Brooks, William Keith

(b. Cleveland, Ohio, 25 March 1848; d. Baltimore, Maryland, 12 November 1908)

zoology, embryology.

William was the son of Oliver Allen Brooks, a Cleveland merchant, and Ellenora Bradbury Kingsley. Shy, retiring, and gentle, and afflicted by a congenital heart defect that became progressively more limiting in later life, Brooks inherited from his mother artistic skill, a studious, idealistic nature, and a dislike for the obvious and the trivial.

A graduate of Williams College in 1870, Brooks enrolled in the first Anderson School of Natural History on Penikese Island, Massachusetts, in the summer of 1873. There he met Louis Agassiz and at once developed the interest in marine organisms that was to determine the course of his career. He completed a thesis under Alexander Agassiz and, in June 1875, received the third Ph.D. conferred at Harvard. He was then appointed associate in biology at the new Johns Hopkins University to serve under H. Newell Martin, who was himself fresh from graduate study in physiology at Cambridge with T. H. Huxley and Michael Foster. Martin and Brooks developed a major new venture in graduate education (modeled on the German university) that led the way in the vigorous growth of American biology during the last quarter of the century. Much of Brooks’s contribution to this venture and the bulk of his research were accomplished at the Chesapeake Zoological Laboratory, a movable marine station established each summer between 1878 and 1906 at various points along the Atlantic coast and in the West Indies. He spent his entire academic career at Johns Hopkins, rising in 1894 to the biology department chairmanship, a position he retained until his death.

Brooks was a descriptive evolutionary morphologist with a strong bias toward studies of whole organisms in their natural environment. A keen observer and an indefatigable amateur philosopher, he was among the late nineteenth-century morphologists who, accepting the transforming power of function in the tradition of Cuvier, added the new insights permitted by the Darwinian concept of the organism as a historical being. Organic form viewed both as a living record of its own ancestry and as dynamically adaptable to new circumstances of life lay at the center of Brooks’s thought. For him, life was adjustment, or it was nothing; fitness, the adaptive response, was paramount: “The thing to be explained is not the structure of organisms, but the fitness of this structure for the needs of living things in the world in which they pass their lives.” For him, nature was a language that a rational being may read. His reading led him to believe with Aristotle that the “essence of a living thing is not what it is made of or what it does, but why it does it”—in a word, its purpose. In thus withholding judgment on analytical, reductionist approaches, Brooks became unable to look beyond the confines of his own generation, and he did not participate in the transformation of morphology from a comparative to a causal science. Yet his more noteworthy students were somehow stimulated to share fully in that transformation; four of them—E. B. Wilson, T. H. Morgan, E. G. Conklin, and R. G. Harrison—laid the groundwork of much of modern cytology, embryology, and genetics.

In both their substance and their manner of presentation, Brooks’s descriptions of the embryology, morphology, and life habits of marine invertebrates were memorable for their scope, their meticulousness, their wealth of illustration, their evolutionary insight, and their charming literary style. His studies of the pelagic tunicate Salpa are classic. He described the complex development of Salpa buds with unexcelled clarity and demonstrated their relation to the buds of the sessile tunicates. He discovered the remarkable fact that the Salpa embryo is first fashioned from follicle cells that are later replaced by regular blastomeres. Brooks’s investigations of crustacea were equally fundamental. In several species he identified and collated the larval stages, a major morphological feat. He was the first to follow an entire crustacean life history from a single egg; this he did for Lucifer, remarkable among the arthropods in having an egg that cleaves totally. Brooks’s descriptions of the life histories of several hydroid coelenterates still stand out for the perceptiveness with which their phylogenetic relationships were assessed. Yet he could also be wrong: his theory of heredity based on pangenesis had only the merit of stimulating others to explore more deeply. And he could be ineffective: an extensive analysis of the practical aspects of conserving the Chesapeake Bay oyster won him only temporary notice despite the accuracy of his warnings.

In his own mind at least, Brooks’s forte was his capacity to identify and reflect on the great issues of biology. His many papers and especially his major book, The Foundations of Zoology, are laden with rhetorical speculation which, as Brooks cautioned his readers, might at first seem obscure but “may, on review, be found consistent and intelligible.”

BIBLIOGRAPHY

I. Original Works. A complete list of Brooks’s works is given in the biographical memoir by Conklin (see below). Collections of his scientific and popular articles are in the Eisenhower Library of the Johns Hopkins University and in the library of the Marine Biological Laboratory, Woods Hole, Massachusetts. His major publications were “The Genus Salpa,” in Memoirs of the Biological Laboratory of the Johns Hopkins University, 2 (1983), 1–303; and The Foundations of Zoology (New York, 1899).

Only fragments of Brooks’s correspondence are known to exist; these are in the Brooks Papers and the D. C. Gilman Papers at Johns Hopkins, and in the Alexander Agassiz Papers at the Museum of Comparative Zoology, Harvard University. No systematic search through other possible locations has been made.

II. Secondary Literature. There is no full biography of Brooks. Brief contemporary accounts of his life were prepared by two former students, E. A. Andrews and E. G. Conklin, the latter’s essay being the more valuable. These and other helpful sources are E. A. Andrews, “William Keith Brooks, Zoologist,” in D. S. Jordan, ed., Leading American Men of Science (New York, 1910), pp. 427–455; E. G. Conklin, “Biographical Memoir of William Keith Brooks, 1848–1908,” in Biographical Memoirs of the National Academy of Sciences, 7 (1913), 23–88; D. M. McCullough, “William Keith Brooks and American Biology in Transition: A Re-evaluation of His Influence” (thesis, Harvard Univ., 1967); C. P. Swanson, “A History of Biology at the Johns Hopkins University,” in Bios, 22 (1951), 223–262; and “William Keith Brooks. A Sketch of His Life by Some of His Former Pupils and Associates,” in Journal of Experimental Zoology, 9 (1910), 1–52.

M. V. Edds, Jr.