CORNELL UNIVERSITY LIBRARY FROM The Estate of ^.H.'^age OK "i n«S°""" ""'™''*'*y l-ibrary pudley memorial volume 3 1924 024 545 273 'M \< Cornell University Library The original of tiiis book is in tine Cornell University Library. There are no known copyright restrictions in the United States on the use of the text. http://www.archive.org/details/cu31924024545273 LELAND STANFORD JUNIOR UNlvfeRSlTY PUBLICATIONS UNIVERSITY SERIES DUDLEY MEMORIAL 'VOLlIMEj CONTAINING A PAPER ,-, ' ■ '■■BY , ';,-; . • , ■ ... WILLIAM RBS^EL: DtJDLEY;: AiJD APPRECIATIONS AND CIONTRIBUTIONS , ' IN, HIS MEM(3Ry ; FRIENDS AND COLLEAGUES (WITH PORTRAIT) STANPQRD' tJNIVERSITY, efpLIFpmffU i^^LISHED BY.THE tfNIVBRSITY . III^IVERSITY SERIES, lisKERiTiNGE OF SilKwqrmS,'!. y-ernon L. Kellogg, Pjrof esspi''; of Ento* ' '" ■:'; mology. - 8# pp.^";4 'pljites. i90k: :Prk:e(,' $lM';; . ^ ' ^ ' '' TheOpISTHOBRANCHIATE MOCLUSCAjO? THE Bi^ANfiER-AGASSIZ ExPEDI' rrioiir to .Brazil. Frank Mace MacFarlandj Professor of liistology..^ 105 pp.^ 19 plates, ; J*6?? Price, $l^p5/v_;:', ' '^^ . ,' -l '' A Study of the Normal; Constitujents; of the; PotAb1.E: Water of - THE San FRANCiscd, PWiNsiTLA, , John Peirce Mitchell, Assistant . ,_ ' .professor of Cheniistry. 7-0 pp., , 1 . map: 1910., Price, "SOc. ^ -■ : ■SYNQi>sis -OF THE';/'t'RUE. , CrABS (iBRACHTlJRA ) , Or i'Pof*rTE^EY;;ii|AY^ iCA'^t- -" 'FORNIA,: Frank Walter Weymouth. 64p^v"l4 plafes. 1910. Price, 5 Qc. ' ' , v -^: ' ;■-.,-.;• , , /■ ,-'■.' •-,,':, The' Osteology of Certain ScpMBRpiji^ .Fishe5.\ Starks, ' Assistant Prof essot of Zoology. 49 pp., 2 plates,;! te^t figure. 1911; ^ Price, SOc- ''' - ', ' .-r^'r"' -■ r.._ ., .- ,-,- ■ A Physical Theory ot' Electrification. Ferjniaii.]4o SanfpM, Professor . '~ of ;'Phy?iicS.. -69-pp., .2 plaites/ 1911. Price, ;SQc. ;^ ■ ' V ■ V- 1 7; The Matzke 'JilEMORiAL ; VolOme. Papers by John, FmSf Matzke^J iiite ■~ ^Professor of RomatiicLangiiagesj and Thirteen Colleagues. 162 pp. '; -r ■:■ ■ ■ 1911, Pttee, $1.00. '> ' ' .;', ' , ' ' _; . , ^ ,,,, , ; , ^ ' ; ' Das- Histqrische' Pra'sens 4;K;'der Xlteren DeutsCH.IIt: SpRac^'e.-';, Bruno, '' ' . Boeziiiger, Assistant Prof essoroif Glblimiami'is Languages. 91 pp.; 1912. Price, 50c. " ;'■,''.-' ' ,' ,'; ' ' ' The E/fect "of a Strictly - VEGfeTABiE. Diet on TIie Spontaneous AcTi'^fiTY, THE Rate '0* /Growth?, a^td the LbSGEyiTt of the ' S: :, Albino Rat. James Rollin, Slonal^er, A^i^tant' Prof essor of Physi- ' '\ „ol6gy.' 36 pp., 1 plate, 1 '5 -text. figures. 1^1'2. Price, SOc. t CA:TA^pGiiJEt'DE- Tous L^/, LITRES' de' Feu ;M; Chapelain. ' ( Biblittth^tie ;■' i ' Natioiiaie, Fonds Fra;n5ais,';Nouv, Acq., No. il?i.) Colbert Searles, ; AWciate^Pi^bfessor; of Romanic taiigijages. ^ 119 pp., 2 plates. 1912. - J- Prijcey; 7Sc. ' LELAND STANFORD JUNIOR UNIVERSITY PUBLICATIONS UNIVERSITY SERIES ■With, ttie oorapliments of H. C. Dudley and of tlie Department of Botany DUDLEY MEMORIAL VOLUME CONTAINING A PAPERj BY WILLIAM RUSSEL DUDLEY AND APPRECIATIONS AND CONTRIBUTIONS IN HIS MEMORY BY FRIENDS AND COLLEAGUES (WITH PORTRAIT) STANFORD UNIVERSITY, CALIFORNIA PUBLISHED BY THE UNIVERSITY 1913 WILLIAM RUSSEL DUDLEY LELAND STANFORD JUNIOR UNIVERSITY PUBLICATIONS UNIVERSITY SERIES DUDLEY MEMORIAL VOLUME CONTAINING A PAPER j BY WILLIAM RUSSEL DUDLEY APPRECIATIONS AND CONTRIBUTIONS IN HIS MEMORY BY FRIENDS AND COLLEAGUES (WITH PORTRAIT) STANFORD UNIVERSITY, CALIFORNIA PUBLISHED BY THE UNIVERSITY 1913 . .;\. ^t^^ ?^ I -.^ ^ t I sr ^ ^ 5 1 V.l^ ^ V\ 11^^^ ^v^- ^ ^^ s ^ ^ ^ \ ^ > z o I. w ° 3 S ° c o z Sc " 2 « _ -^ 3D Pi < n 3 0) H ■< ^ TABLE OF CONTENTS William Russel Dudley Memorial Addresses: John Casper Branner Douglas Houghton Campbell. Appreciations : David Starr Jordan 16 LeRoy Abrams 20 George James Peirce 22 Jared Treman Newman 23 William Franklin Wight 25 List of Publications of W. R. Dudley 27 List of Cornell University Pupils of W. R. Dudley 29 List of Stanford University Pupils of W. R. Dudley 30 Scientific Papers: The Vitality of Sequoia gigantea : 33 William Russel Dudley, Late Professor of Botany The Morphology and Systematic Position of Calycularia RADICULOSA (StEPH.) (TwELVE FiGURES) 43 Douglas Houghton Campbell, Professor of Botany Studies of Irritability in Plants, III, The Formative Influence of Light (One Plate) . . , 62 George James Peirce, Professor of Botany and Plant Physiology The Gymnosperms Growing on the Grounds of Stanford University (Six Plates) 81 LeRoy Abrams, Associate Professor of Botany The Synchytria in the Vicinity of Stanford University (One Plate) iii James McMurphy, Instructor in Systematic Botany The Law of Geminate Species 115 David Starr Jordan, President of Stanford University Some Relations Between Salt Plants and Salt-Spots. . . 123 William Austin Cannon, Desert Laboratory North American Species of the Genus Amygdalus 130 William Franklin Wight, Bureau of Plant Industry WILLIAM RUSSEL DUDLEY was born at Guil- ford, Connecticut, March 1, 1849, and died at Los Altos, California, June 4, 1911. He was educated at Cornell University, graduating with the class of 1874, and taking his master's degree at the same institution in 1876. He was instructor in botany in Cornell Univer- sity from 1873 to 1876, and assistant professor of botany from 1876 to 1892. On leave of absence from Cornell he was acting professor of biology in the University of Indiana in 1880, and he spent the year 1886-1887 studying at Stras- burg and Berlin. He was appointed professor of system- atic botany at Stanford University in 1893, a position which he held from that time until December, 1910, when, on ac- coxmt of ill health, he voluntarily retired and became pro- fessor emeritus. ®mbers(itj> Cfjapel ^eptcmbcrllO. 19U in ilemariam (Emeritus ?^xoteeeov of IBotanp SSorit jUlarci) 1, 1849 3@icb STunc 4, 1911 WILLIAM RUSSEL DUDLEY [An address delivered at the services held in the University Chapel of Stanford University, September 10, 1911.] By John Casper Branner, Vice-President 1 DOUBT if there is any time in men's lives when they come to know each other as well as they do in their college days, especially when they happen to have the same studies, to be in the same classes, and to be much thrown together by any circumstances whatever. Professor Dudley and I belonged to the class of 1874 in Cornell, we had some of the same studies, we belonged to the same fraternity, and as students we had about the same ups and downs. Aside from these mutual interests we were thrown together still more by the fact that Dudley, being a student in the scientific course, had botany in the early part of his studies, while I in the course in Greek and Latin took botany near the end of my college work, and so it came about that in our senior year he was instructor in botany and I was his pupil. As enthusiastic students and as intimate friends we tramped together every hill, explored every gorge and penetrated every swamp for many miles around Ithaca. Under his guidance I came to have a personal acquaintance with and affection for every flowering plant of the region about Cayuga Lake, and for Dudley always a deeper love and a greater esteem. The first piece of scientific work I ever did — a study of the fibro- vascular bundles in the palms — was undertaken and carried through under his guidance. On the slopes of the hills west of Ithaca it was he who pointed out to me for the first time the deep marks cut in the hard rocks by the ice of the glacial epoch. Thus Dudley was not only my first and principal instructor in botany, but he was also, in a way, my first effective instructor in geology. We college professors are more or less given to the discussion of methods of instruction, and it is no uncommon thing to hear this or that man's methods found fault with. I dare say such criticisms are well enough in their way, but after all is said and done there remains one supreme test of a teacher that is often lost sight of in these discussions, and that is his results. I do not speak with a knowledge of the precise number of his students who stand to-day in the front rank of our botanists, but my general impression is that, judged by this standard — ^by results with his 8 DUDLEY MEMORIAL VOLUME Students — Professor Dudley was one of the most successful teachers of botany this country has ever produced. And I am confident that that success is to be attributed to a great extent to the human and personal rather than to the technical part of his methods as a teacher. He was always at the service of his students. No hour of the day or the night was inopportune when a student wanted his advice or direction. His personal influence during his early manhood was the finest and most wholesome that I have ever found among men, whether old or young. Professor W. R. Lazenby of the University of Ohio, who was a classmate, writes of him: "I may say for myself that I owe Dudley a great deal. I roomed with him my first year at Cornell, and he had a great influence for good over my life. I think, all in all, he was one of the best men I ever knew — ^pure gold." Dudley was a warm hearted, genuine lover of nature in all her forms and in all her moods, and this gave him that enthusiasm without which a teacher is not a teacher. No man could have fitted more perfectly into the sentimental side of botany — if botany has any such side. The colors, the beauty, and delicacy of flowers and plants, their lives, their kinships, their histories — all appealed to the artistic side of his nature. This love for and appreciation of nature, however, was his despair as well as his constant delight. His soul overflowed with affection for it all, but he was so sensitive to the defects of language and of other methods of representation that he rarely undertook to give expression to his love for it. But I would not have you imagine that he was a botanist and nothing but a botanist, neither was he a scientific man to the exclusion of other interests. Indeed he was deeply and generally interested in everything human and spiritual. At heart he was a poet. I shall never forget the glow of enthusiasm with which he read to me, when it first appeared, Longfellow's Morituri Saiutamus. He always had about him the works of the best poets and a few pictures and other works of art of the first quality. His was "The love of learning, the sequestered nooks, And all the sweet serenity of books." To be rather than to appear was the steadfast principle of his life. Modesty, gentleness, unobtrusiveness, decorum, and purity of life were his most prominent characteristics. He never did anything for the sake of display; he never courted popularity. His whole life, within and without, was one long, living protest against vulgarity in all its forms. WILLIAM RUSSEL DUDLEY BRANNER 9 He was a man of the finest possible fiber, so fine indeed that the very delicacy of his nature unfitted him for some of the pioneer work he was called upon to do in his lifetime. When Dr. Jordan was President of the University of Indiana, he tried for some time to induce Dudley to go to that institution as professor of botany. And I recall in this connection that Dr. Jordan said to me on one occasion: "Quite aside from his ability as a teacher of botany we need him here on account of his personal influence." But Dudley declined the proflEered position largely because he felt that he was not altogether fitted for the pioneer work required there at that time. With the idea that poverty helps rather than hinders a young man, Dudley did not altogether agree; in fact he entirely disagreed with it in so far as it related to himself. He felt keenly the inconveniences of having to earn his living while carrying on his studies. The necessity of devoting so much time to his teaching and the strictness of the standards he set for himself explain why he was not a writer of books or the publisher of a very long list of scientific papers. Lest some who did not know him well should imagine that so much self-effacement indicated a man with but little force of character I hasten to say that such was very far from being the case. With all his gentleness and sweetness I have never known a man of more decision of character, stronger will power, or of more determination, firmness, and unswerving purpose. In the sxmamer of 1882, I think it was, when Dudley was thirty-three years old, the baccalaureate sermon at Cornell was preached by the Rev. Dr. Heber Newton, who was for a while chaplain here at Stanford Uni- versity. Dudley and I went to chapel together. We found it so crowded that we could not find seats together, and I sat in the row of seats just behind him. Dr. Newton's address was a eulogy upon the life, character, and influence of Ralph Waldo Emerson. You can imagine the tribute he paid to that distinguished writer and lecturer. I recall that when Dr. Newton had finished his eloquent address I said to myself : "Yes, but right here living in our own midst and within the reach of my hand is a man who has every one of the finest traits of character of Emerson." In the latter part of his life certain of his traits became more prominent than during his younger manhood. He was always, and of necessity, a piurist in every sense in which that word can be used. But as he grew older I imagine that his sensitiveness brought him more pain than pleasure. 10 DUDLEY MEMORIAL VOLUME and to this I attribute the rather lonely life he led after coming to California. Unfortunately there are those who knew Professor Dudley only as a name in the university register. I am sure my friend would not thank me to apologize for the modest part he played in this or in any other community, but in closing I am constrained to say a word in behalf of him and of all such men: It behooves us not to lose sight of this blessed truth, that there are fine men and women in this world of ours — and plenty of them, too — who keep out of the limelights, whose names we never see in the headlines of the newspapers, but who lead quiet, sane, and wljolesome lives. Such people always suggest to me the foundations of a great structure. These foundations lie deep beneath the surface of the ground; we never see them; we seldom think of them; they are not decorated with flying flags or written across with gaudy colors or blazing electric lights. But they stand fast and firm, and the stability and the real worth of the entire superstructure depends upon them. One of these foundation-men was William Russel Dudley. WILLIAM RUSSEL DUDLEY [An address delivered at the services held in the University Chapel of Stanford University, September 10, 1911.] By Professor Douglas Houghton Campbell WILLIAM RussEL DUDLEY was bom in Guilford, Connecticut, in 1849 and was one of the earlier students of Cornell University, from which he graduated with the degree of Bachelor of Science in 1874. At that time Cornell University had only been opened for a short time, and I fancy the conditions there were in many respects very much like those of Stanford twenty years ago. The new university at Ithaca had broken away from the traditions of the earlier eastern colleges, and science received far more attention than in most of the other institutions. The opening of the new university with its facilities for scientific work attracted a group of young men who have since attained pre-eminence in their various departments. Among those who are on our own faculty were Dr. Jordan and Professor Branner, with whom Professor Dudley was asso- ciated on intimate terms. Of Professor Dudley's life as spent at Cornell, Professor Branner has just given us a most sjmipathetic account. In Dr.. Jordan's recent sketch of Professor Dudley in Science, he tells us that for a time he was himself instructor in botany, and that Professor Dudley during the early part of his stay at Cornell came under his instruction. However, it was not long before Dudley himself was acting as instructor even in his undergraduate days, and later became attached to the staff of the university. It is hard for us to realize in these days when every college or university of any pretensions whatever has its department of botany well- equipped and well-manned, that during the '70s the number of profes- sorships of botany in the whole United States probably did not exceed half a dozen. Cornell was one of the first of the universities to establish a distinct chair of botany, and at the time that Professor Dudley entered Cornell the chair was held by Professor Albert Prentiss. While a student at Cornell, Dudley attended the summer session of the famous school at Penikese where -Agassiz for the first time instituted a seaside summer school, the model of which has since been repeated in so many places. At Penikese Dudley was associated not only with his fellow students of Cornell but also with a nmnber of other men who laid the foundation of the biological studies which have had such a tremendous influence in the development of science since that time. 12 DUDLEY MEMORIAL VOLUME Professor Dudley very early became interested in the problem of plant distribution. The region about Ithaca is a peculiarly interesting one botani- cally, offering an unusual variety of conditions with a correspondingly varied and interesting flora. Dudley soon became intimately acquainted with the flora of this whole region and the results of his studies were later published under the name "The Cayuga Flora." This was soon sup- plemented by a second similar work on "The Lackawanna and Wyoming Flora." While at Cornell, Professor Dudley also published in collaboration with Professor M. B. Thomas a "Manual of Histology." He also published a number of other shorter papers dealing mainly with the flora of the same region. During the latter part of his stay at Cornell Professor Dudley had charge of the work on the lower plants, especially the fungi, to which he devoted much attention. In connection with this work upon the fungi Professor Dudley made a trip to Europe in 1887, and it was upon this trip that I had the first opportunity of making his acquaintance. I was myself at the time a Situdent at the University of Berlin. My first meeting with Professor Dudley was at Strasburg, where he had gone to study under the famous botanist, De Bary. Somewhat later Professor Dudley went to Berlin, where I was a student, and I had an opportunity of renewing the acquaintance so pleasantly begun at Strasburg. It is seldom that I have had the good fortune to meet a man who has made upon me a deeper impression. The extraordinarily fine quality of Professor Dudley's personality it is not necessary to describe to those who knew him. In every sense of the word he was a gentleman of the finest tjrpe. We little thought then that it was not going to be many years before we should be colleagues in a new university in far-away California, for to us then California seemed very far away indeed. Just twenty years ago a little band of pioneers, to which I had the great good fortune to belong, started our University on its career. Every- thing looked most promising and we were all full of enthusiasm and hope for the future. Two years later Mr. Stanford died, and the university entered upon a period of anxiety and privation, which was only tided over by the noble and self-sacrificing devotion of Mrs. Stanford. Professor Dudley was called to Stanford as profesisor of systematic botany in 1892, but did not come to California until the fall of 1893, just at the time when the outlook was most discouraging. He naturally had expected to have all the necessary equipment for establishing his depart- ment, and of course nobody could have foreseen the unfortunate condition of things which prevailed at the time he took up his duties in the autumn WILLIAM RUSSEL DUDLEY CAMPBELL 13 of 1893. Although it must have been a great disappointment to him, he nevertheless vigorously set to work to make the best of the situation and for several years before the outer quadrangle was built and the present botanical quarters provided, he carried on the work of his department under most discouraging conditions. His laboratories, if such they may have been called, occupied the attic of one of the shop buildings back of the quad- rangle, and were very far from satisfactory either for laboratory or her- barium purposes. However, he began collecting assiduously and before long the nucleus of the fine herbarium which he has left to the university was brought together. The flora of California is a peculiarly rich and interesting one and offers exceptional opportunities to the student of the problems of plant distribution. To Professor Dudley, whose work had been especially along these lines, the opportunities for work in his chosen field must have been very enticing, and doubtless compensated in great measure for some of the drawbacks in other respects which he must have felt keenly when he came to Stanford. From the time of his arrival, almost until his death, he made many trips to all parts of the state, collecting zealously and accumulating an invaluable herbarium which remains to remind future students of our flora of his tireless interest in his work. Professor Dudley paid especial attention to the flora of the Sierras, and was a recognized authority upon it. CalifcSrnia is pre-eminent in its coniferous forests, which are unrivaled in all the world, and Professor Dudley soon became deeply absorbed in a study of the distribution of these magnificent trees. A considerable number of these are peculiar to Cali- fornia and often of very restricted range, like the familiar Monterey cypress. Professor Dudley studied with especial care the habits and distribution of a beautiful fir {Abies venusta) which is only known to grow in the Santa Lucia range. He made a number of trips to this remote region for the purpose of studying this rarest of the Californian firs. His acquaintance however with all of the coniferous trees was most intimate, and he soon became a recognized authority on the distribution of the Cali- fornian conifers. Professor Dudley's interest in the study of the distribution of the forest trees naturally led him to a study of the problems of forestry, which for the past twenty years or so have been arousing so much interest in the United States, and which so deeply concern the welfare of the country. As might be expected, his sympathies were entirely with those who would protect what is left of our magnificent western forests from the reckless exploitation of ignorant or unscrupulous men who have so devastated the 14 DUDLEY MEMORIAL VOLUME forests of the eastern states, and are now threatening the great forests of the Pacific Coast. An intimate friend of Gifford Pinchot, who has been an effective champion of the rights of all the people in our splendid forests, which have, been so wantonly devastated, he always stood for the most enlightened views of forest conservation. The state has never had a more devoted advocate of sound and modern methods in forestry than Professor Dudley. His teaching work in the university, especially in his later years, was to a great extent strongly influenced by his interest in forestry problems, and the students who were intending to devote themselves to forestry as a profession found in his teaching a sound preparation for their future vocation. Professor Dudley's interest in forestry was evinced in a very practical way through his participation in the movement to reserve as a state park the fine body of redwood timber in the Santa Cruz Mountains known as the Big Basin. Largely through his instrumentality this magnificent body of virgin redwood forest was bought by the State as a permanent public park. Until compelled by illness to give up his position, he served as one of the commissioners of the park, in which to the last he took the keenest interest. For many years also Professor Dudley was an active and interested member of the Sierra Club, and accompanied the club in its outings in the Sierras on several occasions. Those who were fortunate enough to be members of the party and thus came to know Professor Dudley in his most congenial surroundings, will always remember with the keenest pleasure their associations with him on those excursions. As a teacher Professor Dudley was devoted to the welfare of his students, who will bear witness to his constant interest in their work and the unfailing assistance always rendered them. Many students both at Cornell and Stanford came under his influence, and the long roll of those who have achieved success in their work after leaving college bears witness to the success of his labors as a teacher. At Cornell, Professor Atkinson, the present head of the department of botany, was one of his students. Professor Trelease, the distinguished director of the Missouri Botanical Gardens, which position he recently resigned, was also a student at Cornell ; and Dr. Coville, head of the National Herbarium at Washington, also claims Professor Dudley as his teacher. Many others, successful both as teachers and investigators), look back with pleasure and gratitude to their student days in his laboratory. On our own faculty Professor Abrams and Mr. McMurphy were both associated with him as students and colleagues, and are carrying on the work which he so well began. WILLIAM RUSSEL DUDLEY CAMPBELL 15 Undoubtedly Professor Dudley's most important scientific work was the collection of the extensive herbarium to which he devoted so much time and labor during all the years that he spent in California. It is doubtful whether any botanist had a more intimate knowledge of the flora of Cali- fornia than he, and the great nvimber of specimens collected by him on his many botanical trips are now the property of the university. And the Dudley Herbarium will remain as a monument to his devoted labors as a student of California plants. A characteristic California genus, Dudleya, has been named in his honor, and will always recall to botanists the name of one of the most devoted students of the flora of our state. WILLIAM RUSSEL DUDLEY* By President David Starr Jordan THE fact that the writer has been intimately associated with Professor Dudley since the day he entered the freshman class at Cornell University, in September, 1870, will perhaps excuse the personal element in this little sketch. The word "instructor" as a technical term, describing a minor assistant to a professor, had just then been invented, and the present writer had just been appointed "instructor in botany" under Professor Albert N. Prentiss. One day Professor Henry T. Eddy, now of Minnesota, brought to me a tall, well-built, handsome and refined young man, older and more mature than most freshmen, and with more serious and definite purposes. Young Dudley had an intense delight in outdoor things and especially in flowers and birds. He wanted to be a botanist, and had turned- from old Yale, to which as a descendant of Chittendens, Griswolds and Dudleys he would naturally have gone, to new Cornell, because Cornell offered special ad- vantages in science. For the rest of my stay at Coriiell, Dudley was my roommate, living in a cottage on the hill, built by students and termed "University Grove." In this cottage was established the boarding-club, known later and appropriately as "The Struggle for Existence," and in later and more economical times as the "Strug." In time he was made botanical collector, and this congenial work he kept up until he became my successor as instructor in botany. In college Dudley was a member of the Delta Upsilon fraternity, and took an active part in holding this society to the high ideals (AiKota yiroOrjKT)) on which it was originally based. He was also a charter member in the honorary scientific society of Sigma Xi (SttouSmi/ UuvS>ves). From 1872 to 1876 he was instructor in botany at Cornell, his eminent knowledge of the eastern flora overbalancing the fact that at first he had not yet received a degree. From 1876 to 1892 he was assistant professor of botany at Cornell, with a year's absence in 1880, in which he served as acting professor of biology in the University of Indiana, in the absence of the present writer, who then held that chair. In 1892, Professor Dudley became professor of systematic botany at Stanford University, which position he held until, in January, 1911, failing * Science, N. S., Vol. XXXIV, 142-145, August 4, 1911. WILLIAM RUSSEL DUDLEY JORDAN 17 health caused his retirement on the Carnegie Foundation, as professor emeritus, his work being then taken by one of his students. Associate Pro- fessor LeRoy Abrams. Many of the leading botanists of the country have been students • of Professor Dudley. H. E. Copeland, Kellerman, Lazenby, Branner were among his associates at Cornell. Atkinson became his successor at Cornell. Abrams, Cook, Elmer, Olssen-Sefler, Cannon, Wight, E. B. Copeland, E. G. Dudley, Greeley, Herre, McMurphy and many others were under his tutelage at Stanford. In Stanford University, Dudley was one of the most respected as well as best beloved members of the faculty. No one could come near to him without recognizing the extreme refinement of his nature; a keen intellect, an untiring joy in his chosen work, and the Puritan conscience at its best, with clear perceptions of his own duties to himself and a generous recogni- tion of the rights and the aspirations of others. Dudley entered with great joy into the study of the California flora. He became especially interested in the study of trees, the evolutionary relations of forms and especially the problems of geographical distribution. The conifers of California were his special delight, and he made many field trips with his students to all parts of the state, notably to the Sierra Nevada and the Sierra Santa Lucia. His extended collections were presented to Stanford University, where with the collections of Dr. Abrams they form the major part of the large "Dudley Herbariiun." A genus of stone-crops, of many species, abounding on the cliffs of California and especially on those which overhang the sea, was named Dudleya by Britton and Rose. Dudleya pulverulenta is one of the most conspicuous plants in California wherever ''sea and mountain meet." Dudley was. instrimiental in inducing the State of California to pur- chase a forest of redwoods {Sequoia sempervirens) , that this, the second of California's giant trees, might be preserved in a state of nature. Two thousand five hundred acres in the "Big Basin" of Santa Cruz county were thus bought and established as the "Sempervirens Park." For several years Dudley served on the board of control of this park. Of the Sierra Club of California, Dudley was a leading member and for some years a director. As an investigator. Professor Dudley was persistent and accurate, doing his work for the love of it. A partial list of his papers is given below. A large work on the conifers of the west was long projected, but still exists only in uncompleted manuscript. 18 DUDLEY MEMORIAL VOLUME Dudley was master of a quiet and refined but effective English style. He was one of those scientific men, too few I fear, who have real love for literature, and who understand what poetry is and what it is about. In his early days he wrote graceful verse. Three of his poems are in print, "The Kaaterskills as seen from the Taconics," "Sunrise on the Kaaterskill" and "A Legend of the Lehigh Valley." The last is the story of the Moravian settlements of "Friedenhiitten, Tents of Peace, and Gnadenhiitten, Tents of Grace." From the first of these, I quote: 'Twas reached at last, with toiling long and weary Taconic's loftiest hill; Then, visions of all visions, stood uncovered The domes of Kaaterskill ! They rose above the lesser hills as sovereigns Above the common herd; They gathered then in conclave grand and solemn ; They breathed no spoken word. But full as anthemed voices of the ocean A soundless song was borne Up from those lips that changeless through the ages Sang on Creation's morn. A mighty calm sits on these silent summits. Time fades, as breath away. O'er all in solemn oceanic pulsings Deep flows — Eternity. From "A Legend of the Lehigh Valley'' I quote the last verses: Full six score years have passed away. Still on the silent smnmer morn, At noon's repose, or evening's gray. O'er Lehigh's vale this dirge is borne. The reaper hears, on far-off hills, And the traveler by the mountain rills. And the fisher in the evening's chills; WILLIAM RUSSEL DUDLEY JORDAN 19 They hear and feel some echo wake Of sorrow slumbering long. A tear Is shed for some sweet lost one's sake, A tear that leaves life's stream more clear. They bless the song and them who sing; They feel the s}mapathy upspring That's bom of hiunan suffering. The air is full of sad-toned bells That never cease their brazen toll; With circling suns their pulsing swells, And in one tireless world-wave roll. But grateful unto sorrow's ear From the Lehigh, far or near, Comes this dirge so sweet and clear — Come these human voices dear. Professor Dudley's health was good until about three years ago, when he set out to study the trees of Persia. In Egjnpt he was attacked by a severe cold or bronchitis which ended in tuberculosis. He never married. PROFESSOR DUDLEY'S WORK FOR STANFORD* By Professor LeRoy Abrams PROFESSOR William Russel Dudley, who became professor emeritus of botany at the opening of the present semester, although born in an old New England town that has been the home of the Dudley family since early colonial times, is essentially a pioneer. Entering Cornell University with its second freshman class, he remained in that young institution after graduation, first as instructor and later as assistant professor of botany, until the foundation of our own university, when at the urgent request of President Jordan, his college mate and intimate friend, he came to Stan- ford as one of the pioneer professors at the opening of its second year. Of Professor Dudley's experience at the very beginning of work in his new field, and of the arduous times during the dark days that en- gulfed the university soon afterward, I have no personal knowledge, for it was some four or five years after his arrival that I came to know him. Upon entering the university I sought out the department of systematic botany with the intention of carrying on some studies in flowering plants. At that time the twelve small buildings which form the inner quadrangle, and three small shop buildings in the rear of them, were the only build- ings available for university work. In my search for the department I was directed to the farthest of the shop buildings, the one situated just back of the new geology building, where I was told that I would find Professor Dudley on the second floor. And here I did find him, tucked away in one end of a loft, in a single room, one corner of which had been partitioned off as an office. In a quiet, reserved manner he talked over my work; then he took me into the main room to select a table and material for study. It was a curious room, this "laboratory," perched high amid the rafters. Three huge beams ran lengthwise of it a good hurdling distance apart, but about five feet and a half from the floor. With an apologetic smile, he warned me of these as he calmly ducked under the first. The table was soon selected and my initial study outlined. Day by day, throughout the course, as he went from student to student directing their studies, he patiently dodged those formidable beams. For ten years this man, one of America's foremost teachers of botany, conducted his classes under such handicaps. Yet with these great obstacles constantly checking the normal growth and development of his cherished plans, he labored on incessantly; his quiet, dignified, courteous manner. * From the Stanford Alumnus, Vol. XII, No. 6, pp. 165-166, February, 1911. PROFESSOR Dudley's work for Stanford — abrams 21 his thoroughness and enthusiasm in his work, his broad interests and scholarly attainments moulding the lives of his students. For none can come under his influence without, at least unconsciously, acquiring higher ideals and more serious purposes. During the summer vacations the pursuit of his botanical studies took him into the mountains and forested areas of the state, where he was con- stantly confronted with the great and shameless waste of our forest resources. He thus became one of the pioneers in the movement toward conservation, and rendered valuable service to the state and nation through suggestions and advice to the Forest Service and other authorities. The establishment of the California Redwood Park, a beautiful tract of forested land in our neighboring mountains, set ciside by the state primarily for the purpose of preserving a forest of the coast redwood in its primitive conditions, was accomplished largely through his eflEorts. And as secretary of the first park commission he labored for its betterment and the establishment of a per- manent policy in its management. But Professor Dudley saw that if the conserving of our forests was to be placed on an intelligent and permanent basis it was essential that young men be trained for the work, and that the people of the states where the forests abound be educated to the necessity of scientific forestry ; he saw that fully nine-tenths of the nation's forests lay west of the con- tinental divide, yet in all this region not one of the educational institutions was training men for the scientific management of this vast wealth. He therefore directed his energies toward the establishment of courses in forestry at Stanford. For a number of years he planned toward this end, and finally, just as success seemed probable, the fateful April 18th wiped out every promising hope of immediate realization. Soon afterward he contracted a serious illness which left him physically weakened. This hampered his work, but not his enthusiasm, and he is now retiring from the regular routine departmental duties in the hope that he may regain his health sufficiently to complete his research studies on the western flora. Professor Dudley's students and his many other friends who have known and followed his courageous and uncomplaining struggle against disheartening obstacles hope that he may not only live to complete his own studies, but that he may yet see young men trained at Stanford for the scientific management of the vast forests of the West. PROFESSOR WILLIAM RUSSEL DUDLEY* By Professor George James Peirce WILLIAM RussEL DUDLEY, professor of systematic botany in Leland Stanford Junior University from 1892 to 1911, died on June 4, 1911, at the age of sixty-two. By ancestry and place of birth a New Englander, a graduate and for twenty years a member of the botanical staff of Cornell University, a student of De Bary's for a time in Strasburg, he brought to California the mature powers of an enthusiastic student and sympathetic lover of nature, the ripe scholarship and the winning personality of the inspiring teacher. At home in the laboratory, he was still more strikingly the gracious host when he was with students and other friends out of doors, in the fields and woods and mountain forests. He knew the forests of middle California as no one else; his acquaint- ance was with individual trees, as his collection of tree portraits testifies. And his studies of their geographical distribution, following and amplify- ing the earlier studies of Asa Gray and others, gave his knowledge a degree of accuracy and detail, as well as breadth, which was very precious. It is to be hoped that his notes and other manuscripts are in such condition that his associates and successor can give them to the world. Professor Dudley's nature was so sensitive, his perceptions so fine, and his ideals so high, that he could but rarely bring himself to publish what he knew. He wished always to add to and improve what he had learned and written. Thus the botanical world had little opportunity to know his accomplishments and achievements. Besides the young men and women whose lives he has enriched, and the Forest Service which he long assisted in various ways, he contributed to the great gift to California and the nation which the state and national forests of California constitute. The "Big Basin Park," the property of the state, will preserve to all time a part of the natural redwood forest of the Santa Cruz mountains. Professor Dudley assisted in securing and preserving as a state park this part of the virgin forest of Sequoia semper- virens. It was his 'interest too which stimulated and directed the federal authorities in the selection of others of the mountain forests of California as national forests. Of courtly manner, cultivated as well as educated, of ripe scholarship and rich in the knowledge of nature, he was a lovable and elevating associate, an inspiring teacher, a devoted man of science, an honor to Stanford Uni- versity of which he was an honored member. *From The Plant World, Vol. XIV, No. 8, pp. 200-202, August, 1911. PROFESSOR WILLIAM RUSSEL DUDLEY* By Jared Treman Newman ONE of the purest and noblest souls — such as one is fortunate to come close to even once or twice in a lifetime — passed to the life beyond yesterday afternoon. Professor Dudley was a prominent scientist, "one of America's fore- most teachers of botany, one of the pioneers in the movement toward conservation," largely instrumental in the establishment of the California Redwood Park, and the secretary of the park commission; yet, it is not of these, nor of his other scientific attainments or accomplishments, that we think chiefly at this time. Of fine New England stock, cultured, with a refinement that was genuine all the way thorough, doing splendid work in his chosen profession and capable of making a great name for himself, his best service to the world was in imparting to other men higher aspirations and nobler ideals. Far back in the early days at Cornell, there was a little coterie of men gathered in close association. It included Jordan and Branner and Nichols and Gage and Fairchild and Kellerman and many others who have deservedly come to high position. Among , them all there was none of finer instincts or more lovable character than Dudley. For many years after his graduation at Cornell, and while he remained a teacher there, he was the guiding and inspiring genius of successive groups of young men. Some were taking his work. Others were attracted to him by his rare personality. Still others he sought out. What he imparted to them, and to all who came close to him, was something of priceless value. It was the very soul of the man. He withheld nothing. Absolutely un- - calculating in his unselfishness, so pure that impurity could not be thought of in his presence, a lover of nature and nature's God, his influence was constantly ennobling. Like many noble souls, he was peculiarly sensi- tive. He was hurt often when no hurt was intended. He was often melancholy, sometimes almost morbid. It has always seemed so strange that one who gave so much and so constantly should not be always happy. Perhaps he made up for it in the intensity of his joys. While he was often misundertood and while the number of persons who came close to him was not relatively large, yet few men have merited, or have known, in so large a degree, the love of their fellows. *From The Palo Alto Times, June 5, 1911. 24 DUDLEY MEMORIAL VOLUME A lover of truth. and imbued with the scieptific spirit, he might have become more famous had he spent more time in research and in publishing the results; but his principal work is of the kind that lives in the hearts of living men, and goes on, and will continue to go on, in a generation of workers who owe to him the touch that makes their work worth while. WILLIAM RUSSEL DUDLEY [Read before the Stanford Alumni Association at Washington, D. C, November 11, 1911.] By William Franklin Wight DURING the early summer one of Stanford's most lovable teachers closed his life's work and found that last long rest which must come to us all. I wish therefore to-night to pay a brief tribute to the memory of Professor William Russel Dudley. His kindly feeling and in- terest in his students made him loved by them all, and he possessed that indescribable quality in a teacher that without thought and without effort instantly arouses enthusiasm in the laboratory and in the classroom. He was an unusual teacher, and it is a sad thought to realize that years before the allotted time of life his voice will be heard no more in the classroom and his charming manner will be unknown to the students who shall fill the halls of Stanford. It was my fortune to be with him on the last day. I had visited him a few weeks earlier, and then he was hopeful that there might still be left to him a few years in which to complete, the botanical work that he had begun almost immediately on coming to California. Nevertheless, those who saw him knew that it was even then too late — that the end must soon come. It was therefore with a sad heart that I went on the morning of June 4th to pay a last visit to my friend and teacher. From the balcony where he lay in the cottage at Los Altos one could look across the valley to the Mount Hamilton range bathed in sunlight, and view the glory of a California landscape. The air was crisp and full of life to the strong. It was indeed a beautiful day in which to live, but there with the vision of nature he loved so well before him, now too far away for his eyes to see, in the midst of a few friends, he calmly awaited the end. It is however of other days that we would keep the memories fresh. We would rather remember him strong and enjoying the activities of a busy life. And I think he took keen pleasure in all his work, for he appeared to go through each year at the university with an enthusiasm equal to that we should expect if the studies and discoveries of the laboratory were as new to him as to the student. But it was on long tramps in the mountains, in the solitude and grandeur of the redwood forest, that one really began to appreciate the fineness of the man, to know how much he saw in mountain and forest, and how much he loved nature in sunshine and in storm. At 26 DUDLEY MEMORIAL VOLUME fifty years there was the freshness and joy of youth in botanical exploration. It was when on such walks too that one came to know the fullness of his knowledge and how perfect was his memory, as every species was recognized and its distribution or other fact of interest was related. In many ways his life at Stanford was a disappointment. He felt the burden of the years of financial stress through which the university passed more than was his share, and very often supplied from his own purse the necessary materials for the laboratory. The hopes and ideals he had for the development of botanical science he could not live to see realized. But whatever was lacking in appropriate rooms and equipment was more than compensated for in the ability and spirit of the teacher. He lived in his work and for his students. His time and energy were so very largely oc- cupied in their interest that he published little, and this is the regret of all who realize the high scientific ideal which guided him in his work, and who appreciate the charm of his literary style. His Flora of Ithaca and of the Wyoming Valley will be regarded as classics and as models of their kind for many years to come. For some it is not given to publish much — ^it is theirs to write in the hearts and minds of men and women, an influence as enduring perhaps as that of printed books. I never heard him speak ill of any person but once, and then he did it deliberately, reluctantly, and as though he felt it a painful duty. It was his habit to see the good qualities in mankind and he did it naturally and without effort. I trust that so long as modesty, thoughtfulness, and a kindly spirit are regarded as evidences of a fine character, that so long the memory of Professor William Russel Dudley will live at Stanford University. PUBLICATIONS OF WILLIAM RUSSEL DUDLEY First Steps in the Study of Botany. Educationist, 5: 7-10. 1880. Leafy Berries in Mitchella repens. Bull. Torr. Bot. Club, 10 ; 1-3. 1883. An Abnormal Orchid. Abstract of a paper read before the American Association for the Advancement of Science. Proc. Amer. Assoc. Adv. Sci. 32: 30. 1883. Sketch of Curtis. Jour. Mycology, 2: 54-59. 1886. Elias Magnus Fries. Jour. Mycology, 2: 91-94. 1886. Charles Christopher Frost. Jour. Mycology, 2: 114-118. 1886. The Cayuga Flora, Part 1. A catalogue of the Phaenogamia growing without cultivation in the Cayuga Lake Basin. Bull. Cornell Univ. Sci. 2: XXX, 1-132. 1886. A Preliminary List of the Vascular Plants of the Lackawanna and Wyoming Valleys. Proc. and Coll. Lackawanna Inst. Hist, and Sci. 1: 32-112. 1887. Strasburg and its Botanical Laboratory. Bot. Gaz. 13: 305-311. 1888. The Death of DeBary. Bot. Gaz. 13: 64-65. 1888. Report of the Cryptogamic Botanist. Rep. Cornell Agri. Exp. Sta. 1-3. 1888-1890. The Strawberry Leaf -Blight. Rep. Cornell Agri. Exp. Sta. 2: 171-182. 1889. Another Disease of the Strawberry. Rep. Cornell Agri. Exp. Sta. 2 : 182-183. 1889. Anthracnose of Currants. Rep. Cornell Agri. Exp. Sta. 2: 196-198. 1889. Leaf -Blight of Quince and Pear. Rep. Cornell Agri. Exp. Sta. 2: 198-199. 1889. The Onion Mold. Rep. Cornell Agri. Exp. Sta. 2: 193-196. 1889. The Clover Rust. J. K. Howell. Footnote by W. R. Dudley. Rep. Cor- nell Agri. Exp. Sta. 3: 129-139. 1890. The Hollyhock Rust. Rep. Cornell Agri. Exp. Sta. 3: 154-155. 1890. Flora of the Lackawanna and Wyoming Valleys; a catalogue of the flower- ing plants and vascular Cryptogams found in and near Lackawanna and Wyoming Valleys. With Charles O. Thurston. Wilkesbarre, Pa. XV, 96. 1892. The Genus Phyllospadix. Wilder Quarter- Century Book, Comstock Pub- lishing Co., Ithaca, N. Y. 403-420. 1893. Botanical Notes. Bull. Torr. Bot. Club, 20: 169-170. 1893. Phyllospadix; its Systematic Characters and Distribution. Zoe, 4: 381-385. 1894. 28 DUDLEY MEMORIAL VOLUME A Laboratory Manual of Plant Histology. With Mason B. Thomas, Craw- fordsville, Ind. viii, 115. 1894. Forest Reservations; With a Report on the Sierra Reservation, California. Sierra Club Bull. 1: 254-267. 1896. The Kaweah Group. Sierra Club Bull. 5:185-191. 1898. Forestry Notes, editor of. Sierra Club Bull. 2-7. 1898-1910. A Short Account of the Big Trees of California, (one of the collaborators). Bull. U. S. Dept. Agri. Div. Forestry, No. 28; 1-30. 1900. The Big Trees of California. Forester, 6.: 206-210. 1900. Lumbering in the Sequoia National Park. Forester, 6: 293-295. 1900. Zonal Distribution of Trees and Shrubs in the Southern Sierra. Sierra Club Bull. 3: 298-312. 1901. Big Basin Redwood Park. Forester, 7: 157-164. 1901. A Notable California Fir. Forestry and Irrigation, 8: 193-198. 1902. Trees along the Tulare Trails. Sierra Club Bull. 4: 153-156. 1902. Trees of Southern California. Los Angeles Saturday Post, 5-6. June 7- August 2, 1902. Near the Kern's Grand Canon. Sierra Club Bull. 4: 301-307. 1903. Notes on California's Uredineae and Descriptions of New Species. (With C. H. Thompson.) Jour. Mycology, 10: 52-55. 1904. The Vitality of Sequoia Gigantea. Read by invitation before the California Alumni Association of Columbia University. Privately printed by mem- bers of the association. San Francisco, 16. 1905. ' Concerning the Vitality of Sequoia Sempervirens. Palo Alto Times, March 17, 1908.- CORNELL UNIVERSITY PUPILS OF PROFESSOR DUDLEY Arthur, Joseph Charles; B.S., M.S., Sc.D. Professor of Vegetable Physiology and Pathology, Purdue University, Lafayette. Atkinson, George Francis; Ph.B. Professor of Botany, Cornell University, Ithaca. Bray, William L.; A.B. A.M., Ph.D. Professor of Botany, Syracuse University, Syracuse. Chester, Frederick Dixon; B.S., M.S. Bacteriologist, (formerly) Director, State Bacteriological Laboratory of Delaware. Corbett, Lee Cleveland; B.S., M.S. Horticulturist, Department of Agriculture, Washington. CoviLLE, Frederick Vernon; A.B. Botanist, Department of Agriculture, Washington. Craig, Moses; M.S. Missouri Botanical Garden, St. Louis. Densmore, Hiram Delos; A.B., A.M. Professor of Botany, Beloit College. Durand, Elias Judah; A.B., Sc.D. Assistant Professor of Botany, University of Missouri, Columbia. Gregory, Emily Lorina. Late Professor of Botany, Barnard College, Columbia University, New York. Henderson, Louis Pourniquet; Ph.B. Formerly Professor of Botany, University of Idaho, Moscow. Hicks, Henry; B.S. Nurseryman, Westbury, Long Island. Hoffman, Harry Natt; B.Agr. Nurseryman, Elmira, New York. Hough, Romyn Beck; A.B. Dendrologist, Lowville, New York. Howell, Jenny Kirk; Ph.B., M.S. Teacher, Plainfield, New Jersey. Kellerman, William Ashbrook; B.S., Ph.D. Late Professor of Botany, Ohio State University, Columbus. Lazenby, William Rane; B.Agr., M.Agr. Professor of Forestry, Ohio State University, Columbus. MooRE, Veranus Alva; B.S., M.D. Director, New York State Veterinary College, Cornell University, Ithaca. Norris, Harry Waldo; A.B. , A.M. Professor of Zoology, Grinnell College, Grinnell, Iowa. Porter, Edna. Botanical Gardens, Buffalo, New York. RowLEE, Willard Winfield; B.L., Sc.D. Professor of Botany, Cornell University, Ithaca. Schrbnk, Hermann Von; B.S., A.M., Ph.D. Consulting Timber Engineer, Plant Pathologist, St. Louis. Smith, Theobald; Ph.D., M. D., A.M., LL.D. Professor of Comparative Pathology, Harvard Medical School, Boston. Snow, Julia Warner; B.S., M.S., Ph.D. Associate Professor of Botany, Smith College, Northampton. Thomas, Mason Blanchard; B.S. Late Professor of Botany, Wabash College, Crawfordville. .Trelease, William; B.S., Sc.D., LL.D. Formerly Director, Missouri Botanical Garden, St. Louis. White, Charles David; B.S. Geologist, Geologic^ Survey, Washington.. Yatabe, Ryokichi; B.S. Late Professor of Botany, Imperial University, Tokyo, Japan. STANFORD UNIVERSITY PUPILS OF PROFESSOR DUDLEY Abrams, LeRoy; M.A., Ph.D. Associate Professor of Botany, Stanford University, California. Aldrich, John Merton; Ph.D. Professor of Biology, University of Idaho, Moscow, Idaho. Anderson, Malcolm Playpair; B.A. Naturalist, British Museum, London, England. Atkinson, William Sackston; B.A. Scientific Illustrator, Stanford University, California. Baker, Charles Fuller; M.A. Professor of Zoology, Pomona College, Claremont, California. Bell, Ruby Green (Mrs.); M.A. M. Albert W. Smith, Ithaca, New York. Berry, Samuel Stillman; M.A., Ph.D. Graduate Student, Stanford University, California. Billings, Frederick Horatio; M.A., Ph.D. Associate Professor of Botany and Bacteriology, University of Kansas, Lawrence, Kansas. Boring, Oramanda. Teacher of Botany and Physiology, Stockton High School, Stockton, Cali- fornia. Bryan, Mary Katherine; B.A. Scientific Assistant, United States Department of Agriculture, Washington, D. C. Burke, Charles Victor; M.A., Ph.D. Graduate Student, Stanford University, California. Burnham, Stewart Henry. Botanist, New York State Museum, Albany, New York. Cannon, William Austin; M.A., Ph.D. Staff Member, Desert Laboratory, Carnegie Institution, Tucson, Arizona. Chapman, Bertha Louise; M.A. Lecturer, Writer, Teacher of Private Nature Study Classes, Kansas City, Missouri, M. Vernon Mosher Cady. Chase, Raymond Eugene. Principal, Reno High School, Reno, Nevada. Cook, Melville Thurston; M.A., Ph.D. Plant Pathologist, Delaware Agricultural Experiment Station, Newark, Delaware. Cooper, Alice Cecilia; M.A. Teacher, Los Angeles Polytechnic High School, Los Angeles, California. Copeland, Edwin Bingham; Ph.D. Dean and Superintendent, College of Agriculture, University of Philippines, Los Banos, P. I. Couch, Mary Juanita; B.A. Teacher, Vacaville, California. Cravens, Mary Ruhama; M.A. Teacher, Sacramento, California. Doane, Renne Wilbur; B.A. Assistant Professor of Entomology, Stanford University, California. Dudley, Ernest Griswold; B.A. Assistant Supervisor, Sierra National Forest, North Fork, California. Elmer, Adolph Daniel Edward; M.A. Botanist and Botanical Collector, Publisher of Botanical Leaflets, Manila, P. I. Fisher, Walter Kendrick; M.A., Ph.D. Assistant Professor of Zoology, Stanford University, California. FuLLAWAY, David Timmins; M.A. Entomologist, United States Agricultural Exp&iment Station, Honolulu, T. H. stanford university pupils of professor dudley 31 Geis, Helen Dudu; B.A. Teacher, Los Angeles Polytechnic High School, Los Angeles, California. Greeley, Arthur White; M.A. Late Professor of Zoology, Washington University, St. Louis. Grinnell, Joseph; M.A. Director, Museum of Vertebrate Zoology, University of California, Berkeley, California. Halsey, Stella Duffield; B.A. Teacher, San Diego High School, San Diego, California. Heller, Edmund; B.A. Naturalist of the Roosevelt Expedition to Africa, 1909-1910, United States National Museum, Washington, D. C. Herrb, Albert William Christian Theodore; M.A., Ph.D. Vice-Principal, Fruitvale High School, Fruitvale, California. HiGLEY, Rose Miriam; M.A. Teacher, San Rafael High School, San Rafael, California. HoLMAN, Richard Morris; B.A. Instructor in Botany, College of Agriculture, University of the Philippines, Los Banos, P. I. Humphrey, Harry Baker; Ph.D. Assistant Professor of Botany, Washington State College, Pullman, Wash- ington. Humphrey, Olive Agatha Mealey (Mrs.); B.S. Pullman, Washington. Jenkins, Hubert Oliver; B.A. Health OfiBcer, Palo Alto, California. Kimura, Tokuza; B.A. Teacher of Biology, Sotokufu Chugakko, Formosa, Japan. Knoche, Edward Louis Herman; B.A. Student of Botany, traveling in Europe until 1912. Kroeck, Louis Samuel; M.A. Teacher, College of the Pa'cific, San Jose, California. Kuwana, Shinkai. Inokichi; M.A. Imperial Agricultural Experiment Station, Tokyo, Japan. McCracken, Mary Isabel; M.A., Ph.D. Assistant Professor of Entomology and Bionomics, Stanford University California. McGregor, Ernest Alexander; M.A. United States Bureau of Entomology, Washington, D. C. Mackay, Minnie Laurie; B.A. Teacher, Santa Clara High School, Santa Clara, California. McMuRPHY, James Ira Wilson; M.A. Instructor in Botany, Stanford University, California. Miller, John Martin. Forest Assistant, United States Forest Service. Morris, Charles Shoemaker; M.A. Teacher of Botany and Zoology, Palo Alto High School, Palo Alto, CaU- fomia. Morris, Earl Leonard. County Entomologist, Santa Clara County, and Field Assistant, University of California, San Jose, California. Nohara, Shigeroku. Assistant in Botany, Agricultural College, Imperial University, Tokyo, Japan. Olsson-Sepper, Pehr Hjalmar; Ph.D. Deceased. Pemberton, Cyril E.; B.A. Field Agent, United States Bureau of Entomology, Lindsay, California. Pemberton, John Rothwell; B.A. Geologist, Argentine Republic, South America. Peterson, Elsa; B-A. Teacher, Normal School, Honolulu, T. H. 32 dudley memorial volume Price, William Wightman; M.A. Fallen Leaf, Lake Tahoe, California. Randall, Josephine Dows; B.A. Sometime Assistant in Botany, Stanford University, California. Randolph, Flora Albertine; M.A. Principal Miss Randolph's School, Berkeley, California. Rose, Jessie Perkins; M.A. Teacher, Fort Klamath, Oregon. Rust, Everett Winder; B.A. Assistant Government Entomologist, Lima, Peru. ScoFiELD, William Launcelot; B.A. Student, Yale Forest School, Yale University, New Haven, Connecticut. Seale, Alvin; B.A. Chief of Department of Fisheries, Bureau of Science, Manila, P. I. Shafer, George Daniel; M.A., Ph.D. Engaged in research work in Entomology, East Lansing, Michigan. Sherfy, Samuel Hash; B.A. Mount Morris, Illinois. Show, Stuart Bbvier; B.A., M.F. Forest Assistant, United States Forest Service. Smith, Charles Piper; M.A. ' Assistant Professor of Botany, Utah Agricultural College, Logan, Utah. Snodgrass, Robert Evans. Artist, New York City. Stark, William Harvey. Nurseryman, Stark City, Missouri. Stokes, Susan Gabriella; M.A. Teacher, Orange Union High School, Orange, California. SwENSON, John Canute; B.A. Professor of History and Economics and Dean of the College, Brigham Young University, Provo, Utah. Thompson, Charles Henry. In charge Department of Succulent Plants, Missouri Botanical Garden, St. Louis, Missouri. Tracy, Hiram Harwood. Wight, William Franklin; M.A. United States Department of Agriculture, Bureau of Plant Industry, Wash- ington, D. C. Williams, Florence; B.A. Sometime Assistant in Botany, Stanford University, California. Williams, Francis Xavier; B-A. Curator of Insects, University of Kansas, Lawrence, Kansas. WiNSLOw, Martha Minerva; M.A. Teacher of Physiology and Botany, Pasadena High School, Pasadena, Cali- fornia. Zschokkb, Theodore Christian; B.A., M.F. Forest Assistant, United States Forest Service. SCIENTIFIC PAPERS THE VITALITY OF THE SEQUOIA GIGANTEA * By William Russel Dudley EXISTING along the western slopes of the Sierra Nevada range in Cali- fornia, in isolated groves from Placer County to Tulare, Sequoia gigantea is a relict of another age and time. In these trees we have — with the Sequoia sempervirens, the redwood of the Coast Ranges — the remnants of a great genus that once spread over all the Northern hemi- sphere, as evidenced by fossil specimens of a considerable number of species found from France and Hungary to Spitzbergen, and from Greenland to Oregon and Nebraska. These fossils are found from the Cretaceous, through the Tertiary, down to recent times. How the species have disappeared and the individuals of the Big Trees have shrunken to the few thousands perched among the high salubrious valleys of our Sierra, cannot be easily answered. The unknown complex of causes which brought about the great ice age, brought Sequoia as a race near to extinction; and conditions surrounding life were so profoundly changed, that their former distribution could never be again restored. Whatever the cause of the present restriction of this species, its comparative rarity, its inaccessibility, its great size, its majesty and the beauty if its coloring have all served to enhance the interest which we all feel in the California Big Tree. Indeed it is so noble, that it has been the subject of a considerable amount of exaggeration and mistaken comment. The statements that its age reaches 4,000 to 6,000 years, and its height exceeds 400 feet do not seem to be based on any actual observation. Nevertheless it is crowned with many titles to greatness, and the most remarkable of all is its relative ap- proach to immortality. The evidence that all living things are finite is so overwhelming that the mind is chastened with the thought of it. But the life of a single great tree of Sequoia gigantea, when known clearly, stirs the imagination again to thoughts of what might be attained, if disease and the crushing weight of physical injury, as factors controlling life, could be eliminated. Certainly the oldest of the Big Trees, such as we see in the Calaveras groves and the forests of the Kings and Kaweah rivers, have the distinction of being the oldest, the longest enduring upon the face of the earth, of any living organism; and this is largely because of their freedom from disease and inherited weakness and, as I propose to show a little later, from their marvelous recuperative power in the face of injury. * Read by invitation before the California Alumni Association of Columbia University, January, 1905. 34 DUDLEY MEMORIAL VOLUME The forests of the Sierra Nevada in October are not dissimilar in aspect to those of the Appalachian mountain ranges. Yellowing oaks lighten the somber conifers, and crimson dogwoods lend an aspect of brilliancy to the forest, unknown to the camper beneath its shade in summer. Even Sequoia exhibits a warm golden tint, due to a thousand small yellow branch- lets which are maturing prepjiratory to the annual natural pruning of the species. It was a pleasure to incidentally note these forest charms when in October, 1900, I made my way into the lumber camp belonging to the Sanger Lumber Company in the Converse Basin near the Kings River. This mill is probably the largest in capacity of any along the forested slopes of the Sierra Nevada. During the previous month of August it cut 200,000 feet of lumber a day, or considerably above 5,000,000 feet for that month. The records for the other working months of 1900 fell some- what short of this amount, but an enormous quantity was flumed for forty miles down to the railroad at Sanger, in the San Joaquin Valley, 6,000 feet below the mill. While some of this was pine and fir, the greater proportion was made from the giant trunks of the California Big Tree. Had a measur- ably large amount of these trunks been utilized for lumber the cutting might have been justified from the Ixmiberman's point of view; but frequently one-half to even three-fourths or seven-eighths of the great trunks were broken and rent beyond use in falling. Not anywhere in the world is there such wasteful lumbering, and this is a species that above all trees, should be saved from the lumberman ! The Converse Basin, before its deforestation — for its forests have now been entirely leveled — ^presented for observation and study the best develop- ment of this rare coniferous species that existed. The trees were large and continuous in area, and this high mountain "basin," like all others on the slopes of the Sierras containing Sequoias, is watered by small brooks of sparkling spring water. Here, too, the streams soon plunge by cataracts into the profound gorge of the Kings River, thus ensuring excellent drain- age and good conditions for growth ; and here a brief visit in the summer before had shown me the great number of cut trees with logs and stumps remaining, which gave an unrivaled opportunity to continue certain ob- servations already begun. My object, while determining the age of the trees by means of the number of their annular layers of wood, was to observe their record year by year, century by century, of their behavior toward nutrition, injury or disease. The age of a tree can only be told by counting the concentric rings of growth on the cross-section of the felled trunk. The question may be asked: Does each ring represent a year's growth? A considerable VITALITY OF THE SEQUOIA GIGANTEA — DUDLEY 35 number of observations on several species of conifers and oaks enables us to answer that it does, approximately, in those observed on the Pacific Coast. If exceptional seasons cause variations from this rule, the variations would be small in number and not greatly affect the totals. During my examina- tion of the felled trees of the Kings River, it was a part of my task to carefully traverse these records of growth; but I will here give you briefly only the results. Of the various trunks of Sequoia gigantea examined rang- ing from 900 years upward, the oldest possessed 2,425 rings, or had begun its existence 525 years before the Christian era. Extended scrutiny un- doubtedly would bring to light trees even older than this, but I do not expect any to exceed 3,000 years of age. It has often been inferred that the size of a Big Tree bears an approxi- mately exact relation to its age. If a tree exists eighty feet in circumference five feet above the base, it was inferred, it would be twice as old as one forty feet in circumference. This was found to be very far from true. The favorite situation of the larger trees is near some hollow, where a tiny perennial spring brook is always flowing. The soil should be good and deep, but with a large amount of mineral matter in it; and above all, I think well drained, though always moist. One tree occupying such a situation and at the confluence of two small Sierra Brooks, was over eighty feet in circumference ten feet from the ground, but was only 1,510 years old, all the rings being measurably thick and uniform. It felt the effects neither of drouth nor of unusual precipita- tion, and it had never been burned beneath its bark. On the other hand, the tree which a little later I shall use as the chief illustration of this paper, was a small tree for one of its age. It stood on a hillside not near a stream; the influence of years of abundant rains and nutrition were shown by rings of fair degree of thickness; the effects of years of scarcity were seen in rings so thin that fifty of them would not cover an inch of the tree's radius. Moreover, from its unprotected situation it had been seriously attacked by forest fires, each burning away portions of its sap-wood and thus assailing the vitality of the plant. This tree was only thirty-nine feet in circumference ten feet from the ground, but had attained the age of 2,171 years and a height approaching 300 feet, although injury and failing strength had resulted in a dead and broken top and reduced the tree to 270 feet at the time of its destruction in 1900. Observations of the greatest interest, however, concerned the Big Tree's behavior toward severe injury; evidences of a remarkable recuperative power being found after examination of the Sequoias of the Converse Basin. The effects of certain tremendous forest fires were registered in the trunks of 36 DUDLEY MEMORIAL VOLUME these trees, but the record was completely concealed by subsequent healthy growth. Among a nimiber of similar cases the most instructive record of these ancient fires was observed in the tree of moderate size — the one of 2,171 years of age above mentioned. This tree, when felled, had an enormous surface burn on one side, occupying eighteen feet of the circumference and with a height estimated at thirty feet. The fire had eaten through the sap-wood and deeply into the heart. It was an immense black scar and an apparently irreparable wound upon a tree already advanced in age, even for a Sequoia. Yet this was not the only similar injury it had suffered, and before we describe the remarkable life of this tree as registered within, let us see how a Sequoia goes about the repair of such appalling wounds. A burn on the bole of a giant Sequoia occurs usually on one side only — the side toward the forest fire. It may be a foot wide or even, as in this case, occupying an enormous area of the trunk surface; it may be of great height, it may decrease the tree's vitality and yet not fatally injure the individual thus attacked. While there is life there is growth. After the wound comes the healing; and there is nothing more insistent (if we may use the word) in the processes of plant life than the attempt of a strong tree to cover a wound on its surface. We have two words in our language in which the pronunciation and even the correct orthography is the same, but each has a different meaning, together with a very distinct, ancient and highly respectable ancestry; I mean the word heal. It is a curious fact, moreover, that one of these words is properly applied to the process of healing seen in animal life, the other describes the process of covering a wound, such as that adopted by the tree. The first and frequently used word heal means to make sound, and implies that, after a wound, new tissue organically connected with the old has been formed, that the muscular, the circulatory and the nervous systems have been extended in a normal way from the old to the new, and conditions in the once injured part have been restored to complete and harmonious working order. The latter word heal means to cover or conceal. It is chiefly observed in the gardener's art in the expression "to heal in" (less correctly "heel") as applied to. nursery seedlings. It is a much rarer word than the former, but good old English, and we are told that it may be traced back through the German, the Gothic and even the Latin. We remark, by the way, that so far as we know, not only the English word, but its whole family-tree clear back to the Latin root, is no older than single individuals of Sequoia gigantea to-day standing in the full vigor of life in the groves of Calaveras and Kaweah. VITALITY OF THE SEQUOIA GIGANTEA — DUDLEY 37 It is the latter word we use in this paper. The effort of an injured tree is not one to re-establish organic connection of the new tissue with the old, injured surface below, but wholly one to complete and re-establish, by extension, the broken circle of growth — the broken annual rings — to round out the tree again to its full circumference, to establish roots below, sup- porting and sap-conducting tissue above. Fortunately for the tree, it has no nervous system connected with delicately organized "nerve centers" ; no circulatory system extending to every point of its surface and connected with and controlled by a small uncertain organ deep within the body. The heart-wood of a Big Tree is imperishable while the tree stands and long after it falls, unless attacked by fire. In the words of the foreman of the logging camp, "nothing hurts the heart of a redwood — nothing; it's always sound." Moreover, it is completely independent of the living zones outside of it, although joined cell by cell with the living tissue. Its cells have ceased to grow or change, and the living juices of the plant have ceased to flow in them. From the point of view of life, whatever tissue in a tree has ceased to grow in every sense, has ceased to be vitally useful. Only that tissue which is in the process of building is living, is a vital part of the great organism, and this life must exist in a complete cylinder forming the outer tissues of the trunk; a circle constantly, and in the case of a Big Tree, indefinitely widening. If the circle is broken, apparently all the energies of life and growth are directed toward closing it again, not toward any useless vivification of the dead cells of the wound below. The burned surface is dead tissue, not differing essentially from that of the tree's heart- wood, and the healing of the latter is only incidental to the tree's supreme effort at the extension of its living tissue over the wound in order to re- unite the margins of the zone that should have remained inviolate and un- broken. The increase of the tree is rhythmical, as we have seen, accompanying the sun and the seasons. The Spring after a wound has occurred, the tree begins its effort toward healing by the formation of a layer of wood and bark along all margins of the burned area. This is repeated the second year, a layer of new wood tissue being superimposed upon that of the previous year along the burned margins. These layers next the wound are much thicker than the ordinary ones, the ring for the same year on the side of the tree opposite the burn being often correspondingly thinner and more attenuated. This process continues with each returning season, and the new tissue reaching inwards from all margins of the injury takes on the form of solid folds of wood growing uniformly broader on each side, the black char narrower, year by year. There is no organic union, however, 38 DUDLEY MEMORIAL VOLUME between the new wood of the folds and the wood of the charred surface underneath them, no healing at this point of contact, in the ordinary sense of the word; but there is effectual covering, or healing in the rarer sense, according to the tree trunk's way. Sometimes, from the attack of insects on the rapidly formed wood of the folds, these folds die. There is no surgeon present to cut away this dead tissue, but the tree patiently begins to form a new fold to cover the dead one. In a species with the ordinary span of life the delay, this waste of effort, might be fatal to the final closing of the wound. Not so with the Big Tree, to whom a score of years is as one. The first fold is overtaken and passed — in one case it took just fifty years to do it — and sooner or later the two folds from opposite sides touch one another ; a few years more and the bark is pinched out, the charred surface is entirely covered, and finally the annular layers become continuous around the entire circumference, each resuming a normal thickness through- out. The process, which has drawn on all the resources of the plant, it may be for scores of years, it may be for centuries, is completed. The wound is healed! This is a momentous event, yet only the spectacle of a perfected cylinder with the splendid circumference of forty or sixty or ninety feet, of living tissue through which the sap of the tree can pass, to a considerable extent laterally as well as vertically, is the result; only con- tinuous healthful growth and unbroken increase, the most inspiring of all spectacles. In the life history of a Big Tree such injury, such prolonged but com- plete and thorough repair may occur not only once but several times, and yet all evidence of the various catastrophies be entirely obliterated except for the thin cavities, each with one charred surface, and the peculiar struc- ture of the repair layers deep within the undecayed heart of the tree. When the tree of 2,171 years of age was cut, in addition to the great burn on its trunk eighteen feet in width, the record of three other fires was revealed. The history of the tree was as follows: It began its existence 271 B. C. At the beginning of the Christian era it was estimated to be already about twelve feet in circumference just above the base. At 516 years of age (A. D. 245) occurred a burning three feet in width on the trunk. One hundred and five years were occupied in healing this wound. One thousand, one hundred and ninety-six years without injury followed. At 1,712 years of age (A. D. 1441) occurred a second burning, making two wounds of one and two feet each in width. Each had its own system of healing. VITALITY OF THE SEQUOIA GIGANTEA DUDLEY 39 One hundred and thirty-nine years of growth followed, including the time occupied by the covering of the two wounds. At 1,851 years of age (A. D. 1580) occurred another fire, causing a burn on the trunk two feet wide, which took fifty-six years to heal. Two hundred and seventeen years of growth followed the fire. When the tree was 2,068 years old (in 1797) a tremendously aggressive fire attacked the trunk (perhaps aided by the burning stem of a neighbor- ing pine or fir) and burned the great scar eighteen feet wide with a height estimated at thirty feet. The 103 years which had elapsed since 1797 had reduced this to fourteen feet in width. If the same rate of growth con- tinued without interruption — a hazardous estimate — and the tree had been in possession of the United States and under its protection, the wound might have been closed in three and one-half centuries more, or about the year 2250. Four centuries and a half to repair in one tree the results of one forest fire ! If the tree had been a younger tree, less the victim of previous fires, we are convinced that such a healing would be possible. In any case Sequoia gigantea practically stands alone, sublime among living objects in its ability to withstand an injury of this magnitude, and to endure a sufficient length of time for its complete recovery. It is to be noted that in the trunk next to each of the three older burns described, there is a thin cavity chiefly occupied by the charcoal of the burned surface (some of that formed in 245 was brought away), and that this produced a pathologically protective covering, no doubt calculated as well as any to prevent decay during the long period consumed in cover- ing the wound with healthy tissue. But this will not account for all this superb resistance to the attack of insect, fungus, ferment or microbe. Burned areas of other trees have the same charred surface, but no oak or sycamore, pine or Douglas spruce under similar conditions would remain so long with- out being attacked in this region by some cause of decay. There is some- thing in the sap of the Big Tree that is an elixir of life, something deposited in the lignified cells of the normally formed layers of wood that resists in an unexampled way the dreaded "tooth of time." The wound is finally covered — not healed, in the surgeon's sense — the new tissue formed above it is thickened, the tree is rounded to its original fullness, bark and wood become continuous about the whole circumference, the latter forming in rings of normal uniformity, the old healthful symmetry of life is re-estab- lished, and no outward sign of distortion exists, or even a scar from the old injury. Nevertheless, well within, and as the centuries pass, deeper and deeper within the heart of the tree the wound exists unchanged and there- fore no source of decay. 40 DUDLEY MEMORIAL VOLUME Again, it is to be observed from the notes of the tree's yearly growth, that after it was out of its first youth, the periods of most vigorous increase were after the successive burnings and during the periods of healing. In part this accounted for the lessened area of the tree's live circumference; but as this thickening of the rings appears to continue after complete healing has taken place, when the tree is again forming tissue over its entire cir- cumference, and as this phenomenon was seen in other trees similarly in- jured, one is led to believe that an increased activity in the tree's life had been occasioned not by the burn, but from the effort at healing and recover- ing from what threatened to be a vital wound; and that this activity led to more vigorous growth. It is a curious fact, moreover, that in the middle of the long period of freedom from fire, from 245 to 1441, a period of nearly twelve centuries, the tree made its least relative increase in diameter. Peace and apparent prosperity had been coincident with a sluggish growth if they had not been the cause of it. About three and one-half inches was the most frequent amount of radial growth during one century, a total increase of about seven inches in the diameter of the stem; but during the first six hundred years its average was five inches (a total increase of ten inches in diameter each century) and during the first and fourth centuries of this tree's existence its radial increase was six inches in each case. During the seventh, eighth, ninth, tenth and eleventh centuries the increase in the radius of the stem was between two and three inches only per century. It was during this period, from the seventh to the twelfth century of its existence — the period of greatest depres- sion in the apparent vitality of the tree — that the rings or annular layers be- came so thin that it was impossible to count them without the aid of a lens : over forty layers were frequently found in one inch of radial line, and in two cases apparently there were fifty^two and fifty-four layers in each inch.' / I cannot help thinking we are here in the presence of one of the most remarkable products of the globe, not excepting those of human civilization. Almost no structure erected by human hands has come down to us intact through the lifetime of a Sequoia; and the few we can admire are hewn from inanimate marble or granite and cannot be compared to a living or- ganism, vast in life and complete in the records of every year of its existence. An empire or republic may be compared to the life of these great trees. But what empire or republic has lived for twenty-five centuries? None worthy of the name, and certainly none among those of the Aryan civilization. Then in the building of a Sequoia, no blood has been shed through all its twenty-five hundred years, no injustice or oppression has secured the means necessary for its construction, no hate or strife has been engendered, VITALITY OF THE SEQUOIA GIGANTEA DUDLEY 41 no accident occasioning pain or suffering or the extinction of human life has left a stain on the history of its growth. Tragedies and great passions, as we have seen, have crossed its silent life, but they have been the elemental passions of fire and storm, clean and wholesome, and the tree has been stimulated by them to a greater and more vigorous growth. Indeed, there is something sublime in the patience of the task and the completeness of its execution when, after centuries of slow rebuilding, we see every outward trace of its injuries eliminated and a robust and uninterrupted life again at- tained. Mr. James Bryce, in his sketch of the life of the late Lord Acton, Professor of History at the University of Cambridge, says: "Twenty years ago, at midnight in his library at Cannes he expounded to me how a history of Liberty might be written and in what way it might be made the central thread of all history. He spoke like a man inspired, seeming as if from some mountain summit high in air he saw beneath him the far winding path of human progress, from dim Cimmerian shores of prehistoric shadow, into the fuller, yet broken and fitful light of the modern time. * * * it was as if the whole landscape of history had been suddenly lit up by a burst of sunlight. I had never heard from any other lips any discourse like this, nor from his did I hear the like again." '^ The impression made was not dissimilar, on that cloudless October afternoon with the crimson leaves of the dogwood and the yellow oak fall- ing silently in the Sierra forests, as one patiently wrought out with lens, measure, pencil and camera the great history of the Sequoia above named, year by year, century by century; centuries of peace, years of tragedy, and again centuries of stimulated growth. It was as if the whole landscape of life, from the dim prehistoric forests until now, "had been suddenly lit up by a burst of sunlight." I had never heard from any other lips any dis- course like this, nor from this fallen seer and patriarch could I hear the like again. During the past ten years hardly a season has passed but I have camped among the Sequoias. I am glad to say I have visited nearly all the groves, but I regret to say that a considerable proportion of them is in private hands; some have been leveled already and the mills are busy in not less than four others, notwithstanding there is little profit in the lumbering. These groves of Sequoias form a question apart from the ordinary questions of forestry. In the heart of the Sierra forests they are, it is true, an im- portant part of the protective forest cover of the headwaters of California rivers; but I believe you will now join me in the assertion that they have an interest for the citizens of California, for the cultivated traveler and the 42 DUDLEY MEMORIAL VOLUME scientific man, far beyond that of the other trees of the forest. The United States should own and properly protect every one of them. Senator Hoar, of Massachusetts, once said if the Calaveras Groves were in Massachusetts, she would herself buy them and not ask the National government to pur- chase them. There are some things, however, that are the natural heritage of a nation, and the Sequoia gigantea is one of them. We would rather see the Yellowstone National Park and the Grand Canyon of the Colorado under the protection of the United States than that of any state, not ex- cepting California or Massachusetts. If the Sequoias are among the most remarkable objects on the globe, if they are the best calculated, as we can show, of any living organism to throw light on certain problems of scientific inquiry, then a nation should own them and their preservation should be a matter of national pride. We make every effort to preserve the manuscript of our great Anglo- Saxon and American charters; nevertheless the ink fades, the parchment crumbles and they disappear, except from the lives of just men. We house the archives of our wars in buildings of great cost, maintained with great care, yet all these are on paper that is more perishable than the parchment of our charters. In these great trees, however, we have, deep in their annual rings, records which extend far beyond the beginnings of Anglo-Saxon peoples, beyond even the earliest struggles for liberty and democracy among the Greeks, the first of the Indo-Europeans to crystallize into national life through the pressure of this struggle. The records are those of forest conilagrations, of the vicissitudes of seasons, of periods of drouth and periods of abundant and favoring rains, and we might find next to the charcoal of some trunk scar, centuries old, the stone inplements belonging to the ancient aboriginal inhabitants of Western America. Practically none of these records have yet been studied. Let the nation purchase these trees of the Cala- veras groves — among the largest of all those still remaining alive — let it take them as a right and a duty, not parleying with the cupidity of an owner who has done nothing to increase their value; let it gradually gather under its protection all the groves of Sequoia, now in alien hands, and care for them all intelligently. When the oldest of trees succumb and die, as from past injuries they must do in time, then let them be felled; and in- stead of being sold or burned with criminal indifference to their real value, as at the present day, may their records be read and recorded by skilled hands and interpreted by the best intelligence ; and finally may their massive timbers, of wonderful fineness, uniformity, luster, color and beauty, be used only in the interior of a nation's buildings, in places which shall the longest endure. THE MORPHOLOGY AND SYSTEMATIC POSITION OF CALYCULARIA RADICULOSA (Steph.) By Douglas Houghton Campbell^ Professor of Botany THE classification of the so-called anacrogynous Jungermanniales, an important group of liverworts, is at present in a very unsatisfactory condition, and much remains to be done before the true relationships of the members of this group can be satisfactorily settled. A recent attempt has been made toward a better classification of the liverworts by Cavers^ and this is a distinct advance upon the classifications which have heretofore been accepted. There are, however, a number of forms whose relationships are still by no means clear, and among these is a rare liverwort from Java originally described as Calycularia radiculosa. Schillner^ speaks of the plant as an extremely rare one, but during a stay in Tjibodas where the plant had been collected before, the writer succeeded in finding it a number of times. The plants did not grow in large masses but were associated with various other liverworts growing on the trunks of trees. The general aspect of the plant (see Figs. 1 and 2) is very much like that of the creeping forms of the genus Blyttia. Pallamcinia (Blyttia) Levieri, a common species Fig. 1. Three male plants of Calycularia radiculosa. Steph. x 3. $ , antheridial recep- tacle. iThe inter-relationships of the Bryophyta. New Fhytologist Rejprint, No. 4, 1911. 2 Die Hepaticae der Flora Von Buitenzorg, 1900. .44 DUDLEY MEMORIAL VOLUME of the same region, much resembles the plant in question, but the latter is readily distinguishable on account of the nimierous dark reddish purple rhizoids. Fig. 2. A, female plant 1x3; the thallus has begun active growth again and developed a second archegonial receptacle, 9^, in the new portion. B, part of one of the involucral scales, slightly magnified. C, female plant bearing a young sporophyte, sp. D, plant with mature sporophyte. E, open capsule with the valves entirely separated at the apex. In, involucre; per, perianth. The material collected by the writer was sufficient to make possible a pretty complete study of the structure and development of the plant, except the earlier stages of the sporophyte which were wanting in the specimens collected. CALYCULARIA RADICULOSA ■ ■ CAMPBELL 45 The genus Calycularia as generally understood comprises four species of thallose liverworts of rather unusual distribution. One of these, C. laxa, occurs in arctic Siberia, two, C. crispula and C. Birmensis, are found in India and Burmah, while the other, C. radiculosa, occurs in Java. The question has been raised whether the latter species really should be united with the others. SchifEner^ after an examination of the plant, concluded that it should be placed in the genus Morkia, a genus sometimes regarded as a section simply of the larger genus Pallavicinia. The material upon which the present account is based was collected by the writer in Java in 1906 while staying at Tjibodas, one of the stations where the plant had originally been collected. GENERAL MORPHOLOGY The plants are dioecious, the male plants (Fig. 1) being decidedly smaller than the females (Fig. 2). Antheridia may be developed while the plants are not more than 5 mm. in length, but the male plants may reach a length of 12-15 mm. The female plants are two or three times as long as the males and may reach a length of about 30 mm., with a breadth of about 14 mm. Tl. A Fig. 3. A, cross-section of the thallus, showing the thickened mid-rib and rhizoids, r, x about 30. B, cells from the ventral side of the thallus, showing the mycorrhiza, m, extending from the rhizoid, r, into the inner cells of the thallus, X about 400. C, an inner cell of the thallus invaded by the mycorrhiza ; the nucleus of the cell is still intact. V, oogonium-like enlargement of a mycorrhizal filament. 3 Osterreichische Botanische Zeitschrift, Feb., 1901. 46 DUDLEY MEMORIAL VOLUME They are usually not branched but may be forked once (Fig. 1, C). There is a deep sinus in front within which lies the growing point of the thallus. A thick midrib is developed strongly, projecting on the lower side where its ventral surface is covered with numerous deep purple-red rhizoids. The margin of the thallus is more or less strongly undulate and folded, but these undulations are hardly distinct enough to be called leaves. The whole aspect of the plant is very much like certain species of Pallavicinia, and also suggests the Japanese genus Makinoa.* A section of the thallus (Fig. 3, A) shows that the midrib comprises about a dozen cells in thickness, but there is no trace of the conducting strands of tissue which are a constant character in Pallavicinia. In Eupallavicinia (Blyttia) there is a single very con- spicuous axial strand, while in Morkia, according to Cavers, there are de- veloped two strands which are however much less strongly developed than in Bl)rttia. In the character of the midrib, therefore, Morkia seems to be somewhat intermediate in character between Ccdycularia radiculosa and Blyttia. The wings of the thallus are composed of a single layer of cells in the marginal region, but toward the midrib the wings are composed of two or Fig. 4. A, vertical section of the thallus apex, in which there are dorsal and ventral segments cut off from the apical cell. B, C, two consecutive sections from a thallus apex, in which a single basal segment is cut off. D, E, two nearly horizontal sections, showing the appearance of the apical cell, x, when seen from above, x 225, h, ventral glandular hairs. * Miyake, K. ; Makinoa, A New Genus of Hepaticae. Bot. Mag., Vol. 13, 1899. CALYCULARIA RADICULOSA ■ ■ CAMPBELL 47 sometimes even of three layers of cells. In this respect Calycularia radiculosa perhaps more nearly resembles Makinoa than it does Pallavicinia. A characteristic feature of the thallus is the presence in the older por- tions of an endophytic fungus or mycorrhiza, very much like that found in the subterranean prothallia of various pteridophytes. A similar mycor- rhiza, however, has been found also by the writer in various green fern pro- thallia, and it is also known to occur in various other liverworts. The fungus both in its structure and manner of growth resembles more closely the mycorrhiza described by the writer® in Ophioglossaceae. As in the Ophioglossaceae there were occasionally found oogoniimi-like enlarge- ments (Fig. 3, D) which may have been perhaps special reproductive bodies, but this could not be positively demonstrated. The genus Calycularia is Fig. 5. A, upper surface of a male plant, x, apex of the thallus. A^, under surface, showing the antheridia, $ , surrounded by the laciniate scales, sc. x 15. B, median section of the thallus, showing the apex, x, and the antheridia, ^ . x 40. C, a horizontal section of the antheridial receptacle, sc, scales. D, scales showing the laciniate margins. 5 Campbell, D. H. The Eusporangiatse. Pub. 140. Carnegie Institution of Washington. 1911. 48 DUDLEY MEMORIAL VOLUME described as having upon the ventral surface leaf -like scales or amphigastria, but Schiffner found that these were not present in Calycularia radiculosa, and the writer's investigations coniirm this. These leaf-like scales are re- placed by multi-cellular hairs (Fig. 4, Bh), such as are common on many other' thallose Jungermanniales. The terminal cell is enlarged and probably secretes mucilage for the protection of the thallus apex. The latter is turned strongly upward (Figs. 5, Bj 8, A) and it is almost impossible to make satisfactory sections parallel with the surface of the thallus. Figures 4, D and 5, B show sections whdch are approximately parallel; but as these are somewhat oblique, the apical cell appears some- what shorter than it really is. In this view it appears somewhat oblong in outline, and it is evident that segments are cut off both from the lateral and from the basal portions. In vertical horizontal sections the apical cell shows certain variations, resembling in this respect the genus Pellia. While Pellia epiphylla has an apical cell with a single basal segment extending the whole length of the thallus, in P- calycina a vertical section shows two sets of seg- ments, dorsal and ventral, such as occur in certain species of Pallavicinia, as ■ well as in the Marchantiales and in Anthocros. In Calycularia radiculosa both of these tjrpes were found. The type found in Pellia epiphylla (See Fig. 4, B, C) were common in the smaller plants, but it was not at all clear whether there really is any definite relation between the thickness of the thallus and the form of the apical cell. The second type is shown in Figure 4, A. Where branching takes place it seems to be a true dichotomy, but whether one of the branches retains the original apical cell or whether two new apical cells are developed, was not investigated. THE MALE PLANT. The male plants (Fig. 1) are usually quite short, often being scarcely longer than broad, and as we have already stated, antheridia are sometimes found upon plants which are not more than 5 mm. in length. The antheridia are in small groups, seldom more than ten together, and are much less nu- merous than is the case in either Morkia or Blyttia. In the restriction of the antheridia to such a limited region the plants suggest Makinoa, but the whole antheridial group is not subtended by a common envelope as in Makinoa, though the antheridia occupy a more or less well-marked depression or cavity upon the dorsal surface of the midrib. Each antheridium is subtended by a much laciniated scale. The scales are often more or less confluent, so that imperfect chambers are formed (Fig. 5, C) about each antheridiiui).' As a rule, only one receptacle occurs upon the plant, but in a few of the larger CALYCULARIA RADICULOSA ■ ■ CAMPBELL 49 ones, where old groups of antheridia were present, a second younger group was occasionally found near the apex ; and in the rare instances where the thallus forks, each branch may bear an antheridial receptacle. (Fig. 1, C.) The antheridia (Figs. 5 and 6) are short-stalked nearly globular bodies and closely resemble those of Pallavicinia, and as usual they are formed in acropetal succession alternately right and left of the apex of the thallus. The earliest stages were not found, so that it is impossible to say whether the early divisions correspond to those observed in other genera, but as the young antheridia resemble so closely those of Pallavicinia, it is to be Fig. 6. Development of the Antheridium. A, median section of young antheridium, x about 225. B, cross-section of the antheridium of about the same age. C, an older antheridium. D, E, two nearly ripe antheridia, x about 90. 50 DUDLEY MEMORIAL VOLUME expected that the early stages would conform to the usual type found in the Jungermanniales. In his description of the genus Calycularia given in Engler and Prantl's Natiirliche Pflanzenfamilien, Schiffner states that the antheridium has a single celled stalk, but in his later description of C. radiculosa, given in his work on the Liverworts of Buitenzorg he says that he did not see the male plants, so that this description would not apply to that species, and as we shall see, the stalk is multicellular, very much like that of Pallavicinia. Figure 6, A shows a longitudinal section of the youngest perfect antheridium that was found. The short stalk shows in sections two rows of cells and the upper portion shows a mass of young spermatogenous cells surrounded by a single layer of sterile cellg. Figure 6 B shows a cross-section of an antheridium of about the same age. Sometimes the stem, of the antheridium is more slender and may have a single basal cell (Fig. 6 E) which often becomes very much elongated. Before the final division of the spermatogenous cells they are polygonal in outline, with dense contents usually more or less contracted, but how far this is normal and how far it is due to artificial shrinkage could not be determined. The cell walls are clearly defined. The nucleus is conspicu- ous and stains strongly. The contents appear somewhat finely granular, the Fig.7. Spermatogenesis. All figures magnified about 750. A, B, spermatogenic cells, just before the final mitosis ; in B can be seen two small bodies, b, which are probably the young blepharoplasts. C, D, stages in the development of the sper- matozoid. E, I, M, show the pair of spermatocytes seen from the side, the others are mostly single spermatocytes seen from above; b, blepharoplast ; c, cilia; n, nucleus of the young spermatozoid. Figs. L, M, show the mature spermatozoids. CALYCULARIA RADICULOSA CAMPBELL 51 nuclear reticulum not being clearly visible, and there are a number of bodies, one of which is probably the nucleus, although it is possible that there may be more than one nucleus (Figs. 7 A, B). In a number of cases shortly before the division to form the spermatocytes, there could be seen two minute granules (Figs. 7 B, b), sometimes surrounded by a colorless area and perhaps representing the young blepharoplasts, but the small size of these, and the presence of other granular bodies in the cytoplasm makes one hesitate to assert positively that these really were blepharoplasts, especially as no division stages were found, and the relation of these bodies to the nuclear spindle could not be determined. The further development of the spermatozoids, which are unusually large in this species and therefore favorable for study, is on the whole much like that of Pellia. After the final nuclear division a definite cell wall is formed between the spermatozoids, the division not running diagonally, as described by Ikeno for Marchantia, but dividing the cell into two approxi- mately hemispherical ones. (Fig. 7 E, I.) In the youngest stages met with ( Figs. 7 C, D ) the nucleus, which now appears somewhat coarsely granular, was still unchanged in form. The cyto- plasm was often contracted but not always so, and in the cytoplasm, some- times at the periphery, but quite as often near the nucleus, could be seen the blepharoplast (Fig. 7, C, D, b), which was already extended into a delicate band. No cases were found where the blepharoplast showed its original round form. A careful examination of the nucleus at this time shows that it is decidedly flattened in the plane of the division wall between the two spermatocytes, so that it appears oval when seen from the side but circular when viewed from above. No signs of any body equivalent to the so-called "Nebenkorper," or the "accessory body" described by Wilson* for Pellia could be seen, and such a body is probably quite wanting in Calycularia radiculosa. In a slightly later sta,ge, however (Figs. E, F, G), there could often be seen what appeared like cytoplasm extending beyond the nucleus and connecting with the ble- pharoplast. Strassburger refers to such a structure in his somewhat brief account of Pellia. The nucleus now begins to elongate and to increase notably in size, hav- ing the form of an almost homogenous crescent-shaped body when seen from the side. The forward end of the crescent is somewhat more prominent than the posterior end and extends into the cytoplasmic prominence, con- necting with the blepharoplast, the exact limits of which are very difificult 6 Wilson, M. Spermatogenesis in the Bryophyta. Ann. Bot., 25, 1911. 52 DUDLEY MEMORIAL VOLUME to determine. The cytoplasm surrounding the nucleus becomes less and less evident as the nucleus increases in size and it can no longer be clearly recognized in the later stages of development, although there probably per- sists a thin envelope of cytoplasm surrounding the posterior coils of the spermatozoid. The blepharoplast at this time forms a short hooped prominence at the forward part of the spermatozoid and merging insensibly into the delicate cytoplasmic prominence which extends beyond the nucleus. The latter con- tinues to elongate and become curved over in the plane of the division wall, so that the older spermatozoid has the form of a flat coil. (Fig. 7, M.) When fully developed the sperm is a slender thread composed of two complete coils and part of a third. In these later stages the double stain of safranin and gentian violet failed to clearly differentiate the different parts, the spermatozoid appearing almost uniformly stained. The two spermato- zoids of a pair are very closely approximated (Fig. 7, M) and present a very characteristic appearance. The cilia could be made out in a number of the older stages (Fig. 7, H), but their exact origin and position could not be determined as accurately as might have been wished. It is probable, however, that as in some other cases which have been investigated, they begin to double up at an early stage and arise somewhat back of the apex. Woodburn' in a recent paper states that in Porella the anterior end of the spermatozoid shows a slight enlargement, which he interprets as the blepharoplast. The cilia in this case arise a short distance back of this enlarged part of the blepharoplast. No trace of it was observed in Calycularia. ARCHEGONIUM. The female plants, as we have already seen, are decidedly larger than the males, and are usually 20-30 mm. in length, with a breadth of from 10-12 rrnn. Like the male plants, they are usually unbranched, but it is not uncommon to find them forked once. The position of the archegonia is much like that of the antheridia, these being grouped on a sort of recep- tacle. (Fig. 2, A.) As a rule, only one archegonial receptacle is formed, but sometimes the thallus will resume its growth and a second one may be formed near the apex. The archegonium appears to agree in all respects with that of other Jungermanniales that have been investigated. (Figs. 8 and 9.) After a short stalk has been formed, the archegonium mother cell divides by the usual three intersecting vertical walls into an axial cell and ' Woodburn, W. L. Spermatogenesis in Certain Hepaticse. Ann. Bet., 25, 1911. CALYCULARIA RADICULOSA — CAMPBELL 53 three peripheral ones. From the axial cell is cut off the cover cell (Fig. 8, B and C) and then follows a series of transverse walls separating the lower part or venter from the upper region or the neck. From the lower of the two primary axial cells the egg and ventral canal cells arise, and from the Fig. 8. A, apex of female plant, showing the position of the archegonia, x about 40. B-E, young archegonia in median section, x about 225. upper ones the series of neck canal cells and the outer cells of the neck. The number of neck canal cells is variable, b^t to judge from the few that were examined, there are first formed four of these neck cells, some or all of which divide again, so that there may be as many as eight. The division, however, is very often not complete but confined to the nucleus (Fig. 9, C ■ and D ) . As usual in the Jungermanniales, the neck of the archegonium has but five peripheral rows of cells (Fig. 9, F). 54 DUDLEY MEMORIAL VOLUME In the ventral region peripheral walls occur in the outer cells, so that at maturity the venter is more or less completely two layered (Fig. 9, D and E). Figure 9, G, shows an abnormal archegonium from a receptacle in which one of the archegonia had been fertilized. In this archegonium there were four axial cells, all of which were a good deal alike and all except one much enlarged, so that they resembled the egg of the normal archegoniimi more than they did the neck canal cells. 1 A _ C / ^ ^ G Fig. 9. A, B, two sections of a young archegonium with five neck canal cells; the ventral canal cell is not yet formed, x about 22S. C, a somewhat older stage, showing the egg, o, and the ventral canal cell, v. D, lower part of a nearly ripe archegonium. E, full grown archegonium which has failed to be fertilized, x about 90. F, cross-section of the neck of archegonium. G, an abnormal archegonium with unusually large axial cells. After fertilization there arises about the group of archegonia a tubular envelope, the perianth, which finally forms a very conspicuous vase-shaped sheath around the sporogonium, inside the involucre, which, like that of the antheridial receptacle, is made up of very much laciniated scales. (Fig. 1, C. D, per.) THE SPOROPHYTE. Only a few very young embryos were found and these were not well fixed, so that it was impossible to make a satisfactory study of the develop- ment of the embryo. The youngest sporogonia of which successful prepara- CALYCULARIA RADICULOSA CAMPBELL 55 tions were made were already far advanced and were differentiated into the various parts. Even before the first division takes place in the young embryo, the venter of the archegoniimi becomes much enlarged, and a calyptra is devel- oped, enclosing the sporogonium until it is far advanced. This attains a thickness of 5 or 6 layers of cells at the base, but is much thinner toward the apex. As the sporogonixmi develops, the upper portion or capsule becomes oval in form, and below consists of a thick seta, which terminates in a large heart-shaped foot very much like that which Cavers describes for Morkia Flowtoudana. (Cavers Loc. cit. Fig. 37). A similar foot has been observed in various other Jungermanniales. Fig. 10. A, median section of a sporogonium at the time of the mitosis of the spore mother cells, x 15. B, apical region of a younger sporogonium, sp. spore mother cells, el. young elator, x 225. C, part of the lateral wall of the sporogonium. The capsule (Fig. 10) has a relatively thick wall which is better de- veloped at the apex than at the sides, this difference becoming still more marked in the later stages. The inner tissue now shows a separation into the roundish spore mother cells and the elongated young elaters. Long before the division of the spore mother cells begins, they show the first indi- cations of the lobing which later becomes so conspicuous. At this stage the walls of both the spore mother cells and the elaters are very delicate, but can be readily demonstrated by suitable stains, e. g. Bismarck brown. 56 DUDLEY MEMORIAL VOLUME The seta at this stage has about the same length as the capsule, and in longitudinal section (Fig. 12, H) shows the cells to be arranged in pretty regular rows. Probably the great elongation of the seta at the time the spores are shed is due to simple elongation of the cells without any cell divi- sions, as has been shown to be the case in other liverworts. The large heart- shaped foot (f) is composed of somewhat irregular cells showing no defi- nite arrangement. Fig. 11. Spore division. A, spore mother cell, shewing the quadripolar spindle, X 750. B, three spore mother cells of about the same age, showing the different arrangement of the lobes of the cell, x about 350. C, young elater. D, spore mother cell, sho>ving the chromosomes. E, a somewhat earlier stage. F-I, successive stages of mitosis with quadripolar spindle. In G and H the chromosomes only are shown. /, mother cell with the lobes in pairs; there are two nuclear spindles at right angles to each other. K, mother cell, showing a bi-polar spindle at the first mitosis. L, first mitosis, showing two nuclei separated by a distinct cell-plate, x about 400. M, mother cell just before the final separation of the spores, x 400. CALYCULARIA RADICULOSA CAMPBELL 57 THE SPORE DIVISION. (Figure 11.) One sporogonium showed the spore mother cells in process of division, all stages occurring in the same sporogonium. The preparation was some- what overstained with haematoxylin, but nevertheless showed pretty well the details of division, which exhibited a considerable amount of variation. The spore mother cells before the first nuclear division are deeply four- lobed, the lobes being usually arranged tetrahedrally, but occasionally placed in pairs at right angles to each other (Fig. 11, J). The nucleus is large, but in this over-stained material the structure was not usually very clear, the nucleus appearing almost homogenous. A few specimens, however, (Fig. 11, E) showed a more or less granular structure, but the reticulum was not clearly evident nor could the nucleolus be seen. A striking feature was noticed in most of the cells, viz. : the extension from the nucleus into each of the four lobes of the cell of a body which was apparently the same as the "quadripolar" spindle described by Farmer for Pallavicinia decipiens^. Often the center of each lobe was occupied by a roundish body which perhaps marked the position of a centrosome, but it cannot be stated positively that centrosomes are present. However, as cen- trosomes occur in Pellia, which in some other respects suggest a relationship with Calycularia, it is possible that centrosomes may have been present in this case also. In the later stages of nuclear division some difiFerences were found to occur. Usually the process seems to agree very closely with that described by Farmer for Pallavicinia decipiens. The nuclear membrane disappears and the separate chromosomes, thick oval bodies, can be made out. (Fig. 11, F.) There are eight of these in Calycularia radiculosa instead of the four found in Pallavicinia decipiens; but it was found that in Pallavicinia radiculosa there were also eight, as there are in Pellia. The eight chromosomes next divide longitudinally (Fig 11, G), and the resulting sixteen chromosomes separate into two groups ( H ) . Usually the chromosomes do not arrange themselves into a new reticulum, but imme- diately undergo a second division, so that there are two groups of 16 chromo- somes (Fig. 11, I), each of which separates into two. secondary groups of eight chromosomes which finally assume the form of resting nuclei. One 8 Farmer, J. B. Studies in Hepaticae Pallavicinia decipiens Mitten. Ann. of Botany, 8, 35-52, 1894. 58 DUDLEY MEMORIAL VOLUME of these nuclei moves to each lobe of the mother cell, but generally remains near the center of the cell, so that the four resulting nuclei are quite close together (Fig. 11, M). Not all of the spore mother cells, however, behave in this fashion, but sometimes after the first division of the chromosomes a conspicuous bi- polar spindle of the usual form was observed. (Fig. 11, K). Later two resting nuclei were seen with a cell-plate between them. (Fig. 11, L). These secondary nuclei then divided again, each developing another bi-polar spindle, these secondary spindles sometimes lying at right angles to each other. (Fig. 11, J). After the nuclei have assumed the resting condition cell walls are formed, simultaneosuly extending inward from the indentations between the lobes and completely dividing the mother cell into its four component parts, the young spores. The ripe spores (Fig. 12, F, G) possess a thick membrane, which in sections shows two well-marked parts, an inner uniform layer and a thick outer one provided with rounded knobs, which give it a very characteristic appearance. The color of the spore is dark purple-brown, like the thicken- ings upon the cells of the capsule wall. It is probable that immediately adjacent to the spore cavity is a thin membrane (intine) of cellulose, but this was not specially investigated and did not show clearly in the sections that were examined. The nucleus of the spore is rather small but fairly conspicuous. The elaters (Fig. 12, C, D and E) show a good deal of variation. They are sometimes very much attenuated, with the spiral bands almost obliterated at the ends, suggesting the elaters of Makinoa, where the spirals are present only in the mid-regSon of the elaters. More commonly, however, they taper more gradually and the double spiral extends to the end. Con- siderable difference in size may be noted (Fig. 12, E). While no basal elaterophore is present, occasionally some of the elaters at the base of the capsule seem to be attached at one end and suggest a rudiment of such an elaterophore, as is said to occur in the other species of Calycularia. The surface markings of the spore in Calycularia radiculosa are strik- ingly different from those of Pallavicinia whether of the section Morkia or Blyttia. In Pallavicinia (See Fig. 12, K, L) the surface markings have the form of a network of delicate ridges, such as also occur in Fossombronia. This marked difference in the character of the spores, together with certain other differences, might be considered to be an objection to uniting Caly- cularia radiculosa with the genus Morkia. The structure of the capsule wall of Calycularia radiculosa, according CALYCULARIA RADICULOSA CAMPBELL 59 Fig. 12. A, apical region of the sporogonium of Calycularia radiculosa, showing the thickenings on the cell walls, x 90. B, lateral wall of the same sporogonium. C, ripe spores and elaters, x about 200. D, very much attenuated elaters, x 350. E, typical elaters, x 350. F, ripe spore, x 350. G, section of spore. H, lower part of seta and foot, f, x 90. /, apex of sporogonium of Pallavicinia (Blyttia) radi- culosa, X 90. /, lateral wall of the same, x 90. K, section of ripe spore, x 350. L, markings on the surface of spore, x 750. 60 DUDLEY MEMORIAL VOLUME to Schiffner, is very different from that of the other species of Calycularia. He examined C. crispula and found that the capsule is much smaller than in C. radiculosa, and was perfectly round instead of being oval. The wall showed quite a different structure, being composed of two layers of cells with somewhat different markings from those found in Calycularia radiculosa. In the ripe sporogonium in the latter species (See Fig. 12, A, B) the wall is much thicker at the apex, where there are five or six layers of cells which form a sort of apical cap, while at the sides there are usually about four layers of which the outer one is composed of much larger cells, the inner layers being made up of much compressed thin walled cells. The radial walls of the outermost layer of cells are marked by conspicuous thick- ened bands, which are sometimes more or less confluent, giving the cells much the appearance of reticulate tracheary tissue. The inner cells have slight thickenings, but these are very irregularly disposed, and are almost wanting upon the inner cells of the lateral walls of the sporogonium. At maturity there may be recognized four valves of equal size, but usu- ally these do not separate completely, but remain together in pairs, the cap- sule opening by two slits. (Fig. 1, D). There may be seen between these two slits, however, a delicate line marking the junction between the two coherent valves. Schiffner states that the valves never separate at the apex but are held together by the apical cap of cells. While this is no doubt often the case (See Fig. 2, D), it may happen that the two pairs of valves separate completely. (Fig. E). In its dehiscence, therefore, Calycularia radiculosa is more like Blyttia than it is like Morkia. In the other species of Calycularia Schiffner states that the capsule at maturity breaks into several (5-6) irregular parts which may break up still further, thus resembling Fossombronia. The seta finally becomes very long ( Fig. 2, D ) and its base is surrounded by the very conspicuous vase-shaped perianth, whose opening is deeply lobed and fringed. Material of Morkia was not available for comparison of the structures of the sporogonium with that of Calycularia radiculosa, but sec- tions of the sporogonium of Pallavacinia (Blyttia) radiculosa were made. In this species the capsule is extremely long, cylindrical and very little thicker than the seta. The foot is pointed and not clearly delimited from the seta. The apex of the capsule is pointed and much more conspicuous than in Calycularia (Fig. 12, I). The capsule wall also differs in the character of the cells. There are about three layers of cells instead of four and the outer cells have the walls uniformly thickened instead of showing the thick- ened bands so conspicuous in Calycularia radiculosa. (Fig. 12, J.) CALYCULARIA RADICULOSA CAMPBELL 61 The spores are very different in their markings, as we have already noted, and this seems to be true also for Morkia. THE AFFINITIES OF CALYCULARIA RADICULOSA. Schiffner, from his study of Calycularia radiculosa concluded that it should be removed from its present association with C. crispula, C. Birmensis and C. laxa and united with Morkia. While there seems to be reason to remove the species from the genus Calycularia, it may be questioned whether its association with Morkia is justified. While Schiffner states that Morkia is without conducting tissue in the mid-rib, Cavers has shown that in M. Flo- towiana there are two strands of conducting tissue, but in Calycularia radicu- losa these are entirely wanting. Moreover, the structure of the sporogonium, i. e., the character of the thickenings of the wall and the markings of the spores are quite different, and more like Makinoa, or some of the forms usually referred to the Codoniaceae. It would probably be better to con- sider this plant as the type of a new genus intermediate in some respects between Morkia and some of the less specialized forms like Makinoa or Pellia. It is hardly likely that the line between the Codoniaceae and Lep- tothceae (or Blyttiaceae, as Cavers has called them) is very well defined, and it is probable that further study of the thallose Jungermanniales will result in decided changes in the accepted arrangement of the genera. STUDIES OF IRRITABILITY IN PLANTS. By George James Peirce^ Professor of Botany and Plant Physiology. III. the formative influence of light. Introduction. IN 1906^ I published a paper, under the above title, recording the results of a series of experiments on the influence of the direction of illumina- tion upon the shape of certain plants. The most striking result reported was that Anthoceros plants grown from the spore on a disc revolving in a horizontal plane, and therefore receiving fairly equal amounts of light on all sides successively, showed no trace of the usual dorsi-ventral form and structure of the thallus but were radial in structure, cylindrical or conical in form. Anthoceros fusiformis, Aust. and A. Pearsoni, M. A. Howe, both native here and growing within a short distance of this laboratory, gave the same results; but the spores of the Marchantiaceous liverwort Fimhriaria ( Asterella) Californica and of the fern Gymno gramme triangularis did not, under the same conditions, give rise to plants round in section. To this extent their dorsi-ventrality failed to show itself — rhizoids grew equally in all directions from their thalli or prothalli, respectively — but the plants were thin plates, though curiously crumpled, as the figures showed. I did not understand this difference in result and have tried in various ways to ascer- tain the reason for it. I have not yet reached a satisfying explanation, but some of the results of these succeeding experiments are interesting enough to record now. THE APPARATUS : A MULTIPLE CLINOSTAT. The apparatus used in the experiments of Czapek^, which suggested mine, consisted essentially of the expensive form of clinostat, the only one generally known and used in botanical laboratories. My experiments were carried on with cheap clocks, modified as described by Ganong^ Such ap- paratus is, however, unreliable. Indeed, cheap apparatus may be the most expensive. Although cheap apparatus may perhaps be well enough for an 1 Annals of Botany, XX, 449-465, 1906. 2 Czapek, F. Weitere Beitrage zur Kenntniss der geotropischen Reizbewegungen. Jahrb. f. w. Bot., XXII, 261, 1898. 3 Ganong, W. F. A laboratory course in Plant Physiology, pp. 120-1, New York, 1901. STUDIES OF IRRITABILITY OF PLANTS PEIRCE 63 experiment lasting only a few minutes or at the most an hour or two, it ought not to be trusted longer. An experiment which lasts a week or even months increases in value as it lasts, if it had any value at the start; and the failure of the apparatus at the end of six months entails a loss much greater than more serious mechanical difficulty within a day or two of the beginning. The cost, unreliability and the wearisomeness of winding a suit- able number of separate instruments drove me to consider other apparatus. And by devising new apparatus I tried to ascertain the dominating reason for the persistent dorsi-ventrality of the thalli of Fimbriaria and of the pro- thalli of Gymno gramme. To Professor W. F. Durand, head of the Department of Mechanical Engineering, is due all the credit for structural details and for supervising the construction and the successive modifications of the apparatus in the Mechanician's Shop of this University. And I take this opportunity to ex- press my most grateful appreciation of his skill in divining what I wanted and his untiring help and unflagging patience in securing it. As a detailed description of the apparatus would be more appropriate elsewhere, I may here content myself with a statement of its essential features. The apparatus may be called a multiple clinostat. As my experiments involved the turning of cultures in a horizontal plane upon a vertical axis, the apparatus began with a set of twenty-five turn-tables on five shelves built into the embrasures of each of three windows, the turn-tables in each window revolving at different speeds, but all the turn-tables in one window turning at the same rate. The actuating mechanism consisted of a clock-work driven by a heavy weight and controlled by a fan governor. This actuating mech- anism was connected by a series of belts and shafts with the batteries of turn- tables. Experience, however, has led to the gradual and final elimination of all belts. Chains and sprockets were first substituted for belts. Finally these were replaced by direct gears. This made possible the consolidation of the cultures into one window, there being five rows of ten turn-tables each on a set of shelves in the window nearest the clock-work and connected with it by a shaft with bevel gears. This shaft is horizontal and runs from the clock-work, bolted to a table, which is itself bolted to the floor, to a vertical shaft at one side of the window. At each shelf this vertical shaft carries a gear which engages a corresponding gear carried on the axle of the nearest turn-table. The margin of this turn-table, and of all the others in the row, is toothed, and the turn-tables are so set that the movement of one of them sets all the others into similar motion. All the turn-tables in a row move at the same rate, but the rate of each row is determined by the ratio of the gear on the vertical shaft to that of the first turn-table in the row. It is possible, 64 DUDLEY MEMORIAL VOLUME therefore, to revolve fifty or more cultures simultaneously, but at five differ- ent rates. The apparatus runs continuously, day and night, and requires to be wound only once in thirty hours. After experience and reflection, I concluded that the only constant and uniform force at my disposal for driving my clock-work was gravity. This is represented by discs of cast iron and of lead, amounting to a weight of two hundred and fifty (250) pounds — approximately 113.4 Kilos — which is hung from a pulley running on a wire cable, the end of which is fastened to a beam in the ceiling of the room. The clock is wound by pulling up the weight nearly to the ceiling, the wire winding upon a drum revolved by a crank. The material and methods of culture have undergone nearly as many changes since I began as the method of revolving the cultures. With the exception to be noted below, I still use small crystallizing dishes. These are about 7 cm. diameter and 3 cm. depth. I have had small square tiles of porous flower-pot clay especially made. These fit into the bottom of the dishes, and, as their upper surface is smooth, the least possible shadow is cast by one part upon another. These porous tile are first boiled in distilled water, to extract, as completely as may be, the soluble matter which they may contain. To this end I boil the tile in distilled water for hours, using three or four waters for this washing and leaching process. The tiles are then allowed to drain and dry. Meanwhile Knopp's Solution is made as follows : Solution A Solution B KNO3 2grs Ca (N03)2 8grs MgS04 2 " Aq. dist 3000 cc K2HPO4 2 " Aq. dist 1000 cc To 1 part of A 3 parts of B are added and the mixture boiled for fifteen minutes in a cotton-plugged flask. The resulting precipitate is either filtered off or allowed to settle in the bottom of the flask. At all events, the clear liquid is poured into the culture dishes to about three-quarters the depth of the porous tiles. The dishes are covered with the lids or bottoms of Petri dishes of suitable size. They are now steam sterilized for an hour or more and are allqwed to cool over night in the sterilizer. As it is necessary to prevent light from falling otherwise than from the side upon the plants to be Cultivated, the lids of the dishes are given, after cooling, a smooth continuous coat of dull black "Japalac," an inexpensive and quickly drying varnish easily applied. It may be wondered why I did not use Plaster of Paris poured into the crystallizing dishes and allowed to set in them as molds. These would have STUDIES OF IRRITABILITY OF PLANTS PEIRCE 65 furnished a fairly smooth substratum of convenient extent. Two considera- tions prevented, namely : the solubility of Plaster of Paris, and its color. The latter could have been modified by lamp-black or any other insoluble pigment, but the constant presence of an undue amount of calcium sulphate in the water which the plants were to absorb seemed to me unnecessary and possibly confusing. Flower-pot tile is certainly more nearly like soil in color and composition than is Plaster of Paris, more convenient to handle, and readily enough obtained in any desired size, if a reasonable number of tiles be ordered at any one time. The spores are sowed as uniformly as possible on the now damp tile, which is standing in sterilized Knopp's Solution in the dishes. The spores are sowed from stiff smooth writing paper by tapping the paper with a pencil or paper knife in such a way as to discharge a fairly, even shower of spores upon the tiles. The culture dishes are then marked and put in place on the turn-tables, and on the shelf beside them as controls, respectively. The speed of the turn-tables is a matter of considerable importance. The greater the speed, the greater the amount of power required. In no case, however, have I used a speed at which the centrifugal force, even at the edge of a culture, could have had any part in the result ; and at the center of revo- lution, which is also the point of most nearly equal illumination, there would be no centrifugal force. The speeds which I have so far used, in addition to those previously reported*, are the following: 10 turn-tables making four revolutions a minute. 20 " " two " " " 10 " " one I have also arranged to have a fifth row of ten turn-tables, so geared as to make a complete revolution in two minutes. I may add that, although I have so far used the turn-tables only on vertical axes, I have, nevertheless, had the shelves so attached to the frame of the shelving in the window that they may be set at any desired angle between the vertical and the horizontal. The positions and structures of the gears of the vertical distributing shafts and of the first turn-tables in each row must be and may be modified accord- ingly. Obviously, if the turn-tables are to be used in a position in which their axes would point obliquely downward, it would be necessary to use cups, into which the axes could be locked; but for my experiments so far, no modi- fication of the cups has been necessary. The cups carrying the axes of the turn-tables are cast steel with a steel ball of suitable size in the bottom of 4 Annals of Botany, XX, 1906. 66 DUDLEY MEMORIAL VOLUME each cup, and screwed to the shelves. So long as the diameter of the bore of the cups remains uniform and the wear of the axis corresponds, the motion of the turn-tables should be uniform. The turn-tables themselves are made of a light alloy, Alzine, which sometimes appears to be too brittle; but if not, teeth of great uniformity may be cut in the edges of the turn-tables. When the regular teeth of adjacent turn-tables are so set that they do not bind or allow too much play, with the inevitable contraction and expansion of the shelves and frame in the changing temperatures and humidities of a laboratory, the clock-work drives them with great regularity. Indeed, next to the very desirable feature of carrying many cultures at once on this multi- ple clinostat, the regularity of revolution is its most valuable feature. It may not be necessary to add that the multiple clinostat now in my laboratory, and thus briefly described, is the product of the experiments, fail- ures and successes, of the last six years. Each improvement has been the fruit of failure. Some of these failures have been very disheartening, for one does not like to lose or to vitiate the accumulated result of six or eight months of work by the clock-work or any set of turn-tables coming to a stand- still for an hour. EXPERIMENTAL WORK. Only one or two of the questions suggested by my previous work and left unanswered in that' paper will be considered in this. The plants which I have experimented upon have been the prothalli of Pteris aquilina and Gym- nogramme triangularis, grown from the spore on tile; plants of Porella Bo- landeri, a foliose liverwort which I brought into the laboratory from rocks and tree-trunks near by and cultivated on the tile in crystallizing dishes; plants of Fimbriaria (Asterella) Calif ornica, also grown from the spore; Anthoceros fusiformis, grown from the spore and used simply as a check, for the results on turn-table and shelf were the same as previously reported; and plants of white mustard and of wheat, raised in two-inch flower pots in good soil from the seed. The results are in the main similar, and I shall discuss them all together after separately describing the experiments on the different sorts of plants. Porella Bolanderi (Aust.) Pearson. On November 14, 1907, I collected plants of Porella Bolanderi growing on rocks and tree-trunks about a half mile from this University. The plants were dry and dormant. I sorted these, after moistening with sterilized water, and selecting clean branches about a centimeter long, placed these upon STUDIES OF IRRITABILITY OF PLANTS PEIRCE 67 Sterilized tiles in crystallizing dishes, containing sterilized Knopp's Solution of 0.35% concentration and covered with blackened lids. These dishes I set on the turn-tables and on the shelf beside the turn-tables. I took all possible pains to select branches clean and healthy-looking, but as no steriliza- tion of the material was possible, it was inevitable that a certain amount of moulding should take place. All of the cultures succumbed to infections sooner or later, but enough of them grew well to justify a record of the experi- ment, although I do not by any means regard it as concluded. The general characteristics of this plant are well known. Detailed de- scriptions may be found in Campbell's "Mosses and Ferns"^ and elsewhere. For our purpose it is sufficient to say that the plant is dorsi-ventral to the extent of having two sets of leaves : foliage leaves, which are green and closely arranged, forming the upper side of the plant, and the so-called amphigastria, leaves or scales not green, and overlapping along the under side of the stem. The plant grows more or less closely appressed to the sub-stratum, whether this is vertical, oblique, or horizontal ; that is, the plant grows at right angles to the direction from which most of the light comes. If, therefore, the plant be put on a horizontal surface and the light be made to fall more or less horizontally upon it, the plant or its leaves should so turn that its foliage leaves would stand mainly at right angles to the incident rays, and the amphi- gastria should be on the side away from the light. This happens with the plants growing on the shelf, receiving light always in one and the same direc- tion, just as it would happen in the case of a Porella plant growing from a horizontal to a more or less vertical sub-stratum out of doors. The case of the plants on the turn-tables, on the other hand, is quite different, for they have no darker side. The position of all the leaves and of the amphigastria on the older parts of these plants changes; they flare more from the stem. On the younger, as well as on the older parts, the amphigastria become less scale-like and grow more and more leaf-like. I have no doubt that this ex- periment, continued with greater freedom from infections than I secured in 1907, would yield results entirely similar to those of Nemec®, but I have not yet been able to repeat it and carry it through. I would suggest here only that experiment seems likely to confirm the opinion of morphologists that amphigastria are modified leaves, and to show that they develop as they do partly because they are on the shaded parts of these plants. = Campbell, D. H. The Structure and Development of Mosses and Ferns. 2d Edition, New York, 1905. 8 Nemec, B. Die Induktion der Dorsiventralitat bei einigen Moosen. Bull. int. de I'Acad. Sci. de Boheme, 1904, 1906. 68 DUDLEY MEMORIAL VOLUME Fimbriaria ( Asterilla) Calijornica, Hampe. Spores of this plant were collected at the beginning of the dry season, late in April or in May, according to the time of ripening, from plants grow- ing on a sandy-loam bank, not far from the laboratory. The spores were kept in envelopes and pasteboard boxes in a case in the laboratory and were, therefore, air-dry and well ventilated. As the humidity of the air runs low during the summer dormant period, both in the laboratory and out of doors, the spores were necessarily inactive for months. With greater humidity or — what would produce this — inferior ventilation, their respiratory activities' would be greater and there would be danger that their germinating and other powers might be impaired. For this reason I avoid keeping spores or seeds which are to be used for germination in tightly-closed jars or bottles. If it be necessary to protect them against mice, I use tins, the lids of which close them loosely enough to permit more or less circulation of air. In this way spore and seed deterioration is delayed and normal dormancy is main- tained. The spores were sowed, as above described, on sterilized tile in black- covered crystallizing dishes, on October 11, 1911. Of these, five were put on turn-tables making two complete revolutions per minute, a,nd three were set on the shelf beside them. The crystallizing dishes standing on the shelf were marked on the side away from the window, so that one might always know the original exposure and the more easily maintain it. As previously shown*, the direction of the plane of division in the ger- minating spores, and of growth in the germ-tube, is determined by the direc- tion from which the light falls upon the spores. On the clocks, therefore, the spores germinate in every direction, and the plantlets are erect from the start. Germination actually begins almost at once, no doubt, but the evi- dences of it are plainly visible within ten days after sowing the spores. In the shelf cultures the plantlets are prostrate, growing toward the light as single chains of chlorophyll-containing cells. In these latter cultures, as in nature, the light falls upon the plantlets mainly from one direction, and the plantlets react accordingly. After the plantlet has become a single chain of several cells, the end cell repeatedly divides in such planes as to change the plantlet to a conical shape. These little cones, with their apices pointing away from the light and obliquely downward, grow both in length and in diameter fairly symmetrical for a few weeks. After a time, however, they 1 Babcock, S. B. Metabolic water ; its production and role in vital phenomena. Research Bull. 22, Univ. Wis. Agric. Exp. Sta., March, 1912. 8 Peirce, G. J. Annals of Botany, XX, p. 45Jf, 1906. STUDIES OF IRRITABILITY OF PLANTS PEIRCE 69 cease to be symmetrical, the side of the cone away from the light growing out in a form more or less shelf-like. This is the beginning of the thallus of the more or less mature form. In cultures I have never carried the plants beyond this beginning — six months or thereabouts after sowing the spores — for although plants of the same species normally survive the summer out of doors^, they do not with- stand the mijich more complete drying in a culture dish, or, if an attempt is made to keep them moist over summer, they succumb to fungus enemies. I do not know what would happen if the cultures were kept continuously on the shelves and on the revolving turn-tables, for it has never been possible for me to stay in my laboratory throughout the long summer vacation, and I have so far been unable to arrange to have my clock-work regularly wound, i: e., daily throughout my absences. It is usually easier, in a laboratory as well as elsewhere, to provide apparatus than to secure assistants. Hence, at the end of the college year, in May, I am obliged to take my cultures from their places on shelf and turn-table and set them away in a dark cupboard. They remain there till September. During these months they have succumbed to mould or drought. Some day, however, I shall be able to carry them along continuously from the spore, to the production of spores again. Turning now once more to the plantlets subjected, on the turn-tables, to light from all directions successively, we find that they maintain the erect position which they assvime immediately on germination. They thicken at the ends away from the spores, and, since they are revolved in a horizontal plane and receive light mainly horizontally, they become vertical. I do not think the force of gravity, or the presence of water below them, or any other influence than light has much to do with the erect position of these plantlets. They become erect cones standing on their apices, and, uniformly on all sides, they develop rhizoids, which attach them to the tiles. The little plants are thus stayed and kept from toppling over. They keep pace in their growth with the plants receiving light from one side only, on the shelf, and after a time exceed them considerably in size. On account of the difficulties pre- viously enumerated, however, I have never been able to carry these plants through the summer or continue the experiment for more than seven months. Though the plants on the shelf are at first and for some weeks radial in structure, they sooner or later go over to the dorsi-ventral form under the influence of light falling upon them from one direction only. This form they maintain throughout all but the very early stages of their existence. The plants revolving on the turn-tables, on the other hand, remain radial in 8 Campbell, D. H. Resistance of drought by liverworts. Torreya, IV, 1904. 70 DUDLEY MEMORIAL VOLUME Structure, conical in form, for a much longer time, and in so doing resemble Anthoceros^". For the reasons given above I cannot say that they would always maintain this radial structure, though experience with these plants and with Anthoceros leads me to believe that dorsi-ventrality, in these two genera at least, is not alone inherited, but that it is a product of circumstances as well as of substance, the continuity of substance and the continuity of influence (direction of illumination) from generation to generation insuring the repetition of this quality in successive generations. It may be suggested that the dorsi-ventrality does not develop in the plants revolving on the turn-tables, not because it is not inherited, but because it is prevented from appearing because one of the conditions for its devel- oping is lacking. I do not care to contribute to a revival of the profitless discussion, wisely dropped, involving a Conceivable if unnecessary distinction between condition (Bedingung) and stimulus (Reiz), for, as will more plainly appear in the next section (pp. 70-74), the influence of the direction of illumination is active rather than passive. When one compares, at the end of an experiment which has lasted for months, the sizes of the plants on the turn-tables with those on the shelf, one realizes the greater size of those more uniformly illuminated, symmetry and size going together. The result of growing Fimbriaria Calif ornica from the spore under con- ditions of equal and of unequal illumination from all directions successively, is the same, so far as the experiments could be continued, as with the two species of Anthoceros previously reported upon, but the longer life-cycle of Fimbriaria makes it necessary to continue the experiment for a longer time than has so far been possible, in order to reach a definitive result and to justify a final conclusion. Fimbriaria grown from the spore does not fruit, at least in my cultures, within the time limits of one natural growing season — that is, between the first rains, say early in November, and the beginning of the dry season, in May. On the other hand, Anthoceros does, but its spores are not equally fertile in successive seasons, and since the wonderful crop of 1905 I have been unable to secure spores of such vigor that I cared to con- tinue experimenting upon the plantlets beyond confirming previous results. Nor is it necessary, as the results reported in the next section will show. Pteris aquilina and Gymnogramme triangularis. Spores of Pteris collected in southern California in June, and of Gym- nogramme collected near the laboratory in September, were sowed on October 11, 1911, under the conditions previously described. Six of the cultures of Pteris were set on turn-tables revolving four times a minute, two on the shelf loPeirce. Ann. Bot, 1906. STUDIES OF IRRITABILITY OF PLANTS PEIRCE 71 by them. A second sowing in other dishes was made on November 13, 1911. About the same number of cultures of Gymno gramme were started on Octo- ber 11 and November 13. The results were similar to those obtained with Pteris but less striking, because of the smaller size of the prothalli. Further- more, the Pteris material was so much freer from contaminating blue-green algae that the cultures were correspondingly more satisfactory. Since steriliz- ing the spores is impossible, it is not usually possible to make a pure culture of fern prothalli directly from the spore. The spores of both ferns germinated well, behaving during the earlier stages on the turn-tables and on the shelf as before described^^ In the ex- periments previously reported, the spores of Gymnogramme germinating on turn-tables developed into thin prothalli, crumpled or waved instead of flat, heart-shaped instead of conical or cylindrical, but with rhizoids, antheridia and archegonia in equal numbers on both sides, and with the prothalli stand- ing erect, but at all possible angles on the tiles. In the shelf cultures, on the other hand, the prothalli were normal in shape and almost linearly placed, in ranks surprisingly regular, at right angles to the incoming light from the window. These results I thought might be due to the slow revolution of my turn-tables and I hoped by using quicker ones I might obtain cylindrical or conical prothalli. I have not yet. Why, I do not know. It may be that the turn-tables, revolving four times a minute instead of once in fifteen as before, are still too slow, or it may be that continuous as well as uniform illumination is necessary. The results of five months' growth on turn-tables are shown in Figures 1 and 2. Figures 3 and 4 show shelf cultures. These two sets of photo- graphs of Pteris cultures sowed on October 11, 1911, were kindly taken for me by Mr. James McMurphy, Instructor in Botany in this University. The culture dishes were uncovered and placed on the horizontal stand of a verti- cally working camera, so focused as to give a picture double natural size.* The magnification was the same for all five photographs. This was made possible by the uniform thickness of the tiles and of the glass of the dishes The four figures are, therefore, perfectly comparable. Figure 1 shows, within the circular line which indicates the bottom of the crystallizing dish, and upon a square porous tile, the corners of which have been knocked off to fit the glass dish, a large number of fern prothalli of two very different sizes. These prothalli are erect or nearly so. Those nearest the center of the tile (the center of revolution) are most nearly erect and most plainly show the copious growth of rhizoids on both sides. The "Annals of Botany, XX, p. 454+, 1906. * Reduced to natural size in the figures. 72 DUDLEY MEMORIAL VOLUME larger prothalli bear archegonia, the smaller prothalli are antheridial. Toward the upper right hand comer is a young sporophyte, showing that conditions in the culture were sufficiently favorable to permit fertilization and subsequent development. Microscopic examination shows that the repro- ductive organs are borne on both sides of the prothallus, as uniformly as the rhizoids. The prothallus has the usual cushion, which bears the rhizoids, archegonia and antheridia. Near the lower edge of the figure is a large archegonial prothallus, irregular in outline but plainly dorsi-ventral in structure. This has rhizoids only on the side toward the middle of the tile. Comparing this prothallus with the two nearest the center of the culture, one sees that the more uniform illumination of the plants in the middle is accompanied by a more uniform growth of rhizoids. The distance between the prothallus at the edge and those in the middle of the tile is about 2 centimeters, but this slight difference in position is accompanied by enough difEerence in illumination to permit the plantlet near the edge to complete its usual dorsi-ventral development, while the plants at the center form rhizoids and reproductive organs equally on both sides. From this one may infer either that the fern prothallus is very sensitive to light, since slight differences in illumination cause such evident differences in behavior, or that it is only slightly sensitive, since exactly uniform illumina- tion is necessary to overcome the usual dorsi-ventrality in any degree. Whether the plant is sensitive or not sensitive can be proved only by experi- ment. One of these experiments would involve constant illumination, and this I hope to try shortly. In Figure 2 the number is larger, the distribution more regular, and the results striking. Figure 3 is that of a shelf culture, the direction of illumination of which is indicated by the arrow. The size, position, form and appendages of the largest archegonial prothalli are quite as usual and normal. Rhizoids and archegonia develop only on the side away from the light, the prothalli are erect in response to the nearly horizontal plane along which the light is re- ceived, but they are nearly flat and extend at right angles to the light and they are of the usual size. The much (and normally) smaller antheridial prothalli correspond in all these respects with the much more striking arche- gonial. I do not know what may be said to be the normal ratio between arche- gonial and antheridial prothalli in Pteris, and I think most botanists would doubt there being any "normal" apart from the conditions or circumstances in nature or in an experimental culture, but the ratio of archegonial prothalli to antheridial in the two turn-table cultures figured is certainly larger than STUDIES OF IRRITABILITY OF PLANTS ■ • PEIRCE 73 in the shelf culture shown in Figure 3. Figure 4 shows another shelf culture, but this was robbed from time to time of some of its prothalli, both arche- gonial and antheridial, for purposes of examination. Pains were taken, how- ever, to remove such numbers of male and female prothalli as to maintain the ratio. We see that this ratio is quite different in the shelf and in the turn-table cultures, there being a larger proportion of female prothalli in the turn-table cultures. Measurements of typical archegonial prothalli from turn-table and. shelf cultures were very kindly made for me by a student in my laboratory, Miss Viola F. Nichols, whom I wish to thank for her help. The data follow : Pteris sowed X, 11, '11. Shelf Turn-Table measured IV, 15, '12. 8 X 4 mm. 10 X 5 mm. 6 X 4 11 X 5 5 X 3.5 9 X 5 5 X 3 10 X 4 2 .5 X 1 10 X 5 4 X 2.5 8 X 4 4 X 3 8 X 4.5 average' 4 . 85 X 2.85 mm. 9.42 X 4.64 mm. measured IV, 15, '12. 5 X 3 10 X 10 X 10 X 9 X 8 X 7 X 6 4 4 4 4 4 average 5 X 3 mm. 9.00 X 4.33 mm. mean of 2 averages 4. 91 X 2.91 9.21 X 4.48 " area 14.28 sq. mm. 41.26 sq. mm. ratio of areas 1 to 2.5 i The difference in size, so apparent to the eye, is thus confirmed by meas- urements. Accompanying this difference in size is a more than correspond- ing difference in the number of archegonia and antheridia on the turn-table prothalli as compared with those in the shelf cultures. This ratio is nearer four or five to one. For this figure I am also indebted to Miss Nichols, but my own observations correspond. These figures, together with an inspection of the photographs, furnish the evidence of differences between Pteris prothalli grown on turn-tables in approximately uniform illumination on all sides successively, and others grown on the shelf with the light always from one side. These differences are of three main sorts : ist, in size of the vegetative parts, the prothalli ; 2nd, the numbers of reproductive organs; 3rd, the proportions of male and 74 DUDLEY MEMORIAL VOLUME female prothalli. These are very surprising, but what I have recorded above for Pteris aquilina sowed X, 11, 1911, is equally true of those started on XI, 13, 1911, allowing for the slight and decreasing differences due to age, and of both sets of cultures of Gymno gramme triangularis. How can one account for these differences? The cultures were sowed all together, the culture solu- tion, the tiles and the dishes had all been treated exactly alike and together ; no selections were made at any time. Some of the cultures were put on the turn-tables at once after sowing and the remainder were placed on the shelf by them. They were thereupon marked. Almost from the moment when the dry spores touched the moist tile they began, either to remain still, or to turn, with the tile on which they had fallen. And so they remained night and day, the diffused sunlight from the window falling through a white Holland shade nearly horizontally upon them by day, darkness enveloping them at night (for I very seldom use artificial light in the room where the multiple clinostat is), turning night and day or staying motionless, according to their position; watered from time to time with fresh Knopp's solution when nec- essary, equally warmed and similarly treated in every respect, so far as I can see, except that in one respect they are not similarly lighted. The light is the same in composition, intensity and duration, not in direction. This dif- ference alone is accompanied by the differences in the vegetative and the re- productive parts above described. It may be easier to gain some insight into this problem, into the reasons or causes of these differences, if we consider the vegetative and the repro- ductive parts separately. Acting on this principle, I proceeded to experiment upon young flowering plants grown from the seed. SEEDLINGS OF MUSTARD AND OF WHEAT. Seedlings of white mustard (Sinapis alba) and of wheat were sowed on sterilized greenhouse soil in 2-inch porous flower pots. I selected these plants because of the promptness with which they germinate and the vigor with which they grow, for a time at least, under laboratory conditions, and because the early growth of the one (mustard) is mainly hypocotyledonary and at the expense of food made by the seedling as well as drawn from the seed, whereas the growth of the other (wheat) is mainly epicotyledonary and the seedling, though well fed, is not self-nourishing for some time. Recalling the well known phototropism of these two seedlings, I thought that by ex- posing the two sets, one on the shelf and one on the turn-tables, to the same light, I could ascertain whether there were any greater stimulus to growth for the one set of plants or the other, whether if an adjustment as to position between light and darkness — that is, between more and less light — cannot be attained, growth will be more rapid than where a plant is able to attain a STUDIES OF IRRITABILITY OF PLANTS PEIRCE 75 position of such adjustment. A geranium, for example, growing on a window sill, turns toward the light. If, after it has accomplished a. bend toward the window, it be turned around, it will reverse the bend or make a new one, again carrying the tip over toward the light. And this process may be repeated indefinitely, with the same result so long as the plant can grow. Such a plant is likely to become longer in the same length of time than one beside it which has not been changed in position. If this is the eifect on stems, the position of which is reversed only at long intervals — say every other day or two — would this also be the case if the intervals were short? The result of an experiment on mustard will throw some light on this question. Seeds of Sinapis alba were sowed on II, 22, 1912, on greenhouse soil in 2-inch flower pots, six of which were put on turn-tables making four revolu- tions a minute, seven on turn-tables making two revolutions each minute, and seven on the shelf, and therefore getting light mainly from one side. These were allowed to grow until the first leaves in the plumule began to show and were thereupon measured, that is, on III, 12, 1912, nineteen days after sowing. The length taken for measurement was that from the surface of the soil to the tip of the plant. The data follow : Average lengfth of 29 seedlings in Pot I on M-minute turn-table. 4 . 08 cm. 3.88 4.13 4.53 4.96 4.38 101 " 4.32 " 16 U ' II 13 u ' III 9 it ' IV 29 " V 5 " ' VI Average length of 24 seedlings in Pot I on J^-minute turn-table. 24 ' II 19 ' III 10 ' IV 25 V 23 ' VI 6 ' VII 131 f 6 seedlings in! Pot I 15 II 20 ' III 12 ' IV 15 V 19 ' VI 14 ' VII 101 3.90 cm. 4.25 3.92 3.20 4.78 4.01 4.09 4.02 " 4.27 cm. 4.80 3.80 3.99 4.89 4.33 3.91 4.28 " 76 DUDLEY MEMORIAL VOLUME ■ From these three averages of the lejigths of mustard seedlings — 4.32 cm., 4.02 cm. and 4.28 cm. — it is clear that, so far as the growth of the hypocotyledonary stem is concerned, it makes no material difference whether the plant is illuminated mainly from one side or on all sides successively. So far as I could see, the seedlings all presented a normal appearance, both in stems and cotyledons, as to size, color and form. This experiment having failed to throw ' any light on the question, I sowed wheat similarly on III, 15, 1912, and put five pots on the quarter- minute turn-tables, five pots on the half-minute turn-tables and five on the shelf beside them. Four weeks after sowing I measured them, from the sur- face of the soil to the tip of the unopened leaf, with the following results : Average length of 10 seedlings in Pot I on M-minute turn-table. 13.2 cm. 7 " " II 11.6 7 " " III 15.2 3 " " IV 16.2 7 " "V 17.3 " " 34 " 14.7 " Average length of 9 seedlings in Pot I on J^-minute turn-table. 17.1 cm. 10 " " II 16.3 12 " " III 9.6 11 " " IV 12.9 4 " "V 14.0 46 " 13.98 " Average length of 6 seedlings in Pot I on the shelf. 9 . 68 cm. 18.63 13.45 12.7 15.3 9 ii ic II 10 a ti III 6 a