Neo-Lamarckism

During the 1890s a revival of Lamarckian ideas became a strong component of the general upsurge in biological thought. Characteristically, the conditions and expressions of the revival varied from one country to another. In Great Britain the Lamarckian scene was dominant by Herbert Spencer's thunderous attack on August Weismann's equally thunderous rejection of the Lamarckian notion of the inheritance of acquired characteristics as a motor of organic evolution. In the United States the situation was dominated by Edward Drinker Cope's heavy reliance on Lamarck's ideas in a frontal attack on the randomness of Darwin's idea of evolution. In France, Edmond Perrier represented a strong group of biologists eager to show that Lamarck's evolutionary theory should be treated not as a rival of Darwin's theory but as an "indispensable base" upon which Darwin built his theoretical edifice.^[12] Alfred Giard made Lamarck's views the basic component of an elaborate evolutionary theory in which the environmental determinants occupied the primary position, and in which Darwin's natural selection was recognized only as one of "the secondary factors."^[13] He pointed out that Bossuet, Montesquieu, and Buffon, the illustrious French thinkers in the eighteenth century, had recognized climate and other physical features of the environment as a direct and primary factor of organic transformation.^[14] Y. Delage and M. Goldsmith noted that many biologists were actually "shy Lamarckists."^[15] Felix le Dantec, the Sorbonne biologist-philosopher, went back to Lamarck to help Darwin's theory become a legitimate part of the Newtonian picture of the world.

In no country was the revival of Lamarckian thought more widespread and intensive than in Germany. Among many strains of resurrected Lamarckism, two stood out because of the unusual strength of their influence and the clear delineation of their interests. One group concentrated on the experimental study of the direct influence of exter-

nal environment on the evolutionary process and on the inheritance of acquired characteristics. This orientation reached the culminating point in the experimental work of Max Standfuss, E. Fischer, Hans Przibram, and Paul Kammerer.^[16] The Russian commentators on modern developments in biology preferred to identify this orientation as mechanical Lamarckism or mechano-Lamarckism, a term used by Ludwig Plate in his classic Selektionsprinzip, primarily because of its goal to reduce vital processes to physicochemical explanations.^[17] The second group elaborated on Lamarck's idea of an internal impulse for perfection as the basic motor of organic evolution. Whereas the first group emphasized the external causation of evolution, the second group attributed the primary significance to internal causation. Whereas the first group sought a physicochemical explanation of evolution, the second group was concerned primarily with psychological explanations.^[18] For this reason it earned the title psychological Lamarckism.

The mechanical orientation in Russia was for a long time inspired by the wide publicity given to Shmankevich's experimental study of the transformation of Artemia salina into Artemia Mühlhausenii . Shmankevich interpreted this unique "transformation" as a result of the direct influence of changes in the salinity and temperature of water in the Odessa lagoons. The experimental research of this kind became so popular that two eminent observers—A. N. Beketov and A. N. Severtsov—went so far as to claim that Lamarckism was the dominant orientation in evolutionary biology during the 1890s.^[19] Beketov and Severtsov did not document their claims, and in the specific reference to Russia, they could not have documented them. During this period Russia did not produce a single Lamarckian scholar capable of attracting the attention of the scientific community. Intense and far-reaching, most Lamarckian thought was submerged in Darwinism.

During the 1890s Shmankevich's reputation waned rapidly. The results of accumulated experimental research contradicted the results of the Odessa studies and helped undermine one of the prize claims of experimental Lamarckism. Challenges to Shmankevich's claims came first from the West. Alfred Russel Wallace argued that Shmankevich's study did not explain "the cumulative effects" of external influences on "the very lowest organisms."^[20] A true Darwinist, he knew that to give in to the Lamarckian emphasis on the "direct influence" of environmental conditions would mean to give up on natural selection as the key factor of organic evolution. As early as 1894 William Bateson claimed that A. salina and A. Mühlhausenii were not different species, as Shmankevich

had claimed, but varieties of the same species.^[21] In 1898 V. P. Anikin, curator of the Zoological Museum of Tomsk University, reported that the "transformations" Shmankevich claimed to have observed were actually pathological effects of a drastic change in the environment. He also found that A. Mühlhausenii appeared in an unusually large number of varieties. Three years later P. N. Buchinskii reported to the Eleventh Congress of Russian Naturalists and Physicians that his long and tedious experiments with the same species and in the same environment did not induce anything resembling the transformation Shmankevich had reported.^[22] Shmankevich came under attack not so much because he was preoccupied with the direct influence of the environment on the transformation of species but because his claims were too extravagant. The rise of Weismann's neo-Darwinism encouraged experimental ventures that contradicted Lamarckian claims. The collapse of the Shmankevich "school" made it much more difficult for the Russian Lamarckists to become a distinct community of scholars.

V. V. Zalenskii, a member of the St. Petersburg Academy of Sciences, noted that the direct influence of the environment on the process of evolution continued to be an open and unanswered question. After surveying a fair sample of contemporary Western studies of the influence of specific environmental factors on the transformation of species, he concluded in 1896 that the paucity of experimental data made it impossible to draw conclusions of a more general nature. He could not point out a single scientific contribution of this type of research which found a practical application in animal husbandry.^[23] He commended Shmankevich for the precision and originality of his research, but he continued to be generally skeptical about relating experimental findings of environmental studies to the laws of evolution.

Psychologically oriented Lamarckism was dominated by the German botanists-philosophers August Pauly and Raoul Francé.^[24] L. Plate stated that the views of these two scholars could be termed "psycho-Lamarckism."^[25] Yves Delage and Marie Goldsmith named this orientation "vitalistic or teleological neo-Lamarckism."^[26] The psycho-Lamarckians built their evolutionary theories on the idea of an innate "striving for progress," characteristic for all animals regardless of the complexity of their organization. It made no difference to them that this claim did not originate with Lamarck but was attributed to him first by Cuvier and then by Darwin.^[27] No doubt, psycho-Lamarckians were attracted to the idea of "inherent impulse" because it invited and legitimated a psychological approach to evolution. They also preferred

Lamarck over Darwin because he presented, they thought, a picture of the living world that was ordered, purposeful, and harmonious with the universal aspects of human society and culture. Lamarck lived in an age in which the line separating biology from metaphysics was neither precise nor fixed.

Theodor Eimer, a noted German zoologist, was generally recognized as an immediate ancestor of the psychological branch of neo-Lamarckism. His studies made many appreciative references to Lamarck's and Cope's statements on the external environment as the primary factor of evolution. He linked a minute empirical study of the wing patterns of two large groups of butterflies to a general theory of evolution. Instead of Darwin's random variation, Eimer presented a picture of organic transformation as a one-directional process, which he named orthogenesis. This theory assumed the existence of an "internal constitution," responsible for making the transformation of organic forms an ordered and purposeful process. It lent support to the cosmology of the day built on two pillars: evolution and progress.

In Russia, Eimer's ideas attracted the attention of popular commentators on the current state of evolutionary theory much more than they attracted the attention of the members of the scientific community engaged in research. They pleased anti-Darwinists without offending Darwinists. Anti-Darwinists appreciated Eimer's firm criticism of Darwin's random variation and close affiliation with the Newtonian world view. Darwinists appreciated the massive proportions of his collection of empirical data sustaining the notion of evolution.^[28]

Psycho-Lamarckians represented one of the best-known wings of a large aggregate of schools of thought that went under the general name of "panpsychism"—an effort to treat every plant and every animal as a full system of intertwined and intricate mental acts. Adaptation, a vehicle of transformation, results, in the eyes of psycho-Lamarckians, from mental activities of living beings. It comes about as a result of the innate ability of organisms to act in accordance with a "principle" expressing definite "purposes" of evolutionary processes.^[29] Behavior makes plants and animals active participants in their own evolution. A typical supporter of psycho-Lamarckism argued that Darwin attributed to plants and animals only a passive role in the process of organic evolution.

In the opinion of Raoul Francé, Darwin's recognition of evolution as a key to the full scientific understanding of the dynamics of life was a great and indelible contribution to modern thought. Darwin's explana-

tion of the inner workings of evolution, however, was inadequate and did not belong to the realm of science. It was too closely tied to mechanistic philosophy—and to philosophical materialism—to penetrate the bewildering complexity of life processes. One of the basic tasks of twentieth-century biology was to modernize Darwinism by freeing it from mechanistic and materialistic controls and by making it an integral part of modern idealism. In the Current State of Darwinism, Francé noted that science had reached the heights of perfection by recognizing the primacy of the psychological factor in living nature. He referred to his ideas as animation theory (Beseelungslehre ), which he presented as the twentieth-century replacement for the nineteenth-century science built on mechanistic principles.^[30] The modern study of evolution shifted the emphasis from external (physicochemical) to internal (psychological) causation.^[31]

Psycho-Lamarckism found little support in the Russian scientific community. A typical biologist viewed the theories presented by Francé, Pauly, and other psycho-Lamarckians as unwholesome mixtures of Lamarckian science and vitalist metaphysics. The defenders of both orthodox and moderate Darwinism in the scientific community were obviously convinced that psycho-Lamarckism was an eclectic movement with little promise for crossfertilization with other orientations in evolutionary biology. Vladimir Vagner argued that all efforts to equate the innate "impulse for progress" with the "vital force" postulated by idealistic philosophers produced a morass of metaphysical speculation and contradicted Lamarck's evolutionary theory in the first place. Metaphysical intrusion, in his view, produced one major result: it exaggerated the difference between the evolutionary views of Lamarck and Darwin. Psycho-Lamarckism was much closer to the vitalistic leanings of Naturphilosophie than to Lamarck's theoretical views.^[32] According to V. V. Lunkevich, Francé's and Pauly's version of panpsychism was no less one-sided and "primitive" than the extreme materialism of Vogt, Büchner and Moleschott.^[33]

While the experimentalist and psychological branches of Lamarckism showed only faint signs of existence in the Russian biological thought, the presence of classical Lamarckism was all too clearly noticeable. In Russia classical Lamarckism had three general characteristics: it found expression in scattered and fragmented ideas rather than in systematic theory and elaborate experimental research; it showed a more conciliatory attitude toward Darwinism than was the case in Germany; and it placed much stronger emphasis on the direct influence of the environ-

ment on the transformation process and on the inheritance of acquired characteristics than on "the internal impulse for perfection."

There was a strong tendency among Russian biologists to minimize the differences between classical Lamarckism and Darwinism. Quite often a Lamarckian scholar worked within a modified framework of Darwinism. Nor was it uncommon to encounter Darwinian scholars who were eager to point out fundamental similarities between Darwin and Lamarck, particularly in interpreting the evolutionary role of the external environment and the inheritance of acquired characteristics. Before becoming distinct entities, the theories of Lamarck and Darwin, according to Menzbir, were made out of similar parts. Menzbir wrote:

Like Lamarck, Darwin finds it impossible to set a limit on variation in plants and animals. He, too, explains variation by resorting to heredity which is responsible for transmitting inborn and acquired characteristics from parents to children. Again like Lamarck, Darwin recognizes the possibility of strengthening individual characteristics in cases where they are equally shared by parents. He, too, looks at variation as a product of changes in the external conditions and of use and disuse of organs, which Lamarck viewed as key factors of evolution. In everything else, however, Darwin appears as a fully independent scientist and follows a path that is completely his own.^[34]

M. M. Filippov, a mathematician who wrote popular articles on scientific, literary, and philosophical themes, was also firmly convinced of the fundamental similarity in the views of Lamarck and Darwin. Both scientists, he wrote, clung steadfastly to the idea of the purposiveness of vital processes as products of natural causes, even though they provided different specifics in explaining it. Neither Darwin nor Lamarck made use of vitalistic conjectures. Both believed in the inheritance of acquired characteristics. And both recognized the direct role of the environment in the evolutionary process, even though not with the same degree of firmness.^[35] Despite these similarities, Filippov did not overlook the critical differences between the two theories. In general, however, he believed that the future of evolutionary biology lay in a firmer and more comprehensive synthesis of Lamarck's and Darwin's views.

V. V. Lunkevich, a careful and informed observer of current developments in biology, diagnosed the situation correctly when he stated that to understand Darwinism and classical Lamarckism was to understand them as complementary, rather than as mutually exclusive, biological orientations.^[36] The botanist V. L. Komarov, with strong Lamarckian leanings, observed that in the study of the origin of species the question of the hereditary transmission of acquired characteristics was not re-

solved. He also noted that it did not make much difference whether the biologists emphasized direct or indirect influence of the environment on the evolution of species. Nor did it make much difference whether they placed the primary emphasis on the emergence of new characteristics or on natural selection.^[37] In their views on the evolutionary role of the environment, Mikhail Menzbir and Aleksei Severtsov, the staunch defenders of Darwinian orthodoxy, showed strong Lamarckian tendencies but only on a most generalized level.

There were also biologists who accepted the idea of a fundamental affinity of Darwinism with neo-Lamarckism, even in its psychological variety. After an analysis of the psychological branch of neo-Lamarckism, M. M. Novikov drew two general conclusions. First, unlike other neovitalist orientations, neo-Lamarckians agreed with Darwinism in accepting evolution as the pivotal notion of biology. A typical neovitalist, by contrast, thought that the time had come to remove evolution from the strategic heights of biological research. In his limited reference to evolution, Driesch, for example, relied exclusively on paleontological and zoogeographical data. Second, neo-Lamarckians and Darwinists accepted the idea of the inheritance of acquired characteristics. In this respect, neo-Lamarckism and Darwinism were united against Weismann's neo-Darwinism, their common enemy.^[38]

All this, however, did not mean that revived Lamarckism was not exposed to growing attacks from varied sources. Neo-Darwinism, the archenemy of environmental determinism in organic evolution, became the leading force in the anti-Lamarckian movement. Kholodkovskii, a moderate Darwinist, wrote in 1891 that it was much easier to show the influence of a specific environmental factor on the emergence of new characteristics than to prove the transmissibility of these characteristics by the mechanism of heredity.^[39] Shmankevich's study of the Artemia and Branchipus, he said, presented a conclusive picture of the active role of the salinity and temperature of Odessa lagoons in inducing morphological changes, but it presented an inconclusive picture of the relationship of these changes to the system of heredity.

Kliment Timiriazev, in his persistent defense of Darwinian theory, did not take the trouble to attack individual branches of neo-Lamarckism: he limited his attack to classical Lamarckism. Lamarck erred, he said, in making generalizations that had no empirical base, particularly in claiming the environment as the direct source of changes in races and species and the inheritance of acquired characteristics. In 1909 Timiriazev went a step further by rejecting Lamarck's notion of adaptation.

Darwin, he wrote, opposed Lamarck's tendency to identify every morphological change as an adaptation to the environment and as a "progressive step" in the evolution of species.^[40] Nor did Lamarck succeed in giving a precise explanation of the work of purposiveness in nature, an effort in which Darwin, according to Timiriazev, was eminently successful. Darwin criticized Lamarck's "unfortunate effort" to treat "mental" or "volitional" activities of animals as a driving force of evolution; he did not criticize Lamarck's claim that the environment is the primary cause of modifications in living forms. Timiriazev saw the future of evolutionary biology in a combination of Lamarck's views on the evolutionary role of the environment and Darwin's views on natural selection.^[41]

In the search for the unity of Lamarckism and Darwinism, A. N. Beketov occupied a particularly strong and appealing position. In this effort he never digressed from the idea that the environment was the primary and direct cause of variation in species.^[42] At the end of the nineteenth century, he wrote, the ideas of Lamarck and É. Geoffrey Saint-Hilaire had taken a "preeminent place" in the theory of evolution. In Beketov's view, only the study of the external causes of variation can raise the theory of evolution to the level of exact science. In other words, only a Lamarckian approach could lead to a "mechanical" (physico-chemical) study of living nature, the only one science admits. Beketov recognized natural selection as a factor of evolution but placed a stronger emphasis on "mutual aid."^[43] Despite his extensive and consistent identification with Lamarck's theory, Beketov made no effort to comment on current developments in various neo-Lamarckian schools. He clearly wanted to be known as a Darwinist with strong Lamarckian leanings. More than any other biologist, he helped nurture a strong Lamarckian tradition in Russia—not as a fully crystallized school of thought but as a reservoir of suggestive ideas that formed a distinct enclave in the mainstream of Darwinian tradition. The rapidly unfolding developments in experimental biology that led to the rise of genetics as a special science and a key discipline concerned with organic evolution found no echo in his scientific contributions.

It was not until 1911 that Lamarck's main work, La philosophic zoologique, was published in a Russian translation. In his introduction V. Karpov made an earnest effort not only to give an unbiased analysis of the translated work but also to discuss its significance within a broader intellectual and historical perspective. He thought that Lamarck viewed organic evolution as a specific ramification of the cosmic evolution, a self-evident process that required no special explanation. A Pri -

roda reviewer noted that "despite the important role Lamarckism has played in modern biology, very few biologists have read Lamarck in the original."^[44] By this time Darwin's major works were available in several Russian translations.

The plant physiologist Ivan Borodin was correct when he asserted in 1910 that the widespread influence of Lamarckism at the turn of the century owed much to the dual appeal of Lamarck's theoretical legacy. The Lamarckian ideas of the direct influence of the environment on morphological modifications and of the inheritance of acquired characteristics appealed to the defenders of mechanistic views in biology. The Lamarckian emphasis on the evolutionary role of an inner—mental—impulse for perfection appealed to many adherents of vitalism and panpsychism.^[45]