Difference between revisions of "Journal of Morphology 22 (1911)"

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THE CHARLES OTIS WHITMAN MEMORIAL VOLUME  
 
THE CHARLES OTIS WHITMAN MEMORIAL VOLUME  
 
 
 
 
  
 
EDITORIAL COMMITTEE  
 
EDITORIAL COMMITTEE  
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Edwin G. Conklin  
 
Edwin G. Conklin  
 
Thomas H. Morgan  
 
Thomas H. Morgan  
 
  
  
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By  
 
By  
  
His Pupils And Associates  
+
His Pupils And Associates
 
 
 
 
  
 
==Contents==
 
==Contents==
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Charles Zeleny. Experiments on the control of asymmetry in the development of the serpulid, Hydroides dianthus. Seven figures 927  
 
Charles Zeleny. Experiments on the control of asymmetry in the development of the serpulid, Hydroides dianthus. Seven figures 927  
 +
 +
{{Ref-Locy1911}}
  
 
William A. Locy. Anatomical illustration before Vesalius. Twenty-three figures 945  
 
William A. Locy. Anatomical illustration before Vesalius. Twenty-three figures 945  
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E. H. Harper. The geotropism of Paramoecium. Five figures 993  
 
E. H. Harper. The geotropism of Paramoecium. Five figures 993  
  
Elliot Rowland Downing. The formation of the spermatophore in Arenicola and a theory of the alternation of generations in animals. Four  
+
Elliot Rowland Downing. The formation of the spermatophore in Arenicola and a theory of the alternation of generations in animals. Four plates and seven text figures 1001
plates and seven text figures 1001  
 
 
 
 
 
  
 
==Biography of Charles Otis Whitman==
 
==Biography of Charles Otis Whitman==

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Journal of Morphology 22 (1911)

J Morphol. : 1 - 1887 | 2 - 1888-89 | 3 - 1889 | 4 - 1890 | 5 - 1891 | 6 - 1892 | 7 - 1892 | 8 - 1893 | 9 - 1894 | 10 - 1895 | 11 - 1895 | 12 - 1896 | 13 - 1897 | 14 - 1897-98 | 15 - 1898 | 16 - 1899-1900 | 17 - 1901 | 18 - 1903 | 19 - 1908 | 20 - 1909 | 21 - 1910 | 22 - 1911 | 23 - 1912 | 24 - 1913 | 25 - 1914 | 26 - 1915 | 27 - 1916 | 28 - 1916-17 | 29 - 1917 | 30 - 1917-18 | 31 - 1918 | 32 - 1918 | 33 - 1919-20 | 34 - 1920 | 35 - 1921 | 36 - 1921-22 | 40 - 1928 | 47 - 1929 | 51 - 1931 | 52 - 1931 |
Historic Journals: Amer. J Anat. | Am J Pathol. | Anat. Rec. | J Morphol. | J Anat. | J Comp. Neurol. | Johns Hopkins Med. J | Ref. Handb. Med. Sci. | J Exp. Zool.
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Charles Otis Whitman

Founded by C. O . Whitman


EDITED BY

J. S. KINGSLEY

Tufts College, Mass. with the collaboration of Gary N. Calkins T. H. Montgomery

Columbia University University of Pennsylvania

W. M. Wheeler William Patten

Bussey Institution, Harvard University Dartmouth College

Edwin G. Conklin

Princeton University


VOLUME 22 1911

THE WISTAR INSTITUTE OF ANATOMY AND BIOLOGY PHILADELPHIA


WAVERLY PRESS BALTIMORE, U S. A.


Prefatory Note

In 1909 a number of friends of Professor Charles Otis Whitman planned a volume of the Journal of Morphology as an acknowledgment of the debt of American science to him as the founder and editor of the Journal.

A committee, consisting of Frank R. Lillie, Edwin G. Conkhn and Thomas H. Morgan, was appointed to receive contributions of articles from his former students and his associates of the Marine Biological Laboratory. It was decided that the numbers for the year 1911 should constitute a 'Whitman Volume.'

Professor Whitman died in 1910 before the first number was issued and the volume becomes a memorial to one who had a wide influence in the elevation of biological science.

As Whitman's ideals were broader than mere morphology, so the volume in the scope of its contents overlaps that field on all sides.

The articles which were accepted by the committee were so numerous and extensive that not all of them could be published during the present year, although the volume is far larger than usual; consequently some of them will appear, with proper acknowledgment, in the next volume of the Journal of Morphology.


THE CHARLES OTIS WHITMAN MEMORIAL VOLUME

EDITORIAL COMMITTEE

Frank R. Lillie Edwin G. Conklin Thomas H. Morgan


To The Memory Of Charles Otis Whitman 1842-1910

This Volume Is Dedicated

By

His Pupils And Associates

Contents

1910

No. 1. MARCH

Bennet M. Allen. The origin of the sex-cells of Amia and Lepidosteus. Twenty-seven figures 1

Leo Loeb. The cyclic changes in the ovary of the guinea pig 37

Edmund B. Wilson. Studies on chromosomes. VII. A review of the chromosomes of Nezara; with some more general considerations. Nine figures. One plate 71

C. B. Davenport. The transplantation of ovaries in chickens HI

W. J. Moenkhaus. The effects of inbreeding and selection on the fertility, vigor and sex ratio of Drosophila ampelophila 123

G. H. Parker. The mechanism of locomotion in gastropods. One figure 155


No. 2. JUNE

C. M. Child. The regulatory processes in organisms 171

Lorande Loss Woodruff. Paramecium aurelia and Paramecium caudatum. One figure ' 223

E. A. Andrews. Male organs for sperm-transfer in the crayfish, Cambarus affinis; their structure and use. Four plates and thirty-one text figures. . 239

Wallace Craig. Oviposition induced by the male in pigeons 299

William Morton Wheeler. The ant-colony as an organism 307

Oilman A. Drew. Sexual activities of the squid, Loligo pealii (Les). Four plates 327

Frank R. LiLLiE. Studies of fertilization in Nereis. I. The cortical "changes in the egg. II. Partial fertilization. One double plate 361

W. E. RiTTER and Myrtle E. Johnson. The growth and differentiation of the chain of Cyclosalpa affinis (Chamisso). Four plates 395

Oscar Riddle. On the formation, significance and chemistry of the white and yellow yolk of ova. Three plates . 455


No. 3. SEPTEMBER

Charles W. Hargitt. Some problems of eoelenterate ontogeny. Three plates and three text figures 493

Victor E. Shelford. Physiological animal geography. Nineteen figures. . . 551

R. M. Strong. On the olfactory organs and the sense of smell in birds. Two plates and four text figures 619

Henry H. Donaldson. On the regular seasonal changes in the relative weight of the central nervous system of the leopard frog. Five charts. . . 663

Ralph S. Lillie. The physiology of cell-division. IV. The action of salt solutions followed by hypertonic sea-water on unfertilized sea-urchin eggs and the role of membranes in mitosis. Three figures 695

ThOs. H. Montgomery, Jr. The spermatogenesis of an hemipteron, Euschistus. Five plates— 147 figures 731

Winterton C. Curtis. The life history of the Scolex polymorphus of the Woods Hole region. Thirteen figures 821

No. 4. DECEMBER

H. H. Newman and J. Thomas Patterson. The limits of hereditary control in armadillo quadruplets : A study of blastogenic variation. Five figures and eight plates 855

Charles Zeleny. Experiments on the control of asymmetry in the development of the serpulid, Hydroides dianthus. Seven figures 927

Locy WA. Anatomical illustration before Vesalius. (1911) J Morphol. 945-

William A. Locy. Anatomical illustration before Vesalius. Twenty-three figures 945

S. J. Holmes. Minimal size reduction in planarians through successive regenerations 989

E. H. Harper. The geotropism of Paramoecium. Five figures 993

Elliot Rowland Downing. The formation of the spermatophore in Arenicola and a theory of the alternation of generations in animals. Four plates and seven text figures 1001

Biography of Charles Otis Whitman

Embryology History - Charles Whitman

Charles Otis Whitman 1842-1910

Five portraits xv


Charles Otis Whitman 1857

Enlaregment from a tin-type. Kindness of Professor E. S Morse

Charles Otis Whitman

Frank R. Lillie


Charles Otis Whitman was born December 14, 1842, in Woodstock, Maine, He died at his home in Chicago, December 6, 1910. On his father's side^ he was descended from Jacob Whitman, who was a resident of Bridgewater, Massachusetts, whence he emigrated to Buckfield, Maine. Three sons of Jacob Whitman settled in Woodstock, Maine, about the beginning of the nineteenth century, among them Joseph (born September 30, 1783), the grandfather of the subject of our sketch. Whitman's father (born February 19, 1821) was the eighth of ten children; his mother (born December 12, 1823) was Marcia Leonard, daughter of Solomon Leonard, also of Woodstock. Whitman married Emily Nunn of Peru, Ohio, in August, 1884, and had two sons, Francis, born in Milwaukee, Wisconsin, and Carroll, born in Worcester, Massachusetts.

His early life was spent in Woodstock, though his father removed to Waterford for a while, subsequently returning to Woodstock. He attended the town schools in Woodstock and Waterford and fitted for college at Norway and other academies, teaching winters to obtain the means for paying his school expenses." "He early developed a taste for natural history, and while here (Woodstock) and a boy, he procured and mounted a very fine collection of the birds of Maine. So artistically prepared where they, and so naturally mounted, that they attracted much attention among ornithological students." ^


^ In the preparation of this brief biography I have been indebted for information to Mrs. Whitman, Dr. Wallace Craig, Prof. H. H. Donaldson, Prof. E. S. Morse, Prof. Cornelia M. Clapp, George T. Little, the librarian of Bowdoin College, Mrs. Cornelia Fletcher Day (Westford, Mass.), Mrs. Sarah H. Trumbull (Beverly, Mass.), Mrs. Helen Keith Frost (Westford, Mass.), Secretary of the Boston School Committee, Headmaster English High School, Boston, Mass., Prof. E. L. Mark, Edward Phelps AUis, Jr., Dr. Reinhardt Dohrn, and others. I must also express my indebtedness to E. G. Conklin and T. H. Morgan, for criticism of the manuscript. The section dealing with Whitman's scientific work was prepared by T. H. Morgan, E. G. Conk in, J. Percy Moore and others — See foot-note p. xlvii.


It would be interesting to know more of his early life but Professor Whitman rarely spoke of it, though he referred at times to work on his father's farm. In reply to the question whether he had been interested in natural history as a boy, he replied to Professor Wallace Craig that he judged he must have been, because of his persistence in getting his grandfather to tell hunting stories. He never tired of the stories, and often walked a mile to have an evening of them; his grandfather was very kind in always telling these when asked. He also said that he kept pigeons as a boy, and was fascinated by them and sat and watched them by the hour, intensely interested in their feeding, their young, and in everything that they did.

We thus get a distinct though undetailed view of a boyhood spent on a New England farm, an education acquired by dint of labor and self sacrifice, and of an original interest in natural history, shown in his observation of pigeons and his collection of the birds of Maine.

He entered Bowdoin College as a sophomore in September, 1865, and graduated with the degree of Bachelor of Arts in July, 1868. The college curriculum of this time was the usual course of required studies with much emphasis on the classical languages, some study of modern languages and of mathematics, the elements of philosophy and a variety of sciences taught no doubt mainly from text-books. The influence of this classical education remained with him all his life, and was no doubt responsible for the views that he entertained in favor of the requirement of Latin for college education. There was certainly little to stimulate his interest in the field in which he subsequently won distinction. His membership in the Greek Letter Society Delta Kappa Epsilon, in the Athenaeum Society (literary), and in the Philologian Society (debating) may help to indicate his social and intellectual interests at this period. At his graduation he ranked about ninth in a class of twenty-three. The title of his commence 2 See Lapham, William B. "History of Woodstock, Me., with family sketches and an appendix." Portland, Stephen Barry, Printer, 1882.


BIOGRAPHICAL SKETCH XVll

ment oration Free Enquiry" indicates already an unfettered mind.

On graduating from Bowdoin, Whitman was appointed principal of Westford Academy in Westford, Massachusetts. He began to teach there on December 16, 1868, and remained until the spring of 1872. He must have taught a great variety of subjects, to judge by the catalogue of 1872, as there was a four years' course involving mathematics, English, Latin, Greek, French, geography, book-keeping, history, natural philosophy, chemistry, mental philosophy, astronomy, physiology, and botany, and there were but two assistant teachers and ninety pupils in 1871-72. However, he continued his interest in birds and taxidermy, and the library of the Academy still has a good collection of Westford birds prepared by a lady whom Whitman instructed in the art while there; it also contains a fine specimen of one of the largest of Maine loons set up by Whitman himself.

During the school year 1871-72 Whitman substituted in the Enghsh High School in Boston, Massachusetts, and was regularly appointed sub-master in September, 1872. At that time the departmental system had not been introduced into the school and he taught general high school subjects. He remained with the school until the summer of 1875.

While in Boston Whitman came under the influence of Louis Agassiz, and was one of the fifty students who, in July and August, 1873, attended the Anderson School of Natural History founded by Agassiz on the island of Penikese. Here he met Professor E. S. Morse, who was an instructor under Agassiz, a circumstance which had a great effect in Whitman's later life, leading to his call to the University of Tokyo as related further on. Professor Morse was much attracted to him by the beautiful and accurate way in which he drew the lower forms of life, particularly the Ascidian Perophora, on which Morse himself was working at the same time. Morse and Whitman remained the best of friends throughout life, and at Whitman's invitation many years later Morse delivered several lectures at the Marine Biological Laboratory.


Louis Agassiz died in December, 1873, and the Penikese school was opened again in 1874 for the last time by his son, Alexander Agassiz. Whitman was again one of the privileged fifty who worked there, though ninety other applicants had to be refused admission for lack of accommodations. The Penikese school started a tide of biological work at the sea-shore in American which ebbed indeed for a while, but began to flow again in the decade of the eighties and has been running stronger ever isnce. No doubt the germ of the Marine Biological Laboratory, Whitman's most significant scientific enterprise, was implanted there in Whitman's heart and in the hearts of others. Doctor Craig states that Whitman felt that he got his first start in scientific zoology from Agassiz, but that he did not really get under way until he worked with Leuckart on Clepsine in Germany. Asked by Doctor Craig (in August, 1910) what he thought of Agassiz's method, Whitman replied that he did not think much of it at first but that as time went on he thought more and more of it. We are apt to do the work for the student too much. What we should do is to set him a problem and let him work it out."

In 1875 Whitman decided to go to Germany to study natural history. Apparently he had not yet decided to abandon his career as teacher in the high school, for he left open the possibility of returning to his position after a year's absence. He sailed in July, 1875, and settled in Leipzig. From there he wrote to his successor in Westford Academy, Mr. William E. Frost, May 28, 1876: Mr. Seaver (the head-master of the Enghsh High School at that time) says he will secure my re-election and another year's absence if possible. I have not much doubt of his ability to do this. At any rate I shall remain another year." But when 1877 arrived he was not yet ready to leave and he decided to remain a part, at least, of a third year. In 1878 he received the degree of doctor of philosophy from the University of Leipzig, and sailed for America in July of the same year, although he still wished "to remain a little longer in Deutschland, but the Fates


5 Letter to Mr. Frost, Jul.v 6, 1878.


In 1878 he published his first scientific paper The Embryology of Clepsine" in the Quarterly Journal of Microscopical Science, vol. 18, pp. 215-315. This was his doctor's thesis; in many respects it was a very notable, indeed epoch-making work. It was the first time that the primordia of any ectodermal organs had been followed to individual cells, and that the cleavage process itself had been adequately interpreted as a process of 'histogenetic sundering.' He laid emphasis on the existence of embryonic axes in the unsegmented egg, and anticipated to a considerable extent views that did not receive adequate recognition until the period of study of ' cell-lineage' began about fifteen years later.

On his return to America he was appointed Junior Master, first grade, of the English High School in Boston, teaching English, and resigned in 1879. It is evident that now for the first time, at the age of nearly thirty-seven, he had irrevocably decided to devote himself entirely to zoology, for by his resignation he burned his bridges behind him. He received an appointment as fellow in biology in Johns Hopkins Uni-^^/?rsity for 1879-80, but he did not enter on the fellowship, having in the meantime accepted the chair of zoology in the University of Tokyo. He sailed for Yokohama, August 21, 1879. On the voyage he made observations on the flight of flying fish which he described in the American Naturalist, vol. 14, 1880, maintaining that their course through the air is actual flight.

WHITMAN IX TOKYO

With his appointment to the University of Tokyo in 1879, begins Whitman's real influence as a teacher and organizer in zoology. He was then nearly thirty-seven years of age and had passed through a most varied preparation for his life-work. He had pushed on without haste but without rest, always carrying with him his original and vital interest in living things since he first studied pigeons as a boy, in spite of the necessity of earning a livelihood by teaching school. He thus came to his chosen life work in full maturity with a mind broadened by varied experiences, yet with actual boyish enthusiasm and interest, that never left him throughout fife.


The work in zoology in the Imperial University of Tokyo was first organized by Professor E. S. Morse, who was invited from abroad in 1877. ^ He remained there two years and was succeeded in 1879 by Professor Whitman. Professor Iwakawa states that Professor Huxley was first invited by Professor Morse to accept the chair as his successor. Professor Huxley wrote that for years he had been desirous of studying biology in oriental countries and that the present call from Tokyo was the best chance he could ever have; however, he regretted that the dechning condition of his health would not allow him to accept. Professor Morse states in a letter that while instructor at Penikese he had known Whitman and was much impressed by the beauty and accuracy of his work ; his experience as teacher in Boston was also a recommendation; so Professor Morse secured Whitman's call to the chair of zoology in Tokyo and it was accepted. Professor Whitman remained in Japan for two years until 1881. He had only four students, but as all became professors of zoology in the Imperial University he may be xu -,tly regarded (as Dr. Takahashi states) as the father of zoolo'^Ty in Japan. Professor Iwakawa says that Professor Whitman's teaching really laid the foundation of modern zoology in Japan.

It is impossible to reproduce the tone of affection and reverence in which these reminiscences are written by two of his original pupils, Iwakawa and Ishikawa, and a later student during the Chicago period, Takahashi. Professor Iwakawa says, "Once he was my teacher while he was in Japan and since then until today I have been paying respects and admiration both for his character and for his work in biology." "I am constrained by what I regard as a duty to him to let others get a glimpse of what I knew him to be while he was with us in Tokyo" — and the whole tenor of his reminiscences is one of affectionate admiration and devotion. Professor Whitman's attitude of mind toward his

  • In the Magazine of Zoolog}-, published by the Zoological Society of Japan,

Tokyo, vol. 23, no. 269, March 15, 1911, there appear three articles on Professor Whitman, the first by Professor Tomotaro Iwakawa, the second by Professor Chiyomatsu Ishikawa, and the third by Dr. Katashi Takahashi. For the translation of these articles I am indebted to Dr. Shigeo Yamanouchi. They form the basis of the following account.



Charles Otis Whitman

1882

From Lapham's History of Woodstock, Maine,


IP


BIOGRAPHICAL SKETCH XXI

pupils was such as a mother toward her son." Professor Ishikawa writes in a similar spirit throughout. He says, "On receiving the tidings of Professor Whitman's death I am very much surprised and bitterly mourned." "I mourned bitterly in the recollection that those delightful days we had together shall never again be realized, but have now become a memory." "The work he has done during his life still remains and will be remembered forever." Takashahi says, "As he was the teacher of our professors, he will be justly regarded as our father of zoology in Japan. I feel as if I had lost my grandfather because of his being the teacher of our professors and because of his cherished kindness shown to me as a father might have shown to his son during my stay in his laboratory in the University of Chicago."

The following incident, as related by Professor Iwakawa and translated by a Japanese friend, is worth quoting:

For the purpose of making bird specimens for the museum, the University secured two government licenses in hunting seasons and the licenses were handed to the Zoological Department for the use of the students.

To make the specimens was one of purposes of hunting and the other end seemed to eat flesh of birds. One Saturday, a number of pigeons was brought to our laboratory and the next day being Sunday, some of us came to the laboratory to have the share of feast. Dr. lijima dissected the birds. A fire shovel was cleansed and put lard on and then flesh; then put into the stove to fry. Salt, sauce, knife and fork were ready and the party waited to have the flesh cooked. Dr. Sasaki had belatedly come. Being he was an hearty eater, the party who were already there refused to add him into the company and so all went to Dr. Whitman's office and locked the door from inside so that he could not get in. Unexpectedly Dr. Whitman came, in spite of that the day was Sunday. He put his slippers on as usual and tried to get in his office by the door he used to enter. To his surprise the door was locked. He came over to our laboratory and there Dr. Sasaki sat alone. Professor Whitman tried to open the door that leads to his office from our laboratory. Again to his surprise the door was also locked. Dr. Sasaki being left very much uneasy, called out, 'Professor Whitman has come!' The party inside the door, including Dr. lijima, believing that Professor Whitman would never come on Sunday and that the warning might be Dr. Sasaki's stratagem to induce the party to open the door, took the alarm easy and were chattering over quite noisily. Then there was heard a voice from outside, 'Who are in the room?' Evidently that was Whitman's. Frightened all at once the party fled


XXU CHARLES OTIS WHITMAN

to a court from the door which leads to the corridor. The party now in the court, sent a spy to look after what Professor Whitman was doing and assured that he, taking a few things, has gone home. As the coast was clear, the party returned to the laboratory. Dr. Sasaki who had been left alone in the laboratory, had already helped himself alone with the fried birds that were left in the stove, and was sitting quite satisfied. He spoke smilingly to the party, when they entered in. 'I was very much embarrassed indeed ! The handle of the shovel was peeping outside the mouth of the stove. Fried birds were making cooking noise inside and a tempting odor was ejecting from within. But our teacher had not asked a bit. Simply he said, ' who are that ran out of the room?' So I answered, 'I do not know.' 'At any event I thank you for your kind feast.' Dr lijima seemed very much disappointed; as yet the case being as such that Dr. Sasaki only cannot be blamed at, and he spoke to Dr. Sasaki, 'You lucky fellow.' The following day we were in a constant concern and reluctantly expected some sort of punishment on our conduct of the previous day. On the contrary Professor Whitman didn't even ask a word about what thus happened the previous day. Professor's Whitman's attitude of mind toward his pupils v/as such as mother toward her son.

There were but two rooms devoted to the department of zoology in the Imperial University in Whitman's day; literature and apparatus were very scanty and Whitman first introduced modern laboratory equipment and methods in microscopical technique. His four students were Ishikawa in the first year class, lijima and Iwakawa in the second year, and Sasaki in the third year. There was an assistant, a janitor and two artists, all of whom were kept busy collecting and drawing leeches. It was characteristic of Whitman that he should set each of his four students to work on a special problem for research, even Ishikawa in his first year of zoology.

Hard work was the order of the day in Whitman's laboratory; he set the example himself, and the students, who lived in a dormitory near by, often worked until midnight. Twice a day Whitman consulted with each student about his work. In the absence of a University biological library Whitman placed his own journals and books at the disposal of the students and aided them in translating German and French. He kept each man close to the study of his individual problem and deplored the wasting of time spent on other subjects. From time to time he delivered lectures on special topics and as a general course he expounded Spencer's Principles of Biology.


BIOGRAPHICAL SKETCH XXHl

At the end of two years each of the four students had a paper ready for pubhcation and Professor Whitman presented them to the Journal of the College of Science of the Imperial University for publication; the officials in charge of the journal replied that, as the journal was organized for the purpose of publishing the researches of professors, any theses of students worth publishing should be published under the name of the professor. This aroused Professor Whitman's indignation, and he withdrew the papers, remarking that he would never again present papers to the University for publication. He then sent three of them to the Quarterly Journal of Microscopical Science where they were published. Sasaki's paper on Salamander was published after Whitman's departure in the Journal of the Science College.

This incident seems to have been the beginning of an estrangement between Whitman and the administration of the University, which was aggravated by the inability or unwillingness of the administration to accede to many of his requests for more adequate equipment for the department. The period of his appointment having come to an end in 1881, the University requested him to remain, but the proposal was refused and in August, 1881, he left Japan without bidding formal farewell to the University. He published a short brochure entitled "Zoology in the University of Tokyo" shortly before he left, but that he had no intention of wantonly hurting Japanese susceptibilities is evident, as Professor Ishikawa states, from a thorough study of it. He had made a close study of the Japanese and he discovered and pointed out their most obvious weak points in an honest and essentially friendly fashion. Professor Whitman loved Japan and sympathized with the Japanese. That his love and sympathy poured forth to the Japanese in a degree far surpassing any ever shown to us was marvelously evidenced at the time of the RussoJapanese War" (Ishikawa). Indeed, those of us who were with Whitman at this time knew that he could not have suffered more keenly in the misfortunes or rejoiced more in the triumphs of his own country. This was fully realized by the Japanese people and the slight unpleasantness of his departure was soon forgiven.


XXIV CHARLES OTIS WHITMAN

IN EUROPE

Professor Whitman left Japan in August, 1881, and from November 11, 1881, to May 2, 1882, he worked at the Zoological Station of Naples as guest of Professor Dohrn. His sojourn in the Zoological Station laid the foundation of an everlasting friendship with Dohrn, and, when he left, Professor Dohrn gave him a testimonial recommending him strongly to some professorship. While at Naples Whitman studied the embryology, life-history, and classification of Dicyemids and wrote a paper on the subject, published in January, 1893, which is still the standard work of reference.

Whitman had thus come under the influence of three of the great leaders of his time in zoology, Agassiz, Leuckart and Dohrn. His original bent in the direction of the natural history of birds was diverted by these experiences towards the study of marine life and lower organisms, but later on he returned to his original interests in birds, particularly pigeons, with a mind deepened by intimate acquaintance with the fundamental problems of biology.

After leaving Naples he went to Leipzig where he remained until the middle of September, engaged among other things in preparing his Naples work for publication. On September 1, 1882, he wrote to Mr. Frost: I leave for America on the 15th of September and shall go to Leonard's (Newton Highlands) and shall hope to see you somehow or somewhere. I have not yet decided where to spend next winter. There is some possibility of my going to Johns Hopkins — though nothing definite yet. Have just finished manuscript of work done in Naples, and a portion is already in print."

AT HARVARD

In the autumn of 1882 he was appointed Assistant in Zoology at the Museum of Comparative Zoology of Harvard University, and held this position until 1886. In the spring of 1883 he went to Key West, Florida, to secure for Mr. Alexander Agassiz material with which to complete Mr. Agassiz's monograph on The Porpitidae and Velellidae." Though he spent six weeks there he did not meet with success. In the summer of 1883 he worked at


BIOGRAPHICAL SKETCH XXV

Mr. Agassiz's Newport Laboratory on the development of pelagic fish eggs. Some of the results, worked up later in Cambridge, were published with Mr. Agassiz in two papers (1884 and 1889). In the first paper the origin of the periblast was correctly described for the first time, a most important contribution in view of the confusion of opinions on this subject. During this summer he met his future wife, Miss Emily Nunn, who was also working at Mr. Agassiz's laboratory.

The years 1883-1886 were productive years; during part of this time Whitman edited the department of Microscopy" of the American Naturalist. He worked out and published his papers on A Rare Form of the Blastoderm of the Chick" ('83), External Morphology of the Leech" ('84), "On the Development of Some Pelagic Fish Eggs" ('84), "Segmental Sense-Organs of the Leech" ('84), "The Leeches of Japan" ('86), "The GermLayers of Clepsine" ('86), and some minor papers. He also prepared and published his book on "Methods of Research in Microscopical Anatomy and Embryology" ('85) (see Bibliography).

THE LAKE LABORATORY AND THE FOUNDING OF THE JOURNAL OF MORPHOLOGY

From 1886 to 1889 Whitman acted as director of the Lake Laboratory at Milwaukee, Wisconsin,^ founded by Edward Phelps Allis, Jr. Mr. Allis had decided to start a laboratory for biological and related research and Whitman was recommended to him as a proper person to take charge. There followed a conference in which the plans and purposes of the laboratory were discussed, and Whitman then presented the need of an American journal for publication of zoological research, pointing out that American workers were obliged either to present their papers to some scientific society or to send them for pubhcation to some one of several European journals. Whitman then asked Mr. Allis if he would consider the publication of such a journal, in connection with the laboratory. He was asked to submit figures and

^ I am indebted to Mr. Allis for some of the information on which the following statements are based.


XXVI CHARLES OTIS WHITMAN

plans, and it was finally arranged that he should come to Milwaukee, take charge of the laboratory, to be known as the Lake Laboratory, «and also edit with the cooperation of Mr. Allis, a journal to be called the Journal of Morphology. The journal was to be a model of publications of the kind.

Whitman may not have been the first to realize the need of establishing a journal of zoological and anatomical science in America, but he was the first to possess sufficient courage, energy and influence to set about realizing the need. He was fortunate indeed to find a man of scientific attainments and enthusiasm with an ample and liberal purse to support him in this undertaking. In the introduction to the Journal Whitman wrote, "The mixed character and scattered sources of our publications are twin evils that have become intolerable both at home and abroad. The establishment of the Journal of Morphology may not be the death blow to these evils; but there is hope that it will, at least, relieve the more embarrassing difficulties of the present situation."

In its make-up both scientific and typographical, the Journal of Morphology was a model of what a research publication should be, and it did much to coordinate zoological research in America, to give it a worthy setting, and to make it better known abroad. Eighteen volumes were published between 1887 and 1903, always at considerable financial loss, and its publication was then suspended for a while in spite of Whitman's efforts to secure the needed support. The American Journal of Anatomy and The Journal of Experimental Zoology, begun in the period of su"spension of the Journal of Morphology, did not, however, suffice for the growing needs of zoological and anatomical science, and the Journal of Morphology was taken up again by The Wistar Institute of Anatomy and Biology in Philadelphia, in 1908, and its publication has continued ever since. As Professor Mall says, "The Journal of Morphology served as a model for many of our scientific journals, both biological and medical, which have come into existence during the past twenty-one years. The importance of sound scientific journals to anatomical and zoological science is now clear to all, and both anatomists and zoologists


BIOGRAPHICAL SKETCH XXVll

owe to Professor Whitman a debt of gratitude for having been the pioneer in this field" (Anatomical Record, vol. 2, 1908, p. 381).

In 1898, realizing the need of some means for more rapid publication than was afforded by the Journal of Morphology, Whitman started the Zoological Bulletin with the cooperation of W. M. Wheeler. The idea was to afford means for the rapid publication of shorter articles and preliminary notices dealing with investigations in zoology which required only simple illustrations. The Bulletin was therefore published monthly. It was intended to be a companion serial to the Journal of Morphology. After the publication of two volumes the name was changed to the Biological Bulletin and it was transferred to the Marine Biological Laboratory as its official publication.

At the Lake Laboratory Whitman was associated with Edward Phelps Allis, the founder, Howard Ayers, William Patten, A. C. Eycleshymer, and some others. The work of the laboratory was research work in morphology, especially embryology. Whitman himself began investigations on Amia and Necturus, but though he carried some of this work quite far, but little of it was ever published. His scientific activity during this time may be inferred from the hst of publications covering the period 1886 to 1889.

AT CLARK UNIVERSITY: 1889-1892

In 1889 Whitman accepted a call to the chair of zoology in the newly founded Clark University of Worcester, Massachusetts. Professor G. Stanley Hall of Johns Hopkins University had sought to establish with the aid of Jonas Clark of Worcester, a strictly graduate and research institution, which should accomplish all that the Johns Hopkins University had set out to do in elevating the standard of scholarship in America, but without the hindrance of undergraduate instruction. Whitman met there with thoroughly congenial conditions and associates. President Hall had assembled a small but remarkable group of scientific men, all animated by the same high ideals of scholarship. They were unencumbered with undergraduate instruction, provided with fairly adequate means for research, and they seemed destined to realize the fine aim that President Hall had set before them.


XXVlll CHARLES OTIS WHITMAN

Whitman's teaching career, interrupted since he left Tokyo eight years before, was now resumed, and continued to the time of his death. A small body of research students was attracted to him, who carried on their work in Worcester during the academic year and at the Marine Biological Laboratory in Woods Hole during the summer. Whitman's laboratory was a paradise to the properly qualified research worker. There was practically no set instruction and the student's liberty was complete in all respects, but a spirit of hard work and complete absorption in the fundamental problems of biology prevailed. The problems of biology were the true topics of the day and, when the zoological club met, such subjects as Darwinism and Lamarckism were discussed with a fire and enthusiasm comparable to the most intense political or rehgious controversies. The main business of each student was his research problem, a secondary business was the preparation of some subject set for presentation at the zoological club, and the animated discussion of fundamental problems of biology prevented too much narrowness. Students read much and thought much because they had both time and inclination, and were not subject to trivial academic demands.

Whitman had a great respect for the intellectual independence of his students. He set them worthy problems but left the working out to the student ; he was at the same time their severest and most friendly critic. He maintained their courage through difficulties, rejoiced with them in their discoveries, and always acknowledged their complete ownership in their results. He required convincing proof of each statement, and one could feel sure that whatever passed him would stand. He was completely loyal to them in all relations, and it is characteristic that the main event which finally induced him to resign and move to Chicago was an act of the administration which he regarded as an injustice to one of his students. He was not alone in his displeasure with the administration, though the causes were various and the departments of physics and chemistry, zoology, anatomy, neurology, and palaeontology of the new University of Chicago were organized by seceders from Clark University in 1892.


BIOGRAPHICAL SKETCH XXIX

PROFESSOR WHITMAN AND THE MARINE BIOLOGICAL LABORATORY

The organization of the Marine Biological Laboratory was a response to the same demand that established and maintained a marine laboratory on the island of Penikese in 1873 and 1874. In his address at the opening of the Marine Biological Laboratory Professor Whitman said:

The Annisquam Laboratory, the immediate predecessor of this, was organized to serve the same ends as the Penikese School, and the forces there engaged have simply been supplemented and transferred to the new Marine Biological Laboratory of Woods Hole, with such changes only as circumstances have rendered necessary. It was through the generous support and active cooperation of the Woman's Education Association of, Boston that Professor Hyatt was able to maintain the Laboratory at Annisquam, and the same Association initiated and carried through the movement that has given us this Laboratory.

In 1886 efforts were made by the Association to place the Annisquam Laboratory on an independent and broader foundation. A circular letter sent to many of the leading biologists of the country received encouraging repHes and accordingly a preliminary meeting was held on March 5, 1887, in the library of the Boston Society of Natural History. A committee was there organized to perfect plans for the organization of a permanent sea-side laboratory, to elect trustees and to devise ways and means for collecting the necessary funds. The committee met with sufficient success for a modest beginning and accordingly in March, 1888, the Marine Biological Laboratory was formally incorporated with ten members. Seven trustees were chosen at a meeting of the Corporation held the same month. In June, 1888, the Trustees issued a circular in which they announced the poUcy of the Laboratory to support instruction as well as research, and invited the cooperation of the universities and colleges of the country. Professor Whitman's appointment as director of the Laboratory was also announced in this circular.

This brief account of some facts in the early history of the Marine Biological Laboratory may suffice to show the origin of Professor Whitman's connection with the institution. He found a local organization that planned to become national in scope, to enlist the cooperation of colleges and universities throughout the country and to provide for research and instruction in biology. The location of the Laboratory was also fixed and the first building erected at Woods Hole. Although the incorporators were all residents of Boston, yet they had provided for a national organization by offering each institution invited to cooperate the privilege of naming five members each of the Corporation during the term of cooperation. Apparently, Professor Whitman had nothing to do with the original statement of these principles, but after his appointment as director, at least, he became their chief exponent and developed them to a much greater extent than the original incorporators had intended, so that the Corporation soon came to have a large and nation-wide membership, and the Board of Trustees was enlarged to include 12 members in 1890, 17 in 1892, and 21 in 1895. The membership of the Corporation grew by leaps and bounds, and rapidly became representative of the entire country, as the practice was followed for some years of inviting all who worked at the Laboratory to become members. The attendance at the Laboratory was 17 in 1888, 44 in 1889, 47 in 1890, 71 in 1891, 110 in 1892, 199 in 1895; and the number of institutions represented was 13 in 1888, 29 in 1889, 32 in 1890, 31 in 1891, 52 in 1892 and 85 in 1895.

The early years of the Laboratory were years of grfeat prosperity; to accommodate the growing tide of workers an L was added to the original building in 1890; in 1892 a building equal to the original Laboratory in size was added to form the third side of a quadrangle, and two separate buildings, one for botany and another for a lecture hall and research rooms were added by 1896.

Whitman's part during this period of rapid material development was to furnish the spirit and develop the ideals of the institution. It is obvious that the idea of cooperation had a primary practical significance in the minds of the original trustees, to secure support for the new institution. Though he did not lose sight of its practical significance, the idea of cooperation was transformed by Whitman into an ideal of a scientific democracy, which furnished a motive for loyalt}^ and devotion such as rarely, if ever, existed in a scientific enterprise, so that the development of the Laboratory became a kind of cult to a large and influential body of naturalists. Whitman not only awakened this spirit, which was compounded of devotion to himself as well as to the ideal which he represented, but he kept it alive, and more than once, by refusing to compromise any fraction of the fundamental idea for immediate practical advantage, he saved the principle from extinction. That the Laboratory today is still a scientific democracy is due entirely to Whitman's uncompromising devotion.

In his first report Professor Whitman states, The new Laboratory at Woods Holl is nothing more, and, I trust, nothing less, than a first step towards the establishment of an ideal biological station, organized on a basis broad enough to represent all important features of the several types of laboratories hitherto known in Europe and America." Thus he formed great plans for the germinal institution. He early maintained that in such an ideal biological station it was essential that all biological interests should be represented, and accordingly successively added departments of botany, physiology and embryology to the original zoology, each with its side of research as well as instruction. But the variety of work that has been welcomed at Woods Hole cannot be included even within these broad divisions. Professor Whitman had most catholic interests in biology and it is remarkable in what fundamental ways he comprehended the problems of each division. The association of workers in different fields of biology has been one of the most helpful and stimulating features of the Station.

The Marine Biological Laboratory was designed for instruction as well as research. The original circular opens with these words: "The Trustees of the Marine Biological Laboratory earnestly desire to enlist your cooperation in the support of a sea-side laboratory for instruction and investigation in biology." Instruction was in fact placed first, not only in the opening sentence but throughout the circular. However, the Laboratory started out at once under Whitman as primarily a research institution, and in his address at the opening of the Laboratory, July 17, 1888, he said:


XXXll CHARLES OTIS WHITMAN

111 every attempt hitherto made to combme the two chief interests here represented, instruction has been the object of first concern. Now the only way to keep the distributive function efficient and active is to unite it in proper relations with the productive function. The Laboratory (i.e., the side of investigation) is the creative agent — -the source of all supplies; the school is merely the receiver and distributor. Any attempt to combine the two which ignores or reverses these relations must end in disappointment and failure.

In the fifth annual report Professor Whitman states:

The two functions of instruction and investigation have worked admirably together, each growing stronger in the success of the other. We have endeavored to keep the two properly balanced, but I think we have nearly reached the limit of our capacity for instruction with our present space and means. We already see that to tax our teaching force much more would not tend to improve the side of investigation.

In the eighth annual report for the year 1895 Professor Whitman again returns to this theme:

Our instruction and investigation have been inspired by a common purpose, and thus kept in such relations that each has added to the strength of the other, and added more and more with every stride forward. If instruction has increased, it is chiefly due to the stimulating influence of investigation; if investigation has gained, it is because instruction has multiplied workers. Mutual service is the bond of union, but the union is not merely one of coordination, in which the two elements are simply balanced one against the other; it is one of a more vital order, in which each is servant and only one is master. All our classes face in one direction — towards original work — and all our activities, sympathies and interests are dominated by the spirit of research. Does that render our instruction less efficient? Just the contrary. It fills with life and purpose, makes students more earnest, dignifies the work of the teachers, and wins their best effort. Moreover, it re-enforces the service of the regular staff by contributions from every member of the investigating departments.

Farther on:

What does instruction mean for us? It means, not wholly, but preeminently, preparation for original work, and much of it is especially designed for the benefit of investigators, not beginners only, but for specialists who are independent workers.

It will be plain,^ I trust, that we are not cultivating two antagonistic functions, between which we have to carefully guard the balance, lest one may prosper at the expense of the other. There can be no excess in either direction, for every gain, whether on one side or the other, is a gain not only for the part but also for the whole.


BIOGRAPHICAL SKETCH XXXlll

These extracts explain Whitman's position with reference to the functions of instruction in a primarily research institution. His ideas seem to have been sound, if we may judge from the experience of twenty-three years, during which the two have existed side by side with mutual advantage.

During the third session of the Laboratory Whitman organized the evening course of Biological Lectures which has proved ever since one of the stimulating features of the Laboratory life. In his report for this session Whitman outlines the idea as follows :

These were not intended to take the place of systematic lectures, such as are given in the regular courses of instruction; they stand rather for the higher and the more general needs of the science. Their leading purpose, if I may be permitted to define it more with reference to the possibilities of its future development than to its present attainment, was to meet the rapidly growing need of cooperative union among specialists. Specialization has now reached a point where such union appears to be an essential means of progress. Specialization is not science, but merely the method of science. For the sake of greater concentration of effort, we divide the labor; but this division of labor leads to interdependence among the laborers, and makes social coordination more and more essential. This is the law of progress throughout the social as well as the organic world. An organism travels towards its most perfect state in proportion as its component cell-individuals reach the limit of specialization, and form a whole of mutually dependent parts. Scientific organization obeys the same law. As methods of investigation improve, specialization advances, and at the same time the mutual dependence of specialists increases. Isolation in work becomes more and more unendurable. Comparison of results, interchange of views and ideas, and a thousand other advantages of social contact, become of paramount importance to the highest development.

In such considerations may be found the leading motive for this course of lectures. While directed in the main to the higher needs of investigators, they deal, as a rule, with subjects of present and quite general interest to beginners. In general, it may be said that the authors undertake to set forth what has been accomplished in their special fields of research, to give the conclusions of the best work and thought, to point out general bearings, and to state the problems that await solution.

The educational value which such lectures may be presumed to have, and the consideration that through them the aims, the needs, and the possibilities of biological work might, in some measure, be made better known to the public, especially to those whose liberal benefactions have enabled the Laboratory to carry forward its work, suggested the propriety of publication.


XXXIV CHARLES OTIS WHITMAN

At various times these lectures, which have sometimes taken on a spirit of some formahty, have been supplemented by informal discussions following lectures dehvered by investigators before classes, especially the class in embryology during the early years, and later in physiology; at other times research seminars have been formed for the distinct purpose of discussing and criticising work presented by the investigators; and at all times in the history of the Laboratory free and informal discussion between investigators of their work in progress has been a characteristic feature in the laboratory life. In all this the steady and sane influence of Whitman was at work. All coveted discussions with Whitman; he had a most sympathetic interest in all work going on in the Laboratory, and deep insight into the fundamental problems. One frequently discovered after unburdening one's self in response to his sympathetic attitude that he had thought out the problem in question more thoroughly. But his courteous and honest attitude always saved such a situation from being painful. He exercised in these ways a steadying influence on the investigations of others, for he was never hurried into following a mere fashion in research.

The social life of the Laboratory in Whitman's time was simple and sincere. He had a horror of all formality and met everybody on a plain and equal footing. His hospitality usually took the form of small dimiers particularly well cooked and served, with not more than half a dozen guests usually. He was a most charming host, gracious and self-effacing. The conversation usually turned on some scientific subject and he had the knack of making the others talk, and it was considered quite a triumph for the others to draw him out. He sustained relations with his students both at Woods Hole and elsewhere, that can only be described as fatherly. He often helped thjem financially, and stood by them with the greatest loyalty in securing positions. To the respect that all his students felt for his scholarship and ability was added the love and devotion that they owed to the best of friends.

No account of Whitman's relations to the Marine Biological Laboratory would be complete which failed to describe his con


BIOGRAPHICAL SKETCH XXXV

duct in various crises of the history of the institution. The essential character of the man comes out better probably in its mingled elements than in any other known relations. But this account must necessarily be incomplete and partial to the extent that Whitman is the subject, and not the Laboratory. Up to about 1895 the relations of the Director and trustees seem to have been on the whole cordial, in spite of minor difficulties. But the rapid growth of the Laboratory imposed financial burdens of no shght amount. In 1890 an 'L' was added to the original building; in 1890 a new wing was built; in 1893-4 a new dining hall and kitchen were erected, and the present botanical laboratory. The expenses of these additions was met by numerous contributions from friends and by a loan of $3,500 secured by a mortgage upon the property of the Laboratory, and an unsecured loan of $3,000 from one of the trustees.

The Boston trustees themselves felt great satisfaction in the rapid growth of the Laboratory. In 1894 they could say: The only serious perplexities of the last year have been the result of its rapid growth and prosperity;" the Laboratory had in fact become self supporting so far as current expenses were concerned. It was important, however, to meet the outstanding loans for new buildings and the following appeal was issued:

Reluctant as the trustees were to incur expenses which would make it necessary, in this time of financial stress, to ask help from the friends of the Laboratory, yet, in the opinion of many, to have checked the growth of the institution at this stage, by turning away desirable students and investigators, would have inflicted a permanent injury. We ask, then, from those whose conviction of the value of such a Laboratory has helped to bring it to its present condition of prosperity, still further aid in its future development (from the Trustees' Report to the Corporation for the year 1894).

But the enlargements, great as they had been, were still inadequate to the growing demand. In proposing the further enlargement which Professor Whitman felt to be necessary to provide for the growth of the Laboratory, he was hampered by the reluctance of some, at least, of the trustees, to incur further indebtedness. A new building was needed of the size of the original laboratory to provide a lecture hall and more rooms for inves


XXXVl CHARLES OTIS WHITMAN

tigators at an estimated cost of $3,000. Professor Whitman organized the investigators of the Laboratory into a Biological Association to work for the needed building. This Association pledged SI, 500 towards the cost of the new building, and the trustees finally agreed to secure an equal sum. The building was erected in 1896, and has been fully occupied ever since, thus justifying Whitman's estimate of the needs of the Laboratory. But this plan left the debt for previous buildings still outstanding. '^ Subsequent events showed that Doctor Whitman raised the whole of the $3,000, besides the money needed for equipment, and the trustees did not as a body raise anything; although a few individuals who were supporters of Doctor Whitman and his policy raised a few hundred dollars" (from A Reply to the Statement of the Former Trustees of the Marine Biological Laboratory," 1897, p. 8).

While it is perhaps undesirable to revive old controversies, yet it seems needful in justice to Doctor Whitman, to state the issues of the years 1896-1897, with the dispassionateness which fourteen elapsed years should furnish. It was never true that a majority of the board of trustees lost confidence in, or were out of sympathy with Doctor Whitman ; but a minority of the board, who nevertheless constituted the governing element by virtue of their original membership and residence in Boston where all the meetings were held, were much displeased with him for not listening respectfully enough to their motives of caution, and for his dominance in Laboratory affairs. The existence of a. small deficit in the operating expenses of the year 1896 led them to declare that the Laboratory should not be opened in 1897, unless a sum of $2,000 were raised not later than April 15. This sum was much in excess of the deficit and the vote was not taken until February 5, 1897. An offer on the part of one of the trustees, Mr. L. L. Nunn, to bear any added deficit resulting from operations of 1897, was refused. The trustees raised the sum of $1,140 by April 12, and the treasurer reported on May 5 that there was a balance in the treasury of $735.55; there was also about $670 accumulated interest in funds available for any purpose the trustees might approve. The deficit in the meantime had melted away. The



October 10, 190S. Photograph by R. AI. Strong



1908. Photograph by KenjiToda

Charles Otis WTiitman


BIOGRAPHICAL SKETCH XXXVll

announcement of the 1897 session was therefore very late and the attendance suffered seriously in consequence of the inimor that had spread that the Laboratory would not be opened that year.

A meeting of the board of trustees was held at Woods Hole on August 6, 1897, and at this meeting a majority of the members present, who were favorable to Whitman, voted to call a special meeting of the members of the Corporation to be held in Boston on August 16 for the purpose of considering changes in the bylaws. The purpose of the proposed changes was (1) to provide that the annual meeting of the Corporation should be held in Woods Hole instead of in Boston, and in August instead of November, and to increase the quorum so as to secure a more representative attendance and avoid local control, and (2) to change the body of the trustees from a body practically self-perpetuating to an elective body, elected by the Corporation in four groups, one such group to be elected each year for a period of four years, and thus avoid the old practice of the simultaneous annual election of all members.

At this meeting about eighty-seven members of the Corporation recorded their names with the clerk, and it was estimated that there were about twenty others present who did not do so. It was the largest and most representative meeting of the Corporation ever held up to that time. The program as outlined was unanimously adopted.

This amounted to no less than a revolution in the government of the Laboratory, and the action was promptly followed by the resignation of seven out of the nine members of the board of trustees resident in Boston and its vicinity. Six of these and one other trustee then drew up a statement which was primarily an attack on the Director, Professor Whitman, which they published in Science, October 8, 1897. To this statement a complete reply was made in the more dignified, but less permanent, form of a separate pamphlet by a committee of three of the trustees who stood by the Director (A Reply to the Statement of the Former Trustees of the Marine Biological Laboratory," Boston, Alfred Mudge and Son, Printers, No. 24 Franklin Street, 1897.) This reply and the facts that two-thirds of the board of trustees stood


XXXVlll CHARLES OTIS WHITMAN

by Professor Whitman, that the places of the 'former trustees' were taken by well-known naturahsts, and that the progress of the Laboratory was not seriously interrupted even by so serious a controversy, constitute a sufficient vindication for Whitman.

This struggle was unfortunately necessary to estabUsh the national, representative and democratic character of the institution, a character that grows with the years and which commands the loyalty and devotion of the present members, both of the Corporation and of the board of trustees.

Once again it was necessary for Whitman to take a firm stand to maintain the fundamental ideals of organization of the Laboratory. This was when the newly organized Carnegie Institution of Washington offered in 1902 to take the Laboratory as a department. This would have permanently solved the difficult problem of maintenance, but Whitman was convinced that it would destroy the representative democratic character of the institution, although every possible concession to the existing form of organization was generously offered by the Carnegie Institution. In this opinion he stood nearly alone, but none the less firmly, and it was his insistence that finally brought about a delay of the decision with an annual grant of $10,000 a year for a period of three years (1903-1905) from the Carnegie Institution in the form of a subscription to twenty work rooms. At the end of this period a very notable petition signed not only by all members of the laboratory, but also by a large number of representative naturahsts, for the continuation of the temporary arrangement was not granted by the trustees of the Carnegie Institution, and the original proposal lapsed. The independence of the Laboratory had been maintained, but it was apparently as far from a stable basis of financial support as ever.

Following this. Whitman gradually withdrew from active participation in the management of the Laboratory, although he retained the title of Dhector until 1908. However, he no longer attended meetings, and was even absent from the Laboratory for two successive seasons, 1904 and 1905. The house which he had occupied at Woods Hole burned down in the winter of 19051906; and, fearing that this would make his return impossible


BIOGRAPHICAL SKETCH XXXIX

his friends raised a sum of S3, 000 by subscription and the property was bought and the house restored and presented to Whitman. This very signal mark of love and appreciation on the part of his friends, indicating as it did so clearly their desire to remove every obstacle that prevented his presence among them, touched Whitman most deeply. He was present again at the Laboratory in the sessions of 1906 and 1907, but never again, except for a brief visit of two or three days in 1909.

His gradually increasing engrossment in the study of heredity and evolution in pigeons may be assigned as the principal cause of his withdrawal from residence at the Laboratory. For many years he transferred his large collection of birds from Chicago to Woods Hole and back again each summer. He always suffered some losses of valuable birds, even when the railroad companies allowed him to take his birds as excess baggage and to attend to them en route. However, when this permission was refused and they had to come by express and might be delayed over an extra night, the losses became more serious. Indeed, the transfer became an intolerable burden, and he relinquished his charge of affairs at Woods Hole rather than curtail his own research, an eminently characteristic choice.

In 1908 he tendered his resignation; his letter and the reply thereto follow:

To the Trustees of the Marine Biological Laboratory, Woods Hole,

Mass.

Gentlemen :

This year has brought the twenty-first birthday of the Marine Biological Laboratory. For these many years you have continued to honor me with the directorship of the Laboratory. In late years I have so far drifted out of office and out of use that a formal resignation at this time can scarcely be more than an announcement of the fact accomplished. The time has arrived, however, when a reorganization seems to be imperatively demanded, and as a prelude thereto, I must ask you to accept this note as a somewhat belated announcement of my resignation of the office of director.

Let me take this opportunity to thank you one and all very heartily for the cordial support you have extended to me.

Respectfully,

CO. Whitman.


Xl CHARLES OTIS WHITMAN

August 13, 1908.

The Corporation and Trustees of the Marine Biological Laboratory, in accepting the resignation of the Director, Professor CO. Whitman, have ordered to be put upon their records and to be forwarded to Doctor Whitman the following minute :

The Corporation and Trustees desire to express to the retiring Director their regret that he finds it necessary to withdraw from the active directorship of the laboratory, and their appreciation of the inestimable value of his services. Since the establishment of the Laboratory at Woods Hole twenty-one years ago, he has been continually its Director and he has to a very large extent guided its growth and development. He has stood for the principles of cooperation and independence which have made the laboratory unique in character and truly national in its reputation and influence. His high ideals and his generous appreciation of the work of others have been an inspiration to the many biologists, who, during these years, have attended the laboratory.

The corporation and trustees desire that the retiring Director niay continue to serve the laboratory as honorary director and trustee and that his presence at the laboratory may continue to be an inspiration in the future as in the past.

Professor Whitman's reply was as follows:

To the Corporation and Trustees of the Marine Biological Laboratory, Woods Hole, Mass.

Ladies and Gentlemen: Your action of August 13, in which you express a desire to have me serve the laboratory as 'honorary director and trustee' is in itself alone an all-sufficient reward for whatever services I have rendered as Director. Your goodwill is the all-important recompense, and no title that you could confer could add to the weight of your approbation. In fact, titles belittle the spirit. Let me have the latter without the former — -without title or office of any kind. Please respect this wish and believe me, as ever, a sincere and devoted friend of the Laboratory.

Respectfully and cordially,

C. O. Whitman.

The report of the trustees to the Corporation bearing on Professor Whitman's resignation and on his services to the Laboratory expresses so well what many others feel that it is appropriate to quote it in large part:

Professor Whitman's resignation as Director of the Marine Biological Laboratory, after twenty-one years of service in that' position, impressively recalls the inestimable value of his services in the establishment and development of this institution. If we have today one of the lead


BIOGRAPHICAL SKETCH xll

ing marine laboratories of the world, we owe it in large part to him. The interest of almost every member of this board of trustees and of the corporation was enlisted through his efforts, and the splendid influence which the Marine Biological Laboratory has had upon the development of biology in this country is traceable ultimately to him.

His connection with the laboratory began at a time when it had neither permanent home, recognized standing, nor scientific ideals. Some of the leading biologists of this country felt that it could not compete as a research station with the U. S. Fish Commission Station, backed as the latter was by the resources of the government, and that its chief field of usefulness must be as a summer school. Whitman thought otherwise, and by his real greatness as a scientist, his untiring energy and enthusiasm, his splendid ideals and his unfailing faith and courage he made it from the start the principal center in America for biological research.

From start to finish his ideals for the laboratory were these: (1) A national center for research in every department of biology; (2) a laboratory founded upon the cooperation of individuals and institutions; (3) an organization independent in its government and free to follow its natural course of growth and development. For these ideals he has labored consistently and persistently year after year, sometimes with a disregard of present advantage, to be gained by the sacrifice of one or the other of these ideals, which cost him friendships which he highly prized. At one particular crisis he wrote : ' If I have made any enemies through unkindness or injustice, I am sincerely sorry for it ; but if I have made any because I have stated my conviction on the question before us I can afford to part with all friends who are made enemies for such a cause.' His faith in the ultimate achievement of these ideals was so great that he chose rather to sacrifice present good than, as he believed, the future welfare of the laboratory; and his plans for the laboratory were so great, while current resources were so small, that he was frequently charged with being impractical. But it is only fair and just to recognize how much was accomplished by adherence to these ideals and to what an extent the spirit and success of the laboratory are due to them.

Woods Hole is indeed a national center for research in several branches, if not in every department, of biology. Whitman had the wisdom to see that biology could progress only as a whole. 'The great charm of a biological station,' he wrote, 'must be the fullness with which it represents the biological system. Its power and efficiency diminish in geometrical ratio with every source of light excluded.' To zoology, which was the only subject represented at first, he added botany and physiology and he strove to make Woods Hole a center in each of these departments. He was one of the first to insist upon adequate provision for experimental work. He was, we believe, the first in this country to plan and plead for a biological farm for the study of problems of heredity and evolution. He desired to make Woods Hole a center for the comparative study of anatomy, pathology and psychology. Some of these lines of work have since been taken up and largely developed elsewhere, but if


Xlii CHARLES OTIS WHITMAN

Whitman could have had the necessary .support in his plans they would have been centered at Woods Hole. This need of a national center of research in every department of biology is still before the laboratory as a living issue, and although this grand concept has so far failed of complete realization, who can say how much the laboratory owes to this catholicity of spirit of its director, how much biology as a whole owes to this splendid ideal?

If the laboratory was to be truly national, Professor Whitman believed that it must be founded upon the cooperation of individuals and institutions; no one man nor institution, however great, could accomplish this purpose. He recognized that common ideals must form the basis of such cooperation, and he sought to bring into close connection with the laboratory every person and every institution that shared these ideals with himself. With these ideals, and by means of his o^vn personal charm and scientific abilities. Whitman secured the cooperation of many of the younger biologists of the country. There was thus developed at Woods Hole a center for research work in biology which has had few equals in the history of the world. By his own work, as well as by his appreciation of the really fundamental problems of biology, he has set a very high standard for the scientific work of the laboratory, and by his kindness, sincerity, and generosity he has called forth similar qualities in others, so that it has been characteristic of Woods Hole, as of few other laboratories at home or abroad, that a spirit of genuine cooperation and mutual helpfulness prevails. Who that experienced it can ever forget the inspiration and enthusiasm of those early years of the laboratory? Who of us can forget the cordial appreciation and generous encouragement which we received from Professor Whitman? Some of us feel that we there incurred a debt of gratitude to him which we can never fully repa3^ Since those early years other laboratories have arisen and other duties have drawn men away from Woods Hole, but the Marine Biological Laboratory never loses its charm for those who have worked there, and this charm will continue as long as the spirit of cooperation, which Whitman instilled into it, prevails.

Finally, Professor Whitman stood for the complete autonomy of the laboratory. Although aid might have been had more than once from universities and institutions by surrendering the independence of the laboratory, he steadfastly and consistently refused to do this, even though in doing so he had to face the opposition of almost all the members of the board of trustees and the corporation. There is still a difference of opinion as to the expediency of this stand, but there is probably no question as to the desirability of the autonomy. If the laboratory can obtain endowments such as to provide for its present and future needs and to insure its independence we shall all greatly rejoice, but whether it shall succeed in this aim or not, we are probably all agreed that this much at least of Professor Whitman's ideal must be maintained, viz : that the laboratory must be left free to grow and develop as its own needs and the interests of science demand.


BIOGRAPHICAL SKETCH xliii

These are the ideals which Professor Whitman succeeded in making part and parcel of the Marine Biological Laboratory and which we count among our most valuable possessions. To those who measure the success of an institution by the size of its buildings or endowments, his efforts at Woods Hole may seem in large part to have failed, but those who realize that ideals are the motive forces of the world, that life consists not in abundance of possessions but in abundance of service, that science is not paraphernalia but knowledge — ^these will not fail to recognize the great value of the work Professor Whitman has done for the Marine Biological Laboratory and for the whole science of biology.

To these words of appreciation there is but little to add. It may be that the Marine Biological Laboratory is Whitman's most enduring monument, as it was his chief work of organization. But the principles will endure eternally, whatever the life of the particular expression they have been given in the Laboratory, and the fact that Whitman was the chief champion of these ideals and that he gave them visible and effective expression is one of his chief claims to affectionate and reverent remembrance.


THE AMERICAN MORPHOLOGICAL SOCIETY

Professor Whitman was the leader in the three most important organizations for the advance of zoology in America during the time of his active life: in 1887 he founded the Journal of Morphology, in 1888 he became director of the Marine Biological Laboratory, and in 1890 he took the leading part in the foundation of the American Morphological Society. A circular was sent out October 16, 1890, calling on those interested to unite in the formation of an Association of Morphologists in connection and affiliation with the American Society of Naturahsts," which shall hold stated meetings during the Christmas vacation, at which special and general morphological problems may be brought forward and discussed. Attention was directed in the circular to the scientific isolation of zoologists in America, and the advantages of their cooperation in such a society. The committee signing this call consisted of C. 0. Whitman (chairman), Henry F. Osborn, E. B. Wilson, E. G. Gardiner, and J. Playfair McMurrich. The first meeting was held in Boston, December 29, 1890. Dr. E. B. Wilson was elected chairman for the meeting. Whitman was then elected president for the next meeting, and was re-elected during, the following three meetings. In 1902 the name of the society was changed to The American Society of Zoologists" and it is still our dominant zoological society.

AT THE UNIVERSITY OF CHICAGO

We have departed from the chronological order of events in thus sketching Whitman's various activities. In 1892 Whitman moved from Clark University to the University of Chicago, taking with him the major part of his department and all his students. Professors Mall, Donaldson and Baur also came at the same time from Clark University to the University of Chicago and took part with others in the formation of a department of biology of which Whitman was head. After the first year, however, the department was broken up into separate departments of zoology, anatomy, neurology, physiology and palaeontology. Concerning this event Professor Mall says (The Resignation of Professor Whitman as Director of the Marine Biological Laboratory at Woods Hole, Mass., Anat. Record, vol. 2, no. 8, November, 1908).

When the University of Chicago was founded in 1893, Professor Whitman was made head of the biological department, which in its organization was unusually strong on the anatomical side. It was planned at the beginning to divide the department as soon as circumstances would warrant, and with the very rapid growth of the University this took place within a year. Then the anatomical department was established coordinate with those of zoology and botany. This proved to be the most important step in the organization of anatomical departments in America, and for it we are largely indebted to Professor Whitman.

Whitman was in fact mainly responsible for the unusually comprehensive organization of the biological departments in the University of Chicago, and for their establishment in a single group of buildings, thus rendering possible a degree of mutual support and cooperation among the biological sciences, the full possibihties of which have not been realized even in Chicago up to this day. Whitman thus carried out in Chicago as far as possible the same form of organization that he planned for Woods Hole, involving the representation of every branch of biological knowledge, so as to bring the conibined forces to bear on the fundamental problems of biology. It was possible in Chicago to proceed along such ideal hues, for the institution was unhampered by history or tradition or by fixed location of departments established in remote neighborhoods.

The department of zoology under Whitman was primarily a research department and the members of his staff were selected primarily for their standing as investigators. Whitman, himself, taught graduate students exclusively, for the most part candidates for the doctor's degree. He lectured but once a week and not always regularly; but each lecture was a finished essay, and in a way, a piece of original work. He never attempted to present what students could find in books. He consulted about once a week with each student on his research problem, and was a very rigorous and strict critic, but he tended more and more as time went on to let each student work out his own salvation. It often became necessary for the student to seek him at his house for consultation about his work, but such a consultation was always well worth while, as Whitman would leave his own work and play the part of host most delightfully, as well as that of teacher.

The details of departmental administration were very irksome to Whitman, but on the fundamental principles of administration of the department he was firm as a rock and quite uncompromising. Even trifling details would often seem to him contrary to correct ideals, and then the matters of administration loomed large and were rigorously decided.

Whitman's students who took the degree of Doctor of Philosophy under his instruction, with their present academic standing, were the following:

1. At Clark University:

Hermon Carey Bumpus, Business Manager, University of Wisconsin. William Morton Wheeler, Professor of Entomology, Bussey Institution, Harvard University.

Edwin Oakes Jordan, Professor of Bacteriology, University of Chicago.


2. At the University of Chicago :

Herbert Parlin Johnson, Associate Professor of Bacteriology, Medical Department of St. Louis University.

Frank Rattray Lillie, Chairman of the Department of Zoology, and Professor of Embryology, University of Chicago ; Director, Marine Biological Laboratory.

Albert Chatjncey Eycleshymer, Professor of Anatomy, Director of Anatomical Department, St. Louis University.

William Albert Locy, Professor of Zoology, Northwestern University.

Howard Stedman Brode, Professor of Biology, Whitman College, Walla Walla, Washington.

Cornelia Maria Clapp, Professor of Zoology, Mount Holyoke College.

Agnes Mary Claypole (Mrs. Robert O. Moody).

Albert Davis Mead, Professor of Comparative Anatomy, Brown University, Providence, R. I.

Charles Lawrence Bristol, Professor of Zoology, New York University.

Samuel J. Holmes, Assistant Professor of Zoology, University of Wisconsin.

John P. Munson, Normal School, EUensburg, Wash.

Emily Ray Gregory, Professor of Biology, College for Women, Constantinople, Turkey.

Aaron Louis Treadwell, Professor of Biology, Vassar College.

Michael Frederick Guyer, Professor of Zoology, University of Wisconsin.

Elliot Rowland Downing, School of Education, University of Chicago.

Wilhblmina Entemann Key, Lombard College, Galesburg, 111.

Ralph Stayner Lillie, Instructor in Comparative Physiology, University of Pennsylvania.

Virgil Everett McCaskill, President State Normal School, Stevens Point, Wisconsin.

John McClellan Prather, Teacher of Zoology, Central High School, St. Louis, Mo.

Eugene Howard Harper, Instructor in Zoology, Northwestern University.

Bennett Mills Allen, Assistant Professor of Anatomy, University of Wisconsin.

William J. Moenkhaus, Professor of Physiology, University of Indiana.

Charles Dwight Marsh, United States Department of Agriculture.

John William Scott, Instructor in Zoology, Kansas State Agricultural College, Manhattan, Kansas.

Charles Zelbny, Associate Professor of Zoology, University of Illinois.

Lynds Jones, Professor of Ecology, Oberlin College.

Horatio Hackett Newman, Associate Professor of Zoology, University of Chicago.

James Francis Abbott, Professor of Zoology, Washington University, St. Louis. Mo.

Victor Ernest Shelford, Instructor in Zoology, University of Chicago.

Oscar Riddle, the University of Chicago.

Charles Henry Turner, Sumner High School, St. Louis, Mo.

Frank Eugene Lutz, Curator, American Museum of Natural History, New York City.


George Washington Tannreuther, Assistant in Zoology, University of Missouri.

Wallace Craig, Professor of Philosophy, University of Maine.

Charles Christopher Adams, Instructor in Ecology, University of Illinois.

James Thomas Patterson, Adjunct Professor of Zoology, University of Texas.

Mary Blount, Teacher, University High School, and Assistant in the Department of Zoology, University of Chicago.

Katashi Takahashi, Professor of Zoology, Gakushinin College, Tokyo, Japan.

Marian Lydia Shorey, Instructor in Zoology, Milwaukee-Downer College, Milwaukee, Wisconsin.

H. L. WiEMAN, Assistant Professor of Zoology, University of Cincinnati.

George William Bartelmez, Associate in Anatomy, University of Chicago.


WHITMAN'S SCIENTIFIC WORK

Professor Whitman's scientific work covered an unusually broad field. He had a distinct predeUction for monographic treatment, and even while engaged on special problems concerning a particular animal or group of animals little escaped his observation or his note-book. Thus his work on leeches was in the first place embryological, but he soon turned to anatomy and taxonomy, and to problems of behavior, all of which are treated in his various publications. Later, in his work on the development of Amia and Necturus, but little of which has been published, he also paid particular attention to problems of anatomy and behavior. And again, his work on pigeons concerned not only problems of heredity and evolution, but he made an exhaustive study of the taxonomy of the group, and the problems of their behavior were constantly in his mind. He also assigned students problems on their development, and a number of papers were published by Guyer, Harper, Blount, Patterson, and Bartelmez in this field. Another illustration was his plan for a monographic study of Arenicola by cooperation of a group of his students, parts of which have been published.

His own publications fall mostly in the subjects of embryology, comparative anatomy and taxonomy, animal behavior, and evolution and heredity, an extraordinary breadth of field for a modern zoologist. But whatever subject he touched he illuminated. He was slow to publish, not because of lack of industry or results, but because he was determined to examine the subject to the bottom, and to be sure of his view-point. He rarely had occasion to correct any published statement, and even less rarely, perhaps, to change in any radical way a point of view to which he had once committed himself. Work of such classical distinction could not be very abundant. He has left a large amount of unpublished material, especially bearing on pigeons, though there is much that dates from an earher period. The statements that follow concerning unpublished material are on Dr. Riddle's authority.


® This section has been prepared by E. G. Conklin, Albert P. Mathews, T. H. Morgan, J. Percy Moore, and Oscar Riddle. The writer of the main body of the biography has merely prepared the introductory paragraphs.



Embryology

In his first scientific pubhcation, The Embryology of Clepsine," Professor Whitman at once took rank as a great zoologist. Although this paper was his doctor's dissertation, it was unusually mature, and showed in striking manner the qualities which characterized all of his later work, viz., patience and accuracy in observation, great power of logical analysis, and a firm hold on large problems. Indeed so fundamental and comprehensive was this work that almost all of Professor Whitman's later embryological work is foreshadowed in it, while it furnished the stimulus for a large amount of work on the organization of the egg and its cell lineage, which was done later by his associates and students at Woods Hole.

Even at this early date (1878) his conclusions as to the organization of the egg and the formation of the embryo were fundamentally the same as his later views. He observed the bilateral symmetry of the egg of Clepsine before cleavage; he studied carefully the cleavage of the egg and observed the formation and subsequent history of the ectoblasts, mesoblasts, neuroblasts, and entoblasts; he described the growth and concrescence of the germ bands of Clepsine, and compared them with the growth and concrescence of the germ ring of fishes. His conception of the fundamental problem of all development is expressed in these words: "In the fecundated egg slumbers potentially the future embryo. While we cannot say that the embryo is predehneated we can say that it is predetermined" — words almost precisely like those used by him twenty years later when deahng with this subject.

In 1887 he pubhshed another paper on this general subject, entitled "A Contribution to the History of the Germ Layers of Clepsine" in which he extended his former observations on the cleavage, orientation of cleavage planes, origin of teloblasts and germ bands, origin of the mesenteron, and the origin of the ectodiferm and its products. And in the same year in a paper on Ookinesis," he came back to the phenomena of maturation and fecundation, which he had treated in his first paper, and gave a very suggestive and comprehensive review and analysis of these phenomena.

Whitman's general point of view regarding the problems of development are made particularly plain in his biological lectures and addresses. In his address before the Zoological Congress of the Worlds Columbian Exposition, on "The Inadequacy of the Cell Theory of Development" he discussed these problems in a striking and suggestive manner. At this time the work of Wilson and others had just shown the possibihty and importance of tracing individual blastomeres throughout the development from the time of their appearance until they give rise to particular portions of the embryo, the cleavage thus constituting "a visible mosaic;" shortly before, Roux had shown that the cleavage of the frog's egg was a mosaic work;" at the same time the work of Driesch and other experimentalists was leading to a directly opposite opinion. In this conflict of opinion Whitman took a strong and independent position, basing his conclusions not merely on comparative embryology but also upon the comparison of protozoa and metazoa. He protested against the view that organization is the product of cell formation, and insisted that "organization precedes cell formation and regulates it." He contrasted the Cell-doctrine with what might be called the Organism-doctrine. He insisted that, "an organism is an organism from the egg onward, quite independent of the number of cells present," that cleavage is not a process by which organization arises, but that organization precedes cleavage. "The test of organization in an egg does not lie in its mode of cleavage, but in subtile formative processes. The plastic forces mould the germ-mass regardless of the way it is cut up into cells."

At the same time he showed clear insight into the results of the experimental embryologists. In this same address he says, "The formation of a whole from a part .... no more disproves the existence of a definite organization in the case of the egg than in the case of hydra." i^nd in the preface to the volume of Biological Lectures for 1894 he strongly contests the view that developmental mechanics" has explained or can explain vital phenomena, without reference to the historical development of the organism. As to mechanism and vitalism he says, "There is no warrant for the assertion that hfe is something different from, and independent of, matter and energy. That is the mistake of vitalism. On the other hand there is no warrant in decomposition for identifjdng dead mechanism with living mechanism." "The ultimate mystery is beyond the reach of both mechanism and vitaUsm; let pretensions be dropped and approximation to truth will be closer on both sides."

The influence of Professor Whitman's work on the science of embryology and on the scientific and philosophical problems connected with development was profound. His work was careful, critical, consistent. He reached conclusions only after most painstaking observations and mature dehberation, and when he had once made up his mind he was not easily moved. Others might' be "tossed about by every wind of doctrine," but he stood unmoved and unshaken, having confidence in his own observations and reflections and refusing to doubt his conclusions until he himself had seen and felt equally strong evidence against them. As a result of this he was unusually stable and consistent, and while his later work shows that his ideas were constantly enlarging with new evidence, yet there was little in his earlier work which needed correction.

Professor Whitman's work on the early development of the teleost egg, published with Alexander Agassiz (1884 and 1889), is a fine example of careful discriminating embryological investigation. The cleavage especially was studied with great care, and the history of the marginal cells led to the solution of "one of the cardinal questions in the early development of the teleostean fishes, namely the precise origin of what His and others have called the 'parablast.' " The authors showed that the nuclei of the periblast, as they termed this layer, are derived entirely from the marginal cells of the blastodisc, and the fallacy of free cell-formation" and of separate origin of the parablast and archiblast" thus received its quietus. Their hope that a similar result would be reached for other meroblastic vertebrate ova has since been reahzed.


Professor Whitman's interest in the Hirudinea, aroused and fostered during his occupany of a table in Leuckart's laboratory at Leipzig, continued unabated until his labors were so tragically terminated. That this interest was no narrow one, but traversed much of the depth as well as the length and breadth of biological science, is much less apparent from a list of the titles of his papers, than from a perusal of their contents. Under such unassuming titles as The Leeches of Japan," or "Description of Clepsine plana," we find treated, not merely specific and faunal details but the much larger questions of the formulation of a new system for standardizing specific descriptions, the morphological basis of generic groupings and the origin, evolution, ecology, adaptations, morphology, classification and paths of dissemination of the land leeches. The latter small group of animals particularly appealed to his naturalist's spirit, not as the disgusting pests that they had seemed to most previous describers and tropical travelers, but as the keen-sensed, facile winners in a competitive struggle for Ufe that must have been of the utmost severity for animals that have departed so widely from the habits and environment of their ancestors.

Although Professor Whitman's contributions to the adult morphology of the Hirudinea were not numerous, numbering only three major papers and seven or eight preliminary sketches, polemics and reviews, they have exerted a great and lasting influence. Taken together they form an exhibit of the catholicity of his interests and exempUfy the soundness of his scientific ideals.


Hi CHARLES OTIS WHITMAN

Whitman was ever keenly alive to the importance of a many sided view of animal life and was equally sympathetic to the work of the field naturalist and student of ecology and habits, of the systematist, morphologist and physiologist in all their multiform specialties, requiring only that their work should be thoroughly and honestly done.

His earlier papers especially indicate a real and vital interest in systematic zoology and the species question. To whatever wide ranging speculations his studies eventually led him, they received their primary impetus through his efforts to find a satisfactory basis for the discrimination and definition of the species of leeches. Indeed, his very first paper after the publication of his Inaugural Dissertation on the "Embryology of Clepsine," was on A new species of Branchiobdella" (B. pentadonta) which he at that time, in common with other zoologists, regarded as a leech.

To such good purpose did he labor that the history of the systematic study of the leeches may fairly be divided into a preWhitmanian and a Whitmanian period. Notwithstanding that he often rode rough-shod over many of the most sacred traditions of systematic zoology, and notwithstanding that his actual determinations of species and genera have sometimes proven unfortunate, nevertheless Professor Whitman discovered the criteria by which evolution and specific radiation in the leeches may best be measured and expressed, and he set a standard for specific description that has since been the guide and model to all the best workers in this field. Out of the chaotic condition that Whitman found order has been wrought, largely through the use of the tools that he devised. It cannot be denied that he imported new light and life to the subject, nor that all later students of leeches have felt the vivifying influence of his ideals.

The key to Whitman's successful analysis of the external morphology of leeches is his discovery (first announced in two prehminary papers pubhshed in 1884 and subsequently repeatedly reverted to and expanded) of the segmental sense organs to which he later appHed Haeckel's term 'sensillae.' These little, whitish, translucent dots, visible on many living leeches, had been noticed occasionally by earlier observers and Ebrard had even suggested


BIOGRAPHICAL SKETCH 1111

that they might have a respiratory function, but their real nature and significance as sense organs had passed quite unsuspected.

Whitman showed that these sensillae are groups of tactile cells which differ from the scattered epidermal sense cells chiefly in their aggregation into groups arranged syimnetrlcally according to a definite plan on one ring (now designated the neural or sensory annulus) of every somite and that they are serially homologous with the eyes, which differ from them chiefly in the acquisition of visual cells (Glaskorper of Leydig) and pigment cup.

During the two years (1879-1881) which he spent in Japan as Professor of Zoology at the University of Toklo, Professor Whitman collected material for the description of the leeches of that country. Under his direction most beautiful and accurate colored drawings were made of the species of the several families, but unfortunately the first part only, treating of the Hirudinidae or ten-eyed leeches," was ever pubhshed. In this paper Whitman's method of analytical description, based primarily upon the metamerlc arrangement of the sensillae and the somite composition thus made evident, was first fully applied to the family Hirudinidae. In the course of somewhat elaborate descriptions of the Japanese land leech (Haemadypsa japonica), the Japanese medicinal leech (Hlrudo nlpponlca) and three species of Leptostoma (earlier named Microstoma and now known as Whitmanla) the somites are analysed successively and compared as respects their constituent elements. Comparisons are made with several other European, American and Asiatic genera and the previously unknown or only vaguely apprehended fact brought to light that the metameres of the different genera of ten-eyed leeches are very differently developed as regards the number and size of constituent rings, particularly toward the extremities of the body. These observed differences in the number of rings constituting the somites are correlated with differences in the mode of life of the animals. In addition to their morphological value, these descriptions and the accompanying illustrations are models of beauty and accuracy, than which no more satisfying have been published before or since.


liv CHARLES OTIS WHITMAN

Much attention is given to the description and illustration of the complex color patterns and, although Whitman perceived that in each species all patterns are variants of a single fundamental one, the structural basis underlying them in the arrangement of the muscles and other organs was left to be discovered by his student Arnold Graf, whose tragic end at Woods Hole, Professor Whitman so keenly felt.

The discovery of the neurology of eyes and sensillae was fully elaborated and the structure of these organs minutely described. They were compared to the lateral sense organs of the Capitellidae and the lateral line organs of fishes. The precise determination of the limits and composition of the somites that his method required, led Professor Whitman to the formulation of definite views regarding the nature and history of the metameres, namely:

1. That the neural or sensory annulus is the first of the somite.

2. That each gangUon of the central nervous system supplies the first three rings of one somite and the last two rings of the immediately preceding somites or equivalent portions of somites having less than five rings, and consequently that there is a lack of correlation between definite neuromerism and definT itive metamerism.

3. That the quinque-annulate or typical complete somite of the middle region is primitive, and that there is a progressive reduction or abbreviation of the somites toward the ends of the body that is correlated with specialization in other respects.

It must not be understood that Whitman neglected the internal anatomy. On the contrary he was assiduous in the dissection and description of the several organ systems, but except in the case of the nervous system, he added comparatively Uttle to our knowledge.

In a later largely controversial article 'Some New Facts about the Hirudinea,' written in reply to criticism. Whitman forcibly reasserts his views and brings many new facts to their support. In this paper his strong leaning toward the annelid theory of the origin of vertebrates, to which reference is frequently made in later papers, is indicated. He also extends his former opinion


BIOGRAPHICAL SKETCH Iv

of the homology of the segmental organs with the lateral line organs to include the ear and even the eye of vertebrates, which he believed to have had their origin in organs similar to the sensillae of leeches.

Three years later ('Description of Clepsine plana,') we find Whitman with all his enthusiasm applying the same criteria of metamerism and the same methods of analysis to the external morphology of the Glossiphonidae. This he designated the type somite and derived from it both abbreviated somites having the number of rings reduced to less than three and supplemented somites with the number multiplied to more than three.

In support of these views he also appealed to the facts of embryology and instanced many cases of ring multiplication, incipient or advanced, in various genera of leeches. Each somite of the leeches' body, to a considerable degree, undergoes an independent individual development, the nature and extent of which is correlated with the physiological demands to which it is subjected. But never did Whitman carry out his view to its logical consequences and see, as others have, that the uniannulate and biannulate somites existing at the ends of the body of nearly all leeches are steps toward the elaboration of the triannulate as the latter is toward the quinque-annulate somite.

The relation of sensillae and eyes became clearer and the proof of their homology was buttressed by many facts. Cases of dual sense organs, composed partly of superficial cells bearing tactile hairs and partly of clear visual cells situated more deeply along the course of the optic nerve, and all gradations from typical sensillae with which one or two visual cells and perhaps a little pigment may be associated, to complete eyes with only a trace of tactile cells were described, the latter leading by easy gradations to the strictly visual organs of Hirudo. In some cases every step in the transition was found in the successive somites of a single leech, as in Clepsine hollensis.

Strong embryological evidence was brought to bear, especially in a paper in the Zoologischer Jahrbiicher for 1893, as showing the common origin of both kinds of sense cells from common proliferations of the ectoderm, also that the segmental sense organs are


Ivi CHARLES OTIS WHITMAN

more primitive than the scattered sense cells (goblet cells, labial sense cells and Bayer's organs) and that they cannot have been derived from aggregations of. the latter, as had been maintained by Maier and Apathy.

It is most fitting that Whitman's last important paper relating to the leeches should have appeared in the "Festschrift zum siebenzigsten Geburtstage" Leuckarts who h^d guided his early interest in the group. This memoir on The Metamerism of Clepsine," is the culmination of ^^^litman's work on metamerism. More than any previous work it is concerned with the nervous system and as an example of complete morphological analysis has few equals among papers dealing with invertebrates. The elements of the central neuromeres and peripheral nerves are correlated one by one with such external features as annuli, sensillae and eyes throughout the body and especially in the simpler somites at the two extremities. The presence in these terminal segments of every morphological element is determined and accounted for and the conclusion reached that complete homodynamy exists throughout. The earlier determination of twenty-six somites in the body of all leeches anterior to the caudal sucker, and of seven in the sucker was confirmed.

Metamerism is traced to the extreme tip of the anterior end, where, however, there is a cephalic region in which the dorsal halves alone of the somites are represented and in which as a consequence, there is a delayed embryonic development. There is no non-metameric residuum at the anterior end and, although acknowledging that embryology furnishes some evidences of the presence of a rudimentary apical organ and remnants of a pair of head kidneys, Whitman denies that there is any element here other than, or added to, the first metamere. There is no prosoma in the sense of Hatschek of an unpaired, non-metameric and premetameric region opposed to the segmented region beginning with the larval mouth. Whitman also contends that these facts confirm the opinion long held by himself in common with Leuckart and others that metamerism originated in multiplication by fission.


Animal Behavior

Two papers were published by Dr. Whitman on Animal Behavior, one as a Woods Hole lecture, the other in the Monist. There was also a short letter, commenting on an article of Professor Lankester's on the origins of intelhgence, pubhshed in the Chicago Tribune. Of these, the lecture entitled "Animal Behavior" is the most important. It shows him at his best and is one of the ablest of his papers, which is equivalent to saying that it is one of the most admirable papers on this subject. Its style is clear, interesting, and direct; and it may be taken as a model by every investigator. Nowhere else is reasoning more solid and sound, or comment more illuminative. No other of his papers illustrates better the qualities of his genius: the selection of a fundamental problem; painstaking study; publication only after years of observation and reflection ; skill in laying bare the simple basis of an apparently complex group of phenomena ; a grasp of the subject in all its bearings; and the use of the comparative or phyletic method of attack.

He considers in this paper the fundamental questions of the origin of instinct and intelligence as illustrated by a study of the behavior of three kinds of animals upon which he worked at different periods of his life: the leech, Necturus, and the pigeon.

He begins with the simplest acts of Clepsine; its deceptive quiet when disturbed, a quiet of intense rigidity; and its rolhng into a ball after feeding when it detaches itself from its host. He shows that the latter instinct can never have been acquired as a habit which has been stereotyped as an instinct, because Clepsine feeds but twice or three times in its life. • If the view here taken be correct," he says, "the instinct of rolhng into a ball is not a matter of deliberation at all, but merely the action of an organization more or less nicely adjusted to special conditions and stimuh. Intelhgence does not precede and direct, but the indifferent organic foundation with its general activities is primary; the special behavior or instinct is built up by slowly modifying the organic basis."


Similarly, the instinct of capturing food exhibited perfectly by the youngest Necturus is innate. The pause before seizing the bait is part of a very old primitive mechanism found is fishes and finally developed into the ^pointing' of a dog. Its object is to fix the aim. The timidity of the young Necturus is also innate and not the result of painful experiences.

"We have taken a very important step in our study when we have ascertained that behavior, which at first sight appeared to owe its purposive character to intelhgence, cannot possibly be so explained, but must depend largely, at least, upon the mechanism of organization. The origin and meaning of the behavior antedate all individual acquisitions and form part of the problem of the origin and history of the organization itself." "We see at once that behavior does not stand for a simple and primary adaptation of a pre-existing mechanism to a special need. As the necessity for food did not arise for the first time in Necturus, the organization adapted to securing it must be traced back to foundations evolved long in advance of the species. The retrospect stretches back to the origin of the vertebrate phylum .

The point of special emphasis here is that instincts are evolved, not improvised, and that their genealogy may be as complex and far-reaching as the history of their organic bases." This passage should be read by all physiologists who, of all biologists, are most given to neglecting phylogeny in their explanations.

While instinct thus comes before intelligence and not after it, as many have believed, some intelhgence was implied by the inhibition of instinctive acts in Necturus by fear. To chnch his argument that instincts are evolved like structures and are not inherited habits, he turns to two instincts cited by Romanes as clear evidence of their origin in habits; the tumbling and pouting of pigeons. An examination shows that the rudiments of these instincts are to be found in all species of pigeons. Again the value of the phyletic method of study is illustrated by the 'brooding' instinct of birds. This he shows to be the evolution of an instinct shown even in fishes, which hover over the nest and drive away


BIOGRAPHICAL SKETCH lix

intruders. Even Clepsine sticks more firmly than usual when it is over its eggs.

He sums up this part of the paper with a few general statements: Instinct and structure are to be studied from the common standpoint of phyletic descent and that not the less because we may seldom, if ever, be able to trace the whole development of an instinct. Instincts are evolved, not involved .... and the key to their genetic history is to be sought in their more general rather than in their later and incidental uses." As the genesis of organs takes its departure from the elementary structure of protoplasm, so does the genesis of instincts proceed from the fundamental functions of protoplasm."

Taking up now the origin of intelligence he says, "Since instinct supplied at least the earlier rudiments of brain and nerve, since instinct and mind work with the same mechanisms and in the same channels, and since instinctive action is gradually superseded by intelligent action, we are compelled to regard instinct as the actual germ of mind." "We are apt to contrast the extremes of instinct and intelligence — to emphasize the bhndness and inflexibility of the one and the consciousness and freedom of the other. It is like contrasting the extremes of Ught and dark and forgetting all the transitional degrees of twihght." "Instinct is blind; so is the highest human wisdom blind. The distinction is one of degree. There is no absolute bhndness on the one side, and no absolute wisdom on the other. Instinct is a dim sphere of light, but its dimness and outer boundary are certainly variable ; intelligence is only the same dimness improved in various degrees. ' '

To show how instinct becomes less fixed and choice appears he cites the behavior of three species of pigeons, the passenger pigeon, the ring dove and the common pigeon. If the egg is removed from under the wild pigeon and placed on the side of the nest, the bird returning to the nest remains a moment or two only and then leaves the nest not to return. The ring dove, on the contrary, after some time of perplexity, will return one egg into the nest leaving the other out; the common pigeon, after a longer period of perplexity and uneasiness, will put both eggs back into the nest. There is here a gradual loss of precision of the instinct to leave the disturbed nest when the rigor of natural selection is relaxed. The common dove is not so much an automaton. Choice begins to appear. "With choice no new factor enters, but only plasticity, so that the pigeon becomes capable of higher action and is encouraged and even constrained by circumstances to learn to use its privilege of choice." This little freedom is the dawning grace of a new dispensation in which education by experience comes in as an amelioration of the law of elimination. This slight amenability to natural educational influences cannot of course work any great miracles of transformation in a pigeon's brain, but it shows the way to the open door of a freer commerce with the external world, through which a brain with richer instinctive endowments might rise to higher achievement."

"Superiority in instinct endowments and concurring advantages of environment would tend to liberate the possessors from the severities of natural selection; and thus nature, like domestication, would furnish conditions inviting to greater freedom of action, and with the same result, namely, that the instincts would become more plastic and tractable. Plasticity of instinct is not intelligence, but it is the open door through which the greater educator, experience, comes in and works every wonder of intelligence."

Evolution

No account of Whitman's work would be complete without reference to his essay on "Evolution and Epigenesis," not only because this essay reveals him in one of his most thoughtful moods, but because the essay defines very sharply Whitman's attitude toward one of the profound questions of the time — a question that was then engaging the best thought and work of all serious biologists. His keen critical sense is here shown to advantage, his independence of thought led him to break some of the idols of the day, and his thorough understanding of what had been written and was being written at the time, all conspire to make the essay a permanent contribution to our knowledge. He succeeds as few others have done in holding the fine balance between the two extremes of thought represented by the terms preformation and epigenesis. He defines his position in the following words: "I should perhaps say at the outset that I have no theory of development either to announce or to defend. It is of more importance just now to have well-defined standpoints and clear ideas of guiding principles. The possibihty, not to say probability, that the egg is from the beginning of its existence as an individual cell definitely oriented has as yet received but little attention." "The drift of opinion, as it seems to me, is neither back to the standpoint of Harvey and WolfT nor to that of Bonnet and Haller, but towards a new standpoint which seeks to avoid the errors and blend the truth of the old hypotheses." Whitman's standpoint is summed up in the two following quotations: The indubitable fact on which we now build is — ^the ready-formed living germ, with an organization cut directly from a pre-existing parental organization of the same kind. The essential thing here is — actual identity of germ organization and stirp organization." And again: Let this organization stand for not more than our neo-epigenesists freely concede, namely, that original constitution of the germ, wh'ich predetermines its type of development — let it stand for nothing more than that and obviously the standpoint rises to an altitude scarcely dreamed of in the philosophy of Harvey and Wolff."

Whitman devoted much time to the study of Bonnet's theory of evolution. If one asks why he should have thought it worth while to give so much attention to a discredited theory of the eighteenth century, the answer is first that he wished clearly to point out the error of those "who imagine that they see in recent theories of development a renaissance of Bonnet's evolution" and, second, that "if our theories of development are carrying us back to the standpoint reached by the evolutionists of the last century it is a matter of more than historical interest." His conclusion is "That the old and the new evolution are based on antithetical conceptions which exclude each other at every point." "The old evolution (preformation) was the greatest error that ever obstructed the progress of our knowledge of development, [f our examination has helped to clear the mist that obscured important distinctions we have not labored wholly in vain."


Dicyemids

In 1882 Whitman published his paper on the Dicyemids. He confirmed much of the earher work of van Beneden, but made out a different relation between the nematogene and the rhombogene individuals. He discovered certain facts that led him to conclude that the germagene of van Beneden arose from fertihzed eggs while all other individuals arose parthenogenetically. Whitman's conclusion in regard to the relationships of the Dicyemids is the same as that to which Hartman has come in his recent monograph.

Pigeons

Those who are familiar with "Whitman's work regard his studies and experiments on pigeons as his greatest achievement. He died at the very moment when he believed that he had reached a point where he was prepared to publish the results of this extensive and exhaustive investigation. Only on a few occasions (see list of publication 1904-1907) has he stated in briefest outline a few of the principal conclusions he had reached. In a paper read at the Universal Exposition in St. Louis in 1904; in his address before the International Congress of Zoologists in Boston, 1907; and in the Bulletin of the Wisconsin Natural History Society, 1907, Whitman has expressed himself clearly and forcibly on certain fundamental questions of evolution. His address of 1907 has not been printed but the substance of that address is found in his other writings.

The dominant feature of Professor Whitman's long and still unpublished work on inheritance and evolution lies in its intensive and extensive attack upon the nature of a specific character. In the 90's he wrote: It is to a comparative and experimental analysis of specific characters that we must look for knowledge of the phenomena of heredity and variation." And again, in 1904, in summarizing the results of many years of study of one such character he wrote as follows :

In tracing the origin and genesis of a single character we meet the leading questions in the evolution of species. First and foremost the question as to the nature of the initial stages. Did the character arise as a variation de novo, or as a progressive modification of a pre-existing character? If de novo, did it spring suddenly forth, with some decisive advantage in the struggle for existence? Or did it appear as one of many minute changes, and by some happy chance get a start that gave it the lead in future development? In other words, did it begin as a discontinuous variation, sport, or mutation? Or did it arise cumulatively, as a continuous development? If it originated by modification of an earher character, was it at first a sudden, sport-like departure? Or was it a slow and continuous transformation, of a progressive or retrogressive nature?

"Then we come inevitably to the deeper question, which natural selection only partially penetrates — the question how variation, multifarious and undirected, without the aid of design or a designer, can advance to such definite and wonderful achievements as specific characters."

Whitman's devotion to the task of learning a specific character knew no bounds; it heeded neither time, personal sacrifice, nor the difficulties which the ensemble of hfe processes creates when a particular process with which the biologist would become familiar is examined. But, he was ever ready and eager to attend to each and every perturbation of the system, from whatever extrinsic source, if its analysis might lead directly or indirectly to a better measure of realities in his own main sphere of study. It thus happens that along the pathway which he has blazed into the central problems of evolution are to be found many discoveries in the fields of instinct, animal behavior, fertility, correlative variation, the nature of sex, etc.

Having selected color-pattern in pigeons as supplying a satisfactorily small group of specific characters easily accessible to study, he first set about determining which patterns are the more primitive and which the higher and more recent ones, the facts being determined through a most painstaking search for the convergent testimony of the most various kinds of evidence. Here his uncompromising ideal of an intensive and extensive study of a character, his own exceptional mastery of the broad field of zoology, the eighteen years of unbroken and devoted study that he gave to this work inevitably led him to results of great importance.

A general survey was made of the color-patterns of nearly six hundred wild species, and of nearly two hundred domestic races of pigeons. Large numbers of genera and species from all parts of the world were brought to the breeding pens of his yard. With indefatigable patience the plumage patterns of the living birds were studied; the sequence of pattern in the plumages from young to old was accurately observed ; and thoughtful experiments were devised to bridge the gap between the moults, and thus displace apparent discontinuities with visually realized continuities. The primitive pattern of many diverse orders of birds was also ascertained, and the general primitive basis of color-marking in all birds — the 'fundamental bars' were discovered.

The direction of the evolution as it was indicated by all these studies was, moreover, again and again retested by evidence of an entirely different sort. Such characters as voice, behavior, and fertility were separately subjected to similar appropriate vigorous comparative and breeding tests to learn whether the resulting data would parallel each other and that furnished by the extensive study of the color-pattern. Only when, by all these means and others, he had accumulated a vast amount of reliable, consistent and convergent testimony as to where the various genera and species stand in the phylogenetic series, did Professor Whitman permit himself to feel that he was reading aright the history of the specific characters of the pattern. And it is a very real monument to his scientific greatness, that, not until he knew all this of the character with which he was working, and much besides, would he write as much as one line concerning it.

In his yards were hybridized nearly forty wild species of pigeons, most of these crosses being made here for the first time. The results of continued breeding of the simple and complex hybrids from these forty pure wild species, and of several domestic races, furnish a mass of most remarkable data. The conclusions from these data being at the same time checked and supported by the results of other lines of study on the same material.


In consequence Professor Whitman's work presents a great body of searchingly self-critical and reliable conclusions, and these conclusions unquestionably lead far into constructive evolutionary theory. For his material he believed he had demonstrated beyond doubt the reahty and regnancy of definitely directed variation, i.e., of orthogenesis, as the method of evolution. He has accumulated and presented the most weighty evidences for continuity as against discontinuity in the phenomena of variation, inheritance and evolution. He has thrown new hght on the nature and meaning of 'mutants;' such 'mutants' at any rate as occur among pigeons. He accomplished in 1903, and continuously since then, the remarkable result which in Mendelian terms may be spoken of as the control or determination of the dominance of sex and color. ^

His work was most bountifully and beautifully illustrated, this feature having occupied many years of the undivided attention of excellent artists. Even in the unfinished parts, however, the outlines of the work are so bold and its details of data are so clear as a result of the polishing process to which he, who was the very spirit of clarity and accuracy, subjected them, that time and care will enable others to arrange most of the results in a form that will still carry conviction to the reader.

Whitman's conception of Orthogenesis, and his attitude toward the mutation theory is stated in the following paragraph:

' ' Among the rival theories of natural selection two are especially noteworthy. One of these is now generally known as orthogenesis. Theodore Eimer was one of the early champions of this theory, basing his arguments primarily upon his researches on the variation of the wall-lizard (1874-81). Eimer boldly announced his later works on 'The Origin of Species' (1888), and the 'Orthogenesis of the Butterflies' (1897), as furnishing complete proof of definitely directed variation, as the result of the inheritance of acquired characters, and as showing the utter ' impotence of natural selection J Elmer's intemperate ferocity toward the views of Darwin and Weismann, coupled with an almost

' Unpublished data.


fanatical advocacy of the notion that organic evolution depends upon the inheritance of acquired characters, was enough to prejudice the whole case of orthogenesis. Moreover, the controversial setting given to the idea of definitely directed variation, without the aid of utility and natural selection, made it difficult to escape the conclusion that orthogenesis was only a new form of the old teleology, from the paralyzing domination of which Darwin and Lyell and their followers had rescued science. Thus handicapped the theory of orthogenesis has found little favor outside the circle of Elmer's pupils."^

The second of the two theories alluded to is the mutation theory of Hugo, de Vries. The distinguished author of this theory .... maintains, on the basis of long continued experimental research, that species originate, not by slow gradual variation, as held by Darwin and Wallace, but by sudden saltations, or sport-like mutations. According to this theory, two fundamentally distinct phenomena have hitherto been confounded under the term variation. In other words, variation, as used by Darwin and others, covers two classes of phenomena, totally distinct in nature, action, and effect. Variation proper is defined as the ordinary, fluctuating, or individual variation, and this is held to be absolutely impotent to form new species.

Granting that the position with respect to the mutants obtained from the evening primrose {Oenothera Lamarckiana) is unassailable, does it follow that all new species have arisen by mutation, and that continuous variation has never had, and never can have, anything to do with the origin of species?

"Plausible as is the argument and impressive as is the array of evidence presented, I can but feel that there are reasons which compel us to suspend judgment for a while on this pivotal point of the mutation theory."

Whitman objected strongly to the implication that a variation tendency must be considered to be teleological because it is not orderless.

  • Whitman — The Problem of the Origin of Species, Congress of Arts and Science,

Universal Exp. 1904.


"I venture to assert that variation is sometimes orderly, and at other times rather disorderly, and that the one is just as free from teleology as the other. In our aversion to the old teleology, so effectually banished from science by Darwin, we should not forget that the world is full of order, the organic no less than the inorganic. Indeed what is the whole development of an organism if not strictly and marvelously orderly? Is not every stage, from the primordial germ onward, and the whole sequence of stages, rigidly orthogenetic? If variations are deviations in the directions of the developmental processes, what wonder is there if in some directions there is less resistance to variation than in others? What wonder if the organism is so balanced as to permit of both unifarious and multifarious variations? If a developmental process may run on throughout life (e.g., the lifelong multiplication of the surface-pores of the lateral-line system in Amia), what wonder if we find a whole species gravitating slowly in one or a few directions? And if we find large groups of species all affected by a like variation, moving in the same general direction, are we compelled to regard such ' a definite variation-tendency' as teleological, and hence out of the pale of science? If a designer sets limits to variation in order to reach a definite end, the direction of events is teleological; but if organization and the laws of development exclude some lines of variation and favor others, there is certainly nothing supernatural in this, and nothing which in incompatible with natural selection. Natural selection may enter at any stage of orthogenetic variation, preserve and modify in various directions the results over which it may have had no previous control."

The particular evidence in favor of orthogenesis on which Whitman rested his case is found in the origin of the bars on the wings of the wild pigeons and on the wings of many domesticated birds.

The rock pigeons (Columba livia) present two very distinct color-patterns; one of which consists of black checkers uniformly distributed to the feathers of the wing and the back, the other of two black wing-bars on a slate-gray ground. These two patterns may be seen in almost any flock of domestic pigeons.


The inquiry as to the origin of these patterns involves the main problem of the origin of species, for the general principles that account for one character must hold for others, and so for the species as a whole Darwin raised the same question, but did not pursue it beyond the point of trying to determine which pattern was to be considered original and how the derivation of the other was to be understood. Darwin's explanation was so simple and captivating that naturalists generally accepted it as final. It is but fair to state that Darwin's conclusions did not rest on a comparative study of the color-patterns displayed in the many wild species of pigeons. Accepting the view generally held by naturahsts, that the rock pigeons must be regarded as the ancestors of domestic races, the question was limited to the point just stated."

Between the checkered and the barred types many intermediate stages may be found in different individuals. But which way is the series to be read, from checkers to bars or from bars to checkers? Whitman finds an answer to this question in the evidence from experiments, from development, and from a comparative study of the Columbidse.

"As an experiment, we may take one or more pairs of pure-bred, typically barred pigeons, and keep them isolated from checkered birds for several years, in order to see if the young ever advance toward the checkered type.

"Another experiment should be tried for the purpose of seeing what can be done by working in just the opposite direction. In this case we take checkered birds, selecting in each generation birds with the fewer and smaller checkers, and rejecting the others, in order to see if the process of reduction can be carried to the condition of three, two, and one bar, and finally, to complete obliteration of both checkers and bars, leaving the wing a tabula rasa of uniform gray color.

"If these experiments are continued sufficiently far, it will be found from the second experiment that a gradual reduction of pigment to the extreme conditions named can be comparatively easily effected, and that the direction of reduction will always be the same, from before backward; while, from the first experiment, it will be seen that it is hopeless to try to advance in the opposite direction, from the bars forward to the checkered condition. No variations will appear in that direction, but such as do appear will take the opposite direction, tending to diminish the width of the bars and to weaken their color. It is in this way that we must account for the existence of some fancy breeds in which the bars have been wholly obliterated. The direction of evolution can never be reversed.

"I have tried both experiments for eight years, and as both tell the same story as to the direction of variation, I am satisfied that further experiments will not essentially modify the results."

After tracing wing bars of diverse kinds to checkers, the origin of the checkers was traced from a still earlier and universal avian character.

It consists of a single dark spot occupying the centre of the exposed part of each feather. In the course of evolution, this spot has been divided into two lateral spots by the disappearance of pigment along the shaft, beginning at the apex of the feather and advancing gradually inward. The old Turtle-Dove character thus passes by a continuous process of division into the Rock Pigeon pattern, consisting of two checkers on each feather, more or less completely separated. The evidences showing such a gradual transmutation are still to be seen, and in such profusion as to wholly exclude doubt. Hundreds of species have been formed in this simple way, leaving no room for the claim of sudden, nontransitional mutations.

. The transitional stages between the Turtle-Dove pattern and the checkered pattern of the Rock pigeons, are exhibited not only as we pass from one species to another, but often as we advance from the juvenal to the adult plumage; and frequently they may be seen in different parts of one and the same individual plumage.

A still older character than the Turtle-Dove spot is seen in the cross-bars, or fundamental bars, that appear to mark all feathers of all species of birds. These bars were first noticed in pigeons in the summer of 1903, and were soon found to be common to all species of pigeons and birds in general. From these fundamental featherbars or their secondary derivatives, a multitude of specific characters have been evolved by gradual modification. The continuity in the evolution of some of these characters can be experimentally demonstrated. The Uttle Diamond Dove (Geopelia cuneata) of Australia, owes its small white spots (two in each feather) to these bars. The transitional stages connecting the spots with the bars are not wholly given in passing from the juvenal to the adult plumage. But if we pluck a few of the juvenal feathers at suitable intervals, their places will be filled by new feathers of different ages, and in this way we may get the stages intermediate between the bars of the young and the spots of the adult. Thus we see that the adult pattern, which normally appears to come in as a striking mutation, by a single jump, is only an endstage in a continuous process of differentiation. So it is everywhere. Suppression of stages in ontogeny looks like saltations; but whenever we can get at the history of the character, we find the continuity comes to light."

PERSONAL CHARACTERISTICS

We have described principally the outward events of a life that was not lacking in incident. Many traits of character shine through these events, but the history of his inner life is far from being comprised in such a sketch, and there is no one competent to write it. With Whitman, more than with most men, one felt that the inner life was the dominant factor, that it was genuine, deep and worth knowing; the outward events were more or less accidental; he would have created similar effects under a totally different set of external circumstances.

In person he was of medium height, of stout build and good color in his maturity, though thin and pale in his last years; his hair was snow white from young manhood, his eyes were direct and piercing, his forehead high, broad and noble; he wore a beard and a heavy mustache that somewhat concealed the rather thicklipped mouth. His bearing was always erect and dignified; his dress was simple and sufficiently conventional, but he entirely eschewed the ceremonial dress and was only once seen in academic cap and gown, and I believe not at all in evening dress.


Whitman's life was simple and studious; it was passed almost entirely between his house and his laboratory. A large part of his work, since 1891 certainly, was done at home, and from about 1895 when he began the study of pigeons, by far the major part. He gradually collected a large number of species of pigeons from all parts of the world, and in the latter part of his life the collection comprised some 550 individuals representing about thirty species. His house was surrounded by pigeon cotes, and he always had some birds under observation indoors, so that the cooing of doves was for years a dominant sound in his house. He took care of the birds for the most part himself, though he usually had the assistance of one or two maids. He thus actually lived with his birds constantly, and very rarely was absent from them even for a single day. He made observations and kept notes on all aspects of the life and behavior of each species, as well as of such hybrids as he was able to produce. He always had one Japanese artist at work continuously drawing pigeons, and for several years two — Hyashi and Toda. Thus he accumulated an immense amount of material for his magnum opus, which, however, he was not permitted to finish. It is hoped that a considerable portion of his work may be available for posthumous publication, owing to the self-sacrificing labor that has been put on it first by Dr. R. M. Strong, and then by Dr. Oscar Riddle, one of his students, who is now devoting his entire time to editing the manuscripts.

For many years Whitman carried his pigeons with him to Woods Hole in June and back again in September, as already related. But the burden became intolerable, especially as he always bore the entire expense of his pigeon work personally. He was finally obliged to relinquish the annual trip to Woods Hole and all the cherished associations of the Marine Biological Laboratory. His work probably flourished better under these conditions, but it is to be feared that his health suffered from too close application to work and from lack of variety in his life.

With all of his close application to his study, he was nevertheless most devoted to his friends; he was always pleased to see them, and would spend hours in conversation with them as though he had no other concern in the world. Although he was no smoker he would always offer cigars and cigarettes, and would frequently light a cigarette himself, — which burned however mostly in his fingers, — to heighten the spirit of hospitality. He rather frequently invited his students or members of his department and other friends to dinner, and then his usually simple meal was changed to a more elaborate repast.

Dr. A. P. Mathews, one of his close friends, writes thus of him,^

It is not, however, of his work as a scientist upon which I wish to dwell, but rather to recall his personality that the memory of it may remain always with us. His white hair; his kindling, eager, but thoughtful eyes; his tender, gentle smile; his reticence of speech; his consideration for others; his generosity and courage; his hospitality and graciousness as a host; these endeared him to us all. We shall never forget his simple, unassuming, modest manner; his encouraging sympathy; his ripe and sane judgment. If when he was alone he lived simply, the absorbed student of science, when with his guests in his home he was the embodied spirit of hospitality.

His great influence as a teacher is due in part to his fine example and noble ideals, and in part to his habit of picking out young men, who showed any love for science, inviting them to his home, drawing them out, encouraging them and giving them his friendship. Many of them he helped financially, and all of those fortunate to work near him owe him a debt of gratitude for his sympathy and inspiration. Probably no teacher in zoology since Louis Agassiz has exerted so great an influence on young men.

He was not a faultless man, but his faults were the outcome of his ardent, ideal, uncompromising disposition. He once said to the writer, about the year 1906, that he felt he had been too uncompromising in his beliefs. But it is questionable whether his life would have been so valuable, had his disposition been more pliable. The mood in which he made this remark was a rare one, and it is to be doubted that even had it been more common he could have overcome his native tendency. This quality of course made him enemies who sometimes did not hesitate to express unfavorable opinions in a more or less open manner. But those, who knew Whitman best, know well that he never sought any small personal advantage, and that any appearance of neglect of small matters was due entirely to his absorption in higher considerations. He had the courage of his convictions and rarely, if ever, avoided an issue, turned from an opponent, or shunned a fight.


9 Science. N. S., vol. 33, no. 837, pp. 56-58, Januarj' 13, 1911.



Although Professor Whitman pubHshed relatively few papers, he nevertheless occupied a commanding position in science. Some of the reasons have already been indicated. His eye was single and his whole body was therefore full of light;" his devotion to scholarship was never open to the slightest shadow of suspicion. He was continuously engaged in his personal research which dealt with the most fundamental problems of biology, and he had accumulated vast stores of data, which we hoped he would live to publish himself. But apparently he could never satisfy himself with reference to the fundamental problems on which his mind was fixed; the grand consummation of his work had not come, and he could not reconcile himself to the publication of more or less fragmentary pieces of work. His published papers, mostly short, are models of condensed thought, written in a fine, polished, characteristic style. No less care was devoted to the form than to the substance, and some of his papers will certainly endure as classics of the biology of his time

It was, however, not only his publications, but also his work with his journal, his laboratory and his students, his constant helpful association with other workers, and the example of his austere and studious life that brought him recognition. He never permitted himself to be distracted by the confusion of modern life, social or academic, nor diverted from his steadfast purpose by clamor for quick results.

SICKNESS, DEATH AND BURIAL

For several years before his death Professor Whitman suffered considerably from indigestion, and lost much flesh. He was, however, in better health than usual in the fall of 1910. A sudden cold wave came on about December 1, 1910, and Whitman spent the entire afternoon in his yard putting his birds into their winter quarters. In his zeal for his pigeons he forgot about himself. The next morning he was found in a state of coma, and pneumonia rapidly developed and brought his life to a sudden and unexpected termination on December 6. He, himself, had looked forward to at least ten more years of active study, for apart from his dyspepsia, which he had learned to control very well, his general health was excellent. He thus died unprepared, with work that he had carried on up to the last moment in an unfinished condition.

Memorial services in his honor were held in Convocation Hall of the University of Chicago on December 8, and the same evening his body was taken to Woods Hole in charge of a committee of four appointed by the University, and his two sons. The interment took place in the lot of the Marine Biological Laboratory in the Episcopal Cemetery at Woods Hole in the presence of a small company of scientific friends and colleagues, who came to Woods Hole for this purpose, and some of his friends of the village. His grave lies almost within sight of the institution which he had loved so well, overlooking the harbor.

At the annual meeting of the Corporation of the Marine Biological Laboratory held in Woods Hole, August 8, 1911, the entire body adjourned and marched to the grave of Professor Whitman, where a memorial address was read and a wreath placed on the grave. Thus those who were unable to attend the interment paid their last respects to the memory of the dead leader.

LIST OF PROFESSOR WHITMAN'S PUBLICATIONS

1878 The embryology of Clepsine. Quar. Journ. Micr. Sci., vol. 18, pp. 215-315.

1878 Ueber die Embryologie von Clepsine. Zool. Anz., Bd. 1, p. 5.

1878 Changes preliminary to cleavage in the egg of Clepsine. Proc. Amer. Assoc. Adv. Sci., vol. 26, pp. 263-270.

1880 Do flying fishes fly? Amer. Nat., vol. 14, pp. 641-653.

1881 Zoology in the University of Tokyo.

1882 Japanese aquatic animals living on land. Amer. Nat., vol. 16, pp. 403-405. 1882 Methods of microscopical research in the Zoological Station in Naples.

Amer. Nat., vol. 16, pp. 697-706; 772-785. 1882 Ibid: Journal de Micrographie, 6, pp. 558-565; pp. 18, 89 and 188, vol. 7 (French translation of preceding paper).

1882 A new species of Branchiobdella. Zool. Anz., pp. 636-637.

1883 A contribution to the embryology, life-history, and classification of the Dicyemids. Mitth. Zool. Sta. Neapel, vol. 4, pp. 1-89.

1883 Treatment of pelagic fish eggs. Amer. Nat., vol. 22, pp. 1204-5.


1883 A rare form of the blastoderm of the chick and its bearing on the question of the formation of the vertebrate embryo. Quar. Journ. Micr. Sci., vol. 23, pp. 376-397, and Proc. Bos. Soc. Nat. Hist., vol. 22, pp. 178-79.

1884 External morphology of the leech. ProcAmer. Acad. Arts and Sci., vol. 20, pp. 76-87.

1884 On the development of some pelagic fish eggs. Proc. Amer. Acad. Arts

and Sci., vol. 20, pp. 23-75 (with A. Agassiz). 1884 Segmental sense organs of the leech. Amer, Nat., vol. 18, pp. 1104r-1109.

1884 The connective substance of Hirudinea. (Review) Amer. Nat., vol. 18, p. 1070.

1885 Methods of research in microscopical anatomy and embryology. VIII + 255 pp. Boston, S. E. Cassino and Co.

1885 Means of differentiating embryonic tissues. Amer. Nat., vol. 19, pp. 11341137.

1886 Osmic acid and Merkel's fluid as a means of developing nascent histological distinctions. Amer. Nat., vol. 20, p. 200.

1886 The leeches of Japan. Quar. Journ. Micr. Sci., vol. 26, pp. 317-416.

1886 Germ layers of Clepsine. Zool. Anz., Bd. 9, pp. 171-176.

1887 A contribution to the history of the germ layers in Clepsine. Jour. Morph., vol. 1, pp. 105-182.

1887 Ookinesis. Jour. Morph., vol. 1, pp. 228-252.

1887 Biological instruction in universities. Amer. Nat., vol. 21, pp. 507-519.

1888 The seat of formative and regenerative energy. Jour. Morph., vol. 2, pp. 27-49.

1888 The eggs of Amphibia. Amer. Nat., vol. 22, p. 857.

1888 Some new facts about the Hirudinea. Jour. Morph., vol. 2, pp. 585-599.

1888 Address at the opening the Marine Biological Laboratory July 17. First Annual Report of the Mar. Biol. Lab. Boston, pp. 24-31.

1889 The development of osseous fishes. 2. The pre-embryonic stages of development, (with A. Agassiz). Mem. Mus. Comp. Zool. Harvard College, vol.. 14, pp. 1-56.

1889 Report of the Director of the Marine Biological Laboratory for the first session, 1888. First Annual Report of the Mar. Biol. Lab. Boston, pp. 14-20.

1890 Report of the Director of the Marine Biological Laboratory for the second session 1889. Second Annual Report of the Mar. Biol. Lab. for the year 1889. Boston, pp. 27-34.

1890 Report of the Director of the Marine Biological Laboratory for the third session, 1890. Third Annual Report of the Mar. Biol. Lab. Boston, pp. 17-23.

1891 Report of the Director of the Marine Biological Laboratory for the fourth session, 1891. Fourth Annual Report of the Mar. Biol. Lab. Boston, pp. 14-29.

1891 Description of Clepsine plana. Jour. Morph., vol., 4, pp. 407-418.

1891 Spermatophores as a means of hypodermic impregnation. Jour. Morph.,

vol. 4, pp. 361-406. 1891 Specialization and organization, companion principles of all progress;

The most important need of American biology. Biol. Lectures, M. B. L.,

pp. 1-26, Boston.


1891 The naturalist's occupation. 1. General survey. 2. A special problem. Ibid., pp. 27-52.

1892 Metamerism of Clepsine. Festschrift Rudolph Leuckart, pp. 385-395. 1892 Artificial production of variation in types. Science, vol. 19, p. 227.

1892 Report of the Director of the Marine Biological Laboratory for the fifth session, 1892. Fifth Annual Report of the Mar. Biol. Lab. Boston, pp. 18^7.

1893 A marine biological observatory. Pop. Sci. Monthly, vol. 42, pp. 1-15. 1893 A marine observatory the prime need of American biologists. Atlantic

Monthly, pp. 808-815.

1893 The inadequacy of the cell theory of development. Jour. Morph., vol. 8, pp. 639-658, and in Biol. Lect. 1893.

1893 A sketch of the structure and development of the eye of Clepsine. Zool. Jahrb., Abth. Anat. u. Ont., vol. 6, pp. 616-625.

1893 The work and the aims of the Marine Biological Laboratory. Biol. Lectures, Woods Hole, pp. 236-241. Boston.

1893 General physiology and its relation to morphology, Amer. Nat. vol. 27, pp. 802-807.

1894 Breeding habits of the three triclads of Limulus. Amer. Nat. vol. 28, pp. 544-545.

Prefatory note Biol. Loct., Woods Hole, 1894, pp. ii-vii.

1894 Report of the Director to the Trustees of the Marine Biological Laboratory on the Work of the Sixth Session, 1893. Sixth Annual Report of the Mar. Biol. Lab. for the year 1893. Boston, pp. 21-31.

1895 Evolution and epigenesis. Biol. Lectures, Woods Hole, 1894, pp. 205-224, Ginn and Co.

1895 Bonnet's theory of evolution. A system of negations. Ibid: pp. 225-240.

1895 The palingenesia and the germ doctrine of Bonnet. Ibid: pp. 241-272.

1896 The egg of Amia and its cleavage, (with Eycleshymer). Jour. Morph., vol. 12, pp. 309-356.

1896 Report of the Director of the Marine Biological Laboratory for the Seventh and Eighth Sessions, 1894-1895. Eighth Annual Report of the Mar. Biol.

Lab. for the year 1895. Boston, pp. 17-83.

1897 The centrosome problem and an experimental test. Science N. S. vol. 5, 1897, pp. 235-236.

1878 Lamarck and a perfecting tendency. Science N. S. vol. 7, p. 99.

1898 Some of the functions and features of a biological station. Science, N. S. vol. 7, pp. 37^4, 1898. (Presidential address toSoc. Amer. Nat. 1897, but not delivered.)

1898 Animal behavior. Biol. Lectures, Woods Hole, pp. 285-338.

1899 Myths in animal psychology. Monist, vol. 9, pp. 524-537. 1899 Apathy's grief and consolation. Zool. Anz., pp. 196-197.

1902 The impending crisis in the history of the Marine Biological Laboratory, Science, vol. 16, pp. 529-533.

1902 A biological farm for the experimental investigation of heredity, variations and evolution, and for the study of life histories, habits, instincts and intelligence. Biol. Bull., vol. 3, pp. 214-224.


1904 Natural history work at the Marine Biological Laboratory, Woods Hole. Science, vol. 13, pp. 538-540.

1904 Hybrids from wild species of pigeons crossed inter se and with domestic races. Research Seminar, M. B. L., Biol. Bull., vol. 6, pp. 315-316.

1904 The origin and relationship of the rock pigeons as revealed in their colorpatterns. Biol. Bull. vol. 6. pp. 307-308.

1906 The problem of the origin of species. Congress of Arts and Sciences, St. Louis Exposition, 1904, Boston, vol. 5, pp. 41-58.

1906 The origin of species. The introduction and abstract of a lecture delivered before the Nat. Hist. Soc. December 20, 1906. Bull. Wis. Nat. Hist. Soc. vol. 5, pp. 6-14.

1907 Cheques and bars in pigeons and the direction of evolution. Agricultural Magazine, vol. 5, no. 6, pp. 174-182.