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ROBERT FRANKLIN PITTS October 24, 908-fune 6, 1977 BY ROBERT W. BERLINER AND GERHARD H. GIEBISCH THE SCIENTIFIC CONTRIBUTIONS of Robert Franklin Pitts have been a major force in moicting the shape of renal physiology in the last half-century. There are few as- pects of kidney function that he did not explore, and his work illuminated each element that came uncler his scrutiny. But his contributions to physiology were not limitect to the study of the kidney. He produced important work in neurophysi- ology early in his career, making contributions that for many would be sufficient to lencl prestige to the work of a lifetime, but that Bob Pitts was able to accomplish in only a few short years. Robert F. Pitts was born in Indianapolis on October 24, 190S, the younger of the two children of John Franklin and Estelle Coffin Pitts. His sister Rebecca, three years his senior, has proviclec! much of the information about the family back- ground, chilc~hoocl, ant! upbringing of her younger brother. Both parents were members of the Society of Friends and traced their ancestry back to the earliest days of Quakerism. They had known each other since childhood anct were mar- riecl in 1899, moving to Inclianapolis the same year. Accorct- ing to Rebecca Pitts: "Because John Franklin Pitts hac] been a farmer's son, he was ill-equipped for city life, and for several years the young couple was very poor. In fact, throughout 323

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324 BIOGRAPHICAL MEMOIRS Robert's childhooct and youth our circumstances were, though not poverty stricken, certainly marked by severely necessary economies anc! occasional periods of real hard- ship.... Until Robert was about fourteen his social life was limited to membership in church youth groups. Our parents had strict notions about keeping us at home in the eve- nings.... Such an environment gives lessons in the oIct Pu- ritan virtues of discipline and industry, economy anct careful planning; and my brother Earned them very well." At Butler University, which he entered as an unclergrad- uate a month before his seventeenth birthday, he ctid very well. According to his sister's account: "He was a member of Phi Delta Theta a fraternity notes! more for football, at least on the Butler campus, than for scholarship. But the Phi Delts were prouct of his scholarship . . . and elected him pres- ident of the chapter.... Although he was never an athlete he was a good tennis player, and in his high school and college years spent many summer afternoons or early mornings on the court." After receiving the Bachelor of Science degree from But- ler at the age of twenty, Bob was awarded a fellowship in biology at Johns Hopkins University. It hac! been his inten- tion, first expressed at the age of four (!) in admiration of the family doctor, to study medicine. It may be assumed that to obtain a Ph.D. in a basic science was, in view of his financial limitations, a practical step toward his long-term goal. In any case, having tract a taste of research in pursuit of his first doctoral clegree, it became clear to him that research rather than the practice of medicine was his real passion. Bob joined the Department of Physiology at the New York University College of Medicine in 1932, fresh from his cloc- toral work in biology at Johns Hopkins. His dissertation work had dealt with physiological processes in amoebae; having obtainer! his Ph.D., however, he closecl the book on that field

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ROBERT FRANKLIN PITTS 325 ant! never returned to it. In the 1930s, the NYU Department of Physiology was the center of an intensive exploration of the function of the kidney. Leadership in renal physiology at that time was clivided between the group uncler Homer Smith at NYU and that led by A. N. Richards at the University of Pennsylvania. The approaches of the two groups were quite separate anal distinct. Richards and his associates were cle- veloping and applying the early and by latter-(lay stan- clards primitive micropuncture techniques. Homer Smith always considered his greatest contribution to experimental physiology to have been the trained, intact, unanesthetize(1 clog. Indeed, except for diversions into work with fish in the summers at the Mount Desert Island Biological Laboratory in Salisbury Cove, Maine, work in the NYU Department of Physiology stuck pretty closely to the intact dog (or occasion- ally man), and Bob Pitts's work was no exception. In fact, his preference for the intact animal (although not necessar- ily unanesthetized) was reflected in his experimental work throughout his career, even when most others had shifted, with greatly improver! instruments and techniques, to the trail of A. N. Richards and micropuncture. From 1932 to 193S, Bob Pitts was an active and productive member of the physiology clepartment. His first paper from his new environment at NYU clealt with the relationship be- tween the excretion of inorganic phosphate anc! the plasma phosphate level in the dog. The subject matter of this work is noteworthy because it was a later anc! more definitive study of phosphate reabsorption by the renal tubules that lecT to what most would consider to be Bob Pitts's most important single contribution to renal physiology: his work on acidifi- cation of the urine. During this first six-year period at NYU, Bob explored the renal mechanisms involvecl in the excretion of a number of substances: creatine, urea, xylose, hexameth- enamine, ammonia, anct phenol red. The work was a signif-

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326 . BIOGRAPHICAL MEMOIRS icant contribution to the state of the art at the time, although it click not leas! to any major new fields of stucly or new ways of looking at renal function. During this period, however, Bob found time to enroll as a medical student and, while continuing his work in the lab- oratory, managed to complete the work for the M.D. degree that he was awarded in 1938. It is clear, moreover, that his completion of medical school was not accomplished in an offhand way: he graduated at the head of his class anct re- ceivecl a mecial for his work in pathology, as well as the senior prizes in both medicine and surgery! Although he never chose to develop his obvious talent for clinical activities, his thorough grounding in medicine influenced all his subse- quent efforts, and he never failed to orient his fundamental physiological work to clinically important problems and to call attention to the relevance of his findings to medicine. Those whose exposure to biomedical science has been lim- ited to the more recent era of relative affluence and avail- ability of research funds may imagine that Pitts's coworkers and technicians kept things running in the lab with only pe- riodic guidance from him, thus allowing him to continue his research in the laboratory while giving unstinted attention to the medical school curriculum. Nothing could be further from the truth. He did all of his own experiments and all of the analyses himself. It is also probable, although unclocu- mented, that he washed his own glassware. Nevertheless, in that six-year period, he published twelve highly creclitable papers, and he was the sole author of eleven of them. Upon graduation from medical school, he chose to launch into a new field and, as a fellow of the Rockefeller Founcia- tion, he spent a year at Northwestern University in the labo- ratory of Magoun and Ranson and the subsequent year in the Johnson Foundation laboratories at the University of Pennsylvania with Detlev Bronk. (Both of these were leading

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ROBERT FRANKLIN PITTS 327 laboratories in what would now be called neuroscience.) Al- though many would hold that one year in a laboratory is har(lly enough time to accomplish anything much, particu- larly in a new field, it is apparent that this rule clid not apply in this instance. The immediate output of that brief period was eight papers; for seven of them, Pitts was the senior au- thor. Moreover, he did not merely fall in step with projects already under way. He built his own electronic equipment and launched into a new field: the study of the meclulIary respiratory centers and related phenomena. The judgment of those familiar with the fielci appears to have been that these were important contributions to neurophysiology. Some six years later, when he had already established himself as the leacling contributor to renal physiology, he was still the author of textbook chapters on the regulation of respiration. In fact, he hell! what must have been a unique distinction in writing, by invitation, a review entitIect "Organization of the Respiratory Center" for Physiological Reviews and the chapter on the kidney in Annual Reviews of Physiology, both in the same year (19461. In 1940 he returned as an assistant professor to the De- partment of Physiology at NYU. For two years he continued his studies on the control of respiration and then moved a short distance up the east sidle of Manhattan to join the Department of Physiology at Cornell University Medical College. It had been his intention to continue his work in neurophysiology, but the expense of establishing a new lab- oratory base(1 on electronic equipment, anti the (1ifficulty in obtaining financial resources to do so, lecl him to abandon that plan. All that was neectect for his work in renal physiol- ogy was a little glassware, some chemicals, and a few trained clogs; so he returned to the stucly of the kidney. The next four years in New York were a period of productivity that incluclecl what Bob himself considered to be some of his best

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328 BIOGRAPHICAL MEMOIRS work. It is easy to concur with his judgment without clispar- agement of the enormous value of his many contributions in subsequent years. The first few papers that Bob Pitts producecT after taking up his position at Cornell clealt with the mechanisms for the reabsorption of amino acicis by the renal tubules. He then returned to work on the reabsorption of phosphate, which was described in a paper with Robert Alexancler as coauthor: "The Renal Reabsorptive Mechanism for Inorganic Phos- phate in Normal ant] Aciclotic Dogs." The essence of this paper was that phosphate was reabsorbed with a saturable transport process and that the capacity of this transport pro- cess was not affecter! by changes in the acid-base status of the animal. Many years later, in a 1971 paper entitlec! "Some Aphor- isms on Research ant} Writing," Bob related how this stucly lee! to his work on the mechanism of urinary acidification. it seems that Bob presented the results of the study at the Cor- nell Research Society, where his statement that phosphate reabsorption was not increased by acidosis was challengect by one of his biochemist colleagues. This colleague stated that inasmuch as it was well known that the urine is rendered acid by the reabsorption of diso(lium phosphate, leaving behinct the more acic! member of the buffer pair, the reabsorption of phosphate must increase in acidosis when the excretion of acid in the urine is increased. Bob thought for several months about that conflict between theory anct data, ant] he con- clucled that the phosphate reabsorption data were not wrong but that the mechanism postulated by his biochemical col- league probably was. In fact, Bob's mentor, Homer Smith, the theoretician and philosopher of renal physiology, had suggester! a different mechanism some ten years earlier: namely, secretion of hydrogen ion by exchange for fixed cat- ion. Neither hypothesis tract ever been tested. Bob decided it

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ROBERT FRANKLIN PITTS 329 was time to do so. The experiment that he clesignect to ex- amine the alternatives was beautifully conceived to provide a definitive answer. According to Bob's account, and the story is confirmed by Robert Alexander, his collaborator and co- author of the resulting paper, Bob wrote the entire paper- minus only the databefore they carried out the first exper- iment! Clearly, the results were all he could have hoped for, showing that hydrogen ion secreted by the renal tubules was indeed responsible for acidifying the urine. The paper, "The Nature of the Renal Tubular Mechanism for Acidifying the Urine" appeared in the American Journal of Physiology in 1945. It was consiclerec] to be absolutely definitive and a landmark of renal physiology. Bob's oIcler sister tells us that their mother was fond of saying, "Plan your work, then work your plan." Bob had learned the lesson well. The paper on acidification of the urine established Robert F. Pitts as the leacling investigator in renal physiology, but it was only the beginning of a series of studies in which he explorer! a number of relater! aspects of renal function: the reabsorption of bicarbonate by the tubules, the factors gov- erning the rate of excretion of titratable acid, anct the renal tubular reabsorption of chIoricle. In acTdition, a paper with William Lotspeich, "The Role of Amino Acids in the Renal Tubular Secretion of Ammonia," introducecI a subject that in later years was to be the focus of Bob Pitts's major line of study. In 1946 Pitts left Cornell to assume the chairmanship ot the Department of Physiology at Syracuse University (the school of medicine that subsequently became the State Uni- versity of New York Upstate Mectical Center). The move was accomplished without any apparent discontinuity in his re- search. With a new group of younger associates, the work on acidification anct bicarbonate reabsorption was extended to the normal human, using the investigators as subjects. The

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330 BIOGRAPHICAL MEMOIRS interests of Bob and his series of younger associates were not, however, limited to acic! secretion by the tubules and related phenomena; while he was at Syracuse, papers appeared deal- ing with the effects of adrenal hormones on electrolyte reab- sorption ant} excretion and on the effects of mercurial di- uretics. The latter were his first venture into the study of diuretics anc! their site anti mechanisms of action, an area of investigation that was to be an important element of his work over the next clecade. In 1950 Pitts returned to Cornell as the chairman of the Department of Physiology, a position he held until 1973, shortly before his retirement from the university. Bob was a conscientious chairman and a devotecI teacher. His lectures were carefully prepared and models of clarity. In acldition, he placed great importance on the teaching efforts of the members of his department. He regularly attencled all the lectures in the course in physiology throughout the period of his chairmanship and participated enthusiastically in the student laboratory exercises. The stream of published reports of first-rate work was never interrupted during the period of his chairmanship. This was the case despite problems of illness, both of his wife and of himself, that began in the midcIle fifties ancT were to plague him through the remainder of his life. His wife was stricken with progressive, incapacitating neurological disease that caused her to be beciriciclen for many years, during which Bob Elevated great personal effort to her care. AncT Bob himself was the victim of at least three ailments that might have lecI a less cledicated and cleterminec] man to give up. In 195S, when one of us was his companion on a mission to the MicIdIe East for the Unitarian Service Committee, his pockets contained a medicine cabinet's assortment of pre- scribec] medications. Nevertheless, through the years Bob not only continued his extraordinarily productive activities but

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ROBERT FRANKLIN PITTS 331 also adheres! to his practice of doing a great clear of the lab- oratory work himself. In fact, he not only diet most of the laboratory work he also built much of his own apparatus, inclucling an early flame photometer for the measurement of sodium and potassium, ancT later an amino-acict analyzer. Throughout his career he washer! his own glassware and in- sistecl that others follow his rigid protocol for achieving ade- quate chemical cleanliness. He later explained his participa- tion in these activities by noting that "some of my best ideas have come to me when I'm performing some routine analyt- ical chore." His work explorecl many areas of renal physiology. He strayed from the kidney only slightly and for a brief period when he, along with Roy Swan and Gerharct Giebisch, cle- fined the extrarenal buffering of acid and base loacts. An excellent series of papers clearing with the site and mecha- nism of action of diuretics, particularly the mercurial ctiuret- ics that were then the therapeutic mainstay, appeared seria- tim from 1950 to 1962. His studies of the potentiation of the diuretic effect by acidifying salts and of the relationship be- tween structure and activity among the mercurial diuretics led him to conclucle that the effect was probably on the trans- port of chloride anti attributable to the intact molecule. This contrasted with the inference of others that dissociation of the mercury was necessary for the effect that was thought to be proclucec] on the transport of the sodium ion. Studies nearly twenty years later with isolated tubules have shown that the Pitts interpretation was correct. Except for a brief dalliance with the stop-flow method in some of his studies of diuretics, Bob stuck pretty much to the intact kidney, often using the intact dog. Even when his own department became one of the leading centers of micro- puncture work, Bob Pitts steered clear of that method. In part, at least, he explainecl this decision in one of his "aphor-

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332 BIOGRAPHICAL MEMOIRS isms on research" in the paper referred to earlier. "Pick an area in which there is no, or at least little current research activity," he advised. The reason: The investigator can cle- velop his ideas without pressure to publish to establish prior- ity. He followed his own precepts studiously, followed his own course unpressured by the work of others, ant! never pub- lished a trivial paper or a wrong one. In the last dozen years of his work, Bob returned to the area that he had opened up in his studies of acidification of the urine anct explored that other element in the regulation of acid-base balance, the excretion of ammonia. Almost everything beyond the initial identification of glutamine by Van Slyke and his associates that we know about the sources of ammonia and the renal processes involved in its excretion is based on the work of Bob Pitts. In 1974 Bob Pitts accepted emeritus status at Cornell ancI moved to the University of Florida in Gainesville where he held the rank of Research Professor in Renal Medicine anc! Physiology. Unfortunately, his health, for a long time far from robust, deterioratect further, ant] he was able to add little to the list of his magnificent accomplishments before his death on June 6, 1977. It might be said that, in the last half- century, if Homer Smith was the high priest of renal physi- ology, then Bob Pitts was surely the builder of its temple.

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ROBERT FRANKLIN PITTS 335 1936 The clearance of hexamethenamine in the dog. Am. }. Physiol., 115:706. The comparison of urea with urea + ammonia clearances in aci- dotic dogs. I. Clin. Invest., 15:571. Excretion of creatine by the marine teleost, the red grouper. In: Annual Report of the Tortugas Laboratory, 1935 - 36, p. 99. Wash- ington, D.C.: Carnegie Institution of Washington. 1938 The excretion of phenol red by the chicken. J. Cell. Comp. Physiol., 11:99. With I. M. Koor. The excretion of urea by the chicken. J. Cell. Comp. Physiol., 11: 117. 1939 The excretion of creatine by the dogfish, Squalus acanthius. 5. Cell. Comp. Physiol., 19:151. With H. W. Magoun and S. W. Ranson. Localization of the medul- lary respiratory centers in the cat. Am. }. Physiol., 126:673. With H. W. Magoun and S. W. Ranson. Interrelations of the res- piratory centers in the cat. Am. J. Physiol., 126:689. With H. W. Magoun and S. W. Ranson. The origin of respiratory rhythmicity. Am. I. Physiol., 127:654. 1940 The respiratory center and its descending pathways. i. Comp. Neu- rol.,72:605. 1941 With M. G. Larrabee and D. W. Bronk. An analysis of hypotha- lamic cardiovascular control. Am. I. Physiol., 134:359. 1942 With D. W. Bronk. Excitability cycle of the hypothalamus-sympa- thetic neurone system. Am. J. Physiol., 135:504. The function of components of the respiratory complex. l. Neu- rophysiol., 5:403.

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336 BIOGRAPHICAL MEMOIRS 1943 A renal reabsorptive mechanism in the dog common to glycine and creatine. Am. I. Physiol., 140: 156. The basis for repetitive activity in phrenic motorneurons. I. Neu- rophysiol., 6:439. 1944 A comparison of the renal reabsorptive processes for several amino acids. Am. I. Physiol., 140:535. The effects of infusing glycin and of varying the dietary protein intake on renal hemodynamics in the dog. Am. J. Physiol., 142:355. With R. S. Alexander. The renal reabsorptive mechanism for in- organic phosphate in normal and acidotic dogs. Am. I. Physiol., 142:648. 1945 With R. S. Alexander. The nature of the renal tubular mechanism for acidifying the urine. Am. {. Physiol., 144:239. The renal regulation of acid-base balance with special reference to the mechanism for acidifying the urine. Science, 102:49. 1946 Organization of the neural mechanisms responsible for rhythmic respiration, pp. 896-912; Regulation of respiration, pp. 913- 32. In: Howell's Textbook of Physiology, ed. I. F. Fulton. Kidney. Annul Rev. Physiol., 8:199. Organization of the respiratory center. Physiol. Rev., 26:609. With W. D. Lotspeich. Bicarbonate and the renal regulation of acid- base balance. Am. J. Physiol., 147: 138. With W. D. Lotspeich. Factors governing the rate of excretion of titratable acid in the dog. Am. I. Physiol., 147:481. 1947 With W. D. Lotspeich and R. C. Swan. The renal tubular reabsorp- tion of chloride. Am. J. Physiol., 148:445. With W. D. Lotspeich. Use of thiosulfate clearance as a measure of glomerular filtration rate in acidotic dogs. Proc. Soc. Exp. Biol. Med., 64:224.

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ROBERT FRANKLIN PITTS 337 With W. D. Lotspeich. The role of amino acids in the renal tubular secretion of ammonia. I. Biol. Chem., 168 :611. With J. L. Ayer and W. A. Schiess. Independence of phosphate reabsorption and glomerular filtration in the dog. Am. I. Phys- iol., 151:168. 1948 With W. D. Lotspeich, W. A. Schiess, and I. L. Ayer. The renal regulation of acid-base balance in man. I. The nature of the mechanism for acidifying the urine. I. Clin. Invest., 27:48. With W. D. Lotspeich, W. A. Schiess, and i. L. Ayer. The renal regulation of acid-base balance in man. II. Factors affecting the excretion of titratable acid by the normal human subject. I. Clin. Invest., 27:57. Renal excretion of acid. Fed. Proc. Fed. Am. Soc. Exp. Biol.,7:418. With I. iahan. Effect of parathyroid on renal tubular reabsorption of phosphate and calcium. Am. J. Physiol., 155:42. 1949 With I. L. Ayer and W. A. Schiess. The renal regulation of acid- base balance in man. III. The reabsorption and excretion of bicarbonate. i. Clin. Invest., 28:35. With O. W. Sartorius and I. C. Roemmelt. The renal regulation of acid-base balance in man. IV. The nature of the renal compen- sations in ammonium chloride acidosis. I. Clin. Invest., 28:423. With I. C. Roemmelt and O. W. Sartorius. Excretion and reabsorp- tion of sodium and water in the adrenalectomized dog. Am. I. Physiol., 159: 124. 1950 With i. i. Duggan. Studies on diuretics. I. The site of action of mercurial diuretics. I. Clin. Invest., 29:365. With J. J. Duggan. Studies on diuretics. II. The relationship be- tween glomerular filtration rate, proximal tubular absorption of sodium and diuretic efficacy of mercurials. I. Clin. Invest., 29:372. With O. W. Sartorius. Mechanism of action and therapeutic use of diuretics. I. Pharmacol. Exp. Ther., 98:161. Acid-base regulation by the kidneys. Am. l. Med., 9:356.

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338 BIOGRAPHICAL MEMOIRS With M. i. Browne and M. W. Pitts. Alkaline phosphatase activity in kidneys of glomerular and aglomerular marine teleosts. Biol. Bull., 99:152. 1951 With W. S. Wiggins, C. H. Manry, and R. H. Lyons. The effect of salt loading and salt depletion on renal function and electrolyte excretion in man. Circulation, 3:275. With D. D. Thompson and M. I. Barrett. Significance of glomer- ular perfusion in relation to variability of filtration rate. Am. I. Physiol., 167:546. Effect of adrenal cortical hormones on renal function. In: Adrenal Cortex, Transactions of the Third Conference, ed. E. D. Ralli, p. 703. New York: Josiah Macy, ir., Foundation. With S. Kupfer and D. D. Thompson. The isolated kidney and its response to diuretic agents. Am. i. Physiol., 167:703. 1952 With K. E. Roberts. The influence of cortisone on renal function and electrolyte excretion in the adrenalectomized dog. Endo- crinology, 50-51. With D. R. Axelrod. Effects of hypoxia on renal tubular function. J. Appl. Physiol., 4:593. With D. D. Thompson. Effects of alterations of renal arterial pres- sure on sodium and water excretion. Am. l. Physiol., 168:490. With D. R. Axelrod. The relationship of plasma pH and anion pattern to mercurial diuresis. J. Clin. Invest., 31:171. With D. R. Axelrod. Anoxia as a factor in resistance to mercurial diuretics. Am. J. Physiol., 169:350. Modern concepts of acid-base regulation. Arch. Int. Med., 89:864. With J. N. Capps, W. S. Wiggins, and D. R. Axelrod. The effect of mercurial diuretics on the excretion of water. Circulation, 6:82. With O. W. Sartorius and D. Calhoon. The capacity of the adre- nalectomized rat to secrete hydrogen and ammonium ions. En- docrinology, 51:444. 1953 With K. E. Roberts and M. G. Magida. Relationship between po- tassium and bicarbonate in blood and urine. Am. I. Physiol., 172:47.

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ROBERT FRANKLIN PITTS 339 With O. W. Sartorius and D. Calhoon. Studies on the interrelation- ships of the adrenal cortex and renal ammonia excretion by the rat. Endocrinology, 53:256. With K. E. Roberts. The effects of cortisone and desoxycorticoster- one on the renal tubular reabsorption of phosphate and the excretion of titratable acid and potassium in dogs. Endocrinol- ogy, 52:324. Mechanisms for stabilizing the alkaline reserves of the body. Harvey Lect. Ser. 48. 1954 With P. l. Dorman and W. I. Sullivan. The renal response to acute respiratory acidosis. I Clin. Invest., 33: 82. With G. Giebisch and H. D. Lauson. Renal excretion and volume of distribution of various dextrans. Am. I Physiol., 178: 168. With R. C. Swan and H. Madisso. Measurement of extracellular fluid volume in nephrectomized dogs. I. Clin. Invest., 33:1147. With P. I. Dorman and W. }. Sullivan. Factors determining carbon dioxide tension in urine. Am. l. Physiol., 179:181. With G. Giebisch and L. Berger. The extrarenal response to acute acid-base disturbances of respiratory origin. J. Clin. Invest., 34:231. With R. C. Swan. Neutralization of infused acid by nephrectomized dogs. J. Clin. Invest., 34:205. Uber active transport Mechanisem in den Tubuli der Niere. Klin. Wochenschr., 33:365. With G. R. Fuller and M. B. MacLeod. The influence of the ad- ministration of potassium salts on the renal tubular reabsorp- tion of bicarbonate. Am. J. Physiol., 182: 111. With R. C. Swan, D. R. Axelrod, and M. Seip. Distribution of so- dium bicarbonate infused into nephrectomized dogs. J. Clin. Invest., 34:1795. With G. Giebisch and M. B. MacLeod. The effects of adrenal ste- roids on renal tubular reabsorption of bicarbonate. Am. I. Phys- iol., 183:377. 1956 With R. R. M. Borghgraef. The distribution of chlormerodrin (Neohydrin) in tissues of the rat and dog. J. Clin. Invest.,35:31. With R. L. Greif, S. J. Sullivan, and G. S. Jacobs. Distribution of

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340 BIOGRAPHICAL MEMOIRS radiomercury administered as labeled chlormerodrin (Neohy- drin) in the kidneys of rats and dogs. J. Clin. Invest., 35:38. With B. K. Ochwadt. Effects of intravenous infusion of carbonic anhydrase on carbon dioxide tension of alkaline urine. Am. I. Physiol., 1 85:426. With R. R. M. Borghgraef and R. H. Kessler. Plasma regression, distribution and excretion of radiomercury in relation to di- uresis following the intravenous administration of Hg203 labeled chlormerodrin to the dog. I. Clin. Invest. 35: 1055. With B. K. Ochwadt. Disparity between the phenol red and the diodrast clearances in the dog. Am. i. Physiol., 187:318. 1957 With P. Poulos. An indirect flame photometric method for calcium in plasma and urine. i. Lab. Clin. Med., 49:300. With R. H. Kessler and R. Lozano. Studies on structure diuretic activity relationships of organic compounds of mercury. I. Clin. Invest., 36:656. With D. D. Thompson, F. Kavaler, and R. Lozano. An evaluation of the cell separation hypothesis of autoregulation of renal blood flow and filtration rate. I. Blood flow, filtration rate and PAH extraction as functions of arterial pressure in normal and anemic dogs. Am. J. Physiol., 191 :494. With R. H. Kessler and O. P. A. Heidenreich. An evaluation of the cell separation hypothesis of autoregulation of renal blood flow and filtration rate. II. Glucose titrations in normal and anemic dogs. Am. I. Physiol., 191:150. 1958 Some reflections on mechanisms of action of diuretics. Am. J. Med., 24:745. With F. Kruck, R. Lozano, D. W. Taylor, O. P. A. Heidenreich, and R. H. Kessler. Studies on the mechanism of action of chloro- thiazide. J. Pharmacol. Exp. Ther., 123:89. With R. S. Gurd, R. H. Kessler, and K. Hierholzer. Localization of acidification of urine, potassium and ammonia secretion and phosphate reabsorption in the nephron of the dog. Am. I. Phys- iol., 194:125. With R. H. Kessler, K. Hierholzer, and R. S. Gurd. Localization of

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ROBERT FRANKLIN PITTS 341 the diuretic action of chlormerodrin in the nephron of the dog. Am. I. Physiol., 194:540. 1959 With R. H. Kessler, K. Hierholzer, and R. S. Gurd. Localization of action of chlorothiazide in the nephron of the dog. Am. J. Phys- iol., 196:1346. The Physiological Basis of Diuretic Therapy. Springfield, Ill.: Charles C Thomas. 1960 With K. Hierholzer, R. Cado, R. Gurd, and R. H. Kessler. Stop flow analysis of renal absorption and excretion of sulfate in the dog. Am. I. Physiol., 198:833. With G. Giebisch and E. E. Windhager. Mechanism of urinary acidification. In: Biology of Pyelonephratis, p. 277. Boston: Little, Brown & Co. The teacher and the ferment in education. (Past president's ad- dress to the American Physiological Society.) Physiologist, 3:20. 1961 With I. L. Brown and A. H. E. Samiy. Localization of amino- nitrogen reabsorption in the nephron of the dog. Am. J. Phys- iol., 200:370. With J. R. Cade, B. Shalhoub, and M. Canessa-Fischer. The effect of strophanthidin on the renal tubules of the dog. Am. I. Phys- iol., 200:373. A comparison of the modes of action of certain diuretic agents. Prog. Cardiovasc. Dis., 3:537. With W. A. Webber and l. L. Brown. Interactions of amino acids in renal tubular transport. Am. I. Physiol., 200:380. 1962 With S. Balagura. The excretion of ammonia injected into the renal artery. Am. J. Physiol., 203: 11.

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342 BIOGRAPHICAL MEMOIRS 1963 With R. i. Shalhoub, W. Webber, S. Glabman, M. Ganessa-Fischer, I Klein, and I. deHaas. Extraction of amino acids from and their addition to renal blood plasma. Am. I. Physiol., 204:181. With I. deHaas and I. Klein. Relation of renal amino and amide extraction to ammonia production. Am. I. Physiol., 204:187. With M. Canessa-Fischer, R. I. Shalhoub, S. Glabman, and I. deHaas. The effects of infusions of ammonia, amides and amino acids on the excretion of ammonia. Am. l. Physiol., 204:192. Physiology of the Kidney and Body Fluids. Chicago: Yearbook Publish- ers. 1964 With G. Denis and H. Preuss. The pNH3 of renal tubular cells. I. Clin. Invest., 43:571. With S. Balagura. Renal handling of cx-ketoglutarate by the dog. Am. I. Physiol., 207:483. Renal production and excretion of ammonia. Am. i. Med., 36:720. 1965 With L. A. Pilkington and J. deHaas. Ni5 tracer studies on the origin of urinary ammonia in the acidotic dog with notes on the enzymatic synthesis of labeled glutamic acid and glutamine. {. Clin. Invest., 44:731. With L. A. Pilkington and J. Welch. Relationship of pNH3 of tu- bular cells to renal production of ammonia. Am. l. Physiol., 208:1 100. With L. A. Pilkington, R. Binder, and J. deHaas. Intrarenal distri- bution of blood flow. Am. J. Physiol., 208: 1 107. With G. Fulgraff. A study of the kinetics of ammonia production and excretion in the acidotic dog. Am. J. Physiol., 209:1206. 1966 With L. A. Pilkington. The relation between plasma concentrations of glutamine and glycine and utilization of their nitrogens as sources of urinary ammonia. I. Clin. Invest., 45:86. The renal metabolism of ammonia. Physiologist, 9:97;

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ROBERT FRANKLIN PITTS 1967 343 With W. I. Stone. Renal metabolism of alanine. I. Clin. Invest., 46:530. With W. l. Stone. Pathways of ammonia metabolism in the intact functioning kidney of the dog. I. Clin. Invest., 46: 1141. With W. i. Stone and S. Balagura. Diffusion equilibrium for am mania in the kidney of the acidotic dog. I. Clin. Invest. 46:1603. 1969 With M. L. Lyon. Species differences in renal glutamine synthesis in viva. Am. I. Physiol., 216: 117. Renal excretion of ammonia. In: Progress in Nephrology, ed. G. Peters and F. Roch-Ramel, p. 75. Berlin: Springer-Verlag. 1970 With A. C. Damian. Rates of glutaminase I and glutamine synthe- tase reactions in rat kidney in viva. Am. I. Physiol., 218: 1249. With A. C. Damian and M. B. MacLeod. Synthesis of serine by rat kidney in vivo and in vitro. Am. I. Physiol., 219:504. Production and excretion of ammonia in relation to acid-base reg- ulation. In: Handbook of Physiology, Renal Physiology, p. 455. Washington, D.C.: American Physiological Society. With L. A. Pilkington and T. K. Young. Properties of renal lumen and antiluminal transport of plasma glutamine. Nephron, 17:51. 1971 The role of ammonia production and excretion in regulation of acid-base balance. N. Engl. l. Med., 284:32. Metabolism of amino acids by the perfused rat kidney. Am. J. Phys- iol., 220:862. Some aphorisms on research and writing. Yale I. Biol. Med., 43:331. 1972 With M. B. MacLeod. Synthesis of serine by the dog kidney in viva. Am. I. Physiol., 222:394. With L. A. Pilkington, M. B. MacLeod, and E. Leal-Pinto. Metab-

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344 BIOGRAPHICAL MEMOIRS olism of glutamine by the intact functioning kidney of the dog. Studies in metabolic acidosis and alkalosis. J. Clin. Invest., 5 1:557. Control of ammonia production and excretion. Kidney Int., 1:297. 1973 With E. Leal-Pinto, H. C. Park, V. F. King, and M. B. MacLeod. The metabolism of lactate by the intact functioning kidney of the dog. Am. I. Physiol., 224:1463.

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