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The idea of immunity: Metchnikoff's metaphysics and science

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Abstract

  1. A. I., Tauber and L., Chernyak, From Metaphor to Theory. Metchnikoff and the Origin of Imaunology (New York: Oxford University Press, 1991); A. I. Tauber and L. Chernyak, “The Birth of Immunology: II. Metchnikoff and His Critics,” Cell. Immunol., 121 (1989), 447–473.

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  2. L., Chernyak and A. I., Tauber, “The Birth of Immunology: Metchnikoff, the Embryologist,” Cell. Immunol., 117 (1988), 218–233.

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  3. W., Bulloch, The History of Bacteriology (London: Oxford University Press, 1938); A. Castiglioni, A History of Medicine (New York: Knopf, 1947); W. D. Foster, A History of Medical Bacteriology and Immunology (London: Heinemann, 1970)

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  4. Max, Neuburger, Die Lehre von der Heilkraft der Natur in Wandel der Zeiten (Stuttgart: Ferdinand Enke, 1926; English trans. by Linn J. Boyd, New York, 1932).

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  5. Neuburger writes: “It states in the book, de natural hominis, by nature is to be understood the combination of the four cardinal humors. These constitute the nature of the body and through them it becomes sick and well. The body of man contains within itself blood and phlegm and two kinds of bile, that is, yellow and black. And these (four elements) constitute in him the nature of the body and because of them he is ill or well. In the same sense, mixture of the four fundamental constitutents of the body the word physis is also used in many places, yet not uniformly in the entire corpus Hippocraticum. Because of this, Galen states (in Hipp, de acutor, morb. victu., Comm. II. 31) that physis by Hippocrates signifies different things. ‘Accordingly, the word “physis” of itself signifies many things.’ Many times by ‘physis’ is to be understood the four qualities (warm, cold, moist, dry), many times the four cardinal humors in their combined effect. Occasionally it is the innate heat, the final cause of all natural effects. (Galen, de placit. Hipp. et Plat. Lib. VIII, cap 7.) In the book, de carnibus, one finds the famous citation borrowed from Heraclitus that warmth is the original basis of all things: ‘It is my opinion that what we call warmth is immortal and perceives and sees and hears and knows all, both that which is and that which is to be.’ ... ‘Natur’ is for Hippocrates at one time, Lawfulness, at another essence and substance; he extends the conception of the power even farther” (ibid. [1932], p. 7nn1,2).

  6. Neuburger writes: “The chief founder of the Stoic physical theology, Chrysippos, sought in the fourth book of this work ‘On Providence’ also to answer the question how the unspeakable misery of bodily distress and diseases could be reconciled with purposefulness in the cosmos, with providence: ‘whether the ills of man arise in accordance with nature.’ From Gellius, Noct. Atticae VIII (VI), 1, we learn the attempted solution of the question from the philosopher: It is in no way intended by the creative power which joins defects and ailments with so many excellencies and uses, the direct work of nature, but it yields itself only by inference, as the unavoidable secondary consequence. Because as for example nature occupies itself with the shaping of the human body, it requires the higher view and the most ultimate purposeful direction in union with its creative work so that the head will be composed of the most delicate and finest bones. But this viewpoint of higher purpose, useful direction, had a certain disadvantage directly as a result, since indeed the head possessed only a weak defense from the outside and relatively slight shocks and impacts against it show it to be very slightly resistant: ‘in a like manner also diseases and illnesses have been caused while health was being secured’” (ibid., p. 10n3).

  7. Rhazes, “A Treatise on the Small-Pox and Measles,” trans. W. A. Greenhill (London: Sydenham Soc., 1848).

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  8. Girolamo, Fracastoro, De contagione et contagiosis morbis et eorum curatione (1546), trans. W. C. Wright (New York: Putnam, 1930), pp. 60–63.

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  9. L., Pasteur, “Sur les maladies virulentes et en particulier sur la maladie appelée vulgairement cholera des poules,” Comp. Rend. Acad. Sci., 90 (1880), 239–248.

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  10. D. E., Salmon and T., Smith, “On a new Method of Producing Immunity from Contagious Diseases,” Proc. Biol. Soc. Wash., 3 (1884–86), 29–33.

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  11. E., Behring and S., Kitasato, “Ueber das Zustandekommen der Diphtherie-Immunitat und der Tetanus-Immunitat bei Thieren”, Deutsch. med. Wochenscher., 16 (1890), 1113–45.

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  12. Pasteur's musing on the topic of possible active self-protection apparently never surpassed this old metaphysical circle of self-supporting “nature”, which he expressed as “life prevents life”. On this topic Émile Duclaux wrote: “Pasteur, who in his heart was indifferent to theories and asked of them only that they suggest experiments to him, held for a long time a purely cellular conception of microbial disease. It was by a struggle between the red blood corpuscles and the bacteridium that he explained in 1878 the resistance of the living fowl to anthrax, and we see him at every instant, in that period, having recourse to vital resistance, and saying: ‘Among the lower forms of life, still more than in the higher species of plants and animals, life prevents life’. Again, it was this same sentiment which guided him in the experiments which we have seen him making, to prevent the development of the anthrax bacteridium by inoculating at the same time with some common bacteria...” (Pasteur: The History of a Mind, trans. E. F. Smith and F. Hedges [Philadelphia and London: W. B. Saunders, 1920], pp. 317).

  13. “The metaphorical idea of ‘warfare against disease’ waged by the forces of the body is not predominant in the Hippocratic-Galenic tradition, if indeed it occurs at all” (L. J., Rather, Addison and the White Corpuscles: An Aspect of Nineteenth-Century Biology [London: Wellcome Institute of the History of Medicine, 1972], p. 188). Elsewhere, Rather writes of the metaphorical war against disease waged by the forces of the human body: “The notion is, however, relatively new. It has little part in the Hippocratic-Galenic tradition.... The idea that disease is or is caused by something that can be ‘cast out’ is, of course, an ancient and ubiquitous one, but I am speaking here of official academic medicine in the West” (L. J. Rather, “On the Source and Development of Metaphorical Language in the History of Western Medicine”, in A Celebration of Medical History, ed. Lloyd G. Stevenson [Baltimore and London Johns Hopkins University Press, 1982], pp. 135–153, quotation on p. 142; see also Peter H. Niebyl's commentary on Rather's paper, on pp. 154–156).

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  14. William, Coleman, Biology in the Nineteenth Century (Cambridge: Cambridge University Press, 1971).

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  15. Rather, “On the Source and Development”: “Riches of metaphor and analogy abound in the medical literature of Germany throughout the Romantic period and far into the first half of the nineteenth century.... In 1845, long before the bacteriological era, Carl Heinrich Schultz (1795–1871), a many-sided naturalist and physician, continued—in what was by that time an outmoded fashion frowned on by scientific physicians such as Jocob Henle and Rudolf Virchow—to formulate much of his pathophysiology in terms of the metaphorical ‘battle against disease’ (Krankheitskampf) and ‘defensive processes’ (Wehrprozesse) of the body. The two key terms in Schultz's metaphorical construct are Heerde and Keim, respectively the ‘focus’ of the body and the ‘germ’ of the disease. The ‘germ’ is a hostile force of undetermined nature; it is not, of course, a bacterial organism....Heerde is the German word for ‘hearth’, the seat of warmth and life of the home, the sun of its little microcosm, a force in its own right” (pp. 143–144). Rather continues: “To physicians of the postromantic school of strict science such language was almost anathema, and its use was vigorously opposed by Henle, Virchow, Gabriel Andral, Rudolph Lotze, and many others” (p. 144). Rather cites Virchow: “We no longer regard the pus corpuscles as gendarmes ordered by the police state to escort over the border some foreigner or other who is not provided with a passport” (p. 145).

  16. “...Addison interpreted inflammation as a healing and protective response, mediated in some way by cell activities, on the part of the organism. As far as the first part of his thesis was concerned, it was well within the medical tradition and was no doubt shared by a large number of his colleagues in England. In Germany on the other hand, attempts to explain biological events in terms of ends and purposes—so-called teleological explanations—had fallen into disrepute among scientifically minded physicians. The essentially teleological interpretation of the inflammatory process as a ‘defense reaction’ had to a considerable extent come under this ban” (Rather, Addison and the White Corpuscles, p. 180) See also 15. above.

  17. For a full discussion, see Chernyak and Tauber, “Birth of Immunology”.

  18. E. Metchnikoff, Sorok Let Iskania Racional'nogo Mirovozzrenia (Moscow, 1913), p. 21.

  19. E., Metchnikoff, “Zakon Zhizni: Po Povodu Nekotorykh Proizvedenii gr. L. Tolstogo” Vestnik Evropy, 9 (1891), 228–260 (reprinted in Sorok Let Iskania, pp. 216–247, quotation on p. 226). In order to comprehend to what an extent Metchnikoff had deserted the optimism of Haeckel's “biogenetic law”, we might compare his opinion with Arthur Keith's summary of Haeckel's point of view, that the apes were but “abortive attempts at man-production” (Arthur Keith, The Construction of Man's Family Tree [London: Watts, 1934], p. 10). Concerning the introduction of juvenile features into adult descendants as a topic of the nineteenth-century discussions of the “biogenetic law”, see S. J. Gould, Ontogeny and Phylogeny (Cambridge, Mass.: Harvard University Press, 1977), pp. 177–184; here Gould cites Cope (1883) (“America's foremost recapitulationist”): “As these characters result from a fuller course of growth from the infant, it is evident that in these respects the apes are more fully developed than man. Man stops short in the development of the face, and is in so far more embryonic. The prominent forehead and reduced jaws of man are characters of ‘retardation” (p. 179). It is obvious how much more disharmonic and “monstrous” is Metchnikoff's vision of man.

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  20. E., Metchnikoff, “Mirovozzrenie i Medicina” [Weltanschauung and medicine], Vestnik Evropy, N1 (1910), 217–235 (reprinted in Sorok Let Iskania, pp. 274–291, quotation on p. 274).

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  21. Metchnikoff, Sorok Let Iskania, p. 21.

  22. E., Metchnikoff, “Vospitanie s Antropologicheskoi Tochki Zrenia” Vestnik Evropy, 1 (1871), 205–235 (reprinted in Sorok Let Iskania, pp. 23–48).

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  23. E., Metchnikoff, “Vozrast Vstuplenia v Brak: Antropologicheskii Ocherk”, Vestnik Evropy, 1 (1874), 232–283 (reprinted in Sorok Let Iskania pp. 48–98).

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  24. E., Metchnikoff, “Ocherk Vozzreniya na Chelovecheskuju Prirodu”, Vestnik Evropy, 4 (1877), 532–560 (reprinted in Sorok Let Iskania, pp. 99–121, quotation on p. 114).

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  25. E., Metchnikoff, “Bor'ba za Suschestvovanie v Obshirnom Smysle”, Vestnik Evropy, 7 (1878) 9–47; 8, 437–483, (reprinted in Sorok Let Iskania, pp. 122–200).

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  26. Metchnikoff, Sorok Let Iskania, p. 20.

  27. E., Metchnikoff, Immunity in Infective Diseases, trans. F. G. Binnie (Cambridge: Cambridge University Press, 1905; reissued New York: Johnson Reprint Corp., 1968); idem The Nature of Man: Studies in Optimistic Philosophy, trans. P. Chalmers Mitchell (New York and London: G. P. Putnam, 1903), pp. 1–16, 166–199; idem, The Prolongation of Life: Optimistic Studies, trans. P. C. Mitchell (London: Heinemann, 1907).

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  28. Metchnikoff, Sorok Let Iskania, p. 20.

  29. Ibid., p. 21.

  30. See Metchnikoff, Nature of Man.

  31. Metchnikoff, Prolongation of Life, pp. 183–187.

  32. See ibid., pp. 193–223.

  33. E. Metchnikoff, “Sovremennoe Sostojanie Nauki o Razvitii Zhivotnykh”, Zh. Minist. Narod. Prosvesch. (March 1869), 158–186 (reprinted in I. I. Metchnikoff, Academic Collection of Works [Moscow, 1953], II, 254–276 quotation on p. 256 [hereinafter cited as ACW])

  34. See Jane M., Oppenheimer, Essays in the History of Embryology and Biology (Cambridge, Mass. and London: MIT Press, 1967). While Oppenheimer dates the change in orientation to the 1870s, we note Darwin's influence in embryology by the mid-1860s.

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  35. For an assessment of Metchnikoff and Kovalevsky in this regard see, for example, B. A., Dogel and A. E., Gaisinovich, “Osnovnye Cherty Tvorchestva I. I. Metchnikova kak Biologa”, in I. I., Metchnikoff, Selected Biological Works (Moscow: Academy of Sciences, USSR, 1950), pp. 677–725 (hereinafter cited as SBW).

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  36. Obeying the temptation to reconstruct history “logically”, Metchnikoff wrote in his essay on Kovalevsky: “Fritz Mueller's little book served as the starting point for a great number of works in the history of lower animal development, among which the studies of Kovalevsky occupy first place” (E., Metchnikoff), “Alexander Onufrievich Kovalevsky: An Essay from the History of Science in Russia”, Vestnik Evropy N 12 (1902), 772–799; reprinted in Stranicy Vospominanii (Moscow: Academy of Sciences, USSR, 1946), pp. 14–443 and in ACW (Moscow, 1959), XIV, pp. 9–39, quotation on p. 13. A. E. Gaisinovich Commented on Metchnikoff's essay: “The question about the influence of F. Mueller's book on the beginning of Kovalevsky's embryological studies is quite unsettled. First of all, Metchnikoff himself points further (p. 21) that during his sojourn in Tuebingen, Kovalevsky drew up for himself an extensive plan of independent work. However, it is known that Kovalevsky was in Tuebingen in 1862 and returned to Russia in 1863. Mueller's book was published, apparently, in the beginning of 1864 (the introduction was written in South America on September 7, 1863). It is doubtful whether Kovalevsky could have known of it before his arrival at Naples, in October 1864. But even aside from these facts, the plan of Kovalevsky's works (taking just the choice of his objects of research) does not reveal any influence of Mueller's book. As is well known, Mueller based his opinions exclusively on the crustaceans. Meanwhile, Kovalevsky writes: ‘during my visit to Naples in 1864, my first concern was to study the history of Amphioxus' development’. And later Kovalevsky did not work on the embryology of crustaceans. The choice of his first research objects obviously reveals some other peculiar reason” (Stranicy Vospominanii, pp. 194–195). (Russian and German translations throughout the text are our own).

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  37. For a discussion of to what degree the scientific climate reflected the actual novelty of Darwin's explanatory principles, see Peter J., Bowler, The Non-Darwinian Revolution: Reinterpreting a Historical Myth (Baltimore: Johns Hopkins University Press, 1988).

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  38. E., Metchnikoff, “Ocherk Voprosa o Proiskhozhdenii Vidov”, Vestnik Evropy, 3 (1876), 68–136; 4, 715–747; 5, 117–149; 7, 158–197; 8, 567–606. Reprinted in SBW, pp. 7–238; quotation on p. 216.

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  39. K. E. von Bear, Ueber Entwicklungsgeschichte der Thiere: Beobachtung und Reflexion (Koenigsberg, 1828–37).

  40. Metchnikoff wrote in 1869, apparently not without some patriotic sentiments: “The first scientific basis of embryology, that is the history of animals' development, was founded by the former St. Petersburg academicians, Pander and Baer” (“Sovremennoe Sostojanie Nauki” [see n. 33 above], p. 254). Von Baer, Metchnikoff asserts here, was the true founder of comparative embryology.

  41. See Metchnikoff, “Sovremennoe Sostojanie” (above, n. 33); Oppenheimer, Essay.

  42. T. H., Huxley, “On Anatomy and Affinity of the Family of the Medusae,” Phil. Trans. Roy. Soc. London, 139 (1849), 413–434. Jane M. Oppenheimer writes that Huxley, as early as 1849, “had appreciated the fundamental relationship between the body-layers of invertebrates and the embryonic layers of vertebrates,” but in Huxley's work of 1869 (An Introduction to the Classification of Animals) he does not mention any relation between coelenterate ectoderm and entoderm on the one hand, and embryonic serous and mucous layers on the other; Oppenheimer concludes: “Yet twenty years later he and all other investigators were still waiting to utilize the generalization in any way, even for pedagogic reasons” (Essays, p. 269). As a generalization, the idea was well suited for its time and need not have awaited the Darwinian era for its utilization. The conclusion is confirmed by the support that Zaddach received from Huxley in the 1850s and 1860s. It seems as if Huxley's modesty in 1869 (in respect to the idea that he first expressed in 1849) is most easily explained by the studies of August Weismann and Metchnikoff, begun in 1864–65, which created a skepticism in respect to (1) the natural-philosophical form of the idea drawing a parallelism between ontogeny and phylogeny, and (2) the old intention to order, in a unilinear pattern, the animal realm. On the other hand, it is our historical prejudice in dealing with a pre-Darwinian idea concerning an affinity of organic forms that inclines us to consider the idea as surpassing its allotted time.

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  43. In one of his first papers (1866) on insect embryology, Metchnikoff noted how profoundly the research of the 1850s had been influenced by Étienne Geoffroy Sainte-Hilaire's teaching: “In those times they tried to follow, as strictly as possible, the analogy with typical vertebrate development. The difference in the positions of the yolk, the difference in the positions of the nervous system, they tried to explain by the correspondence of the back side of Vertebrata to the abdominal side of Arthropoda” (I., Metchnikoff, “Embryologische Studien an Insekten,” Z. wiss. Zool., 16, 1, [1866], pp. 389–390).

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  44. G. Zaddach, Untersuchungen ueber die Entwicklung und den Bau der Gliederthiere, Vol. I, Die Entwicklungen des Phryganiden-Eies (Berlin, 1854).

  45. T. H., Huxley, “On the Agamatic Reproduction and Morphology of Aphis,” Trans. Linn. Soc. London, 22 (1858), 193–220, 221–236; R. Leuckart, “Fortpflanzung und Entwicklung den Pupiparen und Beobachtung an Melophagus ovinus,” Abh. Nat. Gesell. Halle, 4 (1858), 145–226.

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  46. Metchnikoff, “Embryologische Studien an Insekten,” p. 390.

  47. A., Weismann, “Die Entwicklung der Diptera im Ei, nach Beobachtungen an Chironomus Speciosus, Musca vomitoria und Pulex canix,” Z. Wiss. Zool., 13 (1863), 107–220; idem, “Die nachembryonale Entwicklung der Musciden nach Beobachtungen an Musca vomitoria und Sarcophaga carnaria,” Z. Wiss. Zool., 14 (1864), 187–336; idem, Ueber die Entwicklung der Dipteren (Leipzig, 1864).

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  48. Metchnikoff, “Embryologische Studien an Insekten,” pp. 391–392. However, Metchnikoff's first research in embryology was immediately provoked by Nikolay Wagner's discovery of the phenomenon that was later defined by von Baer as “pedogenesis” (parthenogenetic reproduction by insect larvae structurally unable to copulate). (See. E., Metchnikoff, “Ueber die Entwicklung der Cecidomyenlarven aus Pseudoovum,” Arch. Naturgesch., 1 (1865), 304–310. See also von Baer's article — “About Professor Wagner's Discovery of Asexual Reproduction of Larvae,” Zapiski Akad. Nauk, 10, 1 [1866], Appendix, pp. 1–77, —and Metchnikoff's letter to von Baer published as “Issledovanija o Dvukrylykh Nasekomykh,” ibid., pp 78–84 [reprinted in ACW, II, 56–60].) Note that Metchnikoff began his embryological study with the phenomenon that later played a role in undermining the strict parallelism between ontogeny and phylogeny.

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  49. After Olga Metchnikoff's Life of Elie Metchnikoff (trans. E. R. Lankester; London: Constable, 1921), the story dominated Russian literature. See Dogel and Gaisinovich (SBW, pp. 693–694). A similar interpretation was made by Metchnikoff's protégé, Besredka (A. Besredka, Histoire d'une idée: l'Oeuvre de E. Metchnikoff [Paris: Masson, 1921]; trans. Abraham Rivenson and Rolf Gestreicher, The Story of an Idea: E. Metchnikoff's Work [Bend, Oreg.: Maverick, 1979]). Daniel P. Todes, in his comprehensive chapter on Metchnikoff (in Darwin without Malthus: The Struggle for Existence in Russian Evolutionary Thought [New York and Oxford: Oxford University Press, 1989], pp. 82–103), repeats the same version of Metchnikoff's inspiration as being due to the evolutionary hypothesis and Fritz Mueller's ideas, and in our previous paper (see above, n. 2) we are also somewhat guilty of uncritically accepting this version. Only by following Metchnikoff's development from his earliest work and by carefully reconstructing his concept of “disharmony” are we able to recognize his retrospective rationalization.

  50. Before the late 1860s, fixed microsections had not yet been developed, and in many cases Metchnikoff could observe only living larvae. Therefore, he was limited to studying the development of external structures, and only in early development could he observe the emergence of internal organs, as far as transparency allowed. This technical limit was the principal restriction in studying the role of embryonic layers in organogenesis. In fact, the very existence of invertebrate embryonic layers was unresolved for the same reason. See A. D. Nekrasov, “Raboty I. I. Mechnikova v Oblasti Embryologii,” in ACW, III, 401–437.

  51. E., Metchnikoff, “Embryologie des Scorpions,” Z. Wiss. Zool., 21 (1871), 204–232 (submitted to the journal in 1869; published as a separate book in 1870, Leipzig).

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  52. E. Metchnikoff, Dissertation for the master's degree in zoology, St. Petersburg, 1867 (reprinted in ACW, II, 145–177); idem, Dissertation for the doctorate in zoology, St. Petersburg, in Zapiski Akad. Nauk, 13, 1 (1868), Apendix 1, pp. 1–48. (Reprinted in ACW, II, 178–207).

  53. During this period, Metchnikoff did in fact obtain important results, and he was highly productive. Some specialists consider certain of his findings as epoch making — for example, “Studien ueber die Entwicklung der Echinodermen und Nemertinen” (Mem. Akad. Nauk, ser. 7, 14, 8 [1869], 1–73), in which all the basic problems of echinoderm development (enterocoelom, metamerism, metamorphosis) were first correctly formulated. Metchnikoff was the first (1869) to observe coelom development in adult animals without metamerism. He described the basic processes long before the Hertwigs' report in their Coelometeorie (1881). (See Nekrasov, “Raboty I. I. Mechnikova”. R. I. Belkin, “Embriologicheskie Issledovaniia I. I. Mechnikova v Ocenke ego Sovremennikov,” in ACW, III, 438–479.)

  54. Dogel and Gaisinovich state that while “... Weismann (in 1864) had failed in understanding the developmental stages of the embryonic layers in insects, [Metchnikoff in his studies of Hemiptera and Diptera] perfectly [italics ours] demonstrated the differentiation of the two primary embryonic layers” (SBW, p. 694). The assertion is a strange one, and the more so because in 1886 Metchnikoff himself, in recalling the events of twenty years before, wrote quite openly: “among Arthropoda I failed to discover with sufficient clarity the embryonic layers in insects” (see E. Metchnikoff, Embryologische Studien an Medusen: Ein Beitrag zur Genealogie der Primitivorgane [Vienna, 1886]; reprinted in Russian, SBW, pp271–472 [quotation on p. 421]). The two quotations are found in the same book, but the discrepancy can be explained (at least partially) by the impression created by Metchnikoff's own revised version of the events (see n. 49 above).

  55. Oppenheimer, Essays, p. 271.

  56. Ibid., p. 273.

  57. Ibid.

  58. Metchnikoff, “Sovremennoe Sostojanie,” p. 270.

  59. Ibid., p. 256.

  60. See Oppenheimer, Essays.

  61. Metchnikoff, “Sovremennoe Sostojanie,” p. 256 (italics in original).

  62. Fritz, Mueller, Fuer Darwin (Leipzig, 1864; trans. W. S. Dallas, London: John Murray, 1869); Ernst Haeckel, Generelle Morphologie der Organismen, vol. II (Berlin, 1866); Carl Semper, “Reisen in Archipel der Philippen,” in R. Bergh, Malacologische Untersuchungen, pt. 2, vol. I (Leipzig, 1868), chap. 5.

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  63. Mueller, Fuer Darwin (1869), pp. 120–121 (italics added).

  64. For instance, Metchnikoff noted that the jellyfish Aurelia has a long succession of transformations. By analogy with crustaceans, and according to Mueller's rules, the hypothesis predicts that this case might accurately reflect the genealogy of the species and its closest relatives. But such is not the case, for Pelagia noctiluca, a form closely related to Aurelia, develops in a totally different manner: Pelagia's ciliated larva quickly metamorphoses into a form similar to the adult. Again, the parasitic worm Aspidogaster hatches from the egg in a form similar to the adult, without demonstrating its genealogy, but the closely related Distoma demonstrates one of the most intricate patterns of individual development, with a complex reproductive cycle in which two asexual generations alternate with a sexual form.

  65. Metchnikoff offered the following example: The larvae of many turbellarians have life patterns similar to that of the adult forms. The larvae so closely resemble the highest ciliated infusorians that some researchers suggested a close affinity between these two groups. The “Darwinians” using these positive embryological data were eager to link all worms to the ciliated infusorians. But beside these turbellarians there are others (i.e., the order Nemertina) whose development is markedly dissimilar to that of the infusorians, revealing a more complicated succession. If the latter form of development is viewed as the authentic presentation of turbellarians history, then the development through the infusoria-like larva should be recognized as a result of reduction. But why, then, should the similarity between the infusoria-like larva and infusorians be ignored?

  66. See Metchnikoff, “Sovremennoe Sostojanie”; idem, “Ocherk Vo prosa” (above, n. 38).

  67. Metchnikoff, “Sovremennoe Sostojanie”, p. 260.

  68. Metchnikoff, “Sovremennoe Sostojanie”, pp. 258–259

  69. Being unwilling to use Haeckel's terms “phylogeny” and “ontogeny,” Metchnikoff refers correspondingly to “genealogy” and “individual development” (or “history of individual development”).

  70. Metchnikoff, “Sovremennoe Sostojanie,” p. 258.

  71. Ibid., p. 257.

  72. See above, n. 65.

  73. Metchnikoff, “Sovremennoe Sostojanie,” p. 261 Iitalics in original.

  74. See Bowler, Non-Darwinian Revolution (above, n. 37).

  75. Metchnikoff, “Ocherk Voprosa,” in SBW, p. 216.

  76. Metchnikoff, “Sovremennoe Sostojanie,” p. 261.

  77. Ibid., p. 261.

  78. It was von Baer himself who recommended that Metchnikoff submit his work “Embryologische Studien an Insecten” to the St. Petersburg Academy contest, whose prize was established by von Baer and carried his name. Metchnikoff and Kovalevsky won the competition and shared the prize. In his letter to Metchnikoff, von Baer wrote: “I rejoice over your energy and talent and hope that you will be a credit to your fatherland” (K. E. von Baer, letter to Metchnikoff, in Stranicy Vospominanii [above, n. 36], pp. 180–181).

  79. See Metchnikoff, “Sovremennoe Sostojanie.”

  80. Von Baer, Entwicklungsgeshichte (1828), p. 258.

  81. About von Baer's attitude toward the idea of evolution see Metchnikoff, “Ocherk Voprosa in SBW, pp. 63–71; and B. E., Raikov, Karl Ernst von Baer, 1792–1876: Sein Leben und sein Werk (Leipzig: J. A. Barth, 1968).

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  82. Gould, Ontogeny and Phylogeny (above, n. 19), pp. 69–70.

  83. Ibid., p. 73

  84. Ibid., pp. 73–74

  85. Merchnikoff's scientific career began with studies of protozoans, and then he turned to metazoans, studying (between 1865 and 1869) annelids (Polychaeta), Gastrotricha (he was the first to establish the class of lower worms), Turbellaria (Rhabdocoela, Triclada), roundworms, Myzostoma, crustaceans, mollusks, nemertine worms, trematodes, cestodes, insects, lower chordates (Balanoglossus), and Echinodermata, resulting in about thirty published works on the morphology and embryology of invertebrates. The same diversity of research can be noted in Metchnikoff's published papers during the next five years, until 1874.

  86. E., Metchnikoff, “Pozvonochnaja Teorija Cherepa,” Zapiski Novoross. Univ., 7 (1871), 3–20 (reprinted in ACW, I, 189–199).

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  87. Metchnikoff, Embryologische Studien an Medusen, in SBW, p. 423.

  88. Ibid., p. 455.

  89. E., Metchnikoff, “Untersuchungen ueber die intracellulaere Verdauung bei wirbellosen Thieren,” Arb. Zool. Inst. Univ. Wien, 5, 2 (1884), 141–168; reprinted in SBW pp. 239–270; quotation on p. 241.

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  90. E. Metchnikoff, Uchenie ob Organicheskikh Formakh, Osnovannoe na Teorii Prevraschenia Vidov (St. Petersburg, 1869); reprinted in ACW, IV, 23–107.

  91. “The more the [morphological] material is accumulated ... the more the lack is felt of a carefully thought-out plan which would give the possibility of reducing this multitude of unrelated facts to a unity, not to an abstract-scientific one, but to that which, as far as possible, would coincide with living reality. That is what Haeckel has undertaken relying on Darwin's theory” (ibid., pp. 23–24).

  92. E., Metchnikoff, “Studien ueber die Entwicklung der Medusen u. Siphonophoren,” Z. wiss. Zool., 24 (1874), 15–83, quotation on p. 38.

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  93. The immediate impetus, however, for Metchnikoff's critical attitude toward Haeckel's scientific style and strategy, was occasioned by a specific controversy related to a discrepancy between their respective descriptions of larval development in the calcareous sponges: see E., Haeckel, Die Kalkschwaemme, I (Berlin, 1872), 34, 216; and E. Metchnikoff, “Zur Entwicklung der Kalkschwaemme,” Z. wiss. Zool., 24 (1874), 1–4, where Metchnikoff commented about Haeckel's presentation: “It is easy to see the reason why these opinions are so radically different from my observations if we carefully look through the chapter concerning the history of development of the calcareous sponges (pp. 328–338). It turns out that Haeckel never actually observed the postembryonic development of sponges, but only surmised it a priori. Haeckel just concocted the metamorphosis (and concocted it unsuccessfully).” It is interesting to note here L. W. Buss's modern interpretation in light of later findings. (Evolution of Individuality [Princeton: Princeton University Press, 1987], p. 47)

  94. E., Metchnikoff, “Beitrage zur Morphologie der Spongien,” Z. wiss. Zool., 27 (1876), 275–286.

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  95. E., Metchnikoff, “O Pischevaritel'nykh Organakh Presnovodnykh Turbellarii,” Zapiski Novoross. Obsch. Estestv., 5, 1 (1877), 1–12 (reprinted in ACW, I, 252–259; quotation on p. 259).

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  96. As Metchnikoff noted, “Haeckel says himself that ‘the gastraea theory cannot exist without proof of the true homology of two primary embryonic layers in all intestinal animals’” (SBW, p. 233). Haeckel believed that the homology had been fully proved. In contrast, identification of the embryonic layers was for Metchnikoff a most difficult problem. For him, proof of the homology was not a means for reconstruction of the hypothetical metazoan ancestor, but he viewed such a reconstruction as a support in his main task of the identification of developmental structures. Hence, his criticism of the gastraea theory was directed, first of all, against Haeckel's assurance that the homology of the layers in different animals had been proved. Being unconvinced that different gastrulae were homologous, Metchnikoff did not see any reason to accept Haeckel's assumption that the differences in organization and formation of the gastrulae are the result of secondary adaptation and transformation of a single ancestral gastraea. All theoretical reasons (including the explicitly formulated objections to Haeckel's theory) that finally led to the parenchymella hypothesis already had been articulated by around 1876. Many of the most important observations that Metchnikoff laid as the basis of his hypothesis, were already completed in 1877–1879. The very term “parenchymella” appears for the first time no later than 1877.

  97. Metchnikoff writes: “In the early autumn of 1874 my colleague A. Kovalevsky obtained several specimens of Halisarca larvae closely related to H. dujardinii, which I previously called H. pontica. Because I had been studying the development of sponges for a long time, Professor Kovalevsky proposed that I join him in working with the material .... My friend's sharing [in the work] was much more important than my own as a secondary worker” (E., Metchnikoff, “Spongiologische Studien,” Z. wiss. Zool., 33 [1879], 349–387 [submitted 1878]), quotation on pp. 349–350.

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  98. Ibid., p.

  99. N. Lieberkuehn, “Beitraege zur Anatomie der Spongien,” Mueller's Arch., 1857, pp. 376–403.

  100. Metchnikoff, “Spongiologische Studien,” pp. 372–273.

  101. Ibid., p. 374.

  102. Ibid., pp. 377–378.

  103. Metchnikoff, “O Pischevaritel 'nykh Organakh Presnovodnykh Turbellarii”; E. Metchnikoff, “Issledovanie o Razvitii Planarii,” Zapiski Novoross. Obsch. Estestv., 5 (1877), 1–16 (reprinted in ACW, III, 51–60). In Metchnikoff's time, the term “planarians” was used for all turbellarians.

  104. Metchnikoff, “O Pischevaritel 'nykh,” p. 256.

  105. Ibid., p. 257.

  106. E., Metchnikoff, “Vergleichend-embryologische Studien. III. Ueber die Gastrula einiger Metazoen,” Z. wiss. Zool., 37 (1882), 286–313. In relation to this we note that Haeckel himself recognized his debt to Kovalevsky: “For me special value is presented by the outstanding studies of ontogeny of different lower animals published by A. Kovalevsky for the last seven years and which, in my opinion, are the most important and the most fruitful among all modern works in the field of ontogeny” (Mueller-Haeckel, The Basic Biogenetic Law [Moscow, 1940], p. 201). On the other hand, there is an apparent misunderstanding in the historical literature where A. Kovalevsky has been attested as Haeckel's disciple. Thus, in Erik Nordenskioeld's History of Biology (new York: Tudor 1928) we find: “Alexander Kowalewsky (1844–1901), an academician of St. Petersburg, worked in the spirit of Haeckel, encouraged by his commendation” (p. 529). The information is quite misleading (besides, we note that Kovalevsky was born in 1840, not 1844). See Metchnikoff's comparison of Haeckel and Kovalevsky as scientists in Stranicy Vospominanii (above, n. 36), pp. 30–33.

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  107. Kovalevsky attempted to create a theory that encompased a common formation for the embryonic layers in lower chordates (Amphioxus) and invertebrates. He believed that the invaginated gastrula was the stage of development of all animals possessing a segmentation cavity. He thought that the blastopore (the opening of the gastrula) in all gastrulae developed into the anus, but soon he found that this was true only in some cases, whereas in others, the blastopore developed into the mouth, and in yet other cases, the blastopore closed and the anus opened once again. The hypothesis of homology of all gastrulae, which Kovalevsky presented in 1866, was not again repeated — but Haeckel continued to invoke it as a firmly established fact. In order to reconcile these contradictory facts concerning the blastopore, Buetschli (O, Buetschli, “Entwicklungsgeschichtliche Beitraege,” Z. wiss. Zool., 29 [1877], 216–254), and after him Hatschek (B. Hatschek, “Beitraege zur Entwicklungsgeschichte u. Morphologie der Anneliden,” Wien Akad. Sitzungsber., 74 [1877], 443–461), proposed a new hypothesis: the primordial gastraea would have a round opening, which in further evolution developed from an oval form to a slit. This formation would allow the bifurcation of the slit into two openings by closure of its middle by joining opposite edges; one of these openings would then develop into the orifice and the other into the anal opening. The slit shape determined bilateral symmetry. In order to check the hypothesis, Metchnikoff attempted to study gastrulae of many different animal classes. He concluded that most of his observations would not support the hypothesis: e.g., both the sea urchin's anal gastrula and the annelid oral gastrula express radial symmetry, but the sea urchin mouth appears too far from the blastopore; in the nemertines, while the radial gastrula was retained, the transition to bilateral symmetry was conditioned not by the blastopore but by the development of the ectodermal sac; and so on.

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  108. “He [Haeckel] gave to Darwinism the shape in which the theory would be better recognized by nonspecialists, beside the fact that he had dressed it in the form of a natural-philosophical system especially pleasing to Germans. Intending to give the theory perfection and wholeness, he proposed the bravest hypothesis alongside scientific truths and, at last, lost any sense of the basic difference between the latter and the former” (Metchnikoff, “Ocherk Voprosa” [above, n. 38], in SBW, p. 228).

  109. Ibid, p. 230.

  110. Metchnikoff began to ponder the parenchymella hypothesis around 1876–1877. The publication in which he apparently first explicitly expressed his belief in the disharmonic nature of humans is “Upbringing from the Anthropological Point of View” (1871). Olga Metchnikoff (Life of Elie Metchnikoff, p. 63) states that he had been interested in the concept since 1868; as we have seen, around that time Metchnikoff formulated his doubts concerning traditional understandings of recapitulation.

  111. Metchnikoff, “Untersuchungen ueber die intracellulaere Verdauung,” in SBW, pp. 241–242.

  112. Ibid., p. 267.

  113. Metchnikoff, “Ocherk Vozzreniya” (above, n. 24), p. 114.

  114. Metchnikoff, “Zakon Zhizni,” in Sorok Let Iskania, p. 226.

  115. E. Metchnikoff, “Bor'ba za Suschestvovanie Chastei Zhivotnogo Organizma” [The struggle for existence of parts of the animal organism], in Pomosch Gologajuschim: Nauchno-Literaturnyi Sbornik (Moscow, 1892), pp. 321–326 (reprinted in ACW, IV, 364–374). Metchnikoff probably knew of Wilhelm Roux's treatise, Der Kampf der Theile im Organismus [The struggle of parts in an Organism], which had been published in 1881 (Leipzig). To what degree, then, was Metchnikoff's theory of the constituent struggle within the organism influenced by this book? As with many morphologists of his time, who in their discussion of evolutionary issues were interested not in adaptive variations of given structures but rather in evolutionary realization of the processes that led to formation of the structure, Metchnikoff's vision of variability was much more saltative than Darwin's vision of the phenomenon (see Metchnikoff “Ocherk Voprosa” [above, n. 38], in SBW, pp. 137–142). Contrary to Darwin, he believed that crossbreeding does not allow the preservation of small variations (ibid., pp. 139, 191). He supported Fleeming Jenkin (ibid., p. 182) and asserted: “In order to able to start to act, it is not sufficient for natural selection to have merely individual deviations; it is necessary to have a certain sum of individuals who have changed in advance in a similar way” (ibid., p. 183; Metchnikoff's italics). Correspondingly, Metchnikoff believed that the most important and bitter forms of struggle take place between different species, or between a species and its abiotic environment (ibid., p. 146). There is nothing original in this position, which is quite similar to those of Koelliker, Naegeli, Askenasy, and Mivart (all of whom are analyzed at length in the “Essay”). Metchnikoff's originality is in his reinterpretation of the recapitulation idea. From his point of view, recapitulations provide, not a clear-cut parallel between ontogeny and phylogeny, but ontogenic coexistence of different integrities pertaining to different stages of phylogeny. This coexistence creates the basic disharmony of individual development—and this disharmonic relationship Metchnikoff designates as a struggle between the parts of the organism. No semblance of this concept is encountered in Roux; the central thought of Der Kampf is orthodox Darwinism: struggle originates in individual variations. The metaphysical basis of Roux's book is formed by the trivial assumption that there are no two entities that would be equal in every respect; the basis of the struggle between the organism's parts is the principal inequality of those constituents (the “molecules,” the cells, the organs) (Der Kampf der Theile, p. 69). The relation between unequal components then turns into a struggle when the inequality pertains to vital aspects of the organism's life (ibid., p. 67). Roux was apparently comfortable with the fact that, having intended to explain what is vital to the organism in terms of struggle, he then offered an explanation of struggle in terms of vitality. This is not surprising because, as Roux frequently states in Der Kampf, he followed Heraklit in interpreting the struggle concept (“The struggle is the father of everything”), and he believed that Darwin and Wallace simply continued in the same metaphysical tradition (ibid., p. 65).

  116. About Metchnikoff's approach to Darwinism see I. M. Polyakov, “Razrabotka Osnovnykh Problem Darvinizma v Trudakh I. I Mechnikova,” in ACW, IV, 409–468; Todes, Darwin without Malthus (above, n. 49); Daniel P. Todes, “Darwin's Malthusian Metaphor and Russian Evolutionary Thought, 1859–1917,” Isis, 78 (1987), 537–551. Todes follows Olga Metchnikoff in his assertion that Metchnikoff's initial attitude toward zoology and comparative embryology was determined by the goal of establishing evolutionary relations. We have endeavored to demonstrate that the issue was otherwise, and that Metchnikoff in the first years of his embryological career was a strong proponent of the independence of embryological research from evolutionary problematics. From Todes's point of view, Metchnikoff is a typical representative of those Russians who, in comparing the importance of evolutionary factors, paid much more attention to the interspecific struggle than to the struggle between individuals of the same species. While completely true, this interpretation is incomplete. What price did Metchnikoff pay to support this specific attitude? He dipped the interspecies struggle into the individual organism itself and actually considered, as the most important form, the struggle between parts of the same organism. Arguing for the primacy of the interspecies struggle, Metchnikoff made out of it the very soul of individuality. To establish peace among individuals of one species, he proclaimed a war between parts of each individual organism, the war of the individual against itself.

  117. See Tauber and Chernyak, “Birth of Immunology: II” (above, n. 1).

  118. About the varieties of vitalism in the nineteenth century see E., Benton, “Vitalism in Nineteenth-Century Scientific Thought: A Typology and Reassessment,” Stud. Hist. Phil. Sci., 5 (1974), 17–48. About nineteenth-century teleology as a complement to the new reductionist approach see Timothy Lenoir, The Strategy of Life: Teleology and Mechanism in Nineteench-Century German Biology (Dordrecht: D. Reidel, 1982; republished Chicago: University of Chicago Press, 1989).

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  119. Metchnikoff was happily shocked by the idea that the forces of disharmonies (those very forces which were supposed to destroy any other structure as “the other”) could play a harmonizing role. The appeal was not in recognizing a logically developed idea, but rather in discovering a fact (which did not preclude appreciating the logic of the discovery). He did not expect that the disharmonic role of intracellular digestion in higher metazoans must be subjected to a reconciliating evolutionary necessity; he believed that he simply had discovered one fact of such a subjection. As a person who accepted his scientific problems as existential questions of his own life, Metchnikoff developed his pessimistic weltanschauung in an essential degree from his biologically formulated idea of disharmony. Correspondingly, he saw the possibility for an optimistic change in his position, not in a reformulation of metaphysical assumptions, but in what he viewed as the happy end of one evolutionary controversy—adaptation in higher metazoans of the digestive activity of the mesodermic cells.

  120. Neither Darwin himself, nor the theories of the putative Darwinians of the nineteenth century were completely free from traditional teleology; see Bowler, Non-Darwinian Revolution (above, n. 37). An extensive literature deals with the problem of teleology in the modern synthetic theory of evolution, whose teleological nature may be allegedly concealed in some of its (presumably) tautological assumptions. There is a well-supported opinion that the teleological approach to evolution is rooted in the analogy of evolution with individual growth (Bowler, op. cit.). However, this opinion may be correct only to that degree in which the analogy is based on an intuitive assumption that individual development must be of a teleological nature; only when individual development is considered as a realization of a “plan,” can analogy with evolution make the latter teleological. Note that Metchnikoff's view of ontogeny as competition between different types of cells and parts of the organism is basically inconsistent with traditional teleology. In response to Baumgarten's accusations of the phagocytosis theory in teleology (P. Baumgarten, “Referat” Berliner klin. Wochenschr., 21 [1884], 802, 818; discussed in Tauber and Chernyak, “Birth of Immunology:II”), Metchnikoff argued that the theory had nothing in common with teleology and had been completely built “upon the principles of the evolutionary theory” (E., Metchnikoff, “Ueber den Kampf der Zellen gegen Erysipolkokken: Ein Beitrag zur Phagocytenlehre,” Vischow's Archiv., 107 [1887], 209–249). In Baumgarten's next critical paper (“Zur Kritik der Metchnikoff' schen Phagocytentheorie,” Z. klin. Med., 15 [1889], 1–41), he responded ironically to this argument: because of Darwinism's spell, everything that refers itself to that teaching obtains meaning solely due to that reference. However, Metchnikoff consistently assumed that defensive strategy, referring to his own interpretation of “the principles of the evolutionary theory” and the “struggle” as applied to his own understanding of the problem of the organism's integrity. At that time, he did not invoke “Darwinism” in his evolutionary approach.

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  121. Buss, in The Evolution of Individuality (above, n. 93, p. 65), clearly states the issue, arguing that metazoan ontogeny is a sequence of cell lineages progressively denying their own capacity to increase for the collective interest of the individual. This notion is closely related to Metchnikoff's original observations, which were based on the conflict between the potentially opposing processes of various somatic elements and organismal integrity. The existence of harmonious function in favor of the individual (and total) organism in Metchnikoff's original terms is a result of competition between disharmonic parts of the organism. How have organisms evolved so that some cells have abandoned their own capacity to replicate? The strategies are complex, but generally, patterns in cleavage and regulation are adaptations that serve the function of imposing selection at the level of the cell lineage. What Metchnikoff recognized, albeit it in a manner restricted by the then-current level of knowledge, was that evolution must be understood as selective processes that operate on the interactions of cell lineages. As Buss puts it: “Evolutionary pattern has arisen not by selection on individuals alone, but by the interactive effects of selection operating at differing levels of biological organization” (p. 68). See also L. Margulis, Symbiosis and Cell Evolution (San Francisco: Freeman, 1981), where the same general issue is applied to cellular evolution.

  122. There is no real definition of orthobiosis in Metchnikoff's writing. Orthobiosis as an “ideal” way of life is not actually an “ideal,” nor an established or hypothetical norm, but rather a dual definition to (1) continue studies in inherited disharmonies, and (2) elaborate the process of an organism's struggle to resolve disharmony.

  123. It is natural that Metchnikoff did not turn initially to an observation of spontaneous phagocytosis in diseases (which was indeed his following step), but immediately made observations on the role of amoeboid mesodermal cells in embryological development (atrophy and “physiological inflammation”) — the transformation of echinoderm larvae and tadpole metamorphosis (see O. Metchnikoff, Life of Elie Metchnikoff, pp. 120–121). In the echinoderm studies Metchnikoff demonstrated that phagocytic cells were mobilized to eliminate atrophic larval structures, and in the tadpole, phagocytes with the same function arose not from blood, but from the muscular tissue; again, mesodermal diversity revealed primordial function. These studies data from 1892 (see E. Metchnkoff and J. Soudakewitch, “La phagocytose musculaire. Contribution aH94 l'étude de l'inflammation parenchymateuse,” Ann. Inst. Pasteur, 6, 1 [1892]; E. Metchnikoff, “Ueber Muskelphagocytose,” Zentralbl. Bakter., 12 [1892], 294–296).

  124. As early as 1884, Metchnikoff had characterized his immunological approach as comparative-pathological. When first dealing with the phagocyte response in models of infectious diseases, the studies were oriented by the comparative construction (see E., Metchnikoff, “Ueber eine Sprosspilzkrankheit der Daphnien: Beitrag zur Lehre ueber den Kampf der Phagocyten gegen Krankheitserreger,” Vischow's Archiv., 96 [1884], 177–195). Starting with the belief that the general immune response is “a diffuse inflammation,” he began his studies of the phenomenon with Daphnia and then in the next work continued in succession with frogs, lizards, turtles, guinea pigs, and rabbits (see our discussion in Tauber and Chernyak, “Birth of Immunology: II”) We have seen that the problem of the digestive activity of amoeboid cell recapitulation on different “levels” of evolution coincided for Metchnikoff with his primary problem —identification of the second embryonic layer as the mesoderm. Metchnikoff believed that there were enough facts in general pathology and in pathological histology to accept the new phagocytosis theory. On the other hand, the observations from vertebrates offered a complex array of mixed functions of various structures, and the data did not easily reflect the most basic processes constituting the phenomenon of inflammation. But in his preparations of Bippinaria larvae and in other invertebrates, he was able to demonstrate to Virchow (already at Messina in 1883) that he “had set up the phenomena of inflammation without the assistance of nervous or vascular system” (Metchnikoff, Immunity in Infective Diseases [above, n. 27], p. 519). We mentioned above (see n. 115) that Metchnikoff's emphasis on interspecific biological struggle was only external to his evolutionary position, which turned that struggle into conflict within the individual organism. From this point of view, it is understandable why he stressed so often that the struggle of phagocytes with intruders (that is, the interspecies struggle) was only a secondary effect of their normal activity. Thus the essence of inflammation (and immunity) was seen by Metchnikoff not primarily as an expression of interspecies conflict, but as conflicting (sometimes harmonized) relations between parts (cell lineages) of the same body. By the same reasoning, we cannot agree with Todes (Darwin Without Malthus, pp. 91–93) that the “host-parasite” relation played a paradigmatic role for Metchnikoff's phagocytosis theory. Metchnikoff's interest in the phenomenon of parasitism may be traced to his first studies as a morphologist (i.e., to Leuckart's laboratory in the 1860s). In 1874, Metchnikoff published “General Essay on Parasitic Life” (“Obschii Ocherk Paraziticheskoi Zhizni” Priroda, 2 [1874], 33–82; reprinted in ACW, I, 200–234), where he presented parasitism from the morphologist's perspective: Parasites are interesting as the main representatives of regressive evolution. And here he asserted (his usual opinion at that time) that natural selection favors a decrease in organization rather than an increase; he claimed that lower organisms have more opportunities for victory in their struggle for existence than those which are comparatively more organized. In contrast, his idea of immunity hinged upon the organism's ability to mobilize its forces to protect its integrity against intruders with obviously lower organization. In the 1870s and the early 1880s, he saw in the phenomenon of parasitism further proof of his pessimistic belief that inasmuch as the struggle for existence is responsible for a given relationship, the lower forms are likely to triumph over the more complex (see Metchnikoff, “Bor'ba za Suschestvovanie v Obshirnom Smysle” [above, n. 25]). This pessimistic orientation animated Metchnikoff's efforts to fight a southern Russian beet weevil (Anisoplia austriaca) wheat epidemic with the introduction of a parasitic fungal infection. (“Illnesses of the Larvae of the Beet Weevil”) (“Bolezni Lichinok Khlebnogo Zhuka” [1879], in ACW, VI, 339–360). Neither in this paper, nor in the series of papers written on the same topic in 1880 (“Materialy k Ucheniju o Vrednykh Nasekomykh Uga Rossii” [Materials for teaching about the harmful insects of Southern Russia], Zapiski Novoross. Obsch. Estestv., 6, 1 [1880], 1–10 [reprinted in ACW, I, 269–274]; “Zamechania na Sochinenie g. Lindemana o Khlebnom Zhuke” [Notices to the treatise of Mr. Lindeman about the beet weevil], Sel'skoe Khoziaistvo i Lesovodstvo, [1880], section II, pp. 131–149 [reprinted in ACW, I, 275–289]; “Zur Lehre der Insektenkrankheiten,” Zool. Anzeiger, 3, [1880], 44–47), can be found the idea of active host defense. We do not discern any hint of a special system in the host which could play a role in active defense. The only general idea that Metchnikoff expresses in these works on parasitism, and that pertains to interspecies struggle, is that lower forms are the usual victors in the struggle with more complex species. A host protective mechanism appears in no guise and is not suggested by Metchnikoff's understanding of parasitism.

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  125. E. Metchnikoff, “Moe Prebyvanie v Messine,” Russkie Vedomosti, 31/XII, N302, (1908) (reprinted in Stranicy Vospominanii, pp. 70–76). See also Olga Metchnikoff, Life of Elie Metchnikoff, pp. 116–117. For examples of quotations from Metchnikoff's account see James G. Hirsch, “Immunity to Infectious Diseases: Review of Some Concepts of Metchnikoff,” Baltic Rev., 23 (1953), 48–60; idem, “Phagocytosis — Historical Note: A Tribute to Metchnikoff,” Ann. Rev. Microbiol., 19 (1965), 339–350; Alexander Berg, “Elias Metschnikow,” in Geschichte der Mikroskopie, ed. Hugo Freund and Alexander Berg, II, Medizin (Frankfurt: Umschau, 1964), 229–245; Louis Pelner, “Elie Metchnikoff, Ph.D.,” N.Y. J. Med., 69 (1969), 2371–76; Debra Jan Bibel, “Centennial of the Rise of Cellular Immunology: Metchnikoff's Discovery at Messina,” Amer. Soc. Microbiol. News, 48 (1982), 558–560; J. G. Hirsch and I. Hirsch, “Metchnikoff's Life and Scientific Contributions in Historical Perspective,” in Phagocytosis — Past and Future, ed. M. L. Karnovksy and L. Bolis (New York: Academic Press, 1982), pp. 1–12; Robert Herrlinger, “Die historische Entwicklung des Begriffes Phagocytose,” Ergeb. Anat. Entwicklungsgesch., 35 (1956), 334–357; Arthur M. Silverstein, “History of Immunology. Cellular versus Humoral Immunity: Determinants and Consequences of an Epic 19th-Century Battle,” Cell. Immunol., 48 (1979), 208–221.

  126. See Tauber and Chernyak, From Metaphor to Theory (above, n. 1), Chernyak and Tauber, “Birth of Immunology” (above, n. 2), and nn. 13, 15 and 16, above.

  127. We should remember that during the nineteenth century, biology had just started to “gradually emancipate itself from its intellectual and institutional roots in medicine. ... Physiology itself was an ancient science and its students had often turned to animals (but, obviously, rarely to plants) for useful instruction in the workings of the human body. But physiology referred to the study of the functions of the human body and was for the most part a matter of medical concern” (Coleman, Biology in the Nineteen Century [above, n. 14] p. 3). While others viewed the biological phenomenon through a medical perspective, Metchnikoff saw medical (pathological) phenomena through general biological parameters (see Metchnikoff, Lectures on the Comparative Pathology of Inflammation [above, n. 124], p. xiii).

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Chernyak, L., Tauber, A.I. The idea of immunity: Metchnikoff's metaphysics and science. J Hist Biol 23, 187–249 (1990). https://doi.org/10.1007/BF00141470

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