This number of ECOLOGY is dedicated to the memory

of

BURTON EDWARD LIVINGSTON

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms FIG. 1. Burton E. Livingston in his garden at Riderwood, Maryland, August 21, 1944.

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms VOL. 29 JULY, 1948 No. 3

SOME CONVERSATIONAL AUTOBIOGRAPHICAL NOTES ON INTELLECTUAL EXPERIENCES AND DEVELOPMENT: AN AUTO-OBITUARY

BURTON E. LIVINGSTON

INTRODUCTION

(By D. B. Lawrence)

This issue of ECOLOGY is dedicated to would measure the rates at which the the memory of a very influential worker environment supplied materials such as in the field of "physiological ecology," water and oxygen, and rates at which Dr. Burton E. Livingston, who died Feb- the environment removed material such ruary 8th. Livingston was for many as water from the plant. He planned years in charge of the Laboratory of his research in mineral nutrition, al- Plant Physiology and occupant of the ways with an ecological background Chair of Forest Ecology at The Johns which included the fluctuating regimes of Hopkins University. Since his retire- light, temperature, and humidity inherent ment in 1940, he had carried on his re- in the greenhouse conditions at Baltimore. search at his home in Riderwood, Mary- He was not satisfied with constant condi- land. He probably will be remembered tions of environment nor with conditions best by ecologists for his contribution in at any one season of the year so he would atmometry,' "Atmometers of porous por- repeat experiments four times or more to celain, and paper, their use in physiologi- cover the various seasons. Nor was he cal ecology," published in ECOLOGY in satisfied with standard greenhouse bench 1935, and for his publication with For- experimentation which introduced tre- rest Shreve "The distribution of vegeta- mendous differences in microclimatic in- tion in the United States, as related to fluences; nothing short of a tremendous climatic conditions," but his contribution rotating table with flowing solutions, and to physiological ecology was spread widely using very carefully selected plants, chosen over the field of water relations and min- not only for uniform genetic background eral nutrition of plants. His viewpoint but also uniform physiological response, was outstandingly ecological and he con- made him feel content. Livingston was ceived of the environment as something one of the first to carry physiological which had an inherent capacity to supply methods into the field, and to him ecology materials and energy to the dynamic or- is greatly indebted for pointing the way ganism, and also to remove materials and to improvement in method and viewpoint. energy from an organism. He went to Probably Henrik Lundegardh is the only great lengths to devise instruments which other person who can compete with him 1 Readers will be glad to know that the at- for top credit in welding physiology and mometer business which had long been operated ecology together. Livingston had few on a world-wide scale by Dr. Livingston from his home in Riderwood, will be continued at the prejudices, but he had one that was strik- same address by Mrs. Livingston. inor: he had no use for statisticians or 227

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms 228 BURTON E. LIVINGSTON Ecology, Vol. 29, No. 3 their methods and, so far as I know, he fuse was very definitely attached which never went beyond the mean and the ex- took 12 to 15 years to burn. Clements tremes in his mathematical analysis of in 1905 was first in this country to re- scientific data. He felt that if there were spond. In the first chapter of his book important differences to be detected, they "Research methods in ecology" he tried could be seen from a table of figures, and to show that there was essential identity he felt that statistical treatment was too of physiology and ecology and he thought often used as a substitute for adequate that the two fields would become one and planning and careful technique. His lack that the name would probably be physiol- of appreciation of this field was as strik- ogy. Clements seems to have coined the ing as Clements' blind-spot for genetics, term "experimental ecology" in the same and yet this did not deter his students chapter. He considered Bonnier (1890) from interesting themselves in that line. the first to carry on ecological experi- Indeed his antipathy toward statistics mentation out of doors. In December probably contributed strongly to making 1908 at a Botanical Society symposium a student of his one of the U. S. Air in Baltimore entitled "Present problems Forces' top statisticians today. in plant ecology" Livingston was one of Livingston's finest attribute, so far as the five ecologists and physiologists who student training was concerned, was the spoke their minds. All pointed out vig- limitless energy and effort he devoted to orously the need for precise measurement the criticism of a manuscript. Each page of environmental influences 2 and empha- was gone over orally, across the table, sized the fact that ecology could be ex- paragraph by paragraph, line by line, perimental, and had to be if it was to word by word, critically examining each advance to anything that scientists would expression to be sure that no ambiguity mention without a smile. Almost imme- was left, and that there was not some diately thereafter ecologists began apply- more accurate way of expressing the idea. ing experimental methods to the solution He devoted the same energy to answering of ecological problems and this has letters from fellow scientists, and would brought forth fruit abundantly in the often answer a query with a 10-page let- applied fields of forestry, range manage- ter and the apology that he was sorry he ment, plant pathology, horticulture, and didn't have time to write a short letter. wildlife management. The year 1893 Livingston will always occupy an im- should therefore be considered the date of portant place in the history of ecology stimulus and the year 1905 (Clements) because he played a vital role in directing the beginning of the response that brought ecological research toward precise meas- ecology close to the side of physiology, urement of the natural environment, and but it was the year 1908 and the ideas experimentation, during the formative of Livingston, Cowles, Transeau, Spald- years of the development of our field. It ing, and Shaw, that really set ecology and will be remembered that until the late its applied phases off on a new tack with 1800's ecologists, with the exception of the recommended use of experimentation Bonnier in 1890 and 1895, were content in the solution of problems. with superficial observations and untested It seems unnecessary here to list indi- hypotheses, and as late as 1893 the state- vidually the publications that emanated ment was propounded at the Madison, from the Laboratory of Plant Physiology Wisconsin, Convention of Botanists that during Livingston's 31 years of active "ecology was observational and physiol- service as its chief, but a statement of ogy experimental." Nothing could have the total production will reveal the energy been more stimulating to ecologists, and 2 Livingston would not use the word factor in their minds seemed to explode with in- this sense; he preferred to restrict its use to the dignation, though to be sure, a long time- mathematical sense.

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms July, 1948 CONVERSATIONAL AUTOBIOGRAPHICAL NOTES 229 of the man. In all, 269 articles were pre- (white and Norway pine) in the 80's. pared for publication, and 30 students I afterwards mapped that glacial topog- received their Ph.D.'s in that time. It is raphy in the paper on Kent County and doubtful that any other person in the na- described its natural vegetation after a tion has ever demonstrated such a sus- fashion (BOTANICAL GAZETTE, 1903). tained high level of activity in this branch Our home lot was largely in cultiva- of science. It is not the purpose of this tion, with little lawn. There were beds introduction to catalogue the scientific of vegetables, rows of Mexican blue activities of this remarkable man. The maize and potatoes, beds and patches of intent here is to point out the unusual native "wild flowers," currant and goose- fortune of ecologists in having on record berry bushes, grape vines on the "back the following "auto-obituary" as Livings- fence," sumacs, tall tree-like sunflowers ton jokingly called it when he submitted (H. annuus) for the chickens. Around it to me at my request in November, the margins of that lot were large trees 1937. It is important for us because it of sugar maple, silver and Lombardy pop- traces the ecological development of a lar, elm, white cedar, red cedar (Junip- very influential worker of our field of erus virginiana), tamarack, butternut, science during the critical early stages sassafras, horticultural cherries and ap- in its development. ples-both eating and crab. Two of the crab-apple trees had been so grafted that THE AUTO-OBITUARY-NOVEMBER. 1937 each bore three or four kinds and sizes My boyhood was spent in Grand Rap- of fruits, which ripened one after another. ids, Michigan, where our home grounds A single peach tree died before I became were an unusually large town lot near the well acquainted with it. Almost all of edge of the growing city, at the northeast the old-fashioned ornamentals of that re- corner of what are now Sheldon and gion and period were abundant-tulips, First Streets. Across the street to the hyacinths, croci, candidum lily, daffodils, south was the shallow valley of a brook, gladiolus, English violet, primula, colum- which flowed through a treeless mucky bines, various ferns, vinca, euphorbia, stretch where grew mints and iris, Joe perennial pea, flowering currant, lilacs, Pye weed, iron weed and milkweed, snowball, euonymus, roses, German iris, sedges and grasses. In other directions hollyhock, hibiscus, and many annuals. there were neighbors' homes and a few I was something of a specialist in grow- lots still vacant-remnants of upland ing pansies in rock-paved beds when I meadows, where we "studied" bumble was about fifteen. In spring there were bees in one of my early summers, and small flats of seedlings in the house, red clover. The natural forest of this where there were always numerous "house moraine edge was "oak opening," but plants" before the windows. We had ar, there were few native trees excepting a oleander, but no rubber plant. I sup- patch of upland oak forest a block or two pose I planted the seeds of my two potted away. I think I was familiar with the trees of orange and lemon at the age of burr oak and the white oak before I en- four or earlier. They were inconven- tered my fifth year. The other trees of iently large before I went away and left my very early acquaintance had been them to whatever fate finally overtook planted, though most of them represented them. species native in that region. A few Perhaps one of the most influential blocks to the west, in the broad valley portions of my botanical environment in of the Grand River and a tributary creek, very early years was our haphazard there were railways and a variety of plantings of native flowering plants- swampy stretches, where the river over- spring flowers such as two hepaticas, flowed in spring. There were log jams bloodroot, Claytonia and the rest, two

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms 230 BURTON E. LIVINGSTON Ecology, Vol. 29, No. 3 yellow cypripediums, and so on; also the with tools of rusty iron, plows, scrapers common weeds of the garden and road- and the interesting contents of those old side, like stellaria, polygonums, pigweed, tool boxes. The red-chimneyed kerosene amaranth, the smaller ragweed and others. lanterns were of special interest. The last-mentioned was esteemed in mid- In those days, in that region, street summer as furnishing a convenient added grading and paving was all accomplished weight for the end of a kite's tail. We in summer and my father was at home called many of these plants by their most of the time in winter. In that sea- Latin names; I am unable to recollect a son the tools were repaired for use the time when I didn't recognize the two following summer. My father and older hepatica species of that region as triloba brothers were experts at the hand wood- and acutiloba, but I think I had reached work of these operations, and I cannot high-school age before I had any name at remember when I was not familiar with all for Ambrosia artemisiaefolia, and I saws and planes, with draw-shaves and have never called that kite-tail plant rag- spoke shaves, with bolts and screws and weed without feeling bookish and pedantic nails, with wrenches and files. In my to a degree. time we never had a forge, but I played Perhaps I took naturally to plants and about an ingenious shaving horse, which their ways, but parents and older brothers held the work when an axe handle or and sisters had been and were specially pick handle was being shaped from se- interested in plants, and to their interest lected pieces of hickory wood. It is thus was due the peculiar immediate environ- seen that my boyhood surroundings ex- ment into which I came, and those early tended into the realm of simple mechani- surroundings gave what was going to be- cal operations as well as into that of come my mental and philosophical lean- gardening and plant lore. ings an unusual bent towards the field of Still another feature of those early plant science. But there were other in- surroundings lay in a home atmosphere fluences. My parents had grown up on of books, reading and ideas. There were farms and they had the usual knowledge many books about the home and they and interests of farmers, but we lived in continually increased in number. There town and my father had become a street- were generally a number of books from grading and sewer contractor for the rap- the public library. Every one in the idly growing community. In his business family circle read and read. We had the he dealt with soil and gravel and paving complete file of Harper's Magazine and stones, with sand traps, gutters and grav- the subscription was always continued; elled road surfaces, with pick-and-shovel for a number of years we had the old workers and teamsters, with plows, scrap- Century Magazine. Beginning when I ers, paving hammers and an iron road was seven I read the St. Nicholas maga- roller that was drawn by a team of horses. zine pretty thoroughly for the next four He paved several long stretches of streets or five years. The "big dictionary" with cedar blocks, with coal tar poured (Webster's unabridged) was in plain into the joints, but I am sure he never sight on a table in our living room, and it saw a street paved with brick or asphalt was used almost every day. I think my or concrete; those things came later. My first use of it was to elevate a chair seat boyhood surroundings therefore included when I sat at that table to play with letter the grading contractor's equipment; tool cards, author cards, jack-straws or check- boxes, of shovels, picks, saws, hammers, ers. I became acquainted with the names canthooks and such, came home in the of many literary personages and their pic- fall, to remain in our backyard till they tures, also with their main titles, from went out onto the "jobs" the following that old game of "authors." We were spring. My earliest memories are framed never much interested in ordinary playing

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms July, 1948 CONVERSATIONAL AUTOBIOGRAPHICAL NOTES 231 cards, whose peculiar attractiveness has son, Carlyle, and so on. Thoreau's writ- always been far beyond my mental capac- ings may have influenced my develop- ity. Longfellow and Bryant were my ment more than those of any other in first poets. From having them read to those early days. me, I knew by heart many of their pas- From the time I could handle hammer sages before I was able to read. I at- and saw I was forever "building things," tended a Congregational Sunday school following more or less in the path of my regularly for many years, finally ceasing father and older brothers. A treadle when the teacher of my class left town. scroll saw was mine and was in almost Though some of the older brothers and daily use throughout four or five years sisters attended that church, I never before I entered the high school. In the cared for that popular sort of recreation, high-school period I built two canvas and my reading of the Bible was gen- canoes, but never used them because erally confined to the preparation of each available water was too far away. In Sunday-school lesson. I never knew that period I built some glass-covered either of my parents to attend church. trays for our butterfly collection (my Life flowed smoothly in our group, with- older brother Luther's and mine) and out the common religious precepts. I made presses and cases for dried plants. remember a few experiments with pray- Whenever mechanics -could be observed ers when I was very young, but they at work I followed their every move. I always gave negative results. Only when am still greatly impressed by memories a ministerial relative-of whom there of the operations seen in a cooper shop were many, it seems-came to our home and in a wooden-shoe shop passed on the did I hear grace said at mealtime. It way to school when I was about ten. I may be interesting to note that I cannot watched the neighboring blacksmith. A remember to have ever "believed" in relative in a nearby village, whom I some- Santa Claus though we always celebrated times visited, was a cooper as well as a Christmas with many presents and an farmer. In early grade-school days I excess of good things to eat. In the played games with other boys-such as later teens, for interesting information leap-frog and pullaway-and made and about how various peoples have tried to flew kites in summer, built and used set up their backgrounds for thought, I bobsleds, skated and caught rides on dipped now and then into the great re- sleighs in winter. For a number of ligious books: The Koran, the Hindu years I split and stored our firewood, books, the Bible, the Book of Mormon, kept the snow about our home shoveled the Meditations of Marcus Aurelius, in winter and ours and a neighbor's lawn Plato's Dialogues and the like, all in mowed in summer. I earned my first English. After dozens and dozens of money by making toy furniture and by boys' books had been read, including mowing that neighbor's lawn. I still C. C. Coffin's volumes on the American have a set of Shakespeare's works bought Revolution and the Civil War and stories with that lawn-mowing money-but the like Little Lord Fauntleroy, Stockton's morocco-leather backs are now mostly Tinkham Brothers' Tide Mill, Roe's gone. Driven back to Eden, Carrol's Davy and Before I began to attend school I had the Goblin, I think my first serious ven- seen flies' wings and the like under a ture in grown-up reading was Thoreau's small compound microscope. It was in Week on the Concord and Merrimac that same pre-school period that I Rivers. I must have read the latter brought some pieces of woody stems- when I was fourteen, the winter before such as those of raspberry-into our I entered high school. In the next half living room, at a season when the leaves decade I read much of Thoreau, Emer- were off, and cut thin sections of them

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms 232 BURTON E. LIVINGSTON Ecology, Vol. 29, No. 3

to examine under that little microscope. several street-car lines. I thus learned I think I was familiar with the appear- very early to name most of the conspicu- ance of cell structure and with tracheae ous wild plants of that region and to know of such small woody stems before I was where one went to find each sort. We eight years old. I was eight when I specialized a bit on native orchids. I first went to school, where they started remember buckets and pitchers of Cypri- me in the third grade. Somewhat later pedium spectabile, Habenaria psychodes, my brother Luther had a larger micro- H. fimbriata, Calopogon, and so on, scope for a year or more, through which which were arranged about the home on I was allowed to look now and then, but Sunday evenings after such trips. that instrument went away soon-it may In the high-school course in botany have been borrowed-and my real be- each student made an herbarium of one ginnings in microscopy were in high- hundred or so plants. By that time I school days. was botanizing on my own. I added to I suppose I must have learned a great that herbarium for a number of years, deal in the grade schools, which I think till I went to Ann Arbor; and I received were very good in that Michigan town ten hours of advance credit in botany for at that time, but I am unable to recall my herbarium when I entered the Uni- many particulars of what was gained versity of Michigan. Without any ar- there. Orderly development of my con- duous study I came to know many plants scious life seems to have begun with the and their habitats, and the use of books high-school period, which was definitely like Gray's Manual, before I entered col- formative for later things. It was of lege. I never received any instruction in course in that period mainly that botany systematic or taxonomic botany any- (Gray's Lessons and Manual), zoology, where excepting in the half-year course physics, algebra, geometry, chemistry, in high school. rhetoric, grammar, drawing and the like After finishing high school I spent a came really into my conscious field of- week at the first Chicago World's Fair, mental vision. Following an English a good deal of it in the horticultural course throughout, I awoke to the need building; then went to Short Hills, N. J., for other languages at the end of the to work as laborer at Pitcher and Manda's second high-school year and I worked by great nurseries-at six dollars a week. myself at elementary Latin that summer My older brother Luther-a cataloger and at elementary Greek the next sum- and bibliographer-had been making cat- mer, completing in school, as "extra alogs for that nursery and had spent work," the first three years of Latin and a year in Colombia and Venezuela col- the first year of Greek. I never had any lecting Cattleyas and shipping them instruction in these languages beyond the north. He and I lived together at Short reading of Cicero's orations and a little Hills for a year, in a friend's barn, which of Xenophon's Anabasis. Modern lan- we transformed into a passable cottage. guages came later. We did our own cooking and were vege- Almost from the very first I, more or tarians. I worked in most of the various less unconsciously, studied nature as well nursery departments, hardy perennials, as books and people. Two older brothers palms and ferns, seeds and bulbs, orchids were much interested in wild plants in (there were several acres of orchid a sort of quasi-scientific way, and I began houses), packing room. I learned to do to accompany one or the other of them most of the things gardeners do; in the on their Sunday trips to the woods as early autumn I got out the plants for soon as they thought me old enough to orders for hardy perennials. The hardy go along. There were wild areas within grounds comprised about seven acres- easy walking distance of the terminals of just beds and beds and cold frames and

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms July, 1948 CONVERSATIONAL AUTOBIOGRAPHICAL NOTES 233 cold frames ad lib. Everything was la- 2-inch pots. There was a large field of beled with the Latin name. My brother Liliur auratum and L. speciosum, where, Luther was the first to use consistently for several days, I cut tubs full of flowers Latin names and actual photographs for -to go to and be auctioned commercial florists' catalogs, at least in for a cent or two a flower. With a Polish this country. There were many houses boy, I collected a thousand plants of of ferns, palms, anthuriums, chrysanthe- Solomon's seal on the hill west of East mums-a great exhibition of chrysan- Orange, to go to somebody's estate in themums every fall. We were always in- . One afternoon I collected one terested in knowing the regions from thousand stems of golden rod, for some which the plants had come. In my year girl's wedding in New York. there the firm was about the first to Saturday afternoons, Sundays and holi- introduce Japanese iris in this country; days, in this Short Hills period, I spent we were propagating the first twenty botanizing in the neighboring region- varieties received from Japan. Travel- the hills around East Orange, the New- ing collectors sent in plants from various ark marshes, etc. In early summer of parts of the world and there were travel- 1894 I left the nursery and botanized ing salesmen who covered this country. every day till fall, when I went to Ann One man did nothing but take photo- Arbor. The herbarium grew apace, but graphs of plants. Foremen of the vari- I never added to it considerably after ous departments and these traveling men that summer. became known later as heads of nurseries At the University of Michigan there -Manda, Lager, Bobink and Atkins, et was no one who cared especially for the al. Their names are still familiar to taxonomic botany of higher plants, though gardeners. That firm, the United States Prof. V. M. Spalding was sympathetic Nurseries, did a great thing for garden with a plan I had proposed to catalog the and greenhouse botany and ornamental plants of the Ann Arbor region according horticulture. They had a large exhibit to habitat. We knew little or nothing of at the World's Fair-some thirty enor- ecology, the word wasn't known to us, mous tree ferns among other things. but Spalding knew natural vegetation But that nursery never paid, and Mr. very well. There was- a fair local her- Pitcherhadfinancial reverses about 1903- barium. I worked at this plan by myself 4 and ceased to pay the annual deficit. for a year or two. In the meantime I So they tried to curtail, and my year "did courses" in plant morphology, phvsi- there was in the period of curtailment. ology, animal embryology, chemistry, I suppose there were thirty or more men physics, German, French, Italian, and the employed in my time. After I left, to "required" courses in mathematics, psy- go to college, the firm went out of exist- chology, etc.-just one short and unsatis- ence and their large and rare plants were factory course in English composition. sold at auction; the big tree ferns, which I had studied Spanish with my brother had come back from Chicago, were sold at Short Hills-he had recently returned at five dollars each, for the roof of the from South America and liked to speak old Madison Square Garden. In my that language. Because of my plan con- time they were selling off the specimen cerning plant distribution I asked Prof. plants and we were propagating cheap F. C. Newcombe to let me take his lec- stuff, to try to "make some money"- tures in plant physiology, omitting the greenhouse ferns in flats and 2-3-inch laboratory work, but he said that would pots by the thousand, palm seedlings, etc. be only a half-year course and it would I remember working with one other chap do me no harm to know a bit about for two or three weeks doing nothing physiological experimentation; further- but pot tiny fern plants from flats to more, he would predict that after I had

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms 234 BURTON E. LIVINGSTON Ecology, Vol. 29, No. 3 completed that laboratory course I would and others. At that time I first became find physiology more interesting than any acquainted with N. L. Britton and John other field of botany. How he reached K. Small, both of whom exerted a good that prediction I can't tell, but I became deal of influence on my development. his laboratory assistant the next year and Columbia University soon moved to have stuck to physiology ever since. Morningside Heights. In that connec- Newcombe was not interested in ecology, tion, I remember well a rainy Saturday which was then just getting started un- afternoon in the summer of 1894, when der the leadership of Warming and Luther and I explored the woods and Schimper. Their books were not avail- fields where the great university was able to me till I went to the University going to make its home, as had recently of Chicago in the summer of 1899. I been announced in the newspapers. was a confirmed enthusiast for physiol- After leaving Ann Arbor, in June, ogy by then. 1898, I got me a job teaching an array Luther became well known as a bibliog- of sciences (physics, chemistry, human rapher and specialist in rare books. He physiology, physical geography, botany was first with Dodd, Mead and Company and zoology) in the high school at Free- and then-in partnership with one of the port, Illinois. I found time to explore Messrs. Dodd-with Dodd and Livings- the neighboring country (with a bicycle), ton. The last named firm had a store on but did little with botany as such. I Fifth Avenue for several years at a later kept my alga growing in cultures in my time, till Luther became librarian of the bedroom window; the alga had been Widener Collection, which had been given brought from Ann Arbor (discussed more to Harvard University after Harry Wide- fully below). In the late winter of 1898 ner had been lost in the Titanic disaster. I made applications for fellowships in From a wheel-chair, Luther saw the about ten universities and received three famous Harry Widener Memorial Room appointments that spring-at Harvard in the great Harry Widener Memorial University, University of Wisconsin, and Library before the building was finished, University of Chicago. I became a fellow but he died before he occupied it. At and assistant in the last-named institu- the present time the librarian in charge tion, going there about the end of June. of that room is Luther's widow, who Prof. C. R. Barnes had been called from writes as Flora V. M. Livingston. My the University of Wisconsin the year be- eldest brother, Lincoln, was a lawyer, fore, to take charge of plant physiology first in Grand Rapids and then in Denver, and to edit the BOTANICAL GAZETTE. He but he died in middle age, after develop- gave very excellent lectures for elemen- ing a law library in the last-named city. tary and advanced students; he prepared The first summer vacation of my Ann a set of laboratory outlines on the Detmer Arbor period was spent in New York plan, which have become classic through City, where my then widowed mother hisand students, "grand-students," and "great I visited my brother Luther and a sister grand-students." The conducting of the who was living with him, on 21st Street, laboratory work was my own responsi- near Ninth Avenue. Those months I bility; on arrival, I was given the neces- devoted to a library study of plant geog- sary keys and some outlines and was told raphy. Prof. N. L. Britton gave me per- that "the laboratory class would appear mission to study in the library room of next Tuesday." Throughout the next the old botany building of Columbia Uni- four years, with some vacations, I con- versity, on 49th Street, where I was to ducted that laboratory, on the fourth floor be found almost daily that summer, read- of the Hull Botany Building near 57th ing in many of the classic treatises on Street, consulting Professor Barnes at plant geography, De Candolle, Grisebach, times, but in all that period I think my

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms July, 1948 CONVERSATIONAL AUTOBIOGRAPHICAL NOTES 235 chief was never in the laboratory rooms In those times American botany had, in more than perhaps twenty times. When most places, already relegated plant tax- he did appear he showed quite wonderful onomy to the basement, and its main facility in handling experimental appara- quarters were occupied by Strasburger tus, but he always hurried away and he morphology and the beginnings of cytol- never interfered. He left the laboratory ogy; physiology was still generally strug- to his assistant but he always seemed to gling in attic rooms, and ecology-a sort be aware of all that was going on there. of regenerating outgrowth of taxonomy, On an early occasion, when I had asked floristics and plant geography-was just him whether things were going forward beginning to be considered. It was nota- as he wished, he replied, "You may rest ble that three professors of plant physiol- assured that I will let you know if I ogy were appointed about 1909, at Wash- think anything is going wrong." It was ington University (St. Louis), at Harvard through his lectures and personal con- University and at The Johns Hopkins versations that Barnes's students received University. A number of assistant pro- guidance and inspiration from him, and fessorships and instructorships in the new through the sometimes peppery reviews science were already in existence in this that he prepared for the BOTANICAL country and interest was turning in that GAZETTE. I think Barnes's mental proc- direction. esses and his facility in presenting his Throughout my Chicago period I was thoughts were more nearly perfect in greatly attracted toward the field of ecol- precision, clearness and completeness than ogy, which was being rapidly developed those of any other person with whom I there under the general leadership of have ever worked. I was indeed fortu- Professor J. M. Coulter and at the hands nate to be his first active assistant at of the young H. C. Cowles. Like Asa Chicago. His publications in plant physi- Gray, Coulter had realized the possibili- ology were few, but his mind and per- ties of the newer phases of botanical study sonality exerted a profound and lasting that were attracting so much attention in influence on the development of this . A taxonomist himself, he en- science. couraged Chamberlain and Cowles-who At that time Goodale held a professor- were among his earliest Chicago students ship in plant physiology at Harvard Uni- -to devote themselves respectively to versity, but Newcombe, MacDougal and the morphology, histology and cytology Barnes were, I think, the first with that of reproduction and to ecology. Cowles title to devote themselves primarily to was coming into ecology from geology and physiology in this country. In these early plant taxonomy (see ECOLOGY for July, years of the present century, plant physi- 1935) and his main interests lay in the ology was in the incunabula or even em- details of local plant distribution. That bryonic stage of its development here, was exactly the field towards which I although it had already become a distinct had been groping, with the encourage- science in Germany, especially at the ment of Spalding, in the early part of hands of men like de Vries, Sachs, Det- my Ann Arbor period and it inevitably mer, and Pfeffer. Newcombe had been appealed strongly to me when I became inspired by Pfeffer and it seemed to give intimate with Cowles at the time of his the fine old Geheimrat some pleasure classic sand-dune studies. Cowles's great when, in 1908, in his Leipzig laboratory, contribution was toward what he called I remarked to him that I was his "intel- physiographic ecology, which dealt with lectual grandchild." The three volumes local features of distribution in terms of of Ewart's classic English translation of the physiographic characteristics of the Pfeffer's Pflanzenphysiologie appeared in corresponding plant habitats; he and his the very earliest years of this century. students studied vegetation rather than

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms 236 BURTON E. LIVINGSTON Ecology, Vol. 29, No. 3 plant individuals or species, in bogs and differences in those plains might be mainly swamps, on beaches, flood plains and due to differences in soil-moisture con- bluffs, on moraines, in ravines. Glacial tent in summer; sandy uplands were gen- topography was just as important in this erally dry unless the soil was much modi- as was plant taxonomy, and the concepts fied by humus, and clayey uplands were of plant physiology always entered into generally moist, the more so as the humus these studies to a considerable degree. content or covering of the soil was greater. For that period I add the name of Cowles Lowlands were generally moist or wet to those of Newcombe and Barnes as rep- whether the mineral constituents of the resenting the personal contacts that most soil were sandy or clayey-because there influenced my own scientific development. the subterranean water table was at the In my last year at Ann Arbor I had soil surface or not far beneath it. carried out a somewhat elaborate experi- That bog water might perhaps differ mental physiological study on the influ- from the water of drained swamps by ence of the osmotic characteristics of having a much higher osmotic value had liquid culture media on the growth and been deduced from Schimper's brilliant development of a small polymorphic fresh- but cryptic observation that bogs are water alga, and that study was continued "physiologically dry." That deduction throughout my Chicago period. (See was easily shown to be untrue-Thy cryo- "The r6le of diffusion and osmotic pres- scopic tests carried out on samples of sure," Chicago, 1903.) That was also, water from various bogs and swamps of of course, an experimental study in the Michigan and New Jersey. These waters ecology of that alga, and it turned out were then tested by employing my alga that the polymorphism of the latter might as an indicator and it emerged that water be controlled-as we used to say-not from about the roots of typically bog only by the osmotic characteristics of its plants acted on the alga as if it contained environment but also by environmental stimulating or slightly toxic solutes. Bog characteristics that influence water ab- waters were generally more acid than the sorption and water loss; a relatively others (by titration tests; H-ion concen- strong solution of non-toxic solutes acted tration and the now familiar pH did not on the cells as a drying agent and similar appear on the physiological and ecological effects might be produced by exposure to stage till somewhat later), but their stim- the air (evaporation). ulating or toxic influence was apparently It was thus natural and easy to try to not directly related to total acidity. Per- find for ordinary plants relations of vege- haps I was dealing to some extent with tational differences to the drying influ- some cryptotrophic or cryptotoxic solutes ences and water-supplying influences of in bog water, presumably of organic na- air and soil. I had the thought that the ture, and my accounts of these studies physiographic conditions emphasized by now seem to suggest substances more or Cowles might be effective largely in that less resembling some of the hormone-like way. For the pine plains of Michigan, substances now being studied so vigor- which I studied in the summer of 1901 ously in both animal and plant physiology. (see BOTANICAL GAZETTE, 1905), it While at the University of Chicago I turned out that upland vegetation types listened to several series of lectures on were rather nicely correlated to the water- plant morphology given by Coulter, an holding capacity (size of particles) of the inspiring teacher whose talks and writ- surface soil, which appeared to determine ings are fine examples of excellent presen- water supply and soil aeration, for the tation. What I remember of Coulter macro-climate is essentially the same for centers about his fine personality and his the whole of the region considered. This remarkable facility of expression. Of seemed to mean that upland vegetation course his fundamental philosophy or out-

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms July, 1948 CONVERSATIONAL AUTOBIOGRAPHICAL NOTES 237 look on things in general was not physio- Although he was always enthusiastically logical. He always tried to avoid the sympathetic with my problems and find- implications of teleology but one felt that ings, he did not offer me a fellowship in he was perhaps only half-hearted in that. his laboratory, and so I remained in the Barnes's attitude towards the problems of plant physiological field. causation-in plant activity, development From chemical studies of a quasi-re- and evolution, and apparently in every- search nature under the guidance of thing-was similar to that shown in Gomberg (at the University of Michi- Pfeffer's later writings and in those of gan) and of Lengfeld and Stieglitz (at Loeb; light on problems of determination the University of Chicago) I gained much and causation was to be sought in terms in appreciation of the physical and chem- of the properties or characteristics of ical bases of vital phenomena. As is gen- matter and energy, and teleology and erally true for any group of young men anthropomorphism were to be avoided studying in a university-or elsewhere, wherever the nature of language would for that matter-the graduate students permit. I was afterwards much im- and instructors at the University of Chi- pressed and encouraged in my attempts cago exerted great influence on the men- at clear thinking and writing by Ver- tal development of one another. I gained worn's little essay on Kausale und Kon- much from intimate friendship with men ditionale Weltanschauung, but encourage- like H. N. Whitford, W. B. McCallum, ment came to me mainly in those years H. Hasselbring, J. B. Overton, 0. W. from Barnes and Loeb. Caldwell, J. P. Goode, A. W. Greeley, Loeb's general course in physiology and the rest. broadened my field of intellectual and My native instincts were perhaps as philosophical vision very greatly. My much bent toward literary art as toward Weltanschauung seems always to have the intellectual field of science. Language been of the etiological rather than of the and its use have always claimed much of teleological sort, but most writers whom I my attention by the way but, aside from read continually indulged in teleology or elementary high-school and college courses even in out-and-out mysticism. Over- in Latin, Greek, German, French, Italian enthusiastic as he often was, Loeb's char- and English, my education in this general acteristic approach to physiology from field was informal and self-conducted. I the standpoint of physical and chemical have always been wont to devote a good concepts represented a remarkable for- deal of time each week to general reading ward step in the fundamental biological of eminent and less eminent writers. thought of that time. It soon became a Never becoming expert with any foreign sort of vogue to point out where his in- language, I dabbled superficially in many, terpretations and generalizations were far being able to read easily only German two simple to find full support in the and French, however. It has always results of subsequent experimentation, been pleasant to study the dictionary, but what we call his mistakes of over- especially for etymologies and the finer enthusiasm turned out to be valuable con- shades of word meaning. tributions; where he went too far he led A part of one summer of my Chicago others to proceed more slowly in the same period was devoted to teaching botany direction. I think I perhaps received in the Eastern Illinois State Normal somewhat more edification from Loeb School, at Charleston, where I had many than from Barnes, but both men exerted very pleasant and inspiring conversations enormous influence in my formative pe- with the president of that school, L. E. riod. There were times when I decided Lord. that animal physiology should be my field, Toward the end of my Chicago period but Loeb always advised against a change. I spent an autumn at the New York

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Botanical Garden, in an experimental the next December, spent that winter and study of the influence of a number of the early spring in eastern institutions, inorganic salts on the polymorphic green partly in library studies but mostly in alga mentioned above. It was at that experimentation at the Missouri Botani- time that I became acquainted with Mac- cal Garden, at St. Louis, where I gained Dougal, Lloyd, H. M. Richards and C. C. much from pleasant contacts with Wil- Curtis, all of whom were very helpful liam Trelease, H. von Schrenck and J. A. and encouraging. The following summer, Harris. That St. Louis period was de- and a grant from the Carnegie Institu- voted to a study of transpiration in tion, were spent in study at the new greeenhouse-grown cacti. The year 1908 Desert Laboratory, with Cannon and was spent in , mainly at Munich, Lloyd. From the study of desert plants where I learned a great deal from von came many new ideas, especially with Goebel's lectures and field trips and from respect to water supply and water loss. Hegi's botanical-geological excursions. My interest in osmotic pressure and soil Renner, who was studying the physics dryness as environmental features was of plant transpiration at that time, was broadened to include evaporativity and my frequent companion and I owe much the drying influence of sunshine. The to his keen mind. At Munich I was porous-porcelain atmometer was devised engaged in studies on soil moisture and and first used by me at that time-but I it was there that I first devised and tested afterwards learned that it had been inde- the porous-porcelain soil-moisture "me- pendently devised at least twice before, ter," which has recently been independ- by Babinet and by Mitscherlich. After ently developed by Rogers and others. the experiences of that first summer at In the autumn I spent a few weeks at Tucson transpiration and water supply Pfeffer's institute at Leipzig and had the were among the topics that I found most good fortune to have some conversation interesting. They enticed in many direc- with the Geheimrat almost daily. I also tions. visited and became well acquainted with Returning from Tucson in September, Klebs and Gliick, at Heidelberg, and I spent the following autumn at the U. S. made tourist excursions through Ger- Bureau of Soils, in Washington, where I many and Switzerland. In the early gained much from conversations with winter I spent a few days at the Rotham- Whitney, Cameron, Schreiner and a num- sted Station in England, where Hall was ber of other members of the bureau staff. then director. At the end of 1908 I at- We were studying the possibility that tended the Baltimore meeting of the some soils might contain chemicals that American Association and associated so- acted to retard plant growth therein. I cieties on my way back to Tucson. It returned to Chicago for the winter quar- was at the time of that meeting that I ter at the University and then received first saw the late Professor D. S. John- my first appointment to a permanent posi- son's recently established garden, on the tion, in the Bureau of Soils. That posi- newly acquired Homewood tract which tion was not very permanent, however, was to become the home of The Johns for I resigned and entered the staff of Hopkins University. At the close of the Desert Laboratory the following Jan- that meeting Johnson led a day's excur- uary. MacDougal was the newly named sion through the snowy woods and fields director of that laboratory and my studies north of Cockeysville, Maryland, in which in physiological ecology-especially on re- Lloyd and I took part. That was a lations of plants to soil moisture and to memorable occasion; we were almost cold evaporation-were continued with his en- and became more than almost hungry, couragement. I remained at Tucson till but were withal exceedingly enthusiastic

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms July, 1948 CONVERSATIONAL AUTOBIOGRAPHICAL NOTES 239 over many different sorts of botanical Building on Eutaw Street (the place is observation and discussion. now a parking lot) and conducted a lab- Another memorable excursion from oratory course and research in the small which I gained a great deal was from building and greenhouse which already Tucson to the Salton Sea, in the early stood on the Homewood grounds, at the summer of 1909. It was arranged by south side of the new botanical garden. MacDougal partly for several visitors. Among those who studied with me that We spent a day or two around Yuma, a first year were F. H. Blodgett, L. A. day at Yuma Junction and a week in Hawkins, W. D. Hoyt, W. R. Jones, L. camp near the U. S. Weather Bureau's W. Sharp, H. H. York and W. H. evaporation station, where we became ac- Brown. Brown 3 is about to come from quainted with the field studies, in evapo- Manila to be lecturer in botany here. ration, of Bigelow and his associates, Our group was an ideal one, working in who were very busy there with pans and an ideal atmosphere, and I gained as tanks, anemometers, thermometers, and much from those students as they gained psychrometers. We operated atmometers, from me. examined the neighboring desert hills and With the help of Hawkins and others, the newly-formed beach, visited Pelican I soon devised and constructed the first Island and the place on the shore from rotating table for use in the standardiza- which pumice fragments, worn to rounded tion of atmometers. That table, which is pebbles and boulders, were floating away. eight feet in diameter, first stood in the I afterwards referred to those floating attic of the small building just mentioned; stones as notable examples of a remark- it is now on the upper floor of the Lab- ably perfect "adaptation" in the realm of oratory of Plant Physiology (fig. 2), the non-living. where it is still occasionally used. Two While on that trip I received a for- or three years later Shive and I intro- warded telegram asking if I would con- duced rotating tables for series of simul- sider an appointment as professor of plant taneous plant cultures, and such tables physiology at The Johns Hopkins Univer- for equalizing aerial conditions in experi- sity, and my affirmative reply was sent, ments with plants soon came into general on a Sunday evening, from the railway use in many laboratories. After Sam F. office at Yuma Junction (where the Trelease and I had introduced the em- branch for El Central leaves the main ployment of continuously flowing solu- railway line), while the west-bound train tions in solution-culture experimentation halted for a quarter-hour; the appoint- it became desirable to arrange for con- ment was authorized in Baltimore on the tinuous flow in cultures on rotating tables, following day. but that was not actually accomplished In the early fall of 1909 I attended the until 1934, at the hands of Ch. Zinzadze Winnipeg meeting of the British Asso- and Karl A. Grossenbacher. ciation, after which Blakeslee and I made The present laboratory was completed a short trip by boat, north from Kenora, in the early winter of 1911-1912. My on the Lake of the Woods, and I then personal study throughout much of the became a resident of Baltimore. At the preceding year had been devoted to the Winnipeg meeting I became well ac- interior details of this building; all cases, quainted with Yapp, who had been study- tables, sinks, the electric wiring, and so ing evaporation influence on a spiraea on, had been specially designed, to pro- and other plants, also with Gwinne- duce a small, compact and very efficient Vaughan, Wager and others from abroad. working unit. In a great many of the The first year at The Johns Hopkins University I lectured in the old Biology 3Brown died at the end of that academic year,

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_l~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~I

A.> ...... i...... ,...... t FIG. 2. The Johns Hopkins University Laboratory of Plant Physiology at Homewood, Baltimore, Maryland.

arrangements I was aided by students, far as the bridge, at Manitou Rapids, on most of whom were good mechanics. the Nelson River-and we had the bene- The laboratory has served its purpose fit of transportation aid from *the men excellently for a quarter of a century. in charge of construction and fire inspec- For several years after coming to this tion. Without any guide, we traveled University I returned to the Desert Lab- slowly on lakes and rivers from Picotinay oratory each summer for a period of to Split Lake Post and then ascended the three or four months, continuing along Nelson past Grand Rapids to Manitou lines of study that had been opened in Rapids, whence we returned by railway. my Tucson period. Each of these sum- On that trip we gained an excellent gen- mers I had, through small grants of funds eral knowledge of the vegetational con- from the Carnegie Institution of Wash- ditions of that general region. ington, the help of one or more assistants After the new railway had been com- -young men who were about to receive, pleted to Churchill, Mrs. Livingston and or had just received, their Ph.D. degrees, I visited that place, in the summer of either here or elsewhere. Following is a 1935, where we lived for a week in a list of these men, to all of whom I owe a room in Mrs. Riddock's woodshed and great deal: W. H. Brown, J. S. Cald- explored the nearby region, with help well, E. M. Harvey, H. E. Pulling, J. W. from the officials of the new grain port. Shive, A. L. Bakke, H. C. Sampson, E. Although I had for years a desire to visit S. Johnston, J. D. Wilson, L. J. Pessin, the country beyond the northern limit of M. Fraps. forest, I never really passed beyond that The summer of 1915 was spent in the limit, which is apparently not far north region between The Pas and Split Lake, of Churchill. My "farthest north" is the Manitoba, on a canoe trip, in company old Hudson Bay Post at Fort Churchill, with H. E. Pulling. The new Hudson across the river from the new town of Bay railroad was partly constructed-as Churchill. I was much impressed by

This content downloaded from 128.95.104.109 on Fri, 06 Apr 2018 01:22:33 UTC All use subject to http://about.jstor.org/terms July, 1948 CONVERSATIONAL AUTOBIOGRAPHICAL NOTES 241 seeing a large plant of the small white DOCTORS OF PHILOSOPHY IN PLANT lady's-slipper in full bloom (about July PHYSIOLOGY FROM THE JOHNS 6, 1935) at the side of the age-old trail HOPKINS UNIVERSITY, 19909-1939 on the north side of the Churchill River; (Date of degree in parentheses) we also were interested in the gardens of Coggeshall, Mary (1931) residents of The Pas, where, in early Jtily, Darrow, G. M. (1927) we saw the yellow cypripedium and Cy- Edwards, T. I. (1932) pripedium spectabile in full bloom, also Espino, R. B. (1919) the most vigorous plants of the perennial Fawcett, H. S. (1918) delphinium I have ever seen anywhere. Free, E. E. (1917) On that second trip to the North we Gericke, W. F. (1922) Haasis, F. W. (1928) motored from Baltimore to Winnipeg, Hawkins, L. A. (1913) attending the Minneapolis meeting of the Hildebrandt, F. M. (1917) science societies on our way, and, return- Hutchins, L. M. (1924) ing from The Pas, we motored from Johnson, E. S. (1917) Winnipeg to Duluth, took a lake steamer Lawrence, D. B. (1936) to Sault Ste. Marie and then motored LeCompte, S. B., Jr. (1939) back to Baltimore from that point, stop- Mack, W. B. (1929) ping at Mackinac Island and other places. Marshall, Robert (1930) With Mrs. Livingston I spent the sum- Matz, Julius (1932) mer of 1921 at the Desert Laboratory, McCall, A. G. (1916) continuing earlier studies with Forrest McLean, F. T. (1915) Shreve. The book on climates and vege- Norem, W. L. (1936) tational types of the United States, which Pardo, J. H. (1932) had occupied Shreve and me for a period Shive, J. W. (1915) of ten years, was published in that year. Swingle, Charles F. (1927) The summers of 1922 and 1926 were Tang, Pei-Sung (1930) spent at the Desert Laboratory, but I Tottingham, William E. (1917) have not been there since. Trelease, S. F. (1917) To complete this survey I add below Veerhoff, 0. L. (1937) a list of those who have received the Veihmeyer, F. J. (1927) Doctor's degree with me at The Johns Verner, Leif (1934) Hopkins University. Wilson, J. D. (1926)

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