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STERLING BROWN HENDRICKS April 13, 1902-January 4, 1981 BY WARREN L. BUTLER AND CECIL H. WADLEIGH Freedom to inquire into the nature of things is a rewarding privilege granted to a few by a permissive society. Sterling Hendricks The Passing Scene, 1970 TERLl N G B RO WN H EN D R} C KS was born in Elysian ~Fields, Texas, a small village in the eastern part of the state. The family hac! creep roots in the Old South. When Texas seceded from the Union in TS6l, the area around Ely- sian Fields sent a company of men, known as the S. B. Hen- dricks Company, to the Confederate Army uncler the com- mancI of Colonel Sterling Brown Hendricks, Sterling's grandfather. The colonel, a native of Alabama, grew up and studied law in Mississippi ancT moved to Elysian Fields in iS43, where he became a merchant and a farmer. He was also a scholarly man with a large library of books on law, religion, and the classics. Sterling's father, Dr. James Gilchrist Hendricks, was born in Elysian Fields in iS54. He received medical degrees from Louisiana and Tulane universities in New OrIeans and, after interning at Bellevue Hospital in New York City, returned home to practice medicine. Sterling's mother, Martha Daisy (Gamblin) Hendricks, was born in Cactdo Parrish, Louisiana, in 1X73. She graduated from Mansfield Female College in Louisiana as valedictorian of her class. After graduation, she went to Elysian Fields to teach school and met Dr. lames Hen 181

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182 BIOGRAPHICAL MEMOIRS ciricks, who was then a widower. They were marries! in IS93 and had five children; Sterling was the fourth. Sterling received much of his early schooling from his mother. There was no high school in Elysian Fields (only a one-room school house), so Sterling lived with an aunt in Shreveport, Louisiana, during his high school years. Follow- ing his graduation, the family moved to Fayetteville, Arkan- sas, so that several of the children could attend the university there. Sterling graduated from the University of Arkansas in 1922 with a bachelor's degree in chemical engineering. He studier! geology anct chemistry at the graduate level at the University of Iowa in 1923 anc! received a master of science in chemistry from Kansas State University in 1924. Then, in the fall of 1924, he began his doctoral studies at the Califor- nia Institute of Technology. On entering Cal Tech, A. A. Noyes, the director of the Gates Chemical Laboratory, suggested to Sterling that he work on X-ray crystallography in the laboratory of Roscoe C. Dickinson. Dickinson, who four years earlier hac! received the first Ph.D. degree given by Cal Tech, was going to Europe that year, so Sterling worked with Linus Pauling, who tract arriver! in Dickinson's laboratory two years earlier to learn the techniques of X-ray crystallography. Thus began a close friendship that lasted until Sterling's death. Sterling receiver! his Ph.D. degree in 1926, with a major in chemistry and with minors in physics and mathematical physics. Sterling began his Ph.D. research with a reinvestigation of the structure of the minerals corundum, Al2O3, and he- matite, Fe2O3, which hac! been studied earlier by W. H. and W. L. Bragg. He confirmed that the positions previously as- signecI to the aluminum anti iron atoms were correct, but the positions for the oxygen atoms were not. The refined struc- ture provides! a clearer understancTing of the interatomic forces in these crystals. He also cletermined the structure of ~ 7 - OCR for page 180
STERLING BROWN HENDRICKS 183 sodium and potassium azicle, showing that the three nitro- gens were in a linear array rather than in a cyclic structure- as hac! been proposed by some chemists. His Ph.D. thesis also incluclec! the determination of the crystal structures of sev- eral cupric chloride dihydrates. He also worked jointly with Maurier L. Huggins, a postdoctoral fellow in the laboratory, on the structure of pentaerythritol, C(CH2OH)4. Sterling pointed out that the pyramidal structure that tract previously been proposed might be incorrect, ant! that another space group permitted a tetrahectral arrangement of the bonds around the central carbon atom. This latter structure was confirmed a decade later. He continued structure determinations of simple organic compounds cluring two postdoctoral years 1926--27, at the Geophysical Laboratory of the Carnegie Institution of Wash- ington, anc! 1927-28, at the Rockefeller Institute of Medical Research with work that macle important contributions to the chemistry of carbon compounds. In 192X he joined the Fixed Nitrogen Laboratory of the U.S. Department of Agri . culture. He was recruited by F. G. Cottrell, who hoped to benefit mankind by solving the problems of nitrogen fixa- tion. In later years Sterling would often speak of Cottrell. Cottrell, as the inventor of the electrostatic precipitator, clo- nated the returns from his patents to support research through grants from the Research Corporation. It was from Cottrell that Sterling gained an appreciation for practical ap- plications of scientific research. For Sterling, the highest goal of science was to achieve a solution to an important practical problem. In the years that followed, Sterling macle monumental contributions to mineralogy and the stucly of soils. His early Ph.D. research on corundum anc! hematite was followecT by studies of other minerals, including zircon, apatite, gypsum, kaolinite, anauxite, valentinite, alunite, the jarosites, clickite,

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84 BIOGRAPHICAL MEMOIRS halloysite, hydrated halloysite, talc, pyrophyllite, vermiculite, chlorite, montmorillionite, nacrite, cronstedite, glauconite, celladonite, gibbsite, endellite, and the micas. This work re- sulted in an extensive understanding of clays as important components of soils and of the structural basis of ion ex- change of charged groups-central to understanding soil fertility. He used his expertise in X-ray diffraction ant! math- ematical physics to determine the structure of phosphate fer- tilizers and also of bone. In terms of human welfare, one could make a strong case that Hendricks' most important research was in collaboration with soil scientists toward de- termining the structure of soil constituents. In 1930 Hen- dricks and Fry published the results of their research on soil colloids. This paper is now recognized as the most important elucidation of the nature and properties of soils ever pub- lished. A bit of history may be in order. In iS50 a Scottish chemist by the name of I. T. Way pub- lished a paper on the power of soil to absorb manure. He had allowed moderately dilute solutions of neutral salts to seep downward through soil columns. He collected the per- colate and found that its chemical composition was usually different from that of the applied solution. For example, when an ammonium chloride solution was applied, the per- colate contained little ammonium; the percolate was mostly calcium chloride. Way concluded that there was an interac tion between the applied solution and the soil particles. He erroneously concluded that the reaction was irreversible. The distinguished German chemist, tustus van Liebig, looked on Way's report with utter contempt and had no reservations in saying so. For the next three-quarters of a century a vigorous controversy prevailed among soil chemists; some supported Way and others Liebig. These arguments were settled for all time by the publication of the paper by Hendricks and Fry. By using X-ray diffraction procedure, they conclusively

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STERLING BROWN HENDRICKS 185 prover! the crystalline nature of colloidal clay with the prev- alence of negative charges that wouIcT absorb and desorb cat- ions. They showed that Way was on the right roacI. With the exponential increase taking place in world pop- ulation, with the prevalence of famine ant! malnutrition on this planet, and with the finite limitation on the extent of arable soils available for food production, this research by Hendricks and Fry was of inestimable value. These findings openec! the door to an exceedingly important unclerstancling of the chemistry involved in maintaining high potential in soil productivity, and in providing a valid chemical basis for the reclamation of the alkali soils of arid regions. In 1952 Sterling received the Arthur L. Day Meclal awarcled by the Geological Society of America for outstancT- ing work in physics and chemistry advancing the geological sciences. The citation stated: Sterling Hendricks, an able technician and a masterful and imaginative theoretician, has been in the forefront of those who have given us a ra- tional understanding of these most complex and most important minerals. His elucidation of the structure of layered minerals and his demonstrations of the dependence of clay mineral properties upon structural considera- tions have been outstanding. Not only has he provided specific data on the kaolin minerals and, with Ross, on the complex montmorillionite group, but he has at the same time developed fundamentals of broad application, as for example in his studies of the polymorphism of the micas and of the nature of the water layer, and in the determination of minerals with dis- ordered structure and of minerals with random layer sequences. He has never been content merely to explain the well-behaved growths in the mineral world, but has gone on to decipher for us some of nature's "mis- takes." Linus Pauling considers that Sterling's work on the clay min- erals was his most important contribution to knowledge. The work on soils and fertilizers also led to investigations of hydrogen bonds. Hendricks was among the first to use

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86 BIOGRAPHICAL MEMOIRS infrared spectroscopy for the study of molecular structure. Pimente! and McClellan wrote, some twenty-five years later in their book on the hydrogen bond, that this work provides ". . . the most sensitive, the most characteristic and one of the most informative manifestations of the H-bond. From this ho crown tats imm`~nc.e volilm`~ of world ~ ~.~ " Hendricks also became an expert in rauiochemistry, and he showed how fer tilizers tagged with radioactive phosphorus could be used to follow the uptake of phosphorus by the roots of plants. Of course, not all of his scientific endeavors were successful. Sterling tried to obtain a diffraction pattern from crystals of horse hemoglobin some five years before the first successful X-ray crystallographic studies of a protein were made in Cambridge, England. His attempts failed because the protein denatured as the specimen was dried for mounting. He also attempted to obtain a diffraction pattern of a chromosome before it-was known how nucleic acids could be separated in a native state. Thus, in the course of many successes he had some grand failures but even the failures pointed toward forthcoming spectacular successes in biology. Sterling's scientific career took an abrupt change in direc- tion in the early 1940s. A brief history of this period and the subsequent developments is appropriate since it was in these new areas of plant physiology and photobiology that his most ,_ . creative contributions to knowledge lie. In lY/U two scientists in the USDA, H. A. AlIaru and W. W. Garner, discovered that daylength was a critical factor in determining when during the course of the year a given species of plant would flower- a phenomenon which they called photoperiodism. By the middle 1930s the work on photoperiouism was being contin- ued in the USDA by H. A. Borthwick and M. W. Parker, primarily in studies of the flowering of short-day plants (plants that flowered on a short clay long night regime). In the early 1940s they sought out a fellow USDA employee,

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STERLING BROWN HENDRICKS 187 Hendricks, to (liscuss how they shouIcl proceed in their in- vestigation of the effects of light in photoperioclism. It was then known that brief irradiations with light given during the long nights wouIcT inhibit the flowering of short-cl ay plants. They realizect that they might be able to determine the action spectrum (that is, the effectiveness of different wavelengths of light) for this inhibitory effect of light on flow- ering, and they agreed to pool their scientific talents toward this enct. World War I} intervened, anct it was not until 1944 that they began their collaboration. The key to the early successes of this work lay in the ex- perimental (resign of the action spectroscopy. A large spec- trograph was constructed using two exceptionally large glass prisms, which Hendricks had used previously for his infrared studies of hydrogen bonds, and a large second-hand carbon arc lamp like those used in theatres of the time. Absolute energy calibrations were made across the spectrum using a thermopile that was calibratect against a standard lamp. Of equal importance to the success of the work was the knowI- ecige of how action spectra should be measured. Hendricks unclerstood that it was essential to keep the irradiation peri- ods brief to extract the specific characteristics of the photo- reaction from the great complexity of the biological response, which might be assayer! some hours or days later. Borthwick and Parker provided the plants whose flowering response was sensitive to brief periods of irradiation, and Hendricks provicled the irradiation fields of large area, high spectral purity, and adequate intensity. Within a year they had an ac- tion spectrum for the floral inhibition of a short-clay plant, soybean, which showed a pronounced sensitivity to red light. Action spectra were then measured on a number of clif- ferent plants ant] on several different light-sensitive re- sponses, including the floral inhibition of other short-clay plants, the flowering of long-day plants where the night

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188 BIOGRAPHICAL MEMOIRS break irradiation incluced flowering, several growth re- sponses in etiolated plants grown from seed in clarkness, and the germination of lettuce seed. All of these investigations yielded essentially the same action spectrum, with a peak ac- tion in the red near 660 nm. It was concluded that the same pigment was involved in all of these responses. The experiments on seed germination were to provide key observations for elucidating the unusual photochemical properties of the pigment. It was known from earlier long term irradiation experiments (by Flint and McAlister) that rect light promoted the germination of lettuce seed. The USDA group expected to find their typical red action spec- trum for this response. Flint anc! McAlister had also re portect, however, that light in the near infrared region, just beyond the limits of vision, inhibited the germination but . .. . . . . ,% . the significance of the inhibitory effect of such wavelengths of light was generally unappreciated. The USDA group re- cliscoverecI the inhibitory effect of these far-recI wavelengths of light. They clemonstrated that seeds potentiated to maxi- mal germination by a brief irradiation with red light coup! be inhibited to minimal germination by a subsequent brief irradiation with far-reel light, and that these promotive ant! inhibitory effects were repeatedly reversible. The action spec- trum for the photoinhibition of germination showed a max imum at 730 nm. Hendricks cleducect from these experi- ments that the germination of lettuce seer! was controllecT by a pigment that existed in two interconvertible forms: a rec! absorbing form, PR' with an absorption maximum at 660 nm, and a far-red absorbing form, PFR' with an absorption maxi- mum at 730 nm. He concluclect that rect and far-red light caused transformations between the two forms: red PI ~ PER far-red

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STERLING BROWN HENDRICKS 189 After the unusual property of photoreversibility had been founc! in the germination response of lettuce seecT, the other rect-sensitive photoresponses were reexaminecl. They were found to show the same type of photoreversible antagonism between red and far-red light. The unique and unusual pi~- 1 ~ ~ ment system appeared to be ubiquitous in higher plants and to control a number of physiological responses. Hendricks was primarily responsible for the incisive in- sights that penetrated to the molecular level of the photocon- trol process. A given (legree of a physiological display would be used as an endpoint in a titration of responses versus in- cident energy. Whereas most plant physiologists of the time became lost in the great complexity of the biological system, Hendricks designed experiments in such a way that the com- plexities of the dark metabolism canceled out, leaving the pristine properties of the photoreaction to be revealed. The elegance of the approach culminated in a remarkable stucly. The physiological responses of seed germination and inter- nocle elongation of etiolatecl bean plants were titrated from both extremes of the reversible photoreaction, using rect and far-rect light. After making allowances for the light-scattering properties of the biological tissue and the quantum efficien- cies of the photoreactions, Hendricks calculated from the absolute energies required to achieve given degrees of re sponse ant! the first-orcler nature of the photoreactions- that the molar extinction coefficients of the two forms were between i04 and 105 liters mole- cm-~. He concluded on the basis of these high values for molar extinction coefficients and the absence of any visible color in albino mutants of bar- ley, whose growth responses were fully sensitive to recI and far-red light that the pigment system was functional at very low intracellular concentrations. The insight anct clarity of vision that allowed Hendricks to extract a molar extinction coefficient from a complex physiological display were char

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190 BIOGRAPHICAL MEMOIRS acteristic of his approach to science. Unfortunately, the paper reporting these findings was largely ignored. At the time, few workers in the field made the effort to follow the logic of the analysis. Hendricks had deduced the essential molecular proper- t~es of this remarkable pigment system from the physiological studies by the early 1950s. The absorption spectra of the two forms and the reversible nature of the photoreaction were known from the action spectroscopy. It was proposed from the absorption spectrum that the chromophore Of PR was an open-chain tetrapyrole, similar to that of allophycocyanin. It was even proposed, on the basis of the low intracellular con- centrations, that the pigment was an enzyme, and therefore a protein, and that Pi R was the active form of the enzyme. In addition to the photochemical properties, the physiological studies indicated that there was a slow dark transformation of P. ~ to Pa. This clerk transformation of Pl.^ back to PD was A_ _ [K -A - K r row 1' proposed to be the basis of the timing mechanism that en- abled photoperiodic plants to distinguish long nights from short nights. Nevertheless, most plant physiologists of the time did not believe that their subject matter was capable of revealing such molecular detail and, in the absence of direct proof, they were inclined to regard the pigment as a "pig- ment of the imagination." Sterling's group had the good fortune to join another group headed by Karl H. Norris, an agricultural engineer who had developed several spectrophotometers that could accommodate dense, light-scattering materials. From time to time Hendricks or H. W. Siegelman, a plant biochemist who was then associated with Borthwick and Hendricks, would examine these samples in the spectrophotometer for photo- reversible absorbance changes in the red and far-red regions of the spectrum. All of the initial attempts with plant tissues that were known to be sensitive to red and far-red light were

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STERLING BROWN HENDRICKS 203 With W. L. Hill and G. T. Faust. Polymorphism of phosphoric ox- ide. J. Am. Chem. Soc., 65:794-802. With L. Mitchell, G. T. Faust, and D. S. Reynolds. The mineralogy and genesis of hydroxylapatite. Am. Mineral., 28:356-71. With R. A. Nelson. Specific surface of some clay minerals, soils and soil colloids. Soil Sci., 56:285-96. 1944 Polymer chemistry of silicates, berates, and phosphates. I. Wash. Acad. Sci., 34:241-51. 1945 With W. L. Hill, D. S. Reynolds, and K. D. Jacob. Nutritive evalu- ation of defluorinated phosphates and other phosphorus sup- plements. I. Preparation and properties of the samples. J. As- soc. Off. Agric. Chem., 28:105-18. Base exchange of crystalline silicates. Ind. Eng. Chem., 37:625- 30. With W. H. Ross and J. Y. Yee. Properties of granular and mono- crystalline ammonium nitrate. Ind. Eng. Chem., 37:1079-83. With S. S. Goldich and R. A. Nelson. A portable differential ther- mal analysis unit for bauxite exploration. Econ. Geol., 41:64- 76. 1946 With M. W. Parker, H. A. Borthwick, and N. i. Scully. Action spec- trum for the photoperiodic control of floral initiation of short- day plants. Bot. Gaz., 108:1-26. With Sidney Gottlieb. Soil organic matter as related to newer con- cepts of lignin chemistry. Proc. Soil Sci. Soc. Am., 10:117-25. 1947 With W. L. Hill, E. J. Fox, and J. G. Cady. Acid pyro- and meta- phosphates produced by thermal decomposition of monocal- cium phosphate. Ind. Eng. Chem., 39: 1667-72. 1948 With L. A. Dean. Applications of phosphorus of mass thirty-two to problems of soil fertility and fertilizer utilization. Proc. Auburn Conf. on the Use of Radioactive Isotopes in Agricultural Re- search, Auburn, Ala., pp. 76-89.

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204 BIOGRAPHICAL MEMOIRS With H. A. Borthwick and M. W. Parker. Action spectrum for pho- toperiodic control of floral initiation of a long-day plant, wintex barley (Hordeum vulgare). Bot. Gaz., 110: 103-18. With L. A. Dean. Basic concepts of soil fertilizer studies with ra- dioactive phosphorus. Proc. Soil Sci. Soc. Ann., 12:98-100. With C. D. McAuliffe, N. S. Hall, and L. A. Dean. Exchange reac- tions between phosphates and soils: Hydroxylic surfaces of soil minerals. Proc. Soil Sci. Soc. Am., 12:119-23. 1949 With L. A. Dean. Radioactive tracers furnish new help in testing fertilizers. What's New in Crops and Soils, 1~61:14-16. With D. Burk, M. Korzenovsky, V. Schocken, and O. Warburg. The maximum efficiency of photosynthesis: A rediscovery. Science, 110:225-29. 1950 With O. Warburg, D. Burk, and V. Schocken. The quantum effi- ciency of photosynthesis. Biochim. Biophys. Acta, 4:335 -46. With O. Warburg, D. Burk, V. Schocken, and M. Korzenovsky. Does light inhibit the respiration of green cells? Arch. Bio- chem., 23~2~:331-33. With H. T. Hopkins and A. W. Specht. Growth and nutrient accu- mulation as controlled by oxygen supply to plant roots. Plant Physiol., 25: 193 -209. With R. S. Dyal. Total surface of clays in polar liquids as a charac- teristic index. Soil Sci., 69:421-32. With W. L. Hill. The nature of bone and phosphate rock. Proc. Natl. Acad. Sci. USA, 36:731-37. With M. W. Parker and H. A. Borthwick. Action spectrum for the photoperiodic control of floral initiation of the long-day plant, Hyoscyamus niger. Bot. Gaz., 111: 242-52. 1952 With R. S. Dyal. Formation of mixed layer minerals by potassium fixation in montmorillonite. Proc. Soil Sci. Soc. Am., 16:45-48. With M. W. Parker, H. A. Borthwick, and C. E. Jenner. Photo- periodic responses of plants and animals. Nature, 169:242-43. With L. Bramao, I. G. Cady, and M. Swerdlow. Criteria for the .

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STERLING BROWN HENDRICKS 205 characterization of kaolinite, halloysite, and a related mineral in clays and soils. Soil. Sci., 73:273-87. With L. A. Dean. Radioisotopes in soils research and plant nutri- tion. Annul Rev. Nucl. Sci., 1:597-610. With i. C. Brown. Enzymatic activities as indications of copper and iron deficiencies in plants. Plant Physiol., 27:651-60. With C. E. Hagen and V. V. tones. Ion sorption by isolated chlo- roplasts. Arch. Biochem. Biophys., 40:295-305. With H. A. Borthwick, M. W. Parker, E. H. Toole, and V. K. Toole. A reversible photoreaction controlling seed germination. Proc. Natl. Acad. Sci. USA, 38:662-66. With H. A. Borthwick and M. W. Parker. The reaction controlling floral initiation. Proc. Natl. Acad. Sci. USA, 38:929-34. Comments on the crystal chemistry of bone. In: Metabolic Interre- lations with Special Reference to Calcium, ed. E. C. Reifenstein, jr., pp. 185-212. New York: Josiah May, tr., Foundation. 1953 A discussion of photosynthesis. Science, 117:370-73. With H. A. Borthwick and M. W. Parker. Action spectra and pig- ment type for photoperiodic control of plants. Proc. 7th Int. Botanical Congr., Stockholm (1950), p. 785. With T. Tanada. Photoreversal of ultraviolet effects in soybean leaves. Am. }. Bot., 40:634-37. 1954 With H. A. Borthwick, E. H. Toole, and V. K. Toole. Action of light on lettuce seed germination. Bot. Gaz., 115:205 -25. With C. E. Hagen and H. A. Borthwick. Oxygen consumption of lettuce seed in relation to photo-control of germination. Bot. Gaz., 115:360-64. 1955 With E. H. Toole, V. K. Toole, and H. A. Borthwick. Interaction of temperature and light in germination of seeds. Plant Physiol., 30:473-78. With E. H. Toole, V. K. Toole, and H. A. Borthwick. Photocontrol of Lepidium seed germination. Plant Physiol., 30:15-21. Necessary, convenient, commonplace. (The nature of water: Its ba

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206 BIOGRAPHICAL MEMOIRS sic chemical and physical properties). In: U.S. Dept. Agric. Year- book of Agriculture; Water, pp. 9-14. With H. A. Borthwick. Photoresponsive growth. Growth, 19:149- 69. Screw dislocations and charge balance as factors of crystal growth. Am. Mineral., 40: 139-46. 1956 With H. A. Borthwick and R. J. Downs. Pigment conversion in the formative responses of plants to radiation. Proc. Natl. Acad. Sci. USA, 42:19-26. With E. Epstein. Uptake and transport of mineral nutrients in plant roots. Proc. Int. Conf. Peaceful Uses Atomic Energy, Ge- neva, 12:98 - 102. Control of growth and reproduction by light and darkness. Am. Sci., 44:229-47. With C. R. Swanson, V. K. Toole, and C. E. Hagen. Effect of 2,4- dichlorophenoxyacetic acid and other growth-regulators on the formation of a red pigment in Jerusalem artichoke tuber tissue. Plant Physiol., 31 :315 -16. With E. H. Toole, H. A. Borthwick, and V. K. Toole. Physiology of seed germination. Annul Rev. Plant Physiol., 7:299-324. With H. A. Borthwick. Photoperiodism in plants. In: Photoperiodism in Plants and Animals. Proc. Int. Photobiol. 1st Congr., Amster- dam:23-35. 1957 With I. D. Downs and H. A. Borthwick. Photoreversible control of elongation of pinto beans and other plants under normal con- ditions of growth. Bot. Gaz., 118: 199-208. With L. T. Alexander. The basis of fertility. In: U.S. Dept. Agric. Yearbook of Agriculture: Soil: 11-16. Clays. Agron. J., 49:632-36. With H. W. Siegelman. Photocontrol of anthocyanin formation in turnip and red-cabbage seedlings. Plant Physiol., 32:393-98. The clocks of life. Atlantic, 200~0ctober 41:111-15. 1958 With A. T. Jagendorf, M. Avron, and M. B. Evans. The action spectrum for photosynthetic phosphorylation by spinach chlo- roplasts. Plant Physiol., 33:72-73.

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STERLING BROWN HENDRICKS 207 With R. W. Siegelman. Photocontrol of anthocyanin synthesis in apple skin. Plant Physiol., 33:185-90. With A. San Pietro, }. Biovanelli, and F. E. Stolzenback. Action spectrum for triphosphopyridine nucleotide reduction by illu- minated chloroplasts. Science, 128:845. Photoperiodism. Agron. l., 50:724-29. 1959 With H. A. Borthwick. Photocontrol of plant development by the simultaneous excitations of two interconvertible pigments. Proc. Natl. Acad. Sci. USA, 45:344-49. The photoreaction and associated changes of plant photomorpho- genesis. In: Photoperiodism and Related Phenomena in Plants and Animals, ed. R. B. Withrow. Washington, D.C.: American Asso- ciation for the Advancement of Science, Publ. No. 55, pp. 423- 38. With H. A. Borthwick. Photocontrol of plant development by the simultaneous excitation of two interconvertible pigments. II. Theory and control of anthocyanin synthesis. Bot. Gaz., 120: 187-93. With E. H. Toole, V. K. Toole, and H. A. Borthwick. Photocontrol of plant development by the simultaneous excitations of two interconvertible pigments. III. Control of seed germination and axis elongation. Bot. Gaz., 121: 1-8. With H. W. Siegelman. Photocontrol of alcohol, aldehyde, and an- thocyanin production in apple skin. Plant Physiol., 33:409-13. With W. L. Butler, K. H. Norris, and H. W. Siegelman. Detection, assay, and preliminary purification of the pigment controlling photoresponsive development of plants. Proc. Natl. Acad. Sci. USA, 45: 1703-8. 1960 The photoreactions controlling Photoperiodism and related re- sponses. In: Symposium on Comparative Biochemistry of Photoreac- tive Pigments, pp. 303-21. New York: Academic Press. The use of radioisotopes in ion absorption by plants. Proc. Second Annul Texas Conf. on Utilization of Atomic Energy. Tex. Agric. Exp. Stn. Pub. R 72-60, pp. 42-46. With S. Nakayama and H. A. Borthwick. Failure of photoreversible control of flowering in Pharbitis nil. Bot. Gaz., 121~41:237-43.

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208 BIOGRAPHICAL MEMOIRS Basic research in plant nutrition. In: Research Outlook on Soil, Water, and Plant Nutrients. Natl. Acad. Sci. USA Publ. 785, pp. 1-5. With H. A. Borthwick. Photoperiodism in plants. Science, 132(3435): 1223-28. Rates of change of phytochrome as an essential factor determining photoperiodism in plants. Cold Spring Harbor Symp. Quant. Biol., 25:245 -48. .~ a- r--~ With T. E. Leggett. Phosphate and salt uptake by baker's yeast. Na- ture, 183~47531:862 - 63. 1961 With V. K. Toole, E. H. Toole, H. A. Borthwick, and A. G. Snow, in Responses of seeds of Pinus virginiana to light. Plant Physiol., 36(3):285-90. With H. A. Borthwick and S. Nakayama. Failure of reversibility of the photoreaction controlling plant growth. In: Proc. 3rd Int. Congr. on Photobiol.:394-98. 1962 With F. C. Jackson and B. M. Vasta. Phosphorylation by barley root mitochondria and phosphate absorption by barley roots. Plant Physiol., 37( 1 ): 8-1 7. Progress in knowledge of soils. Span, 5~21:84-87. With J. G. Cady and K. W. Flach. Petrographic studies of mineral translocation in soils. Trans. Int. Soil Conf., Comm. IV and V (New Zealand) A 1 (Wellington), p. 7. 1963 Metabolic control of timing. Science, 141~35751:21-27. With H. A. Borthwick. Control of plant growth by light. In: Envi- ronmental Control of Plant Growth, pp. 233-63. New York: Aca- demic Press. With W. L. Butler and H. W. Siegelman. A reversible photoreaction regulating plant growth. i. Physiol. Chem., 66:2550-55. With M. }. Kasperbauer and H. A. Borthwick. Inhibition of flow- ering of Chenopodium rubrum by prolonged far-red radiation. Bot. Gaz., 124~6) :444-51.

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STERLING BROWN HENDRICKS 1964 209 Photochemical aspects of photoperiodicity. In: Photophysiology, ed. E. Geise, pp. 305-31. New York: Academic Press. With H. W. Siegelman. Phytochrome and its control of plant growth and development. In: Advances in Enzymology, ed. F. F. Nord, vol. 26, pp. 1-33. New York: Interscience. With M. i. Kasperbauer and H. A. Borthwick. Reversion of phy- tochrome 730 (Pfr) to P660 (Pr) assayed by flowering in Cheno- podium rubrum. Bot. Gaz., 125~2) :75 -80. Salt transport across cell membranes. Am. Sci., 52~3~:306-33. With W. L. Butler and H. W. Siegelman. Action spectra of phyto- chrome in vitro. Photochem. Photobiol., 3:521-28. 1965 With R. l. Downs, H. W. Siegelman, and W. L. Butler. Photorecep- tive pigments for anthocyanin synthesis in apple skins. Nature, 205 :909-10. With J. E. Leggett and W. R. Heald. Cation binding by baker's yeast and resins. Plant Physiol., 40:665-71. With L. T. Evans and H. A. Borthwick. The role of light in sup- pressing hypocotyl elongation in lettuce and petunia. Planta, 64:201-18. With B. G. Cumming and H. A. Borthwick. Rhythmic flowering responses and phytochrome changes in a selection of Chenopo- dium rubrum. Can. I. Bot., 43:825-53. With H. A. Borthwick. The physiological function of phytochrome. In: Biochemistry o/Plant Pigments, ed. T. W. Goodwin, pp. 405- 36. London: Academic Press. With L. T. Evans and H. A. Borthwick. Inflorescence initiation in Lolium temulentum L. VII. The spectral dependence of induc- tion. Aust. }. Biol. Sci., 18:745-62. With H. W. Siegelman. Purification and properties of phyto- chrome: A chromoprotein regulating plant growth. Fed. Proc. Fed. Am. Soc. Exp. Biol., 24:863-67. 1966 Plant growth. In: McGraw-Hill Encyclopedia oiscience and Technology, pp. 299-302. New York: McGraw-Hill.

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210 BIOGRAPHICAL MEMOIRS With l. C. Fondeville and H. A. Borthwick. Leaflet movement of Mimosa pudica L. indicative of phytochrome action. Planta, 69:357-64. With H. W. Siegelman and B. C. Turner. The chromophore of phytochrome. Plant Physiol., 41: 1289-92. 1967 With A. I. Hiatt. The role of COD fixation in accumulation of ions by barley route. Z. Pflanzenphysiol., 56: S. :220 -32. With i. C. Fondeville, M. }. Schneider, and H. A. Borthwick. Pho- tocontrol of Mimosa pudica L. Leaf movement. Planta, 75:228- 38. Light in plant life. In: Harvesting the Sun, ed. A. San Pietro, F. A. Greer, and T. l. Army, pp. 1-4. New York: Academic Press. With H. W. Siegelman. Phytochrome and photoperiodism in plants. Comp. Biochem., 27:211-35. With H. A. Borthwick. The function of phytochrome in regulation of plant growth. Proc. Natl. Acad. Sci. USA, 58:2125-30. With M. I. Schneider and H. A. Borthwick. Eject of radiation on Hyoscyamus niger. Am. i. Bot., 54:1241-49. 1968 Photoperiodism after 50 years. }. Wash. Acad. Sci., 58:69-74. With l. E. Schiebe. Short communication an observation on the photooxidation of ascorbic acid in strawberry leaves. Phyto- chemistry, 7:31-33. How light interacts with living matter. Sci. Am., 219:175-84. With V. K. Toole and H. A. Borthwick. Opposing actions of light in seed germination of Poa pretensis and Amaranthus arenicola. Plant Physiol., 43:2023-28. With R. P. Burchard. Action spectrum for carotenogenesis in Myxo- coccus xanthus. ]. Bacteriol., 97: 1165-68. 1969 Plant physiology. In: A Short History of Botany in the United States, Eleventh International Botanical Congress, Seattle, Washing ton. With H. A. Borthwick, M. l. Schneider, R. B. Taylorson, and V. K. Toole. The high-energy light action controlling plant responses and development. Proc. Natl. Acad. Sci. USA, 64:479-86.

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STERLING BROWN HENDRICKS 211 With R. B. Taylorson. Action of phytochrome during prechilling of Amaranthus retropexus L. seeds. Plant Physiol., 44:821-25. 1970 The passing scene. Annul Rev. Plant Physiol., 21: 1-10. 1971 With R. B. Taylorson. Changes in phytochrome expressed by ger- mination of Amaranthus retropexus L. seeds. Plant Physiol., 47:619-22. 1972 With R. B. Taylorson. Interactions of light and a temperature shift on seed germination. Plant Physiol., 49:127-30. With R. B. Taylorson. Rehydration of phytochrome in imbibing seeds of Amaranthus retropexus L. Plant Physiol., 49:663 - 65. With R. B. Taylorson. Promotion of seed germination by nitrates and cyanides. Nature, 237:169-70. With R. B. Taylorson. Phytochrome control of germination of Ru- mex crispus L. seeds induced by temperature shifts. Plant Phys- iol., 50:645-58. 1973 With R. B. Taylorson. Promotion of seed germination by cyanide. Plant Physiol., 52:23-27. With R. B. Taylorson. Phytochrome transformation and action in seeds of Rumex crispus L. during secondary dormancy. Plant Physiol., 52:475-79. 1974 With R. B. Taylorson. Promotion of seed germination by nitrate, nitrite, hydroxylamine and ammonium salts. Plant Physiol., 54:304-9. 1975 With R. B. Taylorson. Breaking of seed dormancy by catalase in- hibition. Proc. Natl. Acad. Sci. USA, 72:306-9. 1976 With R. B. Taylorson. Aspects of dormancy in vascular plants. BioScience, 26:95 -101.

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212 BIOGRAPHICAL MEMOIRS With R. B. Taylorson. Variation in germination and amino acid leakage of seeds with temperature related to membrane phase change. Plant Physiol., 58: 7-11. With R. B. Taylorson. Interactions of phytochrome and exogenous gibberellic acid on germination of Lamium amplexicaule L. seeds. Planta, 132:65-70. 1977 With R. B. Taylorson. Dormancy in seeds. Annul Rev. Plant Phys- iol.,28:331-54. 1978 With R. B. Taylorson. Dependence of phytochrome action on membrane organization. Plant Physiol., 61:17-19. 1979 With R. B. Taylorson. Dependence of thermal responses of seeds on membrane transitions. Proc. Natl. Acad. Sci. USA, 76:778- 81. With R. B. Taylorson. Overcoming dormancy in seeds with ethanol and other anesthetics. Planta, 145: 507-10. 1980 With R. B. Taylorson. Reversal by pressure of seed germination promoted by anesthetics. Planta, 149: 108-11. With R. B. Taylorson. Anesthetic effects on seed dormancy an overview. Isr. I. Bot., 29:273 - 80.

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