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Garstang, Walter

GARSTANG, WALTER

(b. Blackburn, England, 9 February 1868; d. Oxford, England, 23 February 1949)

zoology.

Garstang, the son of Walter Garstang, a doctor, intended to read medicine when he entered Oxford in 1884. Influenced by Henry N. Moseley, professor of zoology, he soon changed his mind and took his degree in zoology. In 1888 he was appointed a found er member to the staff of the Marine Biological Laboratory at Plymouth, where he worked off and on until 1901. In 1891 he spent a year at Owens College, Manchester, with Arthur Milnes Marshall, an embryologist and recapitulationist of the Balfour school; and in 1893 he was elected fellow of Lincoln College, Oxford, where he taught zoology until 1897 while continuing to carry out his research at Plym outh during vacations. In 1895 he married Lucy Ackroyd, who shared his biological interests; they had a son and five daughters.

Garstang’s research in this early phase concerned marine biology. His first paper, for instance, was on the warning coloration of nudibranch mollusks; he also wrote descriptive papers on the natural his tory and morphology of various marine organisms, particularly the invertebrate group called tunicates, which was a lifelong interest. By this time he had probably formulated his original views on the relation of embryology and evolution; they are implied, for instance, in a paper of 1894 on the ancestry of the vertebrates; but their main publication would come later.

From 1897 to 1907 Garstang worked full time on the fisheries of the British Isles. He was at Plymouth until 1901, when the International Council for the Exploration of Sea was established; he then founded its Fisheries Laboratory at Lowestoft. He sought the kind of biological knowledge that was necessary for the scientific exploitation of the sea. He thus considered the relation of physical conditions in the sea and plankton numbers, and at Plymouth he in vestigated variation in size among the different stocks of mackerel. this work was a forerunner of a larger study of plaice at Lowestoft. By mark and recapture studies. Garstang worked out the growth rates and migrations of this species. Having noticed that young plaice grow faster on the Dogger Bank (in the North Sea) than off the Dutch coast, he tried catching, marking, and transplanting fry from the latter site to the former. The transplanted fry duly grew faster. Garstang argued that mass transplantations could be commercially possible.

By 1907 the British government had designs upon the Laboratory at Lowestoft. Garstang was not the kind of man to take direction from civil servants, and with the douceur of the new professorial chair in zoology at Leeds University, his resignation from Lowestoft was secured. At Leeds he had less time for research: he was setting up a new school, establishing two laboratories (one at Leeds and a marine station on the coast at Robin Hood Bay), and — as was then customary among professorsm-taking an active interest in his undergraduate students. He helped to found the Leeds Boat Club, assisted with the O.T.C., wrote and produced student entertain ments, and kept open house. Teaching the subject, however, enabled him to return to his interest in fundamental zoology. He turned back to the phylogenetic speculations that had been so important a part of zoology in the late nineteenth century and, in the 1920’s, published his most important work.

In the 1880’s the most influential idea in phylogenetic reconstruction was the theory of recapit ulation—Haeckel’s “biogenetic law” —according to which “ontogeny recapitulates phylogeny”: the phylogenetic relations of a species can be read off by observing, with suitable interpretation, the successive developmental stages of an individual. In the decade before World War I, the vanguard of embryology had moved away from phylogenetic reconstruction to investigate, experimentally, the causes of development itself; and in the 1920’s experimental embryology continued to dominate.

Experimental embryology and genetics implicitly cut away both the factual support and the theoretical coherence of the biogenetic law, but they produced no influential critique of it. A critique was what Garstang par who had little interest in modern experimental biology) provided, together with a more general theory of the relation of evolution and de velopment to replace it. In his “critical retatement” (1922) of the biogenetic law, he pulled no punches. The law still appealed to some, such as the powerful E. W. MacBride, and Garstang’s work was controversial. The fact that, despite his obvious deserts, he was not elected a fellow of the Royal Society, has been attributed to the opposition of the minority who were interested in phylogenetic questions and the indifference of the majority who felt that kind of work was no longer to be counted real science. Nevertheless, there are two important general pa persthe critique of recapitulation (1922) and a study of larval forms (1928)m-and two particular applications—to the ancestry of vertebrates (1928) and of siphonophores (1946.)

According to the theory of recapitulation, evolutionary changes are confined to the adult stage, new forms being added to the end of the organism’s ontogeny. Garstang maintained that changes take place at all stages, and he urged that the larval forms of the marine groups he knew well are adaptive dispersal stages, not recapitulated ancestral forms. Some important phylogenetic transitions, he argued, had taken place through the modification of the ancestral larval form into a descendant adult form, with the loss of the ancestral adult stage. He called this process paedomorphosis. Although the idea was not original with him, he was highly influential in giving it currency.

Garstang applied his theory of paedomorphosis in both of his major phylogenetic papers. The an cestry of the vertebrates was the standard problem in latenineteenth-century phylogenetic speculation; almost no invertebrate group had been left unin vestigated during that imaginative age in the search for the ancestor. Garstang pointed to the similarity of certain echinoderm larvae and the larvae of tuni cates, which in turn are similar to the main chordate groups. The vertebrates, in this theory, originated by paedomorphosis. The theory is still as widely accepted as any other account of vertebrate origins. Garstang likewise suggested that the siphono phoresa group of free-floating colonial marine orgnisms—originated by paedomorphosis, in this case from a larval form, such as the actinula of the common hydroid Tubularia.

But Garstang’s main influence then and later was due not so much to his scientific papers as to his delightful verse. He found time to work out few of his broad-ranging phylogenetic speculations in detail, and he instead wrote down the main ideas amusingly and memorably, in light verse. The verse was written for friends, and little of it appeared in his lifetime; but it no doubt encouraged his graver colleagues not to take him seriously, Garstang’s ideas, how ever, have aged well, and his verse is still read with pleasure by generations that have forgotten the productions of his more fashio–conscious contemporaries.

BIBLIOGRAPHY

I. Original Works.7 “The Theory of Recapitulation: A Critical Re-statement of the Biogenetic Law,” in Journal of the Linnean Society, Zoology, 35 (1922), 81–101 “The Origin and Evolution of Larval Forms.” British Asso ciation. Glasgow, 23 pages, (1928); “The Morphology of the Tunicata, and Its Bearing on the Phylogeny of the Chordata.” in Quarterly Journal of Microscopical Science. 72 (1928), 51–187; and “The Morphology and Relations of the Siphonophora,” ibid., 87 (1946), 103–193.

II. Secondary Literature. Obituaries of Garstang by his so–i–law Sir Alister C. Hardy appeared in Journal of the marine Biological Association of the United King dom, 29 (1951), 561–566, and in Proceedings of the Linnean Society of London, 162 (1950), 99–105. Hardy gives an account of Garstang’s ideas in his introduction to Gar stang’s verse anthology, Larval Forms (Oxford, 1951), which also contains a list of Garstang’s main publications.

More generally, see Stephen Jay Gould, Ontogeny and Phylogeny (Cambridge, Mass., 1977); and Mark Ridley, “Embryology and Classical Zoology in Great Britain,” in T. Horder, J. W. Witkowski, and C. C. Wylie, eds., A History of Embryology (Cambridge, 1986), 35–67.

Mark Ridley

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Garstang, Walter

Garstang, Walter (1868–1949) A comparative anatomist and embryologist who was the first person to propose, in 1928, that vertebrates arose from a neotenous ascidian (neoteny, Ascidiacea) larva. Somewhat diffident about his more radical evolutionary hypotheses, he propounded them in anonymous comic verses that he would leave scattered about the laboratory for his students to find.

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