Sutton, Walter Stanborough
SUTTON, WALTER STANBOROUGH
Sutton was the fifth of the seven sons of William Bell Sutton and Agnes Black Sutton. His father, a farmer, moved from New York to Russell Country, Kansas, when Sutton was ten years old. He was educated in the local public schools and entered the engineering school of the University of Kansas at Lawrence in 1896. The death of a younger brother from typhoid fever in the summer of 1897 decided Sutton on a career in medicine. He transferred to the school of arts (later the college of liberal arts) the following fall and embarked on biological studies. Under the influence of Clarence Erwin McClung, then an instructor in zoology, he began cytologic work. He received the B.A. degree in 1900 and, working as McClung’s first graduate student, earned the M.A. degree in 1901. In the fall of that year he went to Columbia University to work with Edmund Beecher Wilson. Although Sutton never completed a Ph.D. thesis, during the years 1901 – 1903 he formulated the theory of the chromosomal basis of Mendelism, his most noteworthy contribution to science.
After two years as foreman in the oil fields of Chautauqua County in southeastern Kansas (1903 – 1905), Sutton returned to the College of Physicians and Surgeons of Columbia University and completed the requirements of the M.D. degree (1907). He spent the following two years in a surgical house officership at Roosevelt Hospital in New York City. From 1909 until his premature death from a ruptured appendix, he practiced surgery privately in Kansas City, Kansas, and in Kansas City, Missouri.
Sutton was of impressive physical appearance, standing six feet tall and weighting 215 pounds–the basis for his nickname “Bill Taft.” E. B. Wilson  described “his quiet steadfastness and force and a quality of serenity . . .his clear, direct gaze, his self-possessed and tranquil manner.” He was elected to Phi Beta Kappa and to Sigma Xi. At the time of his death, he was an associate professor of surgery at the University of Kansas and a fellow of the American College of Surgeons. He never married.
As a farmboy, Sutton displayed great skill in the repair and operation of agricultural equipment. He also built his own camera, thereby prefiguring his later professional use of photography, and he was adept at drawing. His ingenuity was often evident in the laboratory and in his surgical practice.
In his first publication, “The Spermatogonial Divisions of Brachystola magna" , Sutton used specimens of grasshoppers that he had collected during the summer of 1899 as he rode the “header box” in the wheat fields of his father’s farm. With McClung, he discovered the value of this species for cytologic study, since the large size of its cells made it “one of the finest objects thus far discovered for the investigation of the minutest details of cell-structure.” The paper was the basis for his subsequent deductions about the role of the chromosomes in heredity.
By the fall of 1902 Sutton had been associated with E. B. Wilson for a year; they had spent the preceding summer collecting and studying marine specimens in North Carolina and Maine. Sutton’s intimate familiarity with the meiotic process, together with the expositions of Bateson, the ardent English protagonist of the newly rediscovered Mendelism, crystallized in his mind the relationship between the behavior of the chromosomes at meiosis and Mendelian segregation and assortment. (Bateson himself was slow to accept the chromosomal basis of Mendelism.) Thus, in a second paper on the chromosomes of the grasshopper , published in December 1902, Sutton wrote:
I have endeavored to show that the eleven ordinary chromosomes (autosomes) which enter the nucleus of each spermatic are selected from each of the eleven pairs which make up the double series of the spermatogonia. . . . I may finally call attention to the probability that the association of paternal and maternal chromosomes in pairs and their subsequent separation during the reducing division as indicated above may constitute the physical basis of the Mendelian law of heredity.
Cytologists had been aware for some time that at one phase of meiosis whole chromosomes separate, or segregate (the “reducing division” of August Weismann). In 1901 Thomas Montgomery had concluded that “in the synapsis stage is effected a union of paternal with maternal chromosomes.” It was known, furthermore, that in fertilization chromosomes are contributed in equal numbers by the two gametes (Van Beneden’s law).
On 19 December 1902 Wilson published a short note in Science proposing a relationship between the phenomena of meiosis and Mendel’s laws. He stated the following reason for the note: “Since two investigators, both students in the University, have been led in different ways to recognize this clue or explanation, I have, at their suggestion and with their approval, prepared a brief note in order to place their independent conclusions in proper relation to each other and call attention to the general interest in the subject.” Sutton’s fellow student William Austin Cannon, later professor of botany at Stanford University, was working with fertile hybrid cotton plants and had found separation of paternal and maternal elements in meiosis.
Wilson  later wrote: “I well remember when, in the early Spring of 1902, Sutton first brought his main conclusion to my attention. . . . I also recall that at that time I did not at once fully comprehend his conception or realize its entire weight.” Of their work together in the summer of 1902, Wilson wrote, “It was only then in the course of our many discussions, that I first saw the full sweep, and the fundamental significance of his [Sutton’s] discovery.” A year or so before, the work of several other cytologists had brought them to the verge of the chromosomal theory. “Sutton, however, was the first clearly to perceive and make it known. . . ."
Sutton’s “The Chromosomes in Heredity” (1903) is a major landmark in the biologic literature . Like his 1902 paper, it was intended as a preliminary report. Complete, although concise, it displayed model clarity and logic. Sutton built his argument on six components, of which three were corroborations of the findings or suspicions of his predecessors, while three others were uniquely his own, as was the synthesis. The six points are:
(1) That the somatic chromosomes comprise two equivalent groups, one of maternal derivation and one of paternal derivation;
(2) That synapsis consists of pairing of corresponding (homologous) maternal and paternal chromosomes;
(3) That the chromosomes retain their morphologic and functional individuality throughout the life cycle;
(4)That the synaptic mates contain the physical units that correspond to the Mendelian allelomorphs; that is, the chromosomes contain the genes;
(5) That the maternal and paternal chromosomes of different pairs separate independently from each other– “The number of possible combinations in the germ-products of a single individual of any species is represented by the simple formula 2” in which n represents the number of chromosomes in the reduced series; and
(6)That “Some chromosomes at least are related to a number of different allelomorphs . . . [but] all the allelomorphs represented by any one chromosome must be inherited together. . . . The same chromosome may contain allelomorphs that must be inherited together. . . . The same chromosome may contain allelomorphs that may be dominant or recessive independently”.
Sutton thus predicted genetic linkage and pointed out that Bateson and Saunders, in their experiments with Matthiola, had detected “two cases of correlated qualities which may be explained by association of their physical bases in the same chromosome.” (Bateson himself had an alternative and incorrect explanation.) Although Sutton observed and pictured chiasmata, their specific delineation and the suggestion that parts are exchanged between homologous chromosomes are attributed to F. A. Janssens (1909); the relation to genetic recombination was discovered by Thomas Hunt Morgan and his students.
Cytologists had suspected for fifteen or twenty years before Sutton that hereditary factors are carried by the nucleus and even by chromatin, but Sutton’s demonstration of the relationship between the behavior of the chromosomes in meiosis and Mendel’s two laws was the first strong evidence specifically in support of the theory. Wilson  wrote that subsequent to the appearance of Sutton’s papers, Theodor Boveri had stated that he himself had already arrived at the same general conclusion. Consequently, the chromosomal theory of inheritance is sometimes called the Sutton-Boveri theory.
None of Sutton’s surgical contributions rank with his main contributions to biology. He did, however, introduce colonic administration of ether for surgery of the head and neck and was further responsible for a number of minor technical innovations.
. Sutton, “The Spermatogonial Divisions of Brachystola magna,” in Kansas University Quarterly, 9 (1900).
. Sutton, “Morphology of the Chromosome Group in Brachystola magna,” ibid., 4 (1902).
. Sutton, “The Chromosomes in Heredity,” ibid., 4 (1903), 231–251. repr. in J. A. Peters, ed., Classic Papers of Genetics (Englewood cliffs, New Jersy, 1959).
. A. Hughes, A History of Cytology (London-New York, 1959).
. The anonymous “Walter Stanborough Sutton,” in Journal of the American Medical Association, 67 (1916).
. Walter Stanborough Sutton, April 5, 1877-November 10, 1916 (Kansas City, 1917), published by his family and available from the Library of Kansas University School of Medicine.
. V. A. McKusick, “Walter S. Sutton and the Physical Basis of Mendelism,” in Bulletin of the History of Medicine, 34 (1960), 487–497.
. B. R. Voeller, ed., The Chromosome Theory of Inheritance. (Classic Papers in Development and Heredity) (New York, 1968).
. E. B. Wilson, see .
Victor A. McKusick