Mutationism

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Mutationism, known before 1900 as saltationism, is the set of views of evolution that emphasizes the role of large mutations capable of causing sudden jumps including immediate speciation. It is one of several alternatives to Darwinism that have existed both before and after the publication of Charles Darwin's 1859 book, On the Origin of Species, which was gradualist.[1] Mutation is seen as the source of novelty; if natural selection is considered, it is not seen as creative, and the direction of evolution is understood to reflect both mutation and selection.[2]

Saltationism was the common view before Darwin; in 1822 Étienne Geoffroy Saint-Hilaire argued that species could be formed by sudden transformations. Darwin opposed saltation, insisting on gradualism in evolution as in geology. In 1864, Albert von Kölliker revived Geoffroy's theory. In 1901 the geneticist Hugo de Vries gave the name "mutation" to seemingly new forms that suddenly arose in his experiments on the evening primrose Oenothera lamarckiana, and by 1909, mutationism became a rival to Darwinism supported by early geneticists including William Bateson, Thomas Hunt Morgan, and Reginald Punnett.

Mutationism, along with other alternatives to Darwinism like Lamarckism and orthogenesis, was discarded by most biologists in the modern synthesis of genetics and evolution of the early 20th century; mutation took its place as a source of the variation essential for natural selection to work on. However, in 1940, Richard Goldschmidt again argued for single-step speciation by macromutation, describing the organisms thus produced as "hopeful monsters", earning widespread ridicule. In 1987, Masatoshi Nei argued controversially that evolution was often mutation-limited. Saltation is seen to occur at various scales in various organisms, and some such as polyploidy can certainly create new plant species. However, modern biologists such as Douglas J. Futuyma conclude that essentially all claims of evolution driven by large mutations can be explained within the Darwinian evolutionary synthesis.

History[edit]

Étienne Geoffroy Saint-Hilaire believed that "monstrosities" could immediately found new species.

Geoffroy's monstrosities, 1822[edit]

Prior to Charles Darwin, most naturalists were saltationists.[3] Jean-Baptiste Lamarck was a gradualist but similar to other scientists of the period had written that saltational evolution was possible.

In 1822, in the second volume of his Philosophie anatomique, Étienne Geoffroy Saint-Hilaire endorsed a theory of saltational evolution that "monstrosities could become the founding fathers (or mothers) of new species by instantaneous transition from one form to the next."[4] Geoffroy wrote that environmental pressures could produce sudden transformations to establish new species instantaneously.[5]

Darwin's anti-saltationist gradualism, 1859[edit]

In his 1859 book On the Origin of Species, Charles Darwin denied saltational evolution. He argued that evolutionary transformation always proceeds gradually, never in jumps: "natural selection acts solely by accumulating slight successive favourable variations, it can produce no great or sudden modification; it can act only by very short steps". Darwin continued in this belief throughout his life.[6]

Thomas Henry Huxley warned Darwin that he had taken on "an unnecessary difficulty in adopting Natura non facit saltum ["Nature does not take leaps"] so unreservedly."[7] Huxley feared this assumption could discourage naturalists (catastrophists) who believed that major leaps and cataclysms played a significant role in the history of life.[8]

von Kölliker's heterogenesis, 1864[edit]

Rudolph Albert von Kölliker revived Geoffroy's mutationist ideas, calling his theory heterogenesis.

In 1864 Albert von Kölliker revived Geoffroy's theory that evolution proceeds by large steps, under the name of heterogenesis.[9]

From 1860 to 1880 saltation had a minority interest but by 1890 had become a major interest to scientists.[10] In their paper on evolutionary theories in the 20th century (Levit et al. 2008) wrote:

The advocates of saltationism deny the Darwinian idea of slowly and gradually growing divergence of character as the only source of evolutionary progress. They would not necessarily completely deny gradual variation, but claim that cardinally new 'body plans' come into being as a result of saltations (sudden, discontinuous and crucial changes, for example, the series of macromutations). The latter are responsible for the sudden appearance of new higher taxa including classes and orders, while small variation is supposed to be responsible for the fine adaptations below the species level.[11]

De Vries and Mendelian mutationism, 1901[edit]

The geneticist Hugo de Vries introduced the term "mutation" and a mutationist theory of evolution.

Hugo de Vries's careful 1901 studies of wild variants of Oenothera lamarckiana showed that distinct new forms could arise suddenly in nature, and could be propagated for many generations without dissipation or blending. He gave such changes the name "mutation".[12][13]

By 1909, mutationism was a distinctive rival to Darwinism.[14][15][16] At that time, Darwin's mechanism of natural selection was understood to rely on hereditary blending of abundant continuous variations. However, Wilhelm Johannsen's "pure line" experiments appeared to refute this mechanism. Using true-breeding varieties of beans, each with a different size of seeds, Johannsen showed selection could be used to sort out different varieties, but selection within pure lines would not produce evolutionary changes, even though pure lines continued to generate the kind of abundant variations that Darwinians saw as the fuel for evolution.[17]

These discoveries are often framed as a controversy between the early geneticists—the "Mendelians"—including William Bateson, Wilhelm Johannsen, Hugo de Vries, Thomas Hunt Morgan, and Reginald Punnett, who advocated Mendelism and mutation, and were understood as opponents of Darwin's original view, and the biometricians such as Walter Weldon and Karl Pearson and their allies, who opposed Mendelism and were more faithful to Darwin.[18]

Berg's nomogenesis, 1922; Willis's macromutations, 1923[edit]

John Christopher Willis's The Course of Evolution by Differentiation Or Divergent Mutation Rather Than by Selection, 1940

The botanist John Christopher Willis proposed an early saltationist theory of evolution. He held that species were formed by large mutations, not gradual evolution by natural selection,[19][20] and that evolution was driven by orthogenesis, which he called "differentiation", rather than by natural selection.[21]

A similar combination of mutationism and directed (orthogenetic) evolution was proposed by Lev Berg in 1922 in his book Nomogenesis; or, Evolution Determined by Law. He used evidence from paleontology, zoology, and botany to argue that natural selection had limitations which set a direction for evolution. He claimed that speciation was caused by "mass transformation of a great number of individuals" by directed mass mutations.[22][23]

Goldschmidt's hopeful monsters, 1940[edit]

In his 1940 book The Material Basis of Evolution, the German geneticist Richard Goldschmidt argued for single-step speciation by macromutation, describing the organisms thus produced as "hopeful monsters". Goldschmidt's thesis was universally rejected and widely ridiculed by biologists, who favoured the neo-Darwinian explanations of R.A. Fisher, J. B. S. Haldane and Sewall Wright.[24][25] However, interest in Goldschmidt's ideas has reawakened in the field of evolutionary developmental biology.[26][27][28][29][30]

Modern synthesis, 1930 to 1950[edit]

The eclipse of Darwinism, and the debate between Mendelian geneticists such as de Vries and biometricians such as Pearson, ended with the modern synthesis that took place between about 1930 and 1950. The new population genetics of the 1940s demonstrated the explanatory power of natural selection, and mutationism, alongside other non-Darwinian approaches such as orthogenesis and structuralism, was largely abandoned.[31]

Nei's mutation-driven evolution, 1987[edit]

Contemporary biologists accept that mutation and selection both play roles in evolution; the mainstream view is that while mutation supplies material for selection in the form of variation, all non-random outcomes are caused by natural selection.[32] Masatoshi Nei argues instead that the production of more efficient genotypes by mutation is fundamental for evolution, and that evolution is often mutation-limited.[33][34][35][36][37][38][39][40][41] Nei's book received thoughtful reviews; while Wright,[42] in the conservative journal Evolution, rejected Nei's thinking as mistaken, Galtier,[43] Weiss,[44] Stoltzfus,[33] and Wagner,[32] although not necessarily agreeing with Nei's position, treated it as a relevant alternative view.

Contemporary piecemeal approaches[edit]

Multiple explanations have been offered since the 19th century for how evolution took place, given that many scientists initially had objections to natural selection. Many of these, including mutationism, led to some form of orthogenesis. Better understanding of mutation led instead to the modern synthesis of Mendelian genetics with natural selection, establishing Darwinian evolution throughout biology.

Evidence has been presented for some forms of saltation, often on a small scale,[45][46][47][48] such as in the buccal cavity of the roundworm Caenorhabditis elegans,[49] and the phenotypes of centipedes.[50] Epigenetic inheritance can also produce saltational changes.[51] Saltational evolution may have taken place in Proserpinus sphinx moths,[52] as may pheromone evolution in bark beetles[53] and Bactrocera flies,[54] along with mimicry in butterflies and other insects,[55][56][57] as the geneticist Reginald Punnett argued in 1915.[58] Saltational speciation has taken place in flowering plants, such as in the genus Clarkia (Onagraceae).[59] Calycadenia pauciflora (Asteraceae) may have originated directly from an ancestral race through a single saltational event involving multiple chromosome breaks,[60] while simple homeotic morphs in orchids can lead to new species.[61][62]

The endosymbiotic theory implies rare but major events of saltational evolution by symbiogenesis.[63] Carl Woese and colleagues suggested that the absence of RNA signature continuum between domains of bacteria, archaea, and eukarya shows that these major lineages materialized via large saltations in cellular organization.[64]

Saltation at a variety of scales is agreed to be possible by mechanisms including polyploidy, which certainly can create new species of plant,[65][66] gene duplication, lateral gene transfer,[48] and transposable elements (jumping genes).[67]

Reviewing the history of macroevolutionary theories, the American evolutionary biologist Douglas J. Futuyma notes that since 1970, two very different alternatives to Darwinian gradualism have been proposed, both by Gould: mutationism, and punctuated equilibria.[68][69] Gould's macromutation theory gave a nod to his predecessor with an envisaged "Goldschmidt break" between evolution within a species and speciation. His advocacy of Goldschmidt was attacked with "highly unflattering comments"[68] by B. Charlesworth[70] and Templeton.[71] Futuyma concludes, following other biologists reviewing the field such as K.Sterelny[72] and A. Minelli,[73] that essentially all the claims of evolution driven by large mutations could be explained within the Darwinian evolutionary synthesis.[68]

Historiography[edit]

Historians have interpreted the history of mutationism in different ways.[1][17][74][18] The classical view is that mutationism was an obvious error; the decades-long delay in synthesizing genetics and Darwinism is an "inexplicable embarrassment";[75] genetics led logically to the modern synthesis and mutationism was one of several anti-Darwinian "blind alleys" separate from the main line leading from Darwin to the present.[76] A revisionist view is that mutationists accepted both mutation and selection, with broadly the same roles they have today. At the time of the Darwin centennial in Cambridge in 1909, mutationism and Lamarckism were contrasted with natural selection as competing ideas; 50 years later, at the 1959 University of Chicago centennial of the publication of On the Origin of Species, mutationism was no longer seriously considered.[77]

See also[edit]

References[edit]

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