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Biographical Memoirs: V.59 (1990)

Chapter: William Henry Chandler

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Suggested Citation:"William Henry Chandler." National Academy of Sciences. 1990. Biographical Memoirs: V.59. Washington, DC: The National Academies Press. doi: 10.17226/1652.
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WILLIAM HENRY CHANDLER July 31, 1878{)ctober 29, 1970 BY JACOB B. BIALE B`` ECAUSE OF THIS RESERVE OF DORMANT BUDS, said W. H. Chandler, lecturing during the dark days of World War Il. "a tree is more clependable in a destructive world. It can be broken to pieces pretty badly and will grow new parts to replace the lost ones" (1944,1~. Trees with buds at rest, keeping the secrets of dormancy; trees with buds bursting to bloom and to fruit; trees of dif- ferent climates and of varier! behavior fascinated Chandler and served as his dependable companions throughout a long, productive, and humane life. Delving into their complex functioning, he unraveled the story of their response to in- ternal and external environment. Esteemecl worIdwicle for transforming horticulture from an art into a science, he— with his reservations about the validity of classifying horti- culture or agriculture as distinct sciences wouIct surely have rejected any such claim. But his original research papers and books, filled with knowledge and deep insight, continue to · · · · · . bring norm ~nternat~ona recognition. In addition to advancing the field of horticulture gener- ally, Chandler helped elucidate the mechanism by which frost kills plant tissue. He was the cocliscoverer of the fact that zinc deficiency causes a number of physiological disorders, in- cluding little leaf and mottle leaf. He introcluced a system of 87

88 BIOGRAPHICAL MEMOIRS pruning that resulted in maximum yield. He developer! hy- brids of temperate zone trees that grow, flower, and produce fruits satisfactorily in climates with mild winters. The university community, experiment station workers, and extension staff all valued ChandIer's ideas on research, teaching, and communicating results. He inspired promising investigators, then helped place them where they could con- tribute the most to horticulture and plant physiology. Anyone who had the good fortune to know him—whether profes- sionally or socially—was left with the impression of a man of sturdy character, mild manner, and no pretensions. His con- victions were strong, yet he was open to others' views. He was cultured, appreciating history, poetry and novels. Not blind to human shortcomings, he yet had an idealistic trust in the future of mankind. EARLY LIFE AND EDUCATION Bill ChancIler, the oiliest of eight children, was born in Butler, Missouri, in a little log house where the dog went in ant! out freely through the open door. Many years later he recalled that, during his childhood, all eight children slept in a single room in trundle beds that were stored away under larger beds cluring the clay. His father, who came from the hill country bordering Virginia and Tennessee, clislikecl farming and often allowecl weeds to displace planter] crops. When Bill was ten years old, the family moved to a somewhat larger house and smaller farm, incurring a large debt. They lost the property three years later, and from then on the family was forced to live on rented farms. From the age of fifteen, the responsibility for maintaining his family through farming rested on Bill with the help of a younger brother. Seriously restricted! in the time he could devote to school- ing, Bill attended the country school only during the six

WILLIAM HENRY CHANDLER 89 months of autumn and winter. The remainder of the year he worked full-time on the farm. At eighteen he went to stay with his uncle at the county seat, where he studies! for two semiannual periods at the Academy. From IS98 to 1901 he taught in a single-room country school, while aspiring at the same time to study farming at the University of Missouri Col- lege of Agriculture. When he clivulged his ambitions to his uncles, who were successful farmers, they ridiculed the young dreamer. "It isn't what ~ don't know that loses me money," one toIc! him. "It's what ~ know and don't do." Chandler disregarded the advice of his relatives and en- rolled in agriculture at the University in the fall of 1901. The five-year course led to the B.S. degree in 1905, and a year later he received the M.S. degree. Partly clue to the influence of Dr. I. C. Whitten, then head of the Department, he spe- cializect in tree horticulture, though in later years he re- grettecl that the program of study had not included required courses in physics ant! chemistry. As a student, Chandler was inspired by the teaching of plant physiologist B. M. Duggar. For his doctoral dissertation topic he elected to study the killing of plant tissue by low temperature, a major problem in agriculture in Missouri as well as in many other regions, which continued to interest him throughout later appointments as assistant ~1906-1908), instructor ~~908-l909), and assistant professor ~~9~0-l9~3) in horticulture. Due to a technical regulation, he was not of- ficially awarded the Ph.D. until 1914, when he was no longer afliliatecl with the University of Missouri. In 1913, ChanctIer was invited to join the faculty of the College of Agriculture at Cornell University as professor of Homology. Better pay and research support, the presence of I.iberty Hyde Bailey as clean of the College, and the greater distinction of the University made the over extremely attrac- tive, and he accepted. Once there, he found that the climatic

go BIOGRAPHICAL MEMOIRS conditions and the widespread growing of apples in New York State stimulated his interest in winter injury to fruit trees, and he extended his observations to the relation of winter frost damage to growth responses during the preced- ing summer. ' Local farmers cooperated willingly with the research and extension staff of Cornell's agricultural experiment station. Yet not everyone was equally enthusiastic about the program. "It's the farmer's conservatism that saves him," a skeptical Dean Bailey was reported to say. "If he'd done everything that you Ethe research men] recommend, he'd be ruined." Chandler shared Bailey's respect for the innate intelli- gence and good sense of the farmer. Working on field plots with New York growers, he found their attitude to farm life more wholesome than that of farmers in Missouri, so that the area remained relatively free of land speculation and real estate promotion. "You could not buy a farm at any price," he remarked at the time, "from a man who had a son to take his place." I~ike Bailey, too, he was skeptical about the quality of knowledge imparted by teachers of agriculture and the wor- thiness of certain agriculture research projects. L. H. MacDaniels, a Cornell graduate student at that time, re- ported that when Chandler arrived he was assigned to teach a course in the culture of nut trees that a number of football players took to lighten their load. After delivering a half- dozen lectures, he dismissed the class for the rest of the se- mester, saying that he had covered all that was known about the subject that was backed by evidence. Chandler insisted that the Homology program be related to plant physiology and the basic sciences, arguing that prep- aration for trees research shouIc! lead to a Ph.D. in plant physiology or in another related field that could serve as a background for horticulture. He often directed his graduate

WILLIAM HENRY CHANDLER 9 students to study uncler other professors, ant! though many investigators credit him for inspiring and directing their horticultural or physiological research chaired, in fact, only one cloctoral committee, that of A. I. Heinicke. During his decade at Cornell, Chandler chaired the De- partment of Pomology from 1915 to 1920 and, as vice- clirector for research, administered research funds from 1920 to 1923. This last task, at times frustrating because of the limited funds supporting a number of meritorious pro- jects, allowed him to broaden his contacts with his colleagues. He enjoyed his clearings with members of the general faculty on campus and life in the small, charming community of Ithaca. During this period, he also established his professional standing as the pomologist best able to analyze and under- stand the complex responses of fruit trees. This ability found its fullest expression in Fruit Grozo~ng, a textbook written and reviser! with great care ant} precision during his Cornell years, though published after he left there permanently for the West. In 1922, ChancIler was invited to tour various regions of California in connection with the dedication of a building of the University of California at Davis. Once there, he observed a wealth of horticultural problems that did not exist in New York, where fruit trees had grown for hundreds of years and many of the intricacies of their culture were known. Califor- nia, on the other hand, with its great range of climatic zones and wide spectrum of horticultural materials, was unique. In aciclition to the innate interest to an agriculturalist, C. B. Hutchison, an administrator in the College of Agriculture at Davis who had been ChancIler's associate at Missouri ant! Cornell, also player! a major role in his decision to transfer. Chandler came to California in 1923 as professor of pom- ology and chairman of the Department at both Berkeley and

92 . BIOGRAPHICAL MEMOIRS Davis, with headquarters in Berkeley. He later considered his fifteen at Berkeley highly, both professionally and personally, crediting his accomplishments in part to D. R. Hoagland, professor and chairman of the Division of Plant Nutrition. Hoagland's Division was noted for its research on the nu- tritional requirements of plants, and especially their need for trace nutrients: copper, zinc, molybdenum, manganese, and boron. Using special laboratory apparatus free of contami- nating elements, the Division staff developed a procedure for purifying chemicals to a high degree. In this atmosphere, Chandler investigated physiological disorders known as "little leaf" in peaches, "rosette" in apples and pears, and "mottle leaf" in citrus. His training in both horticulture and plant chemistry enabled him to identify a zinc deficiency as the cause of all of these disorders, thereby solving a problem that had baffled fruit growers since the beginning of the century. Chandler viewer! these zinc- deficiency studies as the most significant economic and scien- tific contribution of his career. He attributed his gratifying results to the combined efforts of his team members, whose diverse talents allowed them to focus on the problem from different angles, and to methodical experimentation using acivancec} procedures of purification and analysis. As a result of this cooperative venture, Chandler and Hoagland established a long-lasting friendship. They shared similar outlooks on research ant! university affairs and act- vocated harmonious interaction between applied and basic research. Both men had unusual personal qualities that in- spired those students ant! colleagues who had the good for- tune to be associated with them. In 193S, with the zinc work partially completed, Chandler was persuader! to accept the assistant deanship of the Uni- versity of California's College of Agriculture and to establish his headquarters on the Los Angeles campus of the Univer-

WILLIAM HENRY CHANDLER 93 sity. Administrative duties held no great attraction for him, but he yielded to the urgent pleas of C. B. Hutchison, then statewide dean of the College. As assistant dean, ChandIer's function was to harmonize relations between the Los Angeles and Riverside Depart- ments of the College and to strengthen UCLAs program in plant science. He also identified profitable directions for re- search in plant biology within the constraints imposed by field work on a campus in an urban setting. He focused on studies not requiring much land that could be conducted in greenhouses and in laboratories, and on plants with rapid growth rates, as the most suitable for graduate thesis work. He was, consequently, instrumental in establishing a Depart- ment of Floriculture and Ornamental Horticulture at UCLA. Knowing, from past experience, the benefits of aciminis- trative association between botany and agriculture, he fur- ther maple a special effort to transfer the Botany Department from the College of Letters and Sciences to the College of Agriculture. This action was later credited! with enriching UCLAs offerings in plant science, particularly at the gradu- ate level. Arranging this transfer was ChandIer's last major administrative act before he relinquished the deanship in 1943. He continued on at UCLA as a professor of horticul- ture until he officially retiree! in 1948. During retirement, Chandler thoroughly revised his two textbooks, Deciduous Orchards and Evergreen Orchards. To col- lect source materials for his books he traveled' to the West Indies, Trinidad, and Central America. UCLAs unofficial acI- visor for campus ~nclscaping, he maintained his interest in plant physiology ant! regularly attendee! seminars. In 1966, the Chandlers moved from Beverly Hills to Berkeley so that they could live closer to their three daugh- ters. In November 1969, he suffered a mild stroke and a year later ctied at the age of 92.

94 BIOGRAPHICAL MEMOIRS Chandler and his wife of sixty-three years, Nancy Caroline, were married in 1905 when he was starting his graduate studies at Missouri. The Chandler home exuded a spirit of tranquility, hospitality, and good comradeship. In his affection for his wife, Chandler named a wisteria after her and dedicated to her several of his books. Mrs. Chandler died in Berkeley in ~ 968. Their son, William Lewis (wife Eleanor), a microbiologist, established his home in Altadena. Their daughters, Carolyn Geraldine Cruess and Ruth Steele I.ewis, live in Berkeley, and Mary Martha Honeychurch has her home in Orinda, California. Chandler is survived by four children, eleven grandchildren, and ten great-grand-children. William Henry Chandler was awarded many honors dur- ing his lifetime. He was elected president of the American Society for Horticultural Science in ~ 92 I, member of the Na- tional Academy of Sciences in 1943, and Faculty Research Lecturer at UCLA in 1944. He won the Wilder Medal of the American Pomological Society in 1948 and, in the same year, was named one of three outstanding American horticultur- ists by the American Fruit Grower magazine. The American Society of Plant Physiologists bestowed on him the Charles Barnes Life Membership in 1951. In 1949 he received the honorary LL.D. degree from UCLA. He held membership in the American Association for the Advancement of Science, American Society for Horticultural Science, American Society of Plant Physiologists, Botanical Society of America, and Sigma Xi. TREES IN TWO CLIMATES On March 2l, 1944, four years before his retirement at the age of seventy, Professor Chandler delivered a talk on this subject as the annual UCLA Faculty Research Lecture (1945,2~. By that time, he had spent two decades in the mild

WILLIAM HENRY CHANDLER . . , 95 climate of California, with the last six years in the subtropical environment of the southern region of the state. Growing conditions and responses of fruit trees in the West differed dramatically from those he had observed during the first two decades of his career in Missouri and New York. The time was ripe for him to summarize his rich experiences with fruit trees of various climatic zones and to analyze the effects of temperature on cellular events as the major factor determin- ing their growth. Death by Freezing Chandler was searching for the mechanism of cellular death by freezing. The killing of plant tissue by low temper- ature had been the subject of his dissertation at the University of Missouri, while in California he had been attracted to the problem of why certain fruit trees required these same chill- ing temperatures to grow. Shortly after transferring from Missouri to Cornell, Chandler began observing the response of deciduous fruit trees to extremely low temperatures. In the early morn- ing hours of January 14, 1914, the temperature of-34°F (-36.7°C) was recorded in an orchard! in upstate New York in which Northern Spy apples were grown. Several days ear- lier ice had begun to form at the outer surfaces of some cells. From his own research and the work of others, Chandler knew that the gradual lowering of temperature facilitated the movement of water from the interior of cells to the intercel- lular spaces where ice crystals were formed. He further clis- covered that, although water expands as it freezes, air in these spaces gave way to ice so that the frozen tree actually shrank. He estimated that seventy to eighty percent of the water in the tree was converted to ice and that a third or more of the weight of the above-ground portion of the tree was ice.

96 BIOGRAPHICAL MEMOIRS Microscopic observation showed, furthermore, that shrunken cells as protoplasm became a thin layer between flattened walls. The pressure of ice particles present in the intercellular spaces appeared to cause distortions of a mag- nitude that suggested severe injury. As judged from the lux- uriant growth during the spring and summer following the severe winter, however, this was not the case, and the tree's survival suggester} that these had been, in fact, the proper conditions for harclen~ng. Through field observations and laboratory tests Chandler discoverecl a decreasing order of resistance to freezing tem- peratures in the various tissues of hardened trees. Most re- sistent was cambium, which, when not well-hardened, turned out to be as sensitive as other tissues; then came bark, sap- wood, ant! pith. He further observed that above-ground por- tions of a tree were more resistant than roots; that flower buds, generally more sensitive than vegetative bucts, were less sensitive when trees were not fully mature; that resistance diminished in some species whose flower buds reached an advanced stage of ctifferentiation by the beginning of winter. With great precision, he described how frost resistance developecI, singling out two ways "maturing" and "hard- ening" deciduous trees and shrubs became resistant to cold. Maturing of wood ant! bucis begins after growth ceases in the summer. It is characterized by the accumulation of carbohydrates, decline of water content, increase in osmotic pressure, thickening of cell walls, and a marked drop in the succulence of newly formed tissues. At the end of the ma- turing process—the time of natural leaf abscission some deciduous trees can withstand temperatures of -l 7° to -25°C. Hardening of mature wood occurs with exposure to freez- ing or near-freezing temperatures, with immature wood re- quiring a longer time to harden. Once hardened, some va- rieties can withstand temperatures ten degrees lower. Even a relatively short warm period can undo this increased resis-

WILLIAM HENRY CHANDLER 97 tance, but it can be regained unless growth started during the warm spell with repeated exposure to low tempera- tures. Chandler found that internal tissue changes during hard- ening included increased osmotic pressure (from starch hy- drolysis) and greater holding capacity for unfrozen water at temperatures above the eutectic point. In some hardy spe- cies, vacuolar sap also contained colloids that held water against freezing and osmotic activity, while, in other species, expansion of cytoplasm and the consequent reduction of the vacuoles might accompany hardening. The most resistant, living cells in well-hardened decidu- ous wood turned out to be nonvacuolated, meristematic cells in leaf buds and cambium. These cells in hardened plants could survive more shrinkage and the loss of a larger pro- portion of their water to ice masses than could cells of un- hardened plants. The protoplasm of hardened cells, further- more, seemed less easily ruptured than that of unhardened cells. These extensive observations on the responses of plants and plant parts to temperature stress led Chandler to probe the fundamental question of how freezing kills plant tissue. Well aware of the variety of centuries-old opinions on the subject, he compiled a list of established facts regarding plant death by freezing. Foremost was the phenomenon of ice formation—in ten- der tissue primarily within cells and, in cold-resistant mate- rial, in intercellular spaces subjected to relatively slow tem- perature fall (the case during a normal cold wave). Rapid temperature drop, on the other hand, caused ice to form within the cells and raised the temperature at which death occurs. Ice formation and death occurred rapidly at killing temperatures, unlike other chemical changes, which were markedly suppressed under such conditions. Death by freezing can best be seen in thawed tissue, which

98 BIOGRAPHICAL MEMOIRS darkens and takes on a water-soaked appearance coupled with a rapid rate of evaporation. In most tissues the rate of thawing floes not influence the level of the killing tempera- ture, but some tissues show symptoms of death before thaw- ing begins. Ice formed in the plant is pure water, while the solution left in the cells is highly concentrated. Sap solutes tend to hold some water unfrozen at temperatures below the eutectic point. Such concentrated sap may be toxic to proto- plasm at room temperature but is a source of protection at freezing temperatures; it also often contains water-binding colloids. Chandler also noted that bacterial spores, seeds, and pollen grains in the proper state of dehydration could with- stanc! temperatures of liquid hydrogen ant! remain viable. Any explanation of the mechanism of freezing to cleath wouIc! have to account for these observations, as well as for supercooling as a means of protecting tissue from injury at freezing temperatures so long as ice formation clid not occur. After examining the various hypotheses concerning the mechanism of death by freezing (including disorganization of protoplasm through water loss and toxicity through con- centration of the sap), he arrived at the conclusion that plants were most probably killed by the pressure of the ice masses on plasma membranes. The Rest Period When he moved to California, ChancIler's concern with low temperature as a limiting factor in the growth of fruit trees took a different turn rather causing losses from freez- ing, low temperatures in fall and winter were necessary to some California plants if they were to develop normal shoot and flower buds the subsequent spring. In a subtropical as opposed to a harsh climate, the limiting factor for growing apples, pears, apricots, peaches, and plums was the absence of sufficient clays at moderately low temperatures to "break

WILLIAM HENRY CHANDLER 99 the rest, 7 a condition known in horticulture as the "chilling requirement." In three of his four textbooks ant! in several special papers Chandler described the phenomenon of the "rest period" precisely. In the spring a hormonal substance produced in the tip prevents newly former! buds along the shoot from growing. Early in the season, if this apical inhibitor is re- moved, these buds will grow. Later in the summer, however, the buds enter the rest period anct absence of the apical in- hibitor does not cause growth. Rest period is, therefore, the period when the plant, or a portion of the plant, will not grow even when temperature, moisture, anc! nutrient conditions are favorable for growth. It is different from "dormancy," a state of inactivity brought on by any cause. An apple tree, for example, might be said to be dormant in February because the temperature is too low for growth, or it might fad! to grow in December, not because of the temperature, but because it is in the rest period. In some fruit trees this rest period! is attained as early as five to seven weeks after the start of spring growth. In the warm winters typical of the coastal regions of California, on the other hancI, buds on some varieties of deciduous trees clo not grow until the micicIle of the following summer, and even then only a small percentage will grow. To demonstrate his point, ChancIler used the striking ex- ample of a Northern Spy apple tree in Berkeley that had experienced a rest period in which no buds grew for two seasons. Yet, ChancIler maintained, if the same tree had been put at 5°C in the fall of the first year, its rest period could have been reduced from two years to six months. Placing a number of branches of a cherry tree at 0°C for two months, he showed that their buds opened a month earlier than buds on the unchilled tree. In another experiment, he subjected peach trees to tem-

100 BIOGRAPHICAL MEMOIRS peratures ranging from-~° to 0°C for two and one-half months cluring the fall, then transferred! them to a warm greenhouse ~15°C), keeping control trees continuously at the higher temperature. The chilled buds grew as much in four- teen clays as the unchilled buds in 133 days. ChandIer's experiments showed that emergence from rest was a function of both temperature and time; that spring growth was more rapid when buds were previously subjected to temperatures of 5° to 10°C for fifty to sixty clays; that the more vigorous ant! later the growth during the preceding summer, the greater the chilling requirement. He found that insufficient chilling caused some buds to open before others, and many to fail to open altogether. Inadequate chilling, furthermore, affected flower buds as much as leaf buds, caus- ing many to fall off before they had fully opened. In some trees, flower initials died in the buds before opening, while apple and pear trees, whose buds are mixed (consisting of both flower ant! shoot initials), insufficient chilling led to the production of leafy shoot only, or of leafy shoots with a re- duced number of flowers. As for the biological role of the rest period, he pointed out that delay in spring bunching less- ened the danger from spring frosts and opening to occur in weather more favorable for pollination and fruit setting. Seeking the cause of the rest period in trees, Chandler suggested that a hormonal substance might be involved and cited changes in ether-extractable auxins in buds upon emer- gence from the rest. Treatments with rest-breaking sub- stances such as ethylene chIorhydrin tended to reduce the auxin levels in plant tissue. Fully acknowledging the lack of verifiable data, ChancIler advanced the idea that a bounc! form of auxin might be responsible for keeping buds from growing during the deep part of the rest, and that the rate of retardation of bud opening was determined by the balance between the bound and free forms.

WILLIAM HENRY CHANDLER FRUIT TREE NUTRITION THE ZINC STORY 101 Regarding the nutritional requirements for optimal growth and yield] in fruit trees, Professor ChandIer's investi- gations ranged from experiments with specific nutrients to analysis of a complex biological system dependent on min- erals derived from a highly variable medium. Before Chandler, experiment station researchers tendec! to concern themselves with annuals. In their orcharct-fertil- izer experiments, they appliecl different quantities and com- binations of required elements over a number of years, then analyzed the results statistically. Chandler questioned the reliability of field trials where experimenters seeking to minimize error increased the size of their samples, necessarily using larger anc! more variable soil plots. He also called attention to errors caused by such frequently overIookec} variables as bud variation, differences in the vigor of seedling stock growth, the cumulative effect of injuries sustained with age, an(1 in measuring growth and yield the number of branches with which a tree started. He pointed out that the outbreak of disease (as happened when mottle leaf blighted certain experimental citrus trees) could vitiate years of carefully penned fertilizer experimen- tation that depended on uniformity of plots to test differen- tial treatments. Cognizant of these clifficulties, Chandler designed a new approach to field testing with fruit trees. Shortly after he arrived in California, orchards in a variety of climatic zones both inland and along the entire Pacific Coast suffered great losses from a tree disease known since the beginning of the century. This disease, affecting both decicluous and ever- green trees (and walnuts and grapes as well), is called "little leaf" in stone fruits almond, apricot, cherry, peach, plum; "mottle leaf" in citrus; and "rosette" in apples and pears.

102 BIOGRAPHICAL MEMOIRS The disease is most dependably characterized by stiff, nar- row leaves about five percent of normal size that appear in the spring. Each of the small tufts, or rosettes, of these abnormally small leaves originates from a bud that wouIct normally produce a shoot. Leaves are also mottled, with yel- low streaks and splashes between veins, while the veins them- seIves, and some adjoining tissue, are green. These symptoms are most conspicuous in spring. Later in the season, healthy shoots may grow from buds lower on the branch. In severe cases, ctistortec! yellow leaves form even late in the summer. Fruit size in all species is reduced and, in some, the fruit is also strikingly clistorted. Moderately affected pome and stone fruits may live for many years producing fruit of inferior quality and yield. In some soils trees grow well for the first few years but then develop symptoms rapidly and die. Chandler undertook to study this problem together with two members of Berkeley's Plant Nutrition Division plant physiologist and soil chemist D. R. Hoagland and chemist P. L. Hibbard. Before starting trials of treatments he care- fully observed conditions in various districts of California. He sought out the experiences of farm advisors, extension spe- cialists, and orchardists. He noted that while trees in creep, well drained, sanely soils with low clay content were the most readily affecter! by the disease, in some regions little leaf also affected trees in loam soils. He paid special attention to or- chards on land formerly used as corrals for livestock. On these soils, with high nitrogen content, the disease was ram- pant. Quickly ruling out deficiencies of nitrogen, phosphorus, potassium, calcium, and magnesium, Chandler proceeded to test for iron. A preliminary mid-winter trial with large quan- tities of a commercial grade of ferrous suIphate resulted in normal leaves in summer. Chandler first thought the iron suIphate worked by re- ducing the alkalinity of the soil, but other pH-Iowering sub-

WILLIAM HENRY CHANDLER 103 stances such as suIphur had no effect. When he then applier! chemically pure ferrous suIphate, the results were equally negative, and it was immediately apparent that an impurity in the commercial grade of ferrous suIphate might account for its effectiveness. Chemical analysis showed that the sul- phate contained one percent of zinc and several other ele- ments in small amounts. Further tests with zinc suIphate gave positive results, though the amounts required varied widely and a broader range of trials seemed called for. Chandler decider! against concentrating his efforts in a single area, opting instead for a wide range of soils twenty- six locations in ten counties. Leaving several severely diseased trees in each locality as controls, he treated some 2,000 oth- ers. It soon became evident to him that the degree of correc- tion was a function of the solubility and dosage of the zinc compounds used. Yet extreme variability in the effectiveness of the treatment also suggested significant differences in the zinc suIphate. To find out whether zinc was essential to fruit tree nutri- tion or had a secondary, soil-related function (such as cor- recting for unclesirable flora), it was necessary to circumvent the soils and apply zinc directly to the trees. This Chandler accomplishecl in a variety of ways. He put dry zinc suIphate in gelatine capsules in holes in tree trunks, getting earlier, longer-lasting benefits than from soil treatments. He found that trees wouIc! absorb zinc from metallic zinc nails driven into the trunk or branches, and—though this treatment caused some injury to the wood- injured areas usually filled with callous tissue if the nails were not too close together. Trees cured of zinc deficiency symptoms by these direct methods, moreover, remained healthy for six years or more after a single application, though with certain citrus and stone-fruit trees, spraying trees with a zinc suIphate solution got the earliest beneficial results. Chandler favored the idea that zinc, a nutrient required

104 BIOGRAPHICAL MEMOIRS in minute amounts, acted as a catalyst for some biochemical process. In view of his observation that the demand! was greatest when respiration was likely to be most rapid, he sug- gestec} that this catalysis might be an essential step in the respiratory pathway. REFLECTIONS, CONVICTIONS, AND FAITH ~ cherish the privilege of having had Professor Chandler as my teacher and mentor during my student days at Berke- ley and as my colleague and frienc} after his move to UCLA. During the decade preceding his retirement, and for a con- siderable time thereafter, he expresse(1 many thoughts (often unorthodox) on matters within ant! outside his immediate professional interests. He was particularly concerned! with the position of the university in society, the role of the inves- tigator and teacher in agricultural schools, and the respon- sibilities of scientists—both as citizens and as members of the human race. Many of these opinions were delivered in speeches to meetings of faculty, students, extension workers, and fruit growers. Copies of Prof. ChandIer's speeches, which ~ was privileged to receive, serve as the main background for the comments in this section. To ChanctIer, work for an institution of higher learning where scholars joined together in the attempt to find truth was a great cause deserving of the highest loyalty. For loyalty to survive the confusing vicissitudes of life, he addecI, its ob- ject had to be too important to be blamed for failures. "I may serve my cause ill," he quoted the philosopher Josiah Royce. "I may conceive it erroneously. ~ may lose it in the thicket of world transient experience. My every human endeavor may involve a blunder. My mortal life may seem one long series of failures. But ~ know that my cause liveth." Chandler singled out universities as the greatest cooper- ative enterprise the world has ever known, for the investi-

WILLIAM HENRY CHANDLER 105 gator—engaged in solving a problem of his choosing—col- laboratec! not only with his contemporaries but with generations of seekers of knowledge from the past as well. Chandler's own work, for instance, depended on that of those brave, "determined souls" of the Dark Ages who recorded unorthodox findings at their own peril. From this historical view of communication's significance to science, Chandler particularly emphasized precise and careful reporting as essential to the great cooperative enter- prise of learning. He remarked that, as methodology be- comes more refined and thinking more rigorous, the presen- tation of ciata becomes more concise. "Where opinions are publisher! in the most words and where there is most argu- ment," he observed, there is the greatest accumulation of ig- norance most likely to be found. Chandler admired the brief, precise reports targeted to a specific audience and unencumbered by lengthy discus- sions common to the physical sciences. By contrast, agri- cultural experimentalists often failed to address their most interested reaclers, being more concerned about a paper's reception in peripheral scientific fields than its usefulness to other horticulturists. They published too often, he main- tained, in too much detail, included exhaustive reviews of the literature, and got lost in warily theoretical explanations. He particularly objected to experimental stations publish- ing special editions of technical papers, which tended to be lengthy, cumbersome, costly, of limited reader access, ant} poorly edited. He favored, rather, publication in society jour- nals, which hac! a wide circulation and were reviewed by peers capable of independent judgment. The issue of priority of authorship in scientific publishing also failed to impress Chandler. Since, he said, investigators were rarely responsible for the same data in a paper, priority played little role in their professional standing among their

106 BIOGRAPHICAL MEMOIRS peers. For all that he aclmonished his colleagues to be espe- cially vigilant in fully crediting their research associates in- cluding assistants and graduate students for their contri- butions. Finally, Chandler cautioned agricultural experiment sta- tions against possessiveness with regard to research projects. While major responsibility ant! funding should go to the best qualified investigators, he contended, others should be en- couraged to test promising leads. Researchers shouIct also welcome the cooperation of county farm advisors and extension specialists. These people, who knew local conditions best, could help by testing labo- ratory results on the farm or arranging for the use of outside growers' field plots. Chandler further adviser! laboratory people to present their findings to farmers through agricultural agents rather than direct contact. He saw no discredit in a researcher at- tending so diligently to his research that he had no time to learn applied aspects of the work necessary for giving the best practical acivice. He himself had intimate personal knowledge of working with trees that yielded publishable data but rarely ant! practical advice for growers even less. In real life, according to Chandler, farmers "harassed by a whole range of nature's reactions" posed chastening ques- tions to horticultural researchers. Yet attempts to solve a problem with fruit trees required the convergence of several disciplines, and those who "discovered" a practical remedy might be no more deserving of credit than the many earlier researchers whose earlier experiences had suggested the so- lution. It was often, he contended, a matter of good fortune to come to a problem when just a few added experiences were needed to supply the solution. In a dinner talk delivered in 1941 to the western section of the American Society for Horticultural Science (1942,2), Chandler reflected on the merits of studying plants.

WILLIAM HENRY CHANDLER 107 "The material we work with has character," he stated, con- siclering himself fortunate in both the trees and the people with whom he had worked. Citing literary references to the sturdy character and earthy beauty of the apple tree, he went on to say that to him fruit and vegetables were not merely a mass of materials but a collection of individuals. Trees and plants, furthermore, were not merely objects worthy of ad- miration, they also exerted an influence on the behavior of the people who tended them. "As the apple tree is among the trees of the wood," he quoted from the Song of Songs, "so is my beloved among the sons. ~ sat down under his shadow and his fruit was sweet to me." Chandler suggester} that Thomas ~e~erson's ability to en- dure the rigors and criticism of political life might be attrib- uted to the comfort and encouragement he derived from the extensive time he spent on his farm working with his trees. Chandler discovered that, in the Scandinavian countries per- haps more than anywhere else, the beauty of flowers and trees, both ornamental and fruit-bearing, was associates! with efforts for the general good that he himself called "effective human love." When he visited Denmark he was told that pref- erence in police recruitment was given to horticultural school graduates who were known for their even tempers. In Sweden, trained agriculturalists were put in charge of urban housing projects in recognition of the importance of plants for social contentment. Chandler expressed his faith in the Tree of Knowleclge anct in humankind in the following words: "The God of Nature reveals his laws, I believe, very rarely to the propa- gandist or to the pompous, or even to the merely zealous, but rather to him who trains diligently in the technique and the records of a system of knowledge, who records his own observations clearly and briefly for the benefit of all workers, who reviews and reorganizes his knowledge fre- quently in the light of new discoveries, who consults as frequently as pos- sible with workers in his field and related fields, hoping for a vision that

108 BIOGRAPHICAL MEMOIRS points to a safe advance in human welfare, and who is meek enough to see a vision unobscured by projects of himself. "Truth discovered by research enters into the lives of the people and its beauty is recorded for all time in literature and art; the drudgery of the laboratory today becomes beauty in the soul of humanity tomorrow. Be- cause our discoveries enter the basic part, the masonry of the soul of hu- manity, we should report them with modest reverence. We want a foun- dation not of spongy lava thrown up by workers each anxious to strut about the biggest pile, even if it is the trashiest but rather of dressed stone, each piece placed carefully where it belongs in the structure. "We can have faith in the triumph of good in humanity in spite of the evil we know exists; in fact, life is richer because of the imperfections in it. I liked the part in one of George Bernard Shaw's plays where the Bishop advised people always to give the devil a chance to state his case, for I have come to believe that the devil has a rather strong case. He stands for self- ishness, and a degree of selfishness is socially necessary for the most dili- gent care of each individual. Furthermore, we need something to struggle against. If in man the instinct of self-preservation, selfishness, and the group instinct, human love, were so nicely balanced that there would be no conflict, so that we could just enjoy our goodness comfortably like pigs enjoy their fatness, would life be very interesting? "Perhaps the richest part of life is knowledge of the great people that have been in it. If selfishness were no problem, we should never have heard of the thundering righteousness of the Hebrew prophets or of Jesus; they would have been just other nicely balanced men. And what use would we have had for Thomas Jefferson or Lincoln or Horace Greeley, or for the thousands of supporters who made their work possible, dormant-bud Jef- fers~ns and Lincolns and Greeleys out among the people? The only changes I want to see in man are those he makes himself struggling upward in response to the soul of humanity and his group instinct. "The emblem of my faith is the tree and its system of dormant buds that can grow only if buds that happen to be in more favorable positions for growth are removed. If ends of branches are removed, shoots will grow out of the older wood from buds that have grown each year only enough to keep their tips in the bark. Then when their opportunity comes, they grow vigorously. Because of this reserve of dormant buds a tree is more dependable in a destructive world. It can be broken to pieces pretty badly and will grow new parts to replace the lost ones. "This condition in the tree symbolizes my faith in humanity, my con-

WILLIAM HENRY CHANDLER 109 victim that society, at least in those countries that have been able to n~ain- tain order without despotism most of the time, cannot long change in any direction except toward a richer life for the average person: For I know there are many dormant buds in human society also." William ChancIler shared his sturdy faith in humanity with the renowned fellow-botanist Liberty Hyde Bailey. Both lived to a ripe and productive old age, and ~ inclucle, in con- clusion, a stanza from "My Great Oak Tree," a poem by Bailey that Chandler greatly cherished: "Ant! thrice since then far over the sea Have ~ journeyed alone to my old oak tree And silently sat in its brotherly shade Anct ~ felt no longer alone and afraid; ~ was filled with strength of its brawny-ribbed bole And the leaves sIow-whispered their peace in my soul."

0 BIOGRAPHICAL MEMOIRS SELECTED BIBLIOGRAPHY 1904 Result of girdling peach trees. West. Fruit Grower, 15: 191. 1907 The winter killing of peach buds as influenced by previous treat- ment. Mo. Agric. Exp. Stn. Bull., 74:1-47. 1908 Hardiness of peach buds, blossoms and young fruits as influenced by the care of the orchard. Mo. Agric. Exp. Stn. Circ.,31: 1-31. Instructions for spraying. Mo. Agric. Exp. Stn. Circ., 34:1-16. 1911 Cooperation among fruit growers. Mo. Agric. Exp. Stn. Bull., 97:3-58. 1912 Combating orchard and garden enemies. Mo. Agric. Exp. Stn. Bull., 102:237-90. 1913 The killing of plant tissue by low temperature. Mo. Agric. Exp. Stn. Res. Bull., 8:141-309. Commercial fertilizers for strawberries. Mo. Agric. Exp. Stn. Bull., 113 :297-305. 1914 Sap studies with horticultural plants. Mo. Agric. Exp. Stn. Res. Bull., 14:491-553. Some problems connected with killing by low temperature. Proc. Am. Soc. Hortic. Sci., 11:56-63. Osmotic relationships and incipient drying with apples. Proc. Am. Soc. Hortic. Sci., 11: 112-16. 1915 Some peculiar forms of winter injury in New York State during the winter of 1914-15. Proc. Am. Soc. Hortic. Sci., 12: 118-21.

WILLIAM HENRY CHANDLER 1916 111 Influence of low temperature on fruit growing in New York State. Cornell Countryman, 13:373-77. 1918 Influence of low temperature on fruit growing in New York State. N.Y. State Fruit Grow. Assoc. Prod., 16:186-94. Winter injury in New York State during 1917-18. Proc. Am. Soc. Hortic. Sci., 15: 18-24. 1919 Pollination. Ind. Hortic. Sci. Trans. for 1918: 11-120, 173 -75. The effect of the cold winter of 1917-18 on the fruit industry. Ind. Hortic. Sci. Trans. for 1918:91-103. Pruning its effect on production. Ind. Hortic. Sci. Trans. for 1918:137-45, 156-61. Some results as to the response of fruit trees to pruning. Proc. Am. Soc. Hortic. Sci., 16:88-101. 1920 Winter injury to fruit trees. Mass. Dep. Agric. Circ., 24:11. Some preliminary results from pruning experiments. N.Y. State Hortic. Soc. Proc., 2:77-84. Some responses of bush fruits to fertilizers. Proc. Am. Soc. Hortic. Sci., 17:201-4. 1921 The trend of research in homology. Proc. Am. Soc. Hortic. Sci., 18:233-40. 1922 The outlook of agricultural research. (Address delivered at the dedication of the Dairy Industry and Horticulture buildings, University Farm, Davis: 24-37.) 1923 Results of some experiments in pruning fruit trees. N.Y. Agric. Exp. Stn. Cornell Bull., 415:5-74.

llY BIOGRAPHICAL MEMOIRS 1924 The advantages and disadvantages of organization and standard- ization in horticultural research. Proc. Am. Soc. Hortic. Sci., 21 :259-63. 1925 Fruit Growing. Boston: Houghton Mifflin Co. Polarity in the formation of scion roots. Proc. Am. Soc. Hortw. Sci., 22:218-22. With A. I. Heinicke. Some effects of fruiting on growth of grade vines. Proc. Am. Soc. Hortic. Sci., 22:74-80. 1926 an- Or - With A. I. Heinicke. The effect of fruiting on the growth of Old- enburg apple trees. Proc. Am. Soc. Hortic. Sci., 23:36-46. 1928 North American Orchards, Their Crops and Some of Their Problems. Phil- adelphia: Lea & Febiger. 1931 Freezing of pollen: evidence as to how freezing kills plant cells. Am. J. Bot., 18:892. With D. R. Hoagland and P. L. Hibbard. Little leaf or rosette of fruit trees. Proc. Am. Soc. Hortic. Sci., 28:556-60. 1932 With D. R. Hoagland and P. L. Hibbard. Little leaf or rosette of fruit trees. II. Effect of zinc and other treatments. Proc. Am. Soc. Hortic. Sci., 29:255-63. With D. R. Hoagland. Some effects of deficiencies of phosphate arid potassium on the growth and composition of fruit trees under controlled conditions. Proc. Am. Soc. Hortic. Sci., 29:267-71. 1933 With D. R. Hoagland and P. L. Hibbard. Little leaf or rosette of fruit trees. III. Proc. Am. Soc. Hortic. Sci., 30:70-86. With W. P. Tufts. Influence of the rest period on opening of bud

WILLIAM HENRY CHANDLER 113 of fruit trees in spring and on development of flower buds of peach trees. Proc. Am. Soc. Hortic. Sci., 30:180-86. 1934 The dry matter residue of trees and their products in proportion to leaf area. Proc. Am. Soc. Hortic. Sci., 31:39-56. With D. R. Hoagland and P. L. Hibbard. Little leaf or rosette of fruit trees. IV. Proc. Am. Soc. Hortic. Sci., 32: 11-19. 1935 With A. S. Hildreth. Evidence as to how freezing kills plant tissue. Proc. Am. Soc. Hortic. Sci., 33:27-35. With D. R. Hoagland and P. L. Hibbard. Little leaf or rosette of fruit trees. V. Effects of zinc on the growth of plants of various types in controlled soil and water culture experiments. Proc. Am. Soc. Hortic. Sci., 33: 131-41. 1936 With D. R. Hoagland and P. R. Stout. Little leaf or rosette of fruit trees. VI. Further experiments bearing on the cause of the dis- ease. Proc. Am. Soc. Hortic. Sci., 34:210-12. With M. H. Kimball, G. L. Philp, W. P. Tufts, and G. P. Weldon. Chilling requirements of opening of buds on deciduous or- chard trees and some other plants in California. Calif. Agric. Exp. Stn. Bull., 611:3-63. 1937 Zinc as a nutrient for plants. Bot. Gaz., 98:625-46. 1938 The winter chilling requirements of deciduous fruit trees. Blue Anchor, 5: 2-5. Our work. (Address to the Synapsis Club of the Citrus Experiment Station at Riverside, California, October 3, 1938, pp. 1-13.) Rolling of leaves on Oriental plum trees, apparently caused by cool summers. Proc. Am. Soc. Hortic. Sci., 26:259-60. 1940 Some problems of pruning, with special application to shade trees. In: Proc. Seventh Western Shade Tree Conference, pp. 50-64.

114 BIOGRAPHICAL MEMOIRS Teaching in a college of agriculture. (Address before the annual conference of the Agricultural Extension Service, January 2, 1940, pp.l-8.) 1942 Deciduous Orchards. Philadelphia: Lea & Febiger. Forty years of helping the farmer with knowledge. Science, 95:563-67. Sermons. (Address before the Western Section of the American Society for Horticultural Science, June 20, 1941.) Proc. Am. Soc. Hortic. Sci., 41:387-97. 1943 Some responses of trees in a few subtropical evergreen species to severe pruning. Proc. Am. Soc. Hortic. Sci., 42:646-51. 1944 Sturdy faith and dormant buds. (Address before joint meeting of the Synapsis Club, Citrus Experiment Station, and the Amer- ican Society for Horticultural Science. January 3, 1944, pp. 1-6.) 1945 Trees in two climates. (Faculty Research Lecture, University of Cali- fornia, Los Angeles, March 21, 1944. Univ. of Calif. Press, Berkeley and Los Angeles, pp. 1-22.) 1946 With D. R. Hoagland and I. C. Martin. Little leaf or rosette of fruit trees. VIII. Zinc and copper deficiency in corral soils. Proc. Am. Soc. Hortic. Sci., 47: 15 -19. With D. Appleman. Little leaf or rosette of fruit trees. IX. Attempt to produce corral injury with constituents of urine. Proc. Am. Soc. Hortic. Sci., 47:25. 1949 Pruning trials on wisteria vines. Proc. Soc. Hortic. Sci., 54:482-84. Evergreen Orchards. Philadelphia: Lea & Febiger.

WILLIAM HENRY CHANDLER 115 1951 Deciduous Orchards, 2d ed. Philadelphia: Lea & Febiger. With D. S. Brown. Deciduous orchards in California winters. Calif. Agric. Ext. Serv. Circ., 179:3-39. 1952 With R. D. Cornell. Pruning ornamental trees, shrubs, and vines. Calif. Agric. Ext. Serv. Circ., 183:1-44. 1954 Cold resistance in horticultural plants: A review. Proc. Am. Soc. Hortic. Sci., 64:552 - 72. 1955 Twenty-five years' progress in California fruit production. (Ad- dress delivered at University of California, Davis, October 29, 1955, pp. 1-15.) 1957 Deciduous Orchards, ad ed. Philadelphia: Lea & Febiger. 1958 Evergreen Orchards, 2d ed. Philadelphia: Lea & Febiger. 1959 Plant physiology and horticulture. (Prefatory chapter.) Annul Rev. Plant Physiol., 10: 1-12. 196 Some studies of rest in apple trees. Proc. Am. Soc. Hortic. Sci., 76:1-10. 1965 Reminiscences. Oral History Program. University of California, Los Angeles, pp. 1-39.

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Biographic Memoirs Volume 59 contains short biographies of deceased members of the National Academy of Sciences.

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