The Lamarckian Line

To be a Darwinian scholar did not require one to make direct references to Darwin. Most biologists made no—or very rare—references to the creator of the theory of natural selection. This was the period of mounting political oppression and of deep conservatism in the high offices of national education. To talk or write about Darwin was to preach heresy. To avoid intimidation by authorities, the evolutionists found it helpful to avoid categorical and blanket endorsement of every idea Darwin put forth and to use a more subtle idiom in expressing their favorable interpretations of general evolutionary views.

Darwinism also competed with a Lamarckian strain in biological theory that emphasized the direct influence of the changing environment on the transformation of plants and animals and made the inheritance of acquired characteristics the main mechanism of evolution. The Lamarckian substratum of the Russian evolutionary theory served as a springboard for the early resistance of a strong segment of Russian biologists to August Weismann's theory, which had no room either for the role of environment in genetic modifications or for the inheritance of acquired characteristics.

A. N. Beketov, known for his work in plant morphology and anatomy, as well as in the popularization of modern scientific ideas, earned an enviable reputation by his effort to give prominence to the Lamarckian

residues of Darwin's theory.^[85] A professor of botany at St. Petersburg University (after having first taught at Kharkov University) and an honorary member of the St. Petersburg Academy of Sciences, he exercised a strong influence on forging a community of Russian biologists and on the flow of modern biological ideas in and outside the academic world. Shortly before the appearance of the Origin of Species, Beketov wrote "Harmony in Nature," which, as he stated in his "Autobiography," emphasized the direct influence of the surrounding environment on the evolutionary role of the inheritance of acquired characteristics. Subsequently, he worked on redefining—rather than on negating—Darwin's theory.^[86]

Despite his strong Lamarckian bias, Beketov was ready to admit that "the great contribution of Darwin and Wallace has been in showing that the struggle for existence and heredity are the leading factors in natural selection: they determine the adaptation of organisms to the surrounding environment and the main course of the evolutionary process."^[87] Beketov treated the struggle for existence as a mechanism of equilibrium—or harmony—in the interaction between organisms and their environment. He viewed competition as a special ramification of cooperation, a result of the constant pull of natural forces toward a state of equilibrium. Translated into sociological terms, equilibrium becomes "harmony," and "harmony," translated into psychological terms, becomes "sympathy." Guided by a loose interpretation of the principles of Newtonian mechanics—and its conceptualization of equilibrium—Beketov built a world made up of Newtonian metaphors and poetic vision, a philosophical view which he chose not to elaborate.

Beketov may be considered the founder of a strong tradition in Russian biology which gave Darwin's theory a strong Lamarckian base. He did not hesitate to make the struggle for existence the keystone of the evolutionary theory; nor did he hesitate to superimpose three Lamarckian ideas upon it: the inexorable evolutionary pull of an "inner impulse for perfection";^[88] the use and disuse of organs as adaptive instruments of organic transformation; and the hereditary transmission of acquired characteristics.

Beketov thought that Malthus's law of discrepancy in the ratios of the growth of population and food resources referred only to "quantitative changes" in nature; only the "inner striving" of organisms for "progressive change" can explain the "qualitative" aspect of evolution. He thought that Alphonse de Candolle, in his Histoire des sciences et savants depuis deux siècles (1873), exaggerated the power of science to

enter the magic world of natural selection.^[89] Science was not yet in a position to tackle the relationship of intellectual development to physical evolution, de Candolle's basic concern. Beketov also opposed the claim of the uniformitarians that the same causes of organic evolution have operated in all geological eras.

In a paper presented before the members of the St. Petersburg Society of Naturalists in 1882 Beketov asserted that "almost all leading contemporary biologists are followers of Darwin."^[90] In his opinion, Darwin was the primary contributor to the elimination of "crude teleological explanations" from modern biology.^[91] He argued, however, that Darwin made the struggle for existence too broad and imprecise and suggested that it be limited exclusively to relations between organisms and the "general physical environment." Under the spell of Kessler's recent pronouncements, he acknowledged the important role of mutual aid in the life of plants and animals.^[92] Loyal to the Lamarckian legacy, Beketov thought that the future work of biologists would concentrate on achieving a full union of evolutionary theory and the general laws of physics.^[93]

While Beketov supported Lamarckism on a theoretical level, V. I. Shmankevich supported it on experimental and practical levels and received wide publicity for it both in Russia and abroad.^[94] In the 1870s Shmankevich, a science teacher in an Odessa secondary school, put forth resounding claims that by changing the salinity of water he was able to observe the transformation of the brine shrimp (Artemis salina ), found in local lagoons, into two different species: whereas increased salinity led to a transformation of Artemia salina into a species similar to Artemia Mühlhausenii, reduced salinity led to the transformation of Artemia salina to animals very similar to various species of the genus Brachipus . Shmankevich found the latter transformation particularly exciting because, according to at least some zoologists, Artemia and Brachipus are different genera.^[95] In all his studies, he combined a phase of observation with a phase of experimental research. In his experiments he depended on two variables: salinity and temperature of the lagoon water. Specific combinations of salinity and temperature led either to progressive or to regressive transformation.

In 1875 the Zeitschrift fur wissenschaftliche Zoologie made Shmankevich's research findings readily accessible to the Western scientific community. During the next two decades, Shmankevich was one of the most cited Russian biologists in western Europe. Even the friendliest supporters mixed guarded praise with strong reservations; the opponents were firm and irreconcilable. In 1893 William Bateson presented

the details of Shmankevich's claims, but he was inclined to think that Artemia salina and Artemia Mühlhausenii were actually the same species.^[96] Especially in Russia and Germany the role of the direct influence of the environment continued to be recognized as a challenge of large proportions. At the Eleventh Congress of Russian Naturalists and Physicians, held in St. Petersburg in 1902, a speaker informed the audience that Shmankevich's experiments and findings continued to be referred to favorably in most university textbooks in zoology.^[97]

Shmankevich noted that it was too early to give a precise description of the relation of his conclusions to Darwin's theory of natural selection. He observed in 1875 that, unlike his (Lamarckian) mode of explanation, Darwin's theory depended exclusively on indirect evidence.^[98] This did not prevent him from expressing high respect for Darwin's contributions and an optimistic view about its future triumph. At no time did he consider Darwin's and Lamarck's theories mutually exclusive. In his view, only the future generations of biologists, armed with the methods of physics and chemistry, would be in a position to shed sufficient light on the process of the evolution of organic forms and on the general validity of the theory of natural selection. In one respect, Shmankevich was very much like Beketov: both praised Darwin without abandoning their primary loyalty to Lamarck. Both contributed to the strength of Lamarckism as a strong undercurrent of the Darwinian tradition.

The 1870s witnessed the rise of yet another form of extreme Lamarckism, presented as an orientation consonant with the spirit and the substance of Darwin's scientific legacy. It was at this time that V. I. Michurin, inspired by the atmosphere created by the diffusion of Darwinian ideas, had barely begun his long-term activity of inducing heritable characteristics in fruit trees by changing the environmental conditions under which they grew. He was guided by the idea of the possibility of adding a new dimension to the Lamarckian theory: the artificial inducement of predetermined and accelerated transformation of characters. His method was the crossing of geographically distant plants; his aim was to produce varieties best adapted to specific environments. Not recognized by the scientific community, mainly because he operated in the realm of folk science, Michurin became part of a popular movement devoted to improving domestic plants in Russia and to extending their cultivation to new areas. With exemplary devotion, Michurin worked on developing new varieties of fruit trees in central Russia. Keeping Lamarckism alive, Michurinism became part of a general cultural setting that encouraged a union of Lamarck and Darwin and that stood in the way of

a faster diffusion of the theoretical ideas of modern genetics. To its chief articulators, Russian Lamarckism of this period represented a modification rather than a negation of Darwinism.

Douglas R. Weiner has suggested that the roots of Michurinism go back to the pre-Darwinian period—to the acclimatization movement of the mid-nineteenth century. In this movement he sees a distinct Russian adoption of the views of Lamarck and É. and I. Geoffroy Saint-Hilaire on the "direct environmental induction of hereditary adaptations." Weiner traces the roots of Lysenkoism to the strong influence of the "French school" on the early acclimatization movement in Russia.^[99]

Darwin did not stand in the way of the current effort to merge his theory with Lamarck's ideas. In fact, he encouraged such an effort. In his preface to the second edition of The Descent of Man he made his position eminently clear:

I may take this opportunity of remarking that my critics frequently assume that I attribute all changes of corporeal structure and mental power exclusively to the natural selection of such variations as are often called spontaneous; whereas, even in the first edition of the Origin of Species, I distinctly stated that great weight must be attributed to the inherited effects of use and disuse, with respect both to the body and mind. I also attributed some amount of modification to the direct and prolonged action of changed conditions of life.^[100]

In recounting the landmarks of his scientific achievement, a surprisingly large number of Russian scientists and ideologues credited Darwin with saving Lamarck from obscurity and with making his evolutionary idea an organic part of the mainstream of modern biology.