ROMANES, GEORGE JOHN

(b. Kingston, Ontario, 2 May 1848; d. Oxford, England, 23 May 1894)

physiology comparative psychology evolution.

Romanes was the third son of Rev. George Romanes, a classical scholar and theologian, and Isabella Gair Smith. In the year of Romanes’ birth his father inherited a considerable fortune, resigned his post as professor of Greek at Queen’s University in Kingston, and moved his family to Britain, finally settling in Regent’s Park, London. Romanes grew up in comfortable circumstances, accompanying his parents on several trips to the Continent. Except for a brief period in a preparatory school (cut short by illness), he was educated at home. This early education was casual and unsystematic, and Romanes apparently showed no early signs of intellectual promise. At the age of seventeen he was sent to a tutor to read in preparation for the university, and in October 1867 he entered Gonville and Caius College, Cambridge.

The six years (1867–1873) that Romanes spent at Cambridge were decisive both in drawing out his talents and in determining the direction of his future work. After reading mathematics for a time, his attention was drawn to science by fellow students; and in 1868 he competed successfully for a natural science scholarship. By the time he received a second class on the natural science tripos of 1870, Romanes had abandoned his intention of entering the Church; and shortly thereafter he began to study physiology under Michael Foster. Foster, at that time just beginning his great reform of British physiology, was a pioneer in the evolutionary approach to the subject; and it was while working with him that Romanes read Darwin’s works for the first time. In the exciting atmosphere of Foster’s laboratory, Romanes’ interest grew rapidly; and a personal note from Darwin praising a piece of his in Nature reinforced his decision to devote himself to scientific research.

In 1874 Romanes took his M.A. and moved to London, where he continued his physiological studies under William Sharpey and John Burdon-Sanderson at University College, and extended the circle of his scientific acquaintances. In the same year he visited Darwin, beginning a personal friendship that remained close until the latter’s death.

Romanes received his training and began his scientific career at a time when evolutionary thinking dominated British scientific circles. Working under Foster and Burdon-Sanderson, and befriended by Darwin himself, he was thoroughly imbued with evolutionary anodes of thought. This influence is clearly reflected in each of the three areas of study to which he devoted himself: physiology, comparative psychology, and the theory of evolution.

In the twenty years following his move from Cambridge and before his death in 1894, Romanes proved to be one of the most brilliant of the second generation of British Darwinists. Freed from professional obligations by family wealth, uninterested in politics, and endowed with tremendous energy and enthusiasm, he was able to put all his resources at the disposal of science. A clear and forceful writer, he was a prolific contributor not only to scientific journals but also to the monthly reviews that then helped to shape educated opinion. His dialectical skill and unshakable good humor made him a formidable opponent in scientific debate and a popular lecturer. Most of his books were translated into French and German, and his work was widely read in the United States. In addition to his writing and lecturing, Romanes also served as zoological secretary of the Linnean Society and secretary of the Physiological Society, and was a member of the Council of University College, London. In 1891 he founded at Oxford the series of lectures that bears his name.

Romanes’ most important early scientific work was in invertebrate physiology. Working at a small marine laboratory set up at his summer home on the Scottish coast, he began by trying to determine whether the medusa (jellyfish) possessed nerve tissue. This was still an open question and one with definite evolutionary interest, since nerve tissue had already been found in all higher organisms.

Excision experiments showed that the marginal tissue of the rim of the medusa’s swimming bell was able to continue its rhythmic contractions when separated from the bell, while the bell itself was paralyzed by the same operation. Romanes concluded that the centers of spontaneous contraction of the medusa were localized in the marginal tissue. Further excision experiments indicated that such centers of spontaneity were concentrated in the small marginal bodies, or lithocysts. At the same time the excised bell retained the capacity to contract in response to mechanical or electrical stimulation. Severe spiral sectioning of the bell with interdigitating cuts failed to destroy its contractile properties. If carried far enough, however, sectioning would eventually produce a “block” of the waves of contraction at a single point. To explain these physiological results, Romanes at first postulated the existence of a fine plexus or dense crisscrossing grid of “lines of nervous discharge,” without, however, being able to describe this plexus histologically. These experiments and their results formed the subject of Romanes’ first paper for the Royal Society, published in 1876.

At Romanes’ request the medusa was examined histologically in 1877 by his friend Edward A. Schäfer (later Sharpey-Schafer), who, using a gold chloride stain, was able to demonstrate the existence of an “interlacement” of nerve fibers on the undersurface of the bell. Schäfer concluded that the individual nerve fibers, though everywhere closely spaced, were anatomically discontinuous. This was a startling result at the time, since physiological continuity was then thought to demand the anastomosis of nervous elements, and Romanes at first hesitated to accept it. In his later papers he did acknowledge it, however, and by 1885 he was arguing that structural discontinuity of nervous elements was entirely compatible with their physiological continuity. This result of the work of Romanes and Schäfer, together with Romanes’ general comparative approach and his skillful coordination of behavioral, physiological, and histological data, had a significant impact on Charles Sherrington’s development of the synapse concept.

As a Student of Michael Foster and inspired by his evolutionary approach to physiology, Romanes had been alert from the beginning to the possible comparative significance of his work on the medusa, especially in relation to Foster’s work on the vertebrate heart. At the time that his first paper on the medusa was published, he thought that both the vertebrate heart and the bell of the medusa might be cases of an intermediate evolutionary stage between primitive, irritable protoplasm and fully differentiated nerve tissue. When Schäfer’s histological results forced him to abandon this idea, he turned, in his second and third papers, to the elucidation of functional parallels. Probably under Foster’s influence, Romanes now gave special attention to rhythmic motion. He began to speak of the lithocysts of the medusa as ganglia, and in his third paper he gave a theory of ganglionic action almost identical to Foster’s theory of the vertebrate heart. According to this theory, the ganglia normally released a continuous impulse to which the muscle tissue could respond rhythmically. At the same time that Romanes was working toward these conclusions, Foster was directing the attention of another student, Walter Gaskell, to the problems of rhythmic motion in the heart. Romanes’ clear and decisive conclusions and the analogy he drew between the cases of the medusa and the heart were important elements in the background of Gaskell’s later demonstration of the myogenic origin of the heartbeat.

While he was carrying out this important work in physiology, Romanes was gathering observations and corresponding with Darwin on the subject of animal intelligence. To both Darwin and Romanes it appeared that the theory of evolution required a fundamental continuity in the spectrum of mental life, extending from the lowest organisms up to and including man. Moreover, the ascending stages of mental development should be susceptible of explanation in terms of natural causes. Romanes set himself the task of demonstrating the fact of this continuous development and of giving an account of psychological processes in the light of their probable historical origins. The means to this end was to be a new science of animal intelligence, which he named, in a hopeful allusion to its successful anatomical counterpart, comparative psychology.

Romanes published three books on the subject. The first and most successful, Animal Intelligence (London, 1882), was entirely devoted to a more or less systematic presentation of a large number of observations of animal activities presumed to be indicative of intelligence. Anecdotal material gathered from diverse sources was included, together with the results of Romanes’ own observations. This was followed by two explicitly theoretical works, Mental Evolution in Animals (London, 1883) and Mental Evolution in Man (London, 1888). Romanes took as part of the goal of comparative psychology the accurate description of the mental states of animals based on inference from their observed behavior. His own account of these mental states was imbedded in the categories of the British associationist tradition. Like Spencer before him, however, Romanes adapted the associationist scheme to allow for the phylogenetic development of successively higher mental levels, outlining a continuous historical evolution culminating in human intelligence.

Romanes’ theoretical books on comparative psychology never exerted much influence. It was rather the observational and experimental side of his work that led to further developments in the field. Romanes’ systematic and relatively critical approach marked a definite advance in the study of animal behavior, hitherto a subject for casual and often sentimental anecdotes. Nevertheless, newer workers in the field, led by his younger friend and colleague C. Lloyd Morgan, criticized the observational methods of Animal Intelligence as lacking in rigor and Romanes’ inferences as too often anthropomorphic. The tendency of later workers in comparative psychology, such as Morgan and Jacques Loeb, was toward greater objectivity in the study of animal behavior, and to reduce any inferences regarding consciousness to the minimum entailed by experiment. The same tendency played a prominent role in the thinking of the early behaviorists in the United States. Yet although his work was soon superseded, Romanes occupies a crucial place in the origins of the modern study of animal behavior.

Romanes was a major participant in the controversies that arose among the Darwinists in the two decades following Darwin’s death. The most general question at issue was whether natural selection was the only factor at work in evolution or whether there were other, subordinate factors, without which many phenomena remained unexplained. The question of the inheritability of acquired characters—especially the effects of use and disuse—became a central issue after August Weismann published his theory of the continuity of the germ plasm in 1883. Weismann denied to the inheritability of acquired characters even the restricted role that Darwin had allotted it. This denial drew a spirited protest from Herbert Spencer, who argued for its retention. Romanes generally concurred with Spencer in rejecting a singlefactor approach to evolution and called for further experimental investigation to decide the particular question of the inheritance of acquired characters. Romanes’ own experimental investigations undertaken to decide the question proved inconclusive, as had his earlier attempts to provide experimental evidence for Darwin’s hypothesis of pangenesis.

A second controversy arose around a proposal made by Romanes himself. In May 1886 he read a lengthy paper to the Linnean Society, “Physiological Selection; an Additional Suggestion on the Origin of Species.” In it he argued that an account of evolution relying solely on natural selection failed to explain three classes of facts: the seeming nonutility of many specific characters, interspecific sterility, and the need for varieties to escape the swamping effects of intercrossing if they were to become established as permanent species. To meet these difficulties Romanes suggested that mutual infertility between two or more portions of a species population might sometimes arise prior to morphological or other distinctions. The varieties thus reproductively isolated would then be free to develop different (and possibly nonuseful) characters independently of one another, even though mixing freely in the same geographical area.

Romanes’ paper provoked a lively but generally negative response from British biologists who, although sometimes acknowledging the difficulties in question, could not accept Romanes’ proposed solution. Interest in the issue waned and was revived only in 1888 with the publication of a paper by the American naturalist John Thomas Gulick. Gulick argued that the main task of evolution theory was to give an adequate account of the causes and conditions of evolutionary divergence. Such an account could not be given in terms of natural selection alone, but required recognition of a wide range of mechanisms by which varieties or incipient species became reproductively isolated from one another. In Gulick’s scheme Romanes’ “physiological selection” appeared as one possible mode of varietal isolation.

Romanes enthusiastically adopted Gulick’s more general approach, and in volume III of Darwin, and After Darwin (1897) he argued vigorously for recognition of the role of isolation in evolution. Although few evolutionists accepted all of Romanes’ arguments and conclusions, the possible roles of isolation and physiological selection had become issues to be debated; and evidence of concern with these issues is reflected in much of the literature on evolution of the first decade of the twentieth century.

BIBLIOGRAPHY

I. Original Works. A nearly complete bibliography of Romanes’ publications may be assembled by consulting four sources: the British Museum Catalogue of Printed Books, CCV, 787–789; the Royal Society Catalogue of Scientific Papers, VIII, 772; XI, 211–212; and XVIII, 281; the Cumulative Author Index for Poole’s Index to Periodical Literature, 373; and Wellesley Index to Victorian Periodicals 1824–1900, 1069.

His most important papers in invertebrate physiology are “Preliminary Observations on the Locomotor System of Medusae,” in Philosophical Transactions of the Royal Society, 166 (1876), 269–313; “Further Observations on the Locomotor System of Medusae,” Ibid., 167 (1877), 659–752; and “Concluding Observations on the Locomotor System of Medusae,” Ibid., 171 (1880), 161–202. An additional study (not mentioned in the text) done with James Cossar Ewart was published as “Observations on the Locomotor System of Echinodermata,” Ibid., 172 (1881), 829–885. The substance of these papers was brought together and published for a wider audience as Jelly-Fish, Star-Fish, and Sea Urchins (London, 1885).

The three books cited in the text contain the major part of Romanes’ work in comparative psychology. The numerous letters to Nature and articles that he contributed on this subject were largely derived from material in these volumes.

Romanes’ skill as an expositor of the theory of evolution for the educated layman is best represented by “The Scientific Evidence of Organic Evolution,” in Fortnightly Review, 36 (1881), 739–758; and Darwin, and After Darwin, I , The Darwinian Theory (London, 1892). Romanes’ principal contributions to the Spencer-Weismann controversy were a review of Spencer’s The Factors of Organic Evolution, in Nature, 35 (1886–1887), 362–364; “The Factors of Organic Evolution,” Ibid., 36 (1887), 401–407; “Weismann’s Theory of Heredity,” in Contemporary Review, 57 (1890), 686–699; “Are the Effects of Use and Disuse Inherited ?” in Nature, 43 (1890–1891), 217–220; “Mr. Herbert Spencer on ’Natural Selection,’ “ in Contemporary Review, 63 (1893), 499–517; “The Spencer-Weismann Controversy,” Ibid., 64 (1893), 50–53; “A Note on Panmixia,” Ibid., 611–612; An Examination of Weismannism (London, 1893); and Darwin, and After Darwin, II, Post-Darwinian Questions, Heredity and Utility (London, 1895). Chapter 1 of this book consists of an important introductory essay, “The Darwinism of Darwin and of the Post-Darwinian Schools,” in which Romanes surveys the issues from his own point of view. Romanes’ original paper on physiological selection appeared in Journal of the Linnean Society, Zoology, 19 (1886), 337–411. His further contributions to the debate on physiological selection and isolation as factors in evolution include “Physiological Selection,” in Nineteenth Century, 21 (1887), 59–80; “Before and After Darwin,” in Nature, 41 (1889–1890), 524–525; “Mr. A. R. Wallace on Physiological Selection,” in Monist, 1 (1890–1891), 1–20; and Darwin, and After Darwin, III, Post-Darwinian Questions, Isolation and Physiological Selection (London, 1897).

Although there is no single repository of Romanes’ MS material, most of it is concentrated in London: archives of the Linnean Society and the Zoological Society; University College (the Burdon-Sanderson papers and the Galton papers); the British Museum (the Wallace papers, the Gladstone papers, and the Croll papers); the Royal Botanic Gardens, Kew (the Hooker papers and the Thiselton-Dyer papers); The Wellcome Institute of the History of Medicine (the Sharpey-Schafer papers); Imperial College of Science and Technology (the T. H. Huxley papers and the Armstrong papers); the Royal Society (paper relating to the early physiological work); and the Passmore Edwards Museum (the Meldola papers) There are several important items in the C. Lloyd Morgan papers at the University of Bristol (relating to Morgan’s editing of Darwin, and After Darwin, III). The American Philosophical Society possesses a large number of letters from Charles Darwin to Romanes (some unpublished) as well as 12 letters from Romanes to James Paget among the latter’s papers and approximately 100 letters from Romanes to John Thomas Gulick.

II. Secondary Literature. The standard biographical source for Romanes is Ethel Romanes, The Life and Letters of George John Romanes (London, 1896). Contemporary obituary notices were published by John Burdon-Sanderson, in Proceedings of the Royal Society, 57 (1894–1895), vii-xiv; E. Ray Lankester, in Nature, 50 (1894), 108–109; C. Lloyd Morgan, in Dictionary of National Biography, XLIX, 177–180; and E. B. Poulton, letter to the Times (London), 19 June 1894.

Romanes’ work in invertebrate physiology and its evolutionary context have been examined in two articles by Richard D. French: “Darwin and the Physiologists, or the Medusa and Modern Cardiology,” in Journal of the History of Biology, 3 (1970), 253–274; and “Some Concepts of Nerve Structure and Function in Britain, 1875–1885: Background to Sir Charles Sherrington and the Synapse Concept,” in Medical History, 14 (1970), 154–165; and in Gerald L. Geison, “Michael Foster and the Rise of the Cambridge School of Physiology, 1870–1900” (Ph. D. diss., Yale, 1970), esp. 459–475.

Literature on Romanes’ work in comparative psychology is of uneven quality and must be used with discretion. Useful standard histories of psychology are Edwin G. Boring, A History of Experimental Psychology, 2nd ed. (New York, 1950), 240–244, 468–476, 620–631; J. C. Flügel, A Hundred Years of Psychology, 2nd ed. (London, 1951), 111–125; L. S. Hearnshaw, A Short History of British Psychology 1840–1940 (London, 1964), 34–46, 86–100; and R. S. Peters, ed., Brett’s History of Psychology, rev. ed. (London, 1962), esp. 694–699, 737–738. Two relevant articles by Philip H. Gray are “The Morgan-Romanes Controversy: A Contradiction in the History of Comparative Psychology,” in Proceedings of the Montana Academy of Sciences, 23 (1963), 225–230; and “Prerequisite to an Analysis of Behaviorism: The Conscious Automaton Theory From Spalding to William James,” in Journal of the History of the Behavioral Sciences, 4 (1968), 365–376.

Romanes’ theory of physiological selection and the reactions that it provoked are discussed in John E. Lesch, “The Role of Isolation in Evolution: George J. Romanes and John T. Gulick,” Isis (in press). An account of the life and scientific work of John T. Gulick, including much of the Romanes-Gulick correspondence, may be found in Addison Gulick, Evolutionist and Missionary: John Thomas Gulick (Chicago, 1932). Some of the problems involved in the recognition of the role of isolation in evolution are treated from the point of view of contemporary biology in Ernst Mayr, “Isolation as an Evolutionary Factor,” in Proceedings of the American Philosophical Society, 103 (1959), 221–230.

Romanes’ lifelong preoccupation with the relationship between religion and science, a side of his thought not considered in this article, is discussed in Frank Miller Turner, Between Science and Religion: The Reaction to Scientific Naturalism in Late Victorian England (New Haven-London, 1974), 134–163.

John E. Lesch