; $2.45 paperbound. A clear, concise, and often stimulating contribution to the history of genetics, presenting Mendel's work as "the culmination of a series of studies which began two centuries ago with the classic hybridisation experiments of Kolreuter." Olby's notes, together with appendices which offer extensive quotes from original source material, help make the book a fine blend of sound scholarship and good reading. Poynter, F. L. N., editor. Medicine and Science in the 1860's. Proceedings of the Sixth British Congress on the History of Medicine. London, England: Wellcome Institute of the History

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of Medicine, 1968; xiii + 324 pp.; ?2.5S. Seventeen papers of varying quality examine biomedical events in the decade 1859-1869, focusing on the application and influence of concepts and research in Britain. Topics include the influences of Darwin, Lister, Pasteur, and Virchow; developments in public health and occupational medicine; and the advent of statesupported medical care and research. Science News Yearbook. 1969/1970. Compiled and edited by Science Service. New York: Charles Scribner's Sons, 1969; xx + 434 pp., illus.; $9.95. The first in a projected series of annual reference works drawing upon material originally published in the weekly newsmagazine Science News. The book's nine divisions cover many facets of science and science policy and should prove pleasurable reading for a wide audience. Thompson, John V. Zoological Researches and Illustrations, 1828-1834. London: Society for the Bibliography of Natural History; vi + 110 pp., illus.; $6.00. The studies on marine biology by the amateur naturalist J. V. Thompson, writes Alwyne Wheeler in his introduction to this facsimile edition, "revolutionized certain aspects of zoological thought." Contemporary neglect of Thompson, Wheeler feels, is due "to the absorption of his discoveries into the mainstream of zoological knowledge, and to the quieting of the controversies which raged around his head . . . and to the scarcity of his principal publications." Veer, P.H.W.A. de. Leven en Werk van Hugo de Vries. Groningen, Netherlands: Wolters-Noordhoff, 1969; viii + 252 pp., illus.; paperbound. A handsomely illustrated, concise study of de Vries's life and his work in genetics and plant cell physiology, liberally utilizing extracts from de Vries's own writings. Weaver, Warren. Scene of Change. New York: Charles Scribner's Sons, 1970; 226 pp., ilius.; $7.50. Few men have participated more widely or intimately in the growth of American science and technology since World War II than Warren Weaver. His work has included administering over $100 million for scientific research as Director for Science of the Rockefeller Foundation and an official of the Sloan Foundation. His highly readable biography will be enjoyed by all students, present and future, of science and its shapers in twentieth-century America. Williams, Greer. The Plague Killers. New York: Charles Scribner's Sons, 1969; x + 345 pp.; $6.95. A fascinating account

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The J. H. B. Bookshelf of the emergence of public health work in the area of infectious disease during the early years of this century. Williams draws upon personal interviews and previously unpublished material in the Rockefeller Foundation archives to examine the origins, conduct, and impact of three of the International Health Commission's major campaigns, against hookworm, malaria, and yellow fever. Young, J. Z., and Tom Margerison, editors. From Molecule to Man. New York: Crown Publishers, 1969; 215 pp., illus.; $19.95. Over 400 superb illustrations and eight well-written chapters fulfill the editors' goal of giving the lay reader a grasp and appreciation of recent and prospective work in areas such as molecular biology, evolutionary and population studies, microbiology, behavioral studies, and medical research.

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