Biographical material - Charles W. Metz

Parents and early days

My father was William Summerfield Metz (1859-1935). He was born in Williamsburg, Pennsylvania; from there he moved with his parents to Odell, Illinois where he attended public and high school. He went to law school in Illinois, receiving his L.L.B. in 1878 from Wesleyan University, Bloomington, Ill. Then he practiced law in Pontiac, Ill., until 1884, when he moved to Wyoming. He descended from one of two brothers who came over from Germany a few generations earlier. My mother, Jennie Gammon Metz, was of colonial New England stock and became a member of the D.A.R., in which she was very much interested. She and my father evidently met during his law school days. Apparently my mother’s family was one of those which migrated to the middle west in the early days, because it was located in Illinois and my mother attended a girl’s seminary there. She was a member of the Gammon family connected with the Gammon and Dearing firm of farm machinery manufacturers well known in the early days.

In 1884, a few years after my parents were married, they moved from Illinois to Sundance, Wyoming, in the Black Hills close to the Devil’s Tower. This move was influenced by my mother’s older brother, John P. Gammon, who had located in or near Spearfish, South Dakota. I was born in Sundance, but both of the families soon moved further west - our family to Sheridan, Wyoming, and the Gammon family to the neighborhood of Buffalo, Wyoming, about 50 miles away. I was two or three years old when we moved. My uncle was a breeder of black percheron horses. His ranch (the T-A) was famous for its connection with the “Cattlemen’s War”. It was here that the battle terminating that conflict took place, and as a boy I was greatly impressed by the numerous bullet holes in the logs of the wall and in the doors. This “war” as, of course, some time before my uncle bought the ranch. My father was a lawyer and for a time a judge. Being a democrat in a strongly republican territory his adventures in politics were not extensive, although he was elected to the Wyoming House of Representatives and was democratic floor leader there. He was well known throughout northern Wyoming and to a considerable extent throughout the state because of his activities as a defense lawyer. The country was pretty wild and thinly populated. Shootings were frequent, often resulting from friction between cattlemen and so-called rustlers, or between cattlemen and sheepherders who were especially resented. In cases involving shooting my father regularly kept the gun used by his client. He acquired a large collection ranging from a small pearl-handled six-shooter to a large 45-70 rifle. For years I was custodian of the collection and I remember especially one of the revolvers which had four notches cut in the handle representing four men shot.

As Sheridan grew from a small village my father and mother were both active in civic affairs including the schools, library development, and cultivation of music and the other arts. My mother aided in the establishment of the Women’s Club and such things as reading clubs in the community. From the time I was a small boy, I was given to understand that a the proper time I would go to college.

Like many of the other boys I practically grew up on a horse. In those days, we always had a team of horses and a cow. And ordinarily one or both of the horses served for riding purposes as well as carriage horses. It fell to us boys of course to care for the horses and cow, including the milking of cow. A flock of chickens also required our attention. My older brother Percy, was five years older than I and my next, Will, five years younger than I. Each in turn had the main responsibility for the animals on the place. So when my older brother went away to college it fell to me to take charge. As soon as I was large enough to run around, I learned to ride horseback. During the summer when school was not in session I was in the saddle a large part of the time.

Although very active in the law, my father had natural interest in ranching and farming, an interest going back to his boyhood. Over the years, he acquired ownership or part-interest in several ranches. One of these was at the foot of the Big Horn Mountains and there I spent a large amount of time in the summers - occasionally the whole summer. This ranch, like the others, was operated by a rancher and his family, with additional men employed as needed. Ordinarily I slept in the hay mow of the barn with the boys of the family and men employees. And of course family and workmen all ate together. One of my most memorable summers was spent on this ranch when I had the job of supervising a herd of cattle, kept at this time of year on top of the mountains. This meant that I lived mostly at the ranch but spent a large part of my time on top of the mountains or going and coming on my horse. My function was to see that our cattle remained in the area allotted to them, and of course to see that none of them got lost. Since I received no pay, I was free to spend a good deal of time at my favorite sport of trout fishing in the mountain streams which at that time were full of trout. The mountains in that region were and still are beautiful as well as rugged.

In those days buffalo heads (skull and horns) and buffalo wallows were common in the mountains (the wallows being depressions from one to three or four feet deep where the buffalo commonly wallowed in the dust). Like many other boys I accumulated buffalo skulls and the antlers of deer and elk for display at home. My great interest in those days, however, was in birds. Very early, I began collecting both eggs and skins for museum specimens. This activity I took very seriously and tried to do as a really scientific enterprise, procuring books and subscribing to bird magazines in order to make first class specimens and to keep proper records, on which I spent a great deal of time. Fortunately my parents approved of this interest, and I was able to get some advice and encouragement locally and even more through an ornithologist with whom I was fortunate enough to make contact. He was the Reverend P.B. Peabody who lived for a time in eastern Wyoming and knew the Wyoming birds. He was exceedingly kind and helpful, both through correspondence and during a visit which I paid, at his invitation, to his home. Among those in Sheridan to whom I am grateful for encouragement and aid I would mention especially Miss Lena Stover, one of my high school teachers, and Mr. Abrahams, the local taxidermist, who gave me a complete freedom in his shop. Most of what I learned about preparing bird skins for museum purposes, however, I had to secure from taxidermy books which I got through the mails. A good many of the bird skins of smaller species were later deposited with the Academy of Natural Sciences in Philadelphia together with the records applying to them. In making collecting trips I often spent entire days on horseback ranging through the open country, much of which was not even enclosed with fences.
Another special interest of mine during high school days centered around the Indians, who were numerous in Wyoming at that time and were frequently seen in Sheridan, either when passing through from one reservation to another or when coming to visit Sheridan and spend short times camping in the outskirts of the town. For the most part the visitors were Crows or Cheyennes, the former usually coming from the Large Crow Reservation in Lodge Grass, Montana.

Through the kindness of Mr. Herbert Coffeen, a friend of our family, I had a good opportunity to make close contacts with the Indians. Mr. Coffeen was a man with artistic leanings and was an unusually attractive and pleasant person. He owned and operated a modest store which served as both an Indian trading post and an art center. Although I visited the Indian Camps and became acquainted with numerous Indians on my own, it was through Mr. Coffeen, a trusted friend of the Indians, who brought me into really close contact with them. While still a small boy in late grammar school or early high school I was enabled through his influence to go to the Indian Reservation in Montana and live with the Indians during one of their annual ceremonies which extended over two or three days. During the period of ceremonies, I lived with the family of “white Arm”, one of the Crow chiefs, with whom I had already become acquainted. Of the various events which occurred during these two or three days one was perhaps the most impressive and another was probably most weird and sensational I have ever experienced. the first of these was a very solemn ceremony extending over a period of perhaps an hour and performed by a group of indian chiefs. It was referred to as “a prayer for the return of the buffalo.” It was performed at a spot some distance away from all the other Indians. the other was a series of ceremonies and dances continuing by torvchlight throughout the entire night in the middle of some dense woods and participated in by what appeared to be the entire tribe living in this locality. the Indians were all decorated and costumed for the occasion. I was the only white person present and like the other boys and men my costume consisted mostly of paint and a few decorations. The women and girls, however, wore elaborate dresses and beadowrk. Needless to say there were times when I felt plenty of chills going up and down my back.

High School
As already indicated I attended high school in Sheridan, Wyoming, entering in the fall of 1904 and graduating in the spring of 1907. The town is located at the junction of Big Goose Creek and Little Goose Creek which flow out of the east side of the Big Horn Mountains and meet at about fifteen miles from the foothills. Most of the town lies between the two streams. For some reason, not altogether logical, the high school was built at the very outskirts of town on top of a hill lying beyond Big Goose Creek. Since the places to cross the Creek were far apart we children had to walk considerable distances and climb a steep grade in order to reach the high school. With no motor vehicles either for transportation or for removing the heavy snows, getting to and from school in winter was a problem. So was the task of drying off shoes, boots and clothing.

Academically, of course, the high school in those days was pretty simple, and the teachers for the most part relatively untrained. Some of them, however, were middle aged women who had had years of experience and were devoted to their work and to their students. We all recognized the superior qualities of these teachers and had a high regard for them. Miss Lena Stover, whom I have already mentioned, was one of the best. Parenthetically let me mentiona also Miss Preston who taught in the last grammar school years and I believe also in the high school freshman year, although she was not located at the high school but rather at the grammar school which was situated in the middle of Sheridan. Other teachers ranged down to relatively young girls who had little if anything more than a high school education and were sometimes of little benefit academically. Considering the circumstances, however, I feel that we got surprisingly good training, at least in learning how to work and in developing aims and ambitions. Any students who were willing to work and anxious to learn were given every help and encouragement, both in and out of the classroom. The fact that the classes were small and we had close contact with the teachers aided greatly. Another feature worthy of note perhaps is the fact that although a frontier town, Sheridan had attracted people of almost all kinds, including some who were highly educated and cultured. An outstanding example was “Professor” von Herbst who taught German in the school and was an excellent musician. He came from a superior German family and evidently from a cultured and sophisticated environment. Why he chose to live in Sheridan was never quite clear, but the presumption was that for some reason it was desirable for him to be far away from Germany. Those of us who studied German with him in high school were very fortunate in getting a sound, basic training in the language and in addition an ability to speak German in the natural way. I learned this latter fact long afterward in Germany by being asked how I obtained my excellent pronunciation of German.

College Days
As already mentioned I learned at an early age that it was taken for granted in the case of each of us boys that when the proper times came we would all go to college. Accordingly, during my last high school year arrangements were made in my case. Having relatives on my mother’s side in and around Pasadena, California, and knowing the region well from visits out there (on one of which I was included), my parents decided that Pomona College in Claremont, California, would be the best for our purposes. Since my academic records were good and in those days there was little competition, I had no difficulty about getting accepted. In addition, arrangements were made for my younger brother, Will, to go at the same time and attend “prep” school since Pomona at that time operated a preparatory school as well as the college. So during the first year I had responsibility for my brother as well as myself in all matters that were not regulated by the school authorities. Having had no contact with colleges and having come from a small town and small school, I went to Pomona with plenty of trepidation and was distinctly uncomfortable, for several weeks at least, until I finally felt assured that I would be able to handle the college work. Pomona had a devoted faculty. Its location was perfect for a boy with my interests, lying as it did out in the foothills surrounded mainly by sagebrush, cactus and boulders with a few orange groves here and there, at the foot of “Old Baldy” Mountain, with crystal clear air and bright sunshine almost all year round (in those pre-smog days). I soon made numerous friends and with them or alone hiked for miles around pomona and up into the mountains. Claremont is located on a long gradual incline and between the town of Pomona and the base of mountains. A road led directly down the slope to the tow of Pomona. Fortunately for me I had a cousin who, with his family, lived on and operated an orange grove in the edge of Pomona town. Thanks to their hospitality this haven made an excellent place for me to visit members of my family (in the broad sense) and occasionally have a wonderful time eating fruits and home-cooked meals. this continued throughout my four years in college. The trip incidentally was made by bicycle. Automobiles were just beginning to appear on the scene in those days. I believe my first automobile ride occurred during my second year in college.

During my first year at Pomona, I took courses which were probably prescribed as parts of a general liberal arts program without anything in the biological field, because at that time I had the idea that I would ultimately go into the law or some other such profession. But before the year was over I found myself spending a good deal of time around the Zoology Laboratories - where I got acquainted with members of the department and the activities going on.

My younger brother, Will, and I roomed in a private house during this first year and if I remember correctly we took our meals in the college dining hall. During my second year, my brother was not there so I operated alone and had a room in another house in which no other students roomed. I joined with a few other students in an eating club which consisted of a tent house - very simple. I believe we used a portable gasoline stove for cooking. We got our own meals and I think we all enjoyed it even though it was very simple living. Of course the southern California climate made this kind of thing relatively easy.

By the beginning of my second year I, of course, knew my way around and was able to plan more carefully than at first. For one thing I knew that I was interested in majoring in Zoology. This brought me under the influence of two men who did more than any others outside my own family to influence or determine the broad aspects of my future professional career and, I might add, my matrimonial future. These two men were professors A.J. Cook and C.F. (Charles Fuller) Baker. Professor Cook was approaching retirement at this time and Professor Baker, who had been brought there by Cook and who was a close personal friend, was middle-aged. The latter was a brother of Ray Stannard Baker who acquired considerable fame as a writer on affairs of national concern and was, if I’m not mistaken, selected to write the biography of President Woodrow Wilson after his death. Both Cook and Baker were high minded, forward-looking men, both in a scientific sense and in the broad general sense. Cook was a mild mannered and kindly person, beloved by all the students. Baker was extremely active, dynamic and stimulating. He made a veritable beehive out of the laboratory - hitting a terrific pace himself and expecting the same from his students, whom he attracted in large numbers and to whom he was devoted, just as they were to him. I do not think I have every known a more stimulating man. He believed in cultivating individual research by students right from the beginning as part of their educational program and he had all of us who majored in Zoology working on individual investigations - which were, so far as possible, designed to explore unknown areas and to lead to publications. My first three published papers were produced during this period. On one of these (a revision of genus Prosopis in North America) I spent a great deal of time and work over a period which I believe covered at least two years. It involved, among other things, the description of numerous new species accompanied by many drawings carefully executed. This project gave me my first direct contacts with problems of evolution and the nature of species in relation to evolution.

So far as I can remember I took all the courses offered at Pomona in Zoology and practically everything in botany. At the same time I tried to include a broad liberal arts program. In looking back I can think of only one area which I should have included and did not. This is astronomy in which subject Pomona had an exceptionally good man in Professor Bracket.

As to activities outside of the classroom, I tried to get some experience in public speaking and I went in vigorously for football during my junior year when I played right half on the college team. To my surprise (for I was concentrating on work in Zoology and was not very socially minded at the time) I was elected editor of the Class Year Book “The Metate” in my junior year and President of the student body (in my senior year).

From the standpoint of benefiting students, one of the great accomplishments of Professor Baker was the establishment of the Pomona College Marine Laboratory at Laguna Beach in 1911. This he did almost single-handedly - with the full encouragement of Professor Cook. Initially, a group of us, with little if any outside support, spent the summer at Laguna completely isolated from other communities, working like beavers under Baker’s direction in quarters consisting of a rented house and two or three tents in which we worked and lived, with an Aunt of one of the students as chaperone - doing our own cooking, housekeeping and laboratory work. the shore had been untouched and abounded in nearly all kinds of marine forms, along the beach or in tide pools of nearly all sizes - an extraordinarily rich marine fauna. Each student had a specific project which resulted in an individual paper suitable for publication. Mrs. Metz (then Blanche Stafford) and I both attended this marine laboratory during the first two years of its existence. My work dealt with the tide pool fishes of Laguna Beach and led to my third publication mentioned above. This work helped to give me an introduction to the field of ecology. And as indicated below it was a primary factor in determining what I would do in my first graduate year.

Although not concerned with fishes himself, Baker foresaw the great importance of the ocean as a future source of food supply and felt that icthyology was a promising field for a young zoologist to enter as a profession. For this reason, he had suggested to me the work on tide pool fishes at Laguna and during the course of this work we discussed the possibility of going on in graduate work, first at Stanford where President David Starr Jordan had built up a department in this field and then, if possible, at Columbia University under Bashford Dean. Other possibilities were of course considered but fortunately we did not need to turn to them. Largely through Baker’s efforts I secured an assistantship at Stanford under Jordan for one year, with the understanding that my aim was to go to Columbia if possible the next year.

If I have not already made it clear, let me emphasize at this point the great importance played by Pomona College in shaping my future and preparing me for it - thanks primarily to the influence and aid of C.F. Baker.

In the fall of 1911, after the close of the summer session at the Laguna Laboratory, I went to Stanford as what might be called personal assistant to President David Starr Jordan - in his research work on taxonomy and distribution of fishes from the general area of Korea - and in his lecture course on “Bionomics” which dealt largely with problems in the area of evolution. President Jordan was a remarkable man as an individual as well as in his scientific activities, his official capacity in the university and his public activities. I got the impression that he regarded his research work in the laboratory as a release from the pressures of his other work. he obviously enjoyed thoroughly these sessions in the laboratory. It was a great privilege and pleasure to me to be able to work with him in this informal and intimate manner, with his conversation ranging over a wide variety of subjects.

It was also a pleasure to work under the direction of Professor Gilbert, Chairman of the Zoology Department and to take courses with the other members of this department and elsewhere in the university, such as Professors Snyder, Stark and Kellogg. But I will not dwell on the year at Stanford because, as I learned more about what was going on in other parts of the country, my plans and desires changed radically, so that much of the work done at Stanford did not lead directly into my later program. The efforts that Professor Baker at Pomona had made in my behalf bore fruit in the offer of a small fellowship at Columbia. Hence, after spending a second summer at the Laguna laboratory (1912) I went to New York with the expectation of working under Professor Dean on fishes, for at least the first year, inasmuch as that was the plan outlined in my fellowship application.

Actually by this time I was anxious to work in the field of experimental zoology and particularly, if possible, in the field represented at Columbia by E.B. Wilson and T.H. Morgan. I had heard glowing accounts of the chromosome work and genetic studies going on there. Needless to say, I was much disturbed by the this situation but, to my great relief, at our first conference Professor Dean immediately asked me if I did not feel that experimental zoology was a more promising field than icthyology, intimating that in his opinion this was the case. He encouraged me to change my plans accordingly and start out at Columbia on that basis. I owe Dr. Dean a tremendous debt of gratitude for this kindness for this hospitality, friendship and continuing generosity in many other ways.

At Columbia I became especially interested in the work going on under Wilson in cytology and under Morgan in genetics. During the first year (1912-3) I came to feel that cytology, heavily slanted toward genetics, would be my choice of fields and that there was an attractive area for cytogenetic studies in Drosophila. With the encouragement of both Wilson and Morgan I took up chromosome work on the Diptera with emphasis on Drosophila, focusing attention at first on the phenomenon of chromosome pairing in somatic cells which had recently been discovered by Nettie K. Stevens of Bryn Mawr. Unfortunately Dr. Stevens pioneer work had been terminated by her untimely death. The subject of chromosome pairing was of particular importance in those days because of its bearing on the emerging concepts regarding the individuality and continuity of chromosomes and the chromosome theory of heredity in general, both of which were in dispute. Work for my doctor’s dissertation in 1916 consisted of the comparative study of chromosome pairing in somatic cells of various representatives of the order Diptera. It was designed to find out (1) whether somatic pairing actually occurred in the Diptera (which was in dispute), (2) if so, the detailed nature of the process, especially whether it involved homologous chromosomes and was similar to synapsis in germ cells, and (3) whether it was found throughout the order, thus distinguishing the Diptera from other cytologically known organisms. All these questions were answered in the affirmative, thus lending support to the major hypothesis being tested.

It is hardly necessary to add that my graduate student experiences at Columbia were most inspiring, stimulating and pleasant. to work under and in close contact with Wilson and Morgan and to associate intimately with fellow graduate students such as A.H. Sturtevant, C.B. Bridges and H.J. Muller (to mention only the three with whose work mine overlapped most) was a rare opportunity whose importance has become more and more evident to me over the years. I only regret that I did not have the benefit of continued graduate work at Columbia for at least one or two more years. But I compensated for this lack to some extent by keeping in close contact with the Columbia group while I was in the Carnegie Institution Laboratory at Cold Spring Harbor, where I received an appointment and went directly from Columbia in the summer of 1914. In the Carnegie Laboratory, then known as the Station for Experimental Evolution, with C.B. Davenport as director, I was able to continue my chromosome studies, including my Ph.D. dissertation work, without interruption until after the United States became involved in World War I.

A year before the move from Columbia to the Carnegie Laboratory, Mrs. Metz (Blanche Elizabeth Stafford) and I were married at her home in Ramona, California (August 20, 1913). We had been classmates at Pomona College, both zoology majors graduating in 1911. During the year 1911-1912, while I was at Stanford, she remained at Pomona as a graduate student and assistant instructor in zoology, receiving her M.A. in June 1912. Then she went to Cornell as an assistant to Professor James Needham, where she spent the year before our marriage. After moving to Cold Spring Harbor we spent most of the winter of 1914-1915 in Cuba on a very profitable trip collecting Drosophila and other Diptera for my cytogenetic studies. In 1916, our first child was born; Charles Baker Metz, named after our Pomona professor Charles Fuller Baker.

During this period, 1913-1916, my research activities centered around three main areas. The studies dealing with somatic chromosome pairing were rounded out for the publication of my dissertation in 1916. The others, however, were continued into the 1920’s, except for a break during the war years, 1917-1918. It will be convenient to consider them as a unit and then return briefly to the war years. In connection with the work on somatic pairing, as well as separately, extensive comparative studies were made on chromosome behavior in germ cells, especially spermatogonia and spermatocytes, in a wide variety of Diptera. These emphasized particularly the aspects that bear on genetics (synapsis, crossing over, segregation, chromosome continuity, etc.) and on evolution (comparison of chromosome groups in different species and genera). In the genus Drosophila a highly interesting series of morphologically interrelated chromosome groups were found, suggesting that a basic genetic continuity is maintained by chromosomes through long periods of evolutionary time but that alterations may occur by end to end fusions of chromosomes or segments of chromosomes and conversely by the transverse breakage of a chromosome to form two chromosomes. On this hypothesis corresponding chromosomes, or chromosome arms, in related species should have similar sets of genes. To test the hypothesis a complementary set of genetic studies was required. For this purpose two favorable species were selected for comparison with the well-known Drosophila melanogaster.
The activities described above were interrupted in 1917 after we entered the war. At that time I went into anti-malaria work around army camps in the south. I was put on a deferred basis in selective service because of the heart lesion resulting presumably from rheumatic fever I had had when a child. The anti-malaria work, first in the Red Cross then in the U.S. Public Health Service continued throughout the remainder of the war. Most of it consisted of purely practical measures for control of malaria mosquitos, but I was encouraged to include simple observational and experimental studies where feasible. The most interesting of these dealt with the flight range of Anopheles crucians - a study made possible by the presence of a pond contaminated by wastes from a chemical plant to a degree favorable only to A. crucians. This provided an abundant source in a region where no other source of this species could be found; hence determining the flight range was relatively simple.

Following the close of the war I resumed my work in the Carnegie Institution. through arrangements kindly made by Professor Morgan at Columbia and Professor Davenport at Cold Spring Harbor I was enabled to spend the first winter with the Columbia University Drosophila group. At this time I was fortunate also in being authorized by the Carnegie Institution to transfer my work during the summers to the Marine Biological Laboratory at Woods Hole thus benefitting from wider associations there. Thenceforth, spending the summers in Woods Hole has been our custom.

In continuing the work after the war, and during all the later work in the Carnegie Institution, I had able assistants and associates to whom, as the publications indicate, much of the credit is due for whatever was accomplished. At first considerable emphasis was put on the studies on chromosome evolution in Drosophila. In this area we secured increasingly convincing evidence that the postulated chromosome continuity in an evolutionary sense, mentioned above, actually exists. Similar or identical mutant characters obtained in different species were found to be due to genes in apparently homologous chromosomes or chromosome arms, thus supporting the view that the basic genic make-up of chromosome tends to persist over long periods of time, and from species to species in evolution.
However, this evidence only established a probability. The only way to prove that a gene in one species was homologous with the one in another species would be cross the two species and show that the genes were allelomorphic. To carry out a program on this basis did not appeal to me, even assuming that it could be done, which was very uncertain at that time. It would almost certainly be a large and complicated undertaking and, more importantly, it would divert my activities away from cytological studies (my primary interest).
Consequently I gave serious consideration to the possibility of switching to a different type of program. Since I had long had a natural leaning toward comparative studies and exploration, I felt that someone should investigate the cytogenetics of some organism or group of organisms, showing conspicuously different characteristics from those of Drosophila. With this in mind I set about investigating the possibilities of various organisms which could be handled satisfactorily in the laboratory. Many forms were tested “on the side” while continuing our regular work. Among these were some specimens of Sciara, collected first in the pigeon houses of Dr. Oscar Riddle at the Carnegie Station. Almost immediately we detected striking cytological and genetic peculiarities in this genus. It seemed clear that some of these violated supposedly well-established principles of chromosome and genetic behavior. Further study verified this inference, with the result that Sciara work gradually replaced most of our other activities as the individual studies could be rounded out and satisfactorily terminated. As a genus representative of the lower Diptera, Sciara was well suited for purposes of comparison with Drosophila, which is a representative of the higher Diptera. But our studies soon showed that the phenomena exhibited by Sciara were primarily of interest in relation to basic problems of chromosome behavior rather than simply for comparison with Drosophila.
For example it was found that in the first spermatocyte division an accurate segregation of homologous chromosomes occurred in a unipolar mitotic figure and without any previous synapsis of the homologs. Furthermore, the segregation was highly selective in that the maternal homologs all moved toward the single pole, whereas the paternal moved in the opposite direction away from the pole, and the latter chromosomes moved backward in direct opposition to the retarding influence of the spindle fibers. All three of these aspects were upsetting, and indeed still are. None of the well-known theories as to the mechanism of mitosis or meiosis will account for the phenomena in the light of what we have subsequently learned as to their precise nature. The nature and complexity of the problem may be illustrated to some extent by considering the chromosomes which move backward away from the pole. In the first place, these chromosomes are indistinguishable in genic make-up from their homologs which move toward the pole. Secondly, these retreating chromosomes all came from the father, and int he father they were the chromosomes which went to the pole -- in other words they moved in exactly the opposite manner from their movement here. In the father they were the maternal chromosomes. Their type of movement is not determined by their genetic composition, but by the sex of the parent from which they came. They have retained, through embryonic development, some influence exerted upon them in the father which causes them at this time to respond in a manner which is the opposite of that shown by their homologs from the mother. And the material thus reversibly influenced is apparently not the ordinary genic material. As to the mechanism of the chromosome movement backward away from the pole we are equally in the dark. The chromosomes are obviously not attracted toward anything. The spindle fiber, which would normally indicate the direction and path of movement, serves here as a retarding agent that is or reflects the presence of, something that impedes the progress of the chromosome as the latter moves backward directly away from the pole. As indicated in our papers, the behavior suggests that the chromosome itself is responsible for the unusual type of movement. We have also suggested that in ordinary cells the chromosome may exert a similar activity, but oriented in the same direction as the activity represented by the spindle fiber so that the two together, and possibly other untested activities, cooperate in propelling the chromosome toward the pole. These points are noted here simply to give an example of what led us to work on Sciara. Many other phenomena exhibited by these flies likewise bear on fundamental cytogenetic problems. The years from 1924 or 25 to 1930 were very exciting ones, as the various new and entirely unexpected aspects of Sciara reproduction, sex determination, development and inheritance unfolded. By 1930 we had obtained a general picture of what the basic phenomena are like and how they are interrelated, but in some cases we still had little idea, even in a descriptive sense, as to how they were brought about.

For example we found that the egg and the sperm together transmitted certain chromosomes to the fertilized egg which were not found later in the developing larvae. It was evident that certain chromosomes were lost from the somatic line, and that one particular chromosome was regularly lost from the germ line. From analogy with Ascaris and Miastor cases, we suspected that the loss occurred during cleavage stages in the developing egg. But much work was required lated to determine the time and nature of eliminating events. The story in the soma line was revealed by Dr. Anne Marie Du Bois who came to work with us after I met her in professor Goldschmidt’s laboratory in the Kaiser Wilhelm Institute in Berlin in the summer of 1930 when I was in Europe attending two scientific congresses. Dr. Du Bois was with us in the Baltimore laboratory for two years (see bibliography for her publications). The elimination in the germ line was later identified by R.O. Berry (as part of his doctoral dissertation with me at Johns Hopkins University) and was found to be one of the most unexpected and presently inexplicable processes found in Sciara.

From the brief statement above, it is evident that in this work we were faced with a whole array of distinct but interrelated problems, any one of which could keep all of us busy for a long time. Therefore, one of our great difficulties was to select and investigate the most appropriate or promising aspects of the enterprise. It was absolutely necessary to carry on genetic work and cytological work at the same time. And in order to get at the broader significance of some of the unique phenomena it was desirable to study different species of Sciara comparatively. Numerous aspects had to be ignored or only partially investigated. And a great deal of information which did not apply directly to the work in hand had to recorded and laid aside, to be followed up later if possible. In some ways the situation was frustrating because we could concentrate sufficiently. Understanding of one aspect required knowledge of from one to several other aspects. Each chromosome had to followed, both cytologically and genetically, not only through the stages of gametogenesis but through fertilization and cleavage stages, early development within the egg including both germ line and soma, and on through later development.

1930 to 1940
In 1930 we transferred our work from Cold Spring Harbor to the Department of Embryology of the Carnegie Institution in Baltimore. This came about quite unexpectedly, as an indirect result of an invitation to me from Washington University in St. Luis to come to the zoology department there for a year or two to fill the position of Professor Frank Blair Hanson who had been invited to the Rockefeller Foundation in New York. Apparently Dr. Hanson (already a good friend of mine) did not want to commit himself until he had tried out for a year or two the work involved in the New York office of the Rockefeller. Inasmuch as I had spent approximately fifteen years at the Cold Spring Harbor laboratory and felt the need of getting into a less isolated situation, at least temporarily, I was interested in the invitation from St. Louis. Not knowing much about the department at Washington University, it occurred to me to ask the advice of Professor H.S. Jennings of Johns Hopkins with whom I am well acquainted. To my astonishment, I received in reply a suggestion from Dr. Jennings that I come to Baltimore on an arrangement whereby I would be a member of the Department of Embryology of the Carnegie Institution and at the same time would be a visiting professor at Johns Hopkins - teaching a graduate course in cytology while carrying on my research work in the Carnegie. I believe he indicated in this first letter that he had already discussed the matter with Dr. Streeter, chairman of the embryology department of the Carnegie Institution, and received a favorable response.
The upshot was that we moved on a trial basis to Baltimore in the late summer of 1930. This was a hectic time for me because I was already scheduled to attend a congress in London and then the closing sessions of another in Amsterdam, on the way to Berlin where I was slated to visit the Kaiser Wilhelm Institute for a week or two - especially the section headed by Professor Richard Goldschmidt. I was barely able to get back from Europe in time to move to Baltimore and be ready to start teaching as well as getting the research work established. The research work was practically at its height at that time. I had three assistants (one of them Dr. DuBois from Germany already mentioned). What was worse, however, was the fact that I had never done any teaching, either undergraduate or graduate, and I had to start a course in cytology with practically no time to prepare. Furthermore, in view of the fact that there had been no course in cytology available at Hopkins, the students in my first class were practically all advanced, and two of them (T.H. Sonneborn and Daniel Raffael) already had their Ph.D.’s. I was scared stiff at the beginning, but was soon relieved somewhat when Sonneborn came to me and asked if I could not take up more elementary aspects at the beginning - saying that I was in fact talking over the heads of the class. This indicates, by the way, how relatively new chromosome cytology was at that period. I practically had to begin with what is now routine first year college aspects.
Teaching and working with the graduate students proved to be stimulating and exhilarating, as I had anticipated. And of course I found that teaching cytology greatly improved my own knowledge and perspectives in the field. Believing that, in cytology, laboratory work is of prime importance, I handled the laboratory sessions myself and worked closely with the students in the laboratory - encouraging discussion there as well as in the lecture periods. To further stimulate activity on the part of the students themselves, I saw to it that each individual student once or twice each semester worked up a special topic (usually from original sources) and reported it to the class - with each report followed by free discussion.
Despite the heavy work load involved in teaching and carrying on a full research program, this first year in Baltimore proved to be very pleasant. Since it was also satisfactory to the Carnegie Institution and Johns Hopkins, the arrangement was continued. Mrs. Metz and I both found the Baltimore environment enjoyable and the Carnegie group and Johns Hopkins group of people most congenial. It seemed like an ideal situation. But unfortunately as time went on, I found myself carrying two full time jobs, because my research work in the Carnegie Institution called for full time attention and at the university I found that supervision of doctoral candidates in cytology, committee work and other university activities, together with my regular classwork, represented another full time load. Further, the fact that I was working in two widely separated laboratories meant that I had to use up a great deal of time and energy in traveling back and forth. The Carnegie Institution laboratory was housed in the Johns Hopkins Medical School on the opposite side of the city from the zoology department in the University proper and I had assistants and had work going on continually in both places. After seven years, therefore, I found the strain too great and regretfully gave up the university work, confining myself thereafter to the research work in the Carnegie. This continued very pleasantly for three years and gave me a chance to catch my breath, so to speak.
Then in 1940, I was offered the chairmanship of the zoology department at the University of Pennsylvania. Not knowing that a second world war would actually occur and that my research program would be completely upset and remain so for an indefinite time if I took on an administrative university job of this kind, I accepted that offer and we moved to Philadelphia in the summer of 1940. I might add that this was a very difficult decision to make and I consulted numerous friends before finally making it.
For the first few years after moving to Pennsylvania, with the aid of research grants from the Carnegie Institution and others, which enabled one of my assistants (Dr. Martha Bozeman) to accompany us to Philadelphia, we continued some of the research work already underway. But as we became deeply involved in the second world war, our activities were more and more disrupted. The army headquarters on the university campus was located in our building and before long we were devoting ourselves to activities adjusted to the needs of the war effort either directly or indirectly. As in other universities, retrenchments of various kinds were required and financial problems increased.
From the standpoint of research work there is little to add concerning the period up to 1955. Following the war we had the same reconstruction problems as other universities, including financial difficulties brought on by inflation, difficulty of securing competent employees for non-academic as well as academic, and other required to adjust to the changed conditions brought about by the war. I had no difficulty about securing research grants from outside the university, but my time and energy were so fully taken up by my university duties (which I regarded as my first responsibility) that I could not devote sufficient attention to research work to make suitable progress - especially when it was next to impossible to secure sufficient trained research assistants, as was the case at that time. though I had the hearty cooperation and support of the university administration, my research activities unavoidably became greatly restricted as time went on. This bothered me more and more despite the fact that I enjoyed teaching my class in cytology and advising Ph.D. candidates who chose to work with me. In 1955 I asked to be relieved of the chairmanship and its duties so I could get back to some of my research, with a salary reduction such that I would not feel under any pressure. This request was kindly granted, on a much more liberal basis than I had suggested and with a provision that I was to be free to teach or not to teach, as I chose, during the remainder of the time until my retirement in 1959. For the first year I did not teach, although of course I continued supervision of my graduate students. During the following year I taught my regular course and supervised doctoral candidates. Then in 1957, with the approval of the university, I transferred my work to the Marine Biological Laboratory in Woods Hole and we moved from Philadelphia to Woods Hole, where we remained on a year-round basis until and after my retirement at seventy as Professor emeritus. Subsequently I have continued my research work as much as possible here at the Marine Biological Laboratory under very favorable conditions. This has consisted entirely of trying to finish or round out, as many as possible of the numerous partly completed pieces of work mentioned previously in this account. These are mainly studies in which the material is already prepared and preliminary observations recorded.
Although it is obviously not feasible to mention here the many people whose cooperation, support or counsel was of great value to our department at Pennsylvania during the years I was there, I feel impelled to take this opportunity to pay tribute to a man who had no direct official connection with the department and therefore might not be thought of in relation to it. He is Dr. A.N. Richards, who was vice-president of the university in charge of medical affairs (a man well known to members of the National Academy). From the time of my arrival until my retirement many years later, his counsel, encouragement and support were invaluable to me. We were brought into close contact in many ways during the war and in numerous university activities after the war. His wisdom, judgment and keen insight into almost every matter that came up were remarkable. Combined with his equally remarkable personal qualities and his experience, these made him a most valuable and trusted advisor. I always came away from a conference with him invigorated by a feeling of benefit and uplift.