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Biographical Memoirs: Volume 58 (1989)

Chapter: David Nachmansohn

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Suggested Citation:"David Nachmansohn." National Academy of Sciences. 1989. Biographical Memoirs: Volume 58. Washington, DC: The National Academies Press. doi: 10.17226/1645.
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Suggested Citation:"David Nachmansohn." National Academy of Sciences. 1989. Biographical Memoirs: Volume 58. Washington, DC: The National Academies Press. doi: 10.17226/1645.
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DAVID NACHMANSOHN March 17, 1899-November2, 1983 BY SEVERO OCHOA DA V ~ D N A C H M A N S O H N S scientific lifepath was strongly influenced by his early studies on the biochemistry of muscle in Otto Meyerhof's laboratory. This experience led to an interest in the biochemistry of nerve activity, a field of study to which he wouIcl devote most of his scientific life. In so doing, he contributed perhaps more than any other in- vestigator to our understanding of the molecular basis of bioelectricity. David Nachmansohn was born in Jekaterinoslav, Russia (now Dnjetropetrowsk, USSR). His parents came from mid(lle-cIass families among whom were many lawyers, phy- sicians, anc! other professionals. Before David and his two sisters reacher! school age, the family moved to Berlin where they had many relatives. Thus, Davicl's background and eclu- cation were essentially, if not exclusively, German. His college education was strongly humanistic, with Latin, Greek, liter- ature, and history as mainstays, some mathematics, and the rudiments of physics. Through his readings, perhaps pri- marily through his reacting of the second part of Goethe's Faust when he was only seventeen years of age, he became interested in philosophy—so much so that he continued to attend courses anc! seminars in philosophy even while a med- ical student at Heidelberg in 1920. 357

358 BIOGRAPHICAL MEMOIRS When he entered the University of Berlin in the spring of HIS, Davict was strongly oriented toward the humanities. After Germany's defeat in World War I, however, the newly establishe(1 republic face(1 grave social, political, and eco- nomic problems, ant] David was advisect to study medicine, a profession that could provide economic indepenclence. He accepted this advice and became a medical student; but as time went on, he became more and more interested in biol- ogy through his avid readings about the lives and scientific accomplishments of Bernard, Pasteur, Helmholtz, Ehrlich, and others. Eventually, he deciclect to devote his life to bio- mectical research and after his graduation in 1924 joined the laboratory of Peter Rona at the Charite for training in bio- chemistry. The Charite was the university hospital of Berlin Univer- sity Medical School in whose Department of Pathology Rona directed a division of biochemistry. There, Nachmansohn joine(1 an exceptional company of bright young people: among them, Fritz Lipmann, Hans Adolph Krebs, Ruclolph Schoenheimer, Ernst Chain, Karl Meyer, and Hans H. Weber. Nachmansohn's first paper, "Vital Staining and Adsorption," was publisher! in collaboration with Krebs, an endeavor that began a lifelong friendship between the two. Nachmansohn also (lic1 some collaborative work with Weber that lecl to the publication of a paper entitIec3, "The Independence of Pro- tein Hydration anti lonisation." At Rona's, he became familiar with the work of the great Dahiem biochemists Meyerhof, Warburg, and Neuberg, which he found fascinating. Weber adviser! Nachmansohn to go to Otto Meyerhof at the Kaiser-Wilhelm Institut fur Biol- ogie in Berlin-DahIem for further training. But when Nach- mansohn approached Meyerhof, the eminent researcher in- formec! him abruptly that he die] not accept beginners a position he reversed after speaking with the young Nach-

DAVID NACHMANSOHN 359 mansohn awhile. In Meyerhof's laboratory, Nachmansohn's postcloctoral contemporaries included Fritz Lipmann, Her- mann Blaschko, Francis O. Schmitt, and this author. Karl Lohmann, who later discovered ATP, was Meyerhof's assist- ant, and Dean Burk was a visiting scientist. Hans Krebs was also in the same building, in Otto Warburg's laboratory. Nachmansohn often mentioned that it was Meyerhof who had had the most profound! impact on his later work anct scientific outlook. Nachmansohn joined the Meyerhof laboratory in ~ 926. At that time, Grace and Philip Eggleton at the Cambridge biochemical laboratory had recently discovered a new phos- phorylated compound in muscle they called "phosphagen" because it liberated inorganic phosphate cluring contraction. Soon thereafter, Fiske anc! Subbarow at Harvard Medical School showed the new compound to be phosphocreatine in which phosphate is linked to creatine through a phosphoam- icle boncI. During this period, Meyerhof was interested in the ener- getics of muscular contraction. He worked to determine, as he had previously with various hexose phosphates, the heat of hy(lrolysis of phosphocreatine. It proved to be very high- of the order of 10,000 to 12,000 calories per mole- which contrasted with the low heat of hyclrolysis of hexose phos- phates (1,500 to 3,000 calories per mole). This finding en- ablec! researchers to ctistinguish between high- and low- energy compounds in metabolism. (Some years later, it was shown that the breakdown of ATP to ADP and inorganic phosphate was the energy-yielding process more immedi- ately related to muscular contraction, whereas the break- down of phosphocreatine served to resynthesize the AT P. Lactic acid formation, most of which took place after con- traction, was like phosphocreatine breakdown—a recovery process aimed at restoring rapidly the relatively small ATP

360 BIOGRAPHICAL MEMOIRS stores of resting muscle. Finally, the glycogen that gave rise to the lactic acid was resynthesized from lactic acid using the energy released by officiation of a fraction of the lactic acic! produced). These clevelopments fascinated the young David Nach- mansohn and greatly influenced his later work. During his early years in Meyerhof's laboratory, the function of phos- phocreatine was unknown, and interest in this compound was very strong. It is therefore not surprising that Nachman- sohn was given the assignment of looking for the relations among phosphocreatine breakdown, lactic acict formation, and the tension cleveloped by muscle during isometric con- traction in anaerobiosis. He also compared the phosphocrea- tine content of cli~erent kinds of muscle, especially muscles slivering in the rapidity of their contraction. He found that rapidly contracting muscles contained much more phospho- creatine than slowly contracting ones, a fact that was consis- tent with, and in a way foretold, the function of phospho- . . . creating In muscular contraction. Nachmansohn viviclly clescribed the atmosphere at Dah- lem in the 1920S2 when several Kaiser-Wilhelm research in- stitutes were concentrated in a relatively small area: the In- stitute of Physical Chemistry, with Haber, l~adenburg, Polanyi, Freun~llich, and Bonhoeffer; the Institute of Chem- istry, with Beckmann, WilIstatter, Otto Hahn, and Lise , . 1 ~ ~ ~ ~ . ~ ~ . ... . ~ ~ . ~ Meatier; the Neuberg Institute of Biochemistry; and the In- st~tute ot Biology, with Meyerhof, Warburg, Gol(lschmidt, Correns, and Hartmann. The young Nachmansohn was par- ticularly stimulates! by the "Haber Colloquia" in which Fritz Haber, the discoverer of the process for conversion of nitro- gen and hydrogen into ammonia, attempted to bridge the ' Nachmansohn Ascribed these influences in an unpublished manuscript entitled "Molecular Aspects of Bioelectricity: An Autobiography." 2 David Nachmansohn, "Molecular Aspects of Bioelectricity"; "Biochemistry As Part of My Life," Annual Review of Biochemistry 41(1972):1-27.

DAVID NACHMANSOHN 361 gap between physicists, chemists, and biologists so as to pro- mote better unclerstancling and cooperation among them. Nachmansohn credited these monthly colloquia, which were regularly attencled by many members of the various insti- tutes, with having greatly expanclect his scientific and spiri- tual horizons. Like so many others of Jewish origin, Nachmansohn left Germany when Hitler came to power. He was offerer! the opportunity of working at the Sorbonne, and in 1933 estab- lishecl himself in Paris with his wife, Edith, ant! their baby daughter, Ruth. From Paris, Nachmansohn made several vis- its to London, only a few hours away, to attend meetings of the British Physiological Society. As he explainect in the 1972 autobiographical article in the Annual Review of Biochemistry, he could never have anticipated that, by attending those meetings, his scientific interests wouIc] take a new, unex- pectec! turn. He conic! also not have preclictect that this new turn wouIc! determine the direction of his scientific work for the rest of his life. At that time, one of the main topics of discussion in the London meetings was the role of acety~choline in nerve ac- tivity. Following the pioneering work of Otto Loewi and of Dale and his colleagues, Dale proposed that acety~choline acts as a transmitter of nerve impulses across junctions (synapses) between neurons or between nerve and muscle, in contrast to the electric currents that propagate impulses along nerve ant! muscle fibers. This idea was supported by two main lines of observations: (~) the release of acety~choline at synaptic junctions, as judges! by its appearance in the perfusion fluid of certain ganglia, or striated muscle motor enclplates, upon electrical stimulation of the adherent nerves; and (2) the pow- erful stimulating action of acety~choline when applied locally to synaptic junctions, which was in striking contrast to its failure to elicit a response when applied to nerve fibers. Acety~choline was known to be rapidly hy(lrolyzecl by an

362 BIOGRAPHICAL MEMOIRS enzyme, acetylcholine esterase, which is strongly inhibited by the alkaloid, eserine. In fact, no acetylcholine was found in the perfusion fluid of stimulated ganglia unless the fluid con- tained eserine, an indication that the acetylcholine released by electrical stimulation was rapidly hydrolyzed. It seemed to Nachmansohn that much more knowledge was needler} on the nature, distribution, and concentration of acety~choline esterase in various tissues and that such infor- mation might provide clues to the role of this enzyme in nerve activity. He began work on this problem in Paris in 1936 and soon fount! that acetylcholine esterase was present at high concentrations in many different types of excitable fibers of nerve and muscle ant! in brain tissue, in both ver- tebrates and invertebrates; it was hardly detectable, however, in such organs as the liver or kidney. In aciclition, the con- centration appeared! to be several-fold higher at the neuro- muscular junctions than in the nerve fibers. In his study of the literature on the neuromuscular junc- tion, Nachmansohn came across an article by I. Linhard in which the electric organs of fish were clescribect as moclified muscle fibers, comparable to motor en(lplates, in which the muscular elements were either missing or present only in rudimentary form. He thought it would be of interest to cle- termine the acety~choline esterase content of electric tissue. Nachmansohn had happened to see live Torpedo at the 1937 Paris Worlds Fair; he manager! to procure some for study and found the concentration of acety~choline esterase in the electric organ to be exceedingly high. In his own words, "The result was simply stunning: ~ g of electric tissue (fresh weight) hyclrolyzecl 3-4 g of acetylcholine per hour, although the tissue is 92% water ant! only 3% protein."3 3 David Nachmansohn, "Biochemistry As Part of My Life," Annual Review of Bio- chemistry 45(1972):1-27.

DAVID NACHMANSOHN 363 The importance of this discovery, which opened the way for the elucidation of the molecular mechanisms involved in the generation of bioelectricity, can hardily be overestimated. In collaboration with Egar Lederer, Nachmansohn soon used the electric organ of the Torpedo fish to purify acety~choline esterase. (This work was reported in a 1939 paper publishect in the Bulletin de la Societe de Chimie Biologique EParis].) In ad- clition, Nachmansohn carries] out experiments on the same organ in June 1939 at the Marine Biological Station at Ar- cachon, near Bordeaux. Together with W. Feldberg (a phar- macologist from Dale's group) ant! A. Fessarc! (an electro- physiologist at the Sorbonne), Nachmansohn provider} the first unequivocal evidence for the electrogenic action of ace- ty~choline; the results were published in 1940 in the Journal of Physiology. His next paper on electric tissue, prepared in colIabora- tion with C. W. Coates and R. T. Cox, was publishecT from Yale in 1941 in the journal of General Physiology. This paper dealt with the correlation between the electrical potential and the acety~choline esterase content of different sections of the electric organ of the electric eel. The use of electric tissue later made possible the crystallization and biochemical char- acterization of acety~choline esterase in Nachmansohn's lab- oratory as well as the isolation of choline acetylase anct the acety~choline receptor. In 1939, John Fulton invitec! Nachmansohn to join his department at Bale University. He stayed in New Haven until 1942, when he mover! to Columbia University and became associated with the Departments of Neurology and Biochem- istry at the College of Physicians and Surgeons. In New Ha- ven, he hac] aIreacly begun to work with the electric organ of the electric eel (which he obtainer! from the New York Aquar- ium) and found not only that the acety~choline esterase con- centration was as high as in Torpedo but that the electric tissue

364 BIOGRAPHICAL MEMOIRS contained phosphocreatine and ATP in concentrations com- parable to those in striated muscle. Furthermore, the electri- cal discharge was accompanied by phosphocreatine break- down. These observations suggested to him that the energy required for resynthesis of the acety~choline hydrolyzect clur- ing the electrical discharge was supplied by the same pro- cesses that provide the energy required for muscular con- traction, namely ATP ant! phosphocreatine breakdown, lactic acid formation, anti, in the last instance, carbohydrate oxi- clation. Soon after Nachmansohn moved to Columbia, he tested the Plea that electric tissue contains enzymes capable of uti- lizing the energy of ATP for the acetylation of choline, an idea that indeed prover! to be the case. This was, in many respects, key because it was the first time that ATP was found to drive a synthetic reaction other than through phosphory- lation. Nachmansohn soon found that choline acetylase, the enzymeks) responsible for the acetylation reaction, requires! a coenzyme because the activity of the acetylase-containing extracts was lost after dialysis ant! was restorer! by the addi- tion of boiled enzyme. The identity of this coenzyme re- mainec! obscure, however, until Lipmann and coworkers found that an enzyme catalyzing the formation of acetyIsul- fonamicle from ATP, acetate; and sulfonamicle also required a coenzyme (coenzyme A, or CoA for short) for activity. They elucidates! the structure of this coenzyme in 1947. The discovery of choline acetylase was published by Nach- mansohn ant! Machaclo in the journal of Neurophysiology in ~ 943. Ironically, three journals (Science, Journal of Biological Chemistry, and Proceedings of the Society for Experimental Biology and Medicine) refused to publish this eminent and trail- blazing biochemical paper. The reviewers apparently could not believe that ATP would participate in reactions other than phosphorylations. In retrospect, they cannot be blamed

DAVID NACHMANSOHN 365 for their skepticism because Nachmansohn's finding was to- tally unexpected. Acetylation was eventually found to result from the coupling of two reactions: (~) ATP + acetate + CoA ~ AMP + inorganic pyrophosphate + acetyI-CoA; and (2) acety! CoA + choline (or sulfonamicle) ~ CoA + acetyI- choline (or acetyI-sulfonamide). Work proceeded in a number of laboratories on the To- calization of acety~choline esterase using biochemical assays (e.g., of the extrucled axoplasm and the sheath of the giant axon of the squid) and electron microscopic observations. The results of these studies macle it appear highly probable that the enzyme was a component of excitable membranes everywhere not only of synaptic membranes but also of the membranes of axons anct conducting fibers in general. In his Harvey Lecture entities! "Metabolism and Function of the Nerve Cell" (cleliverec} in 1953 and publishecl in 1955), Nach- mansohn acivanced the view that acety~choline acts as a signal recognizes! within the membrane by an acety~choline recep- tor protein; this results in a conformational change that leacis to increased local permeability to ions and membrane depo- larization, thus generating an action potential an idea that prover! to be correct. Ernest Schoffeniels, in Nachmansohn's laboratory, was able to isolate the electroplax, the single- celled elementary unit of electric tissue, which was found to be extremely rich in acety~choline esterase ant] receptor protein. If one considers that receptors are now recognized as the initial elements in the response of all cells to specific stimuli and that the concept originated with the acety~choline recep- tor, it becomes evident that Nachmansohn set a biological lancimark. This was also the first neurotransmitter receptor to be characterizes! biochemically, thanks to its accessibility in the vertebrate muscle endplate and its abundance in the spe- cialize{l electric organ of electric fish.

366 BIOGRAPHICAL MEMOIRS The finding that acety~choline esterase activity is very high in excitable membranes including nerve fiber mem- branes ant! that the localization of the acety~choline recep- tor is the same as that of the esterase led Nachmansohn to postulate that the nerve impulse is generated through a de- polarization of the membrane by acety~choline releaser! by the stimulus from an inactive complex with protein. The ac- tion potential thus generated wouIc! give rise to the release of acety~choline in adjacent sites leading to propagation of the current along the fiber by successive acety~choline bursts. Rapid hycirolysis of acety~choline by the esterase anct the ion pump mechanism coupled to the breakdown of ATE wouIc! restore membrane polarization at each point as the impulse travelled down the fiber. Nachmansohn's theory, aIreacly suggested in earlier pub- lications, was presenter! in cletai! in his book, Chemical and Molecular Basis of Nerve Activity, first publisher} in 1959. A revised edition appeared! in 1975 with considerably more ex- perimental support for his Fleas. The revised edition also container! two supplements, one by Nachmansohn on the properties and functions of proteins of the acety~choline cycle in excitable membranes anct one by E. Neumann that presented a molecular mode} for bioelectricity. Nachmansohn's ideas, however, were not accepted by neu- rophysiologists. His molecular theory of nerve conduction is still highly controversial, despite the fact that a variety of experiments by Nachmansohn and others (detailec! in the 1975 edition of his book) would appear to nullify objections to his theory. The fact, for instance, that acety~choline when appliecl locally stimulates at synaptic junctions or motor end- plates but has no effect on axons, may be explainec! by im- permeability of the intact axonal membrane to quaternary ammonium ions. Acety~choline, therefore, stimulates axons when applied at the Ranvier node sites where the myelin

DAVID NACHMANSOHN 367 sheath is much thinner. It also stimulates when applied to areas of a nerve fiber where the phospholipids of the myelin sheath have previously been hydrolyzed by treatment with phospholipase. By the same token, curare which competes with acety~choline for binding to the receptor blocks trans- mission of the nerve impulse across synapses but does not affect conduction when applier! locally to the surface of nerve fibers. It cloes, however, block conduction when applied at Ranvier nodes or to the surface of fibers previously treated with phospholipase. Physostigmine (eserine), a tertiary ammonium base, and prostigmine, a quaternary ammonium base, are both inhibi- tors of acety~choline esterase, anti the former- but not the latter can depress conduction when appliecl to a single frog sciatic nerve fiber. Moreover, the excitable membrane of cer- tain axons (e.g., those of the walking leg of the lobster) ap- pears to be incompletely protectecl; these axons can be stimulated by the local application of acety~choline. Organo- phosphates such as cliisopropyIfluorophosphate (DFP) or tetraethy~pyrophosphate (TEPP) are irreversible inhibitors of acety~choline esterase anti block conduction across syn- apses and along nerve fibers. Both the inhibition of acety~choline esterase and the conduction block can be reversed by pyricline-2-aldoxime (PAM) an organophosphate antagonist developed in Nach- mansohn's laboratory by Irving Wilson as a war gas antidote. (It may be mentioned parenthetically that organophosphates are used commercially as insecticides. PAM has found a non- military application in the systemic treatment of insecticide poisoning. Some local anesthetics are structural analogs of acety~choline and compete with the latter for receptor bind- ing, blocking electrical activity in the conducting and synaptic parts of excitable membranes. Despite these results, the current belief is that the acety]

368 BIOGRAPHICAL MEMOIRS choline system is intercellular anct not intracellular. AcetyI- choline is thought to be liberated only at cholinergic nerve enclings in the synaptic cleft ant! to bind to the receptor on the postsynaptic side, functioning, therefore, exclusively as a synaptic transmitter. Axonal conduction is believecI to involve primarily electric field effects on confirmational transitions of protein-ion channels. The high concentration of acetyI- choline esterase and acety~choline receptor in axonal mem- branes is nevertheless a remarkable fact that remains unex- plained. Nachmansohn's work attracted a great number of stu- dents and investigators, and his laboratory at the College of Physicians anct Surgeons was for many years a place of much excitement and feverish activity. Nearly four hunclre(1 pa- pers, the majority original research papers, were publishecl from his laboratory between 1947 and 1977. In acldition, Nachmansohn was an indefatigable traveler ant! lectured ex- tensively in the United States and abroad. In the spring of 1980, former students, collaborators, and friends of David Nachmansohn organizer! an international symposium at the University of Liege to honor him on his eighty-first birthclay.4 It was apparent at this meeting that the field of endeavor he had pursued so vigorously for many years hac} been expancled in many directions by his former associates and their students. Particularly noteworthy was the tremendous progress in our knowlecige of the molecular structure of acety~choline esterase and of the acety~choline receptor. The importance of Nachmansohn's acety~choline receptor 4 Molecular Aspects of Bioelectricity: Festschrift and Proceedings of the International Sym- posium and Poster Session in Honor of David Nachmansohn on the Occasion of his 81st Birthday, Liege, May 25 - 27, 1980 under the Auspices of the Universite de Liege, Belgium, and the Max-Planck-Institut fur Biochemie, Martinsried, Munchen, Germany. Ed., Ernest Schoffeniels and Eberhard Neumann. Oxford: Pergamon Press, 1980.

DAVID NACHMANSOHN 369 idea for our understancling of the generation of bioelectricity in molecular terms may be gauged from the review by Chan- geux (a former collaborator with Nachmansohn) and his as- sociates,5 anc! a recent reports prepared for the National In- stitute of Mental Health by panels of scientists in various areas of neurobiology ancI relater! fields. In these publica- tions, the monomeric form of the receptor is described as a transmembrane, allosteric protein with an approximate mo- lecular weight of 250,000, containing two acety~choline (ag- onist) binding sites and consisting of four types of polypep- ti~le chains of apparent molecular weights: 39,000 (ax), 4S,000 All, 5S,000 (-y), and 64,000 (~) in a ratio Of ~2939~8 The re- ceptor has several functional states: In the resting state, it has low affinity for agonists, and the ion channel is closed; in the active state, the binding sites are occupied by agonist anc! the channel is open. On binding two molecules of acety~choline, the receptor undergoes rapict transitions (on a submilli- second time scale) between the resting and the active state. These fluctuations last a few milliseconds until hydrolysis of the acety~choline causes its dissociation from the receptor. After his retirement in 1967, Nachmansohn continued to work, travel, anct lecture extensively. He was an enthusiastic supporter of the Zionist cause and macle many visits to Israel. He was very active on behalf of the Hebrew University and the Weizmann Institute and was for many years a member of the board of governors of the latter institution. Nachman- sohn was a firm believer in the worIc3 fraternity of science and was among the first scientists of German-lewish origin to visit Germany after the war, working with strong deter- mination for the reestablishment of scientific ties between 5 ].-P. Changeux, A. Devillers-Thiery, and P. Chemoulli, "Acetylcholine Receptor: An Allosteric Protein," Science 225(1984):1335 - 45. 6 The Neuroscience of Mental Health. U.S. Department of Health and Human Ser- vices Publication no. (ADM)84-1363. Rockville, Md.: 1984.

370 BIOGRAPHICAL MEMOIRS Germany and the West. He also promoted intensely scientific rapprochement between Germany and Israel. In the 1970s Nachmansohn devoted himself to the study of the role played by German-iewish scientists in the explosion of scientific knowledge that took place in the first quarter of this century. This effort culminated in the publication of his book, German- fewish Pioneers in Science: 1900-1933.7 Because of his interest in art and history, David Nach- mansohn was a stimulating travel companion. My wife and ~ enjoyed his company on many a visit to Israel, Italy, Sicily, and Greece, profiting from his scholarly knowledge of the classical world. David had strong convictions and defended them stubbornly, but he never let scientific preoccupations interfere with his enjoyment of life. He was refined in his tastes and gentle and understanding with his friends. ~ AM GREATLY INDEBTED to Arthur Karlin (Columbia University) and Jean-Pierre Changeux (Institut Pasteur) for helpful sugges- tions. 7 David Nachmansohn, German-Jewish Pioneers in Science: 1900-1933 (New York: Springer, 1979).

DAVID NACHMANSOHN HONORS 371 Nachmansohn became a member of the National Academy of Sciences in 1965. He was also a member of the American Academy of Arts and Sciences, the German Academy of Natural Sciences (Leopoldina), and an honorary member of the Weizmann Institute of Sciences of Israel and the Berlin Medical Society. He was a re- cipient of the Pasteur Medal (Paris), the Neuberg Medal (New York), the Medal of the Societe de chimie biologique (Paris), the Albrecht van Graefe Medal of the Berlin Medical Society, the Nicloux Medal (Paris), and the Gold Medal of the Spanish Council for Scientific Research. He received an honorary M.D. degree from the Free University of Berlin and honorary D.Sc. degrees from the University of Liege (Belgium) and Tufts University (Boston). An international symposium on the molecular basis of nerve activity was held at the Free University of Berlin, in October 1984, in memory of David Nachmansohn. That this symposium was sponsored jointly by the Max-Planck-Gesellschaft zur Forderung der Wissenschaften, the Societe franchise de chimie biologique, the Weizmann Institute of Sciences, the Deutsche Forschungsgemein- schaft, the Senator for Science and Research of the City of Berlin, the Free University of Berlin, and the Gesellschaft fur Biologische Chemie, attests to the high esteem in which David Nachmansohn was held by his colleagues and friends.

372 BIOGRAPHICAL MEMOIRS SELECTED BIBLIOGRAPHY 1927 Zur Frage des "Schlafzentrums." Eine Betrachtung der Theorien uber Entstehung des Schlafes. Z. Gesamte Neurol. Psychiatr., 107:342-401. With H. A. Krebs. Vitalfarbung und Adsorption. Biochem. Z., 186:478-84. With P. Rona and H. W. Nicolai. Uber den Fermentstoffwechsel der Bakterien. Biochem. Z., 187:328-43. 1928 Die Entstehung des Schlafes. Med. Klin. (Munich), 31:1192-95. Uber den Zerfall der Kreatinphosphorsaure in Zusammenhang mit der Tatigkeit des Muskels. Biochem. Z., 196:73-97. With O. Meyerhof. Neue Beobachtungen Uber den Umsatz des "Phosphagens" im Muskel. Naturwissenschaften, 16:162. 1929 Sur la relation de la chronaxie musculaire avec la decomposition du phosphate de creatine. C. R. Seances Soc. Biol. Paris, 101: 1086-87. Uber den Zerfall der Kreatinphosphorsaure in Zusammenhang mit der Tatigkeit des Muskels. Biochem. Z., 208:237-56. Uber den Zerfall der Kreatinphosphorsaure in Zusammenhang mit der Tatigkeit des Muskels. Biochem Z., 213:262-300. Uber den Zusammenhang des Kreatinphosphorsaurezerfalls mit Muskelchronaxie und Kontraktionsgeschwindigkeit. Med Klin. (Munich), 42:1-8. With H. H. Weber. Die Unabhangigkeit der Eiweisshydratation von der Eiweissionisation. Biochem. Z., 204:215-52. 1930 Die Guanidinophosphorsauren ("Phosphagene") des Muskels. In: Handbuch Biochemisches des Menschen und der Tiere, Erganzungs- band, pp. 162-74. Uber die Synthese der Kreatinphosphorsaure im lebenden Mu- skel. Biochem. Z., 222:1-20.

DAVID NACHMANSOHN 373 1934 Lactic acid formation in the muscles of adrenalinectomized ani- mals. ~. Physiol. (London), 81:36-37. With R. Debre, G. Semelaigne, and E. Gilbrin. Les hepatomegalies polycoriques. Bull. Mem. Soc. Med. Hop. Paris, 50:1023-41. 1936 With R. Debre, }. Marie, and T. Bernard. Diabete insipide. Etude de ltelimination des chlorures et du pouvoir concentrateur du rein. Bull. Mem. Soc. Med. Hop. Paris, 52:967-75. With R. Debre and }. Marie. Etude chimique du muscle preleve par biopsie dans la myopathic. C. R. Acad. Sci. Paris, 202: 520-22. With A. Dognon and B. S. Levin. Sur la difference de la radiosen- sibilite du foie et du rein isoles du cobaye. C. R. Seances Soc. Biol. Paris, 122: 1083 -84. With R. Debre, ~. Milhit, }. Marie and P. De Font-Reaulx. Accidents nerveux graves et troubles profondes de la glycore gulation chez l'enfant. Bull. Mem. Soc. Med. Hop., Paris, 52:1653-63. With {. Wajzer and R. Lippmann. Action des substances sympatho- et parasympathomimetiques sur les processus chimiques four- nissant ltenergie de la concentration musculaire. Bull. Soc. Chim. Biol., 18: 1207-31. 1937 With A. Marnay. Action des substances sympatho- et parasympa- thomimetiques sur les processus chimiques fournissant l'ener- gie de la contraction musculaire. Bull. Soc. Chim. Biol., 19: 446-52. Action des substances sympatho- et parasympathomimetiques sur les processus chimiques fournissant l'energie de la contraction musculaire. Bull. Soc. Chim. Biol., 19:453-59. With R. Debre, J. Marie, and S. Bidou. Remarques sur deux ob- servations de nephrite chronique de ltenfance avec troubles du developpement ou nanisme renal. (Latence clinique, impor- tance de la polydipsie, troubles des glucides). Bull. Mem. Soc. Med. Hop. Paris, 53:62-70. With A. Marnay. Cholinesterase in voluntary frog's muscle. J. Phys- iol. (London), 89:359-67.

374 BIOGRAPHICAL MEMOIRS With Z. M. Bacq. Cholinesterase in invertebrate muscles. I. Physiol. (London), 89:368 - 371. With A. Marnay. Cholinesterase dans le muscle strie. C. R. Seances Soc. Biol. Paris, 124:942 - 44. With A. Marnay. Sur la repartition de la Cholinesterase dans le muscle couturier de la grenouille. C. R. Seances Soc. Biol. Paris, 125:41-43. With A. Marnay and B. Minz. Cholinesterase dans les terminaisons nerveuses du muscle strie. C. R. Seances Soc. Biol. Paris, 125:43-47. With A. Marnay. Cholinesterase dans le muscle de lezard. C. R. Seances Soc. Biol. Paris, 125:489-90. With A. Marnay. Cholinesterase dans le nerf du homard. C. R. Seances Soc. Biol. Paris, 125:1005-7. Cholinesterase in the central nervous system. Nature (London), 140:427. La transmission de ['influx nerveux dans le systeme nervoux cen- tral. C. R. Seances Soc. Biol. Paris, 126:783-85. With A. Marnay. Cholinesterase dans le muscle strie apres dege- nerescence du nerf moteur. C. R. Seances Soc. Biol. Paris, 126:785-87. 1938 Cholinesterase dans les tissue embryonnaires. C. R. Seances Soc. Biol. Paris, 127:670-73. With A. Marnay. Choline esterase in voluntary muscle. t. Physiol. (London), 92:37-47. Cholinesterase dans le tissu nerveux. C. R. Seances Soc. Biol. Paris 127:894-96. Distribution de la cholinesterase dans le cerveau humain. C. R. Se- ances Soc. Biol. Paris, 128: 24-25. La transmission de ['influx nerveux dans le systeme nervoux cen- tral. Presse Med., 48:942-43. Cholinesterase dans les fibres nerveuses. C. R. Seances Soc. Biol. Paris, 128:516-18. Transmission of nerve impulses in the central nervous system. }. Physiol. (London), 93:2-3. Changements de la cholinesterase dans le muscle strie. C. R. Se- ances Soc. Biol. Paris, 128:599-603. La cholinesterase dans les cultures du coeur de ltembyon chez la poule. C. R. Seances Soc. Biol. Paris, 128:577-79.

DAVID NACHMANSOHN 375 With R. Couteaux. Cholinesterase at the end-plates of voluntary muscle after nerve degeneration. Nature (London), 142:481. Cholinesterase dans le ganglion cervical sympathique superieur du chat. C. R. Seances Soc. Biol. Paris, 129:830-33. Sur ['action de la strychnine. C. R. Seances Soc. Biol. Paris, 129: 941-43. 1939 With E. Lederer. Sur quelques proprietes chimiques de la cholin- esterase. C. R. Seances Soc. Biol. Paris, 130:321-24. Cholinesterase in voluntary muscle. J. Physiol. (London), 95: 29-35. Cholinesterase dans le systeme nerveux central. Bull. Soc. Chim. Biol., 21 :761-96. With E. Lederer. Sur la biochimie de la cholinesterase. I. Prepa- ration de ltenzyme. Groupements-Sh. Bull. Soc. Chim. Biol., 21 :797-808. Sur ['inhibition de la cholinesterase. C. R. Seances Soc. Biol. Paris, 130: 1065-68. 1940 With R. Couteaux. Changes of choline esterase at end plates of voluntary muscle following section of sciatic nerve. Proc. Soc. Exp. Biol. Med., 43:177-81. With W. Feldberg and A. Fessard. The cholinergic nature of the nervous supply to the electrical organ of the torpedo (Torpedo marmorata). J. Physiol. (London), 97:3-5. Choline esterase in brain and spinal cord of sheep embryos. J. Neu- rophysiol., 3:396-402. With E.J. Boell. Localization of choline esterase in nerve fibers. Science, 92:513-14. On the physiological significance of choline esterase. Yale {. Biol. Med., 12:565-89. Action of ions on choline esterase. Nature (London), 145:513-14. Electricity elicited by an organic chemical process. Science, 91. 405-6. 1941 Does acetylcholine act specifically as "synaptic transmitter"? Am. J. Physiol., 133:395-96. .

376 BIOGRAPHICAL MEMOIRS With E. C. Hoff. Choline esterase in the spinal cord of cats after section of dorsal roots. Am. I. Physiol., 133:331. With B. Meyerhof. Relation between electrical changes during nerve activity and concentration of choline esterase. I. Neuro- physiol., 4:348-61. With C. W. Coates and R. T. Cox. Electric potential and activity of choline esterase in the electric organ of Electrophorus electricus (LinnaeusJ. ]. Gen. Physiol., 25:75-88. Electrical potential and activity of choline esterase in nerves. The Collecting Net, 16. With H. B. Steinbach. On the localization of enzymes in nerve fibers. Science, 95:76 -77. 1942 Electrical potential and activity of choline esterase in nerve. Fed. Proc. Fed. Am. Soc. Exp. Biol., 1:62. With H. B. Steinbach. Localization of enzymes in nerves. I. Suc- cinic dehydrogenase and vitamin B.. J. Neurophysiol., 5:109- 20. On the mechanism of transmission of nerve impulses. The Col- lecting Net, 17: 1-6. With T. H. Bullock. Choline esterase in primitive nervous systems. I. Cell. Comp. Physiol., 20:1-4. With R. T. Cox, C. W. Coates, and A. L. Machado. Action potential and enzyme activity in the electric organ of Electrophorus electr?- c~s (Linnae~bs). I. Choline esterase and respiration. l. Neuro- physiol., 5 :499-516. 1943 With R. T. Cox and C. W. Coates. Phosphocreatine as energy source of the action potential. Proc. Soc. Exp. Biol. Med., 52:97-99. With J. F. Fulton. Acetylcholine and the physiology of the central nervous system. Science, 97:569-571. With H. B. Steinbach, A. L. Machado, and S. Spiegelman. Locali- zation of enzymes in nerves. II. Cytochrome oxidase. J. Neu- rophysiol., 6:203-11. Acetylcholine and the mechanism of nerve activity. Exp. Med. Surg., 1:273 - 77. With R. T. Cox, C. W. Coates, and A. L. Machado. Action potential and enzyme activity in the electric organ of Electrophorus electri-

DAVID NACHMANSOHN 377 cus. II. Phosphocreatine as energy source of the action poten- tial. J. Neurophysiol., 6:383-96. With A. L. Machado. The formation of acetylcholine. A new en- zyme "choline acetylase." T Neurophysiol., 6:397-404. With H. M. John and H. Waelsch. Eject of glutamic acid on the formation of acetylcholine. J. Biol. Chem., 150:485-86. With H. Waelsch. On the toxicity of atabrine. Proc. Soc. Exp. Biol. Med., 54:336-38. 1944 With E. C. Hoff. Effects of dorsal root section on choline esterase concentration in spinal cord of cats. I. Neurophysiol., 7:27-36. On the energy source of the nerve action potential. Biol. Bull., 87:158. With M. A. Rothenberg. On the specificity of choline esterase in nervous tissue. Science, 100:454-55. With H. M. John. Inhibition of choline acetylases by cx-keto acids. Proc. Soc. Exp. Biol. Med. 57:361-62. 1945 With H. M. John. Studies on choline acetylase. I. Effect of amino acids on the dialyzed enzyme. Inhibition of cx-keto acids. I. Biol. Chem., 158:157-71. With M. A. Rothenberg. Studies on cholinesterase. I. On the spec- ificity of the enzyme in nerve tissue. J. Biol. Chem., 1 58: 653-66. The role of acetylcholine in the mechanism of nerve activity. In: Vitamins and Hormones, ed. R. S. Harris and K. V. Thimann, vol. 3, pp. 337-77. New York: Academic Press. Chemical mechanism of nervous action. In: Currents in Biochemical Research, ed. D. E. Green, pp. 335-36. New York: Interscience Publishers, Inc. With H. M. John. On the formation of acetylcholine in the nerve axon. Science, 102:250 -51. With H. Schneemann. On the effect of drugs on cholinesterase. J. Biol. Chem., 159:239-40. 1946 With T. H. Bullock and M. A. Rothenberg. Effects of inhibitors of choline esterase on the nerve action potential. J. Neurophysiol., 9:9-22.

378 BIOGRAPHICAL MEMOIRS On the role of acetylcholine in the mechanism of nerve activity. In: Recent Progress in Hormone Research: Proceedings of the Laurentian Hormone Conference, vol. l, pp. l-26. New York: Academic Press. With C. W. Coates and M. A. Rothenberg. Studies on cholinester- ase. II. Enzyme activity and voltage of the action potential in electric tissue. I. Biol. Chem., 163:39 -48. With H. M. John and M. Berman. Studies on choline acetylase. II. The formation of acetylcholine in the nerve axon. J. Biol. Chem., 163:475-80. Chemical mechanism of nerve activity. Ann. N.Y. Acad. Sci., 47:395-428. With M. A. Rothenberg. Chemical aspects of the transmission of the nerve impulses. Prog. Neurol. Psychiatry, 1:59-75. With T. H. Bullock, H. Grundfest, M. A. Rothenberg, and K. Ster- ling. Effect of all-isopropyl fluorophosphate (DFP) on action potential and choline esterase of nerve. J. Neurophysiol., 9: 253-60. With C. W. Coates, M. A. Rothenberg, and M. V. Brown. On the energy source of the action potential in the electric organ of Electrophoru~s electricus. J. Biol. Chem., 165:223-31. With M. Berman. Studies on choline acetylase. III. On the prepa- ration of the coenzyme and its effect on the enzyme. l. Biol. Chem., 165:551-63. With R. Couteaux, H. Grundfest, and M. A. Rothenberg. Effect of all-isopropyl fluorophosphate (DFP) on the action potential of muscle. Science, 104:317. Effects of drugs on axonal conduction and synaptic transmission. Proc. Rudolf Virchow Med. Soc. City N.Y. (memorial issue- Leopold Lichtwitz), Vol. 5: 95 -103. 1947 With T. H. Bullock, H. Grundfest, and M. A. Rothenberg. Gener- ality of the role of acetylcholine in nerve and muscle conduc- tion. J. Neurophysiol., 10:11-21. With T. H. Bullock, H. Grundfest, and M. A. Rothenberg. Effect of all-isopropyl fluorophosphate (DFP) on action potential and cholinesterase of nerve. II. J. Neurophysiol., 10:63-78. With M. Berman and M. S. Weiss. Presence of choline acetylase in striated and cardiac muscle. J. Biol. Chem., 167:295-96. With H. Grundfest and M. A. Rothenberg. Effect of all-isopropyl

DAVID NACHMANSOHN 379 fluorophosphate (DFP) on action potential and cholinesterase of nerve. I I I . J. Neurophysiol., 10: 155 - 64. With M. A. Rothenberg. Studies on cholinesterase. III. Purification of the enzyme from electric tissue by fractional ammonium sul- fate precipitation. I. Biol. Chem., 168:223 -31. With M. A. Rothenberg and E. A. Feld. The in vitro reversibility of cholinesterase inhibition by all-isopropyl fluorophosphate (DFP). Arch. Biochem., 14: 197-211. With E. A. Feld. Studies on cholinesterase. IV. On the mechanism of all-isopropyl fluorophosphate (DFP) action in viva. J. Biol. Chem., 171:715-24. Difference between drug effects on axonal conduction and synaptic transmission. Trans. Am. Neurol. Assoc., 72:42 - 46. 1948 With M. S. Weiss. Studies on choline acetylase. IV. Effect of citric acid. I. Biol. Chem., 172:677-97. With M. A. Rothenberg and E. A. Feld. Rate of penetration of elec- trolytes into nerve fibers. I. Biol. Chem., 172:345-46. Effect of inhibitors of cholinesterase on conduction in nerve and muscle. In: Proceedings of the Seventeenth International Congress on Physiology, Oxford, England. With M. A. Rothenberg and D. B. Sprinson. Site of action of acetyl- choline. J. Neurophysiol., 11: 111-16. With E. A. Feld, H. Grundfest, and M. A. Rothenberg. Effect of all-isopropyl fluorophosphate (DFP) on action potential and cholinesterase of nerve. IV. l. Neurophysiol., 11: 125-32. With M. A. Rothenberg and E. A. Feld. Studies on cholinesterase. V. Kinetics of the enzyme inhibition. I. Biol. Chem., 1 74: 247-56. The role of acetylcholine in conduction. Bull. Johns Hopkins Hosp., 83:463-94. 1949 With K.-B. Augustinsson. Substrate concentration and specificity of choline ester-splitting enzymes. Arch. Biochem., 23:111-26. With K.-B. Augustinsson. Studies on cholinesterase. VI. Kinetics of the inhibition of acetylcholinesterase. J. Biol. Chem.., 179: 543-59.

380 BIOGRAPHICAL MEMOIRS With K.-B. Augustinsson. Distinction between acetylcholinesterase and other choline ester-splitting enzymes. Science, 110:98-99. With S. Hestrin. The reaction of acetylcholine and other carboxylic acid derivatives with hydroxylamine and its analytical applica- tion. I. Biol. Chem., 180:149-61. With S. Hestrin and H. Voripajeff. Enzymatic synthesis of a com- pound with acetylcholine-like biological activity. }. Biol. Chem., 180:875-87. With S. Hestrin. Acylation reactions mediated by purified acetyl- choline esterase. i. Biol. Chem., 180: 879-81. With S. Middleton and H. H. Middleton. The acetylcholine-like action of a product formed by an acetylating enzyme system derived from brain. Proc. Soc. Exp. Biol., 71:523-26. 1950 With S. R. Korey. Some factors influencing the contracility of a non-conducting fiber preparation. Biochim. Biophys. Acta, 4:48 - 57. (Also in: Metabolism and Function: Otto Meyerhof Anni- versary Volume. New York: Elsevier.) Chemical control of nervous activity, A. Acetylcholine. In: Hor- mones, vol. 2, ed. G. Pincus and K. V. Thimann, pp. 515-99. New York: Academic Press. Studies on permeability in relation to nerve function. I. Axonal conduction and synaptic transmission. Biochim. Biophys. Acta, 4:78-95. (Also in: Metabolism and Function: Otto Meyerhof Anni- versary Volume. Amsterdam: Elsevier.) With M. A. Rothenberg. Studies on permeability in relation to nerve function. II. Ionic movements across axonal membranes. Biochim. Biophys. Acta, 4:96-114. (Also in: Metabolism and Function: Otto Meyerhof Anniversary Volume. Amsterdam: Else- vier.) With S. Hestrin. Acylation reactions mediated by purified acetyl- choline esterase. Biochim. Biophys. Acta, 4:310-21. (Also in: Metabolism and Function: Otto Meyerhof Anniversary Volume. Am- sterdam: Elsevier.) Electric currents in nerve tissue and in electric organs. Electr. Eng., 69:231-34. With I. B. Wilson and F. Bergmann. Studies on cholinesterase. VII. The active surface of acetylcholinesterase derived from effects of pH on inhibitors. J. Biol. Chem., 185 :479-89.

DAV I D N A C H M A N S O H N 381 With I. B. Wilson and F. Bergmann. Acetylcholinesterase. VIII. Dissociation constants of the active groups. l. Biol. Chem., 186:683-92. With F. Bergmann and I. B. Wilson. Acetylcholinesterase. IX. Structural features determining the inhibition by amino acids and related compounds. I. Biol. Chem., 186:193-203. With F. Bergmann and I. B. Wilson. The inhibitory effect of stil- bamidine, curare and related compounds and its relationship to the active groups of acetylcholinesterase. Action of stilbami- dine upon nerve impulse conduction. Biochim. Biophys. Acta, 6:217-24. With I. B. Wilson and F. Bergmann. Acetylcholinesterase. X. Mechanism of the catalysis of acylation reactions. I. Biol. Chem., 186:781-90. 1951 With S. R. Korey. Effect of dilantin and mesantoin on the giant axon of the squid. Proc. Soc. Exp. Biol. Med., 76:297-99. With S. R. Korey and B. de Braganza. Choline acetylase. V. Estar- ifications and transacetylations. I. Biol. Chem., 189:705-15. With I. B. Wilson. Acetylcholinesterase. XI. Reversibility of terra- ethyl pyrophosphate inhibition. I. Biol. Chem., 190:111-17. With I. B. Wilson. Mechanism of enzymic hydrolysis. I. Role of the acidic groups in the esteratic site of acetylcholinesterase. Biochim. Biophys. Acta, 7:466-740. With I. B. Wilson. The enzymic hydrolysis and synthesis of acetyl- choline. In: Advances in Enzymology, vol. 12, pp. 259-339. New York: Interscience. Energy sources of bioelectricity. In: Phosphorus Metabolism, vol. 1, ed. W. D. McElroy and B. Glass, pp. 568-85. Baltimore: The Johns Hopkins University Press. With I. B. Wilson. Mechanism of hydrolysis. II. New evidence for an acylated enzyme as intermediate. Biochim. Biophys. Acta, 7:520-25. With S. R. Korey and R. Mitchell. Studies on permeability in rela- tion to nerve function. III. Permittivity of brain cortex slices to glycine and aspartic acid. Biochim. Biophys. Acta, 7:507- 19. Otto Meyerhof 1884-1951. Proc. Rudolf Virchow Med. Soc. City N.Y., 10:89 - 91.

382 BIOGRAPHICAL MEMOIRS 1952 Chemical mechanism of nerve activity. In: Modern Trends of Physi- ology and Biochemistry, ed. E. S. G. Barron, pp. 229 - 76. New York: Academic Press. With I. B. Wilson, S. Levine, and I. Freiberger. Effects of electrical charge upon the activity of liver esterase. I. Biol. Chem., 194:613-17. With I. B. Wilson, S. R. Korey, and R. Berman. Choline acetylase. VI. Substitution of ATP-acetate by thiolacetate. I. Biol. Chem., 195:25-36. With H. Grundfest, C. Y. Kao, and R. Chambers. Mode of blocking of axonal activity by curare and inhibitors of acetylcholinester- ase. Nature (London), 169: 190. With S. Ochoa and F. A. Lipmann. Otto Meyerhof: 1884-1951. Science, 115:365 -69. The neuromuscular junction. In: Le Muscle: Etudes de Biology et de Pathologic (compte rendu du collogue tenu a Royaumont, France, 8/31 to 9/6, L'expansion), pp. 121-72. With I. B. Wilson. Acetylcholinesterase. XII. Further studies of binding forces. I. Biol. Chem., 197:215-25. With S. Korkes, A. Del Campillo, S. R. Korey, I. R. Stern, and S. Ochoa. Coupling of acetyl donor systems with choline acetylase. I. Biol. Chem., 198:215-20. Nerve function and irradiation effects. I. Cell. Comp. Physiol., 39: 137-78. With I. B. Wilson. Acetylcholinesterase the mechanism of en- zyme activity. Baskerville Chem. J., 3:7-12. Metabolisme et fonction de la cellule nerveuse. Bull. Soc. Chim. Biol., 34:447-65. With I. B. Wilson. Acetylcholinesterase. XIII. Reactivation of alkyl phosphate-inhibited enzyme. J. Biol. Chem., 199: 113 -20. With S. R. Korey. Studies on permeability in relation to nerve func- tion. IV. Effect of glutamate and aspartate upon the rate of entrance of potassium into brain cortical slices. Biochim. Bio- phys. Acta, 9:633-35. With I. B. Wilson. Preparation of acetyl coenzyme A. I. Am. Chem. Soc., 74:3205-6. La conduction de ['influx nerveux et la transmission synaptique.

DAVID NACHMANSOHN 383 In: Estratto Dai Rendiconti Dell'Istituto Superiore de Sanita, vol. 15, pp. 1 267-30 1. 1953 With I. B. Wilson and M. Cohen. The essentiality of acetylcholin- esterase in conduction. Biochim. Biophys. Acta, 11:147-56. Transmission of nerve impulses across the neuromuscular junc- tion. In: Proceedings of the First and Second Medical Conferences (1951-1952) of the Muscular Dystrophy Association of America, Inc., ed. A. T. Milhorat, pp. 2-15. New York: Muscular Dystrophy Assn., Inc. With M. Altamirano, C. W. Coates, and H. Grundfest. Mechanisms of bioelectric activity in electric tissue. I. The response to indi- rect and direct stimulation of electroplaques of Electrophorus electricus. ]. Gen. Physiol., 37 :91-1 10. With I. B. Wilson and E. K. Meislich. Reactivation of acetylcholin- esterase inhibited by alkylphosphates. J. Am. Chem. Soc., 75: 4628. With R. Berman and I. B. Wilson. Choline acetylase specificity in relation to biological function. Biochim. Biophys. Acta., 12: 315-24. 1954 With I. B. Wilson. The mechanism of enzyme hydrolysis studied with acetylcholinesterase. In: The Mechanism of Enzyme Action, ed. W. C. McElroy and B. Glass, pp. 642-57. Baltimore: The Johns Hopkins University Press. With I. B. Wilson. The active surface of the serum esterase. I. Biol. Chem., 208:123-32. With R. Berman-Reisberg. Sulfl~ydryl groups of choline acetylase. Biochim. Biophys. Acta., 14:442-43. With I. B. Wilson. The generation of bioelectric potentials. In: Ion Transport Across Membranes, ed. H. T. Clarke, pp. 35-64. New York: Academic Press. With I. B. Wilson and E. Cabib. Is acetylcholinesterase a metallo enzyme? i. Am. Chem. Soc., 76:5154. With S. L. Friess, I. B. Wilson, and E. Cabib. On the Mg (II) acti- vation of acetylcholinesterase. I. Am. Chem. Soc., 76:5156.

384 BIOGRAPHICAL MEMOIRS 1955 With M. A. Eisenberg. The acetate-activating enzyme of Rhodospi- 7illum rubrum. Biochim. Biophys. Acta, 16:58-65. Metabolism and function of the nerve cell. In: Harvey Lectures, 1953-1954, pp. 57-99. New York: Academic Press. Acetylcholine and energy transformations in nerve cells. In: A Text- book of Physiology, 17th ea., ed. J. F. Fulton, pp. 192-204. Phila- delphia: W. B. Saunders Co. With M. Altamirano, W. L. Schleyer, and C. W. Coates. Electrical activity in electric tissue. I. The difference between tertiary and quaternary nitrogen compounds in relation to their chemical and electrical activities. Biochim. Biophys. Acta, 16:268-83. Stroffwochsel und Funktion der Nervenzelle. Dtsch. Med. Woch- enschr., 80: 196-98. With W. L. Schleyer. Electrical activity in electric tissue. II. Evalu- ation of esterase activity in intact electroplax. Biochim. Biophys. Acta., 16:396-403. With I. B. Wilson and S. Ginsburg. Reactivation of acetylcholines- terase inhibited by alkylphosphates. Arch. Biochem. Biophys., 54:569-71. With A. Weber. Value of models for understanding of muscular contraction. Am. J. Phys. Med., 34:19-32. Mechanisms of impulse transmission across neuromuscular junc- tions. Am. J. Phys. Med., 34:33-45. With W. Hasselbach and A. Weber. Models for the study of the contraction of muscle and of cell protoplasm. Pharmacol. Rev., 7:97-117. With M. Altamirano, C. W. Coates, and H. Grundfest. Electrical activity in electric tissue. III. Modifications of electrical activity by acetylcholine and related compounds. Biochim. Biophys. Acta, 16:449-63. With I. B. Wilson. Molecular basis for generation of bioelectric po- tentials. In: Electrochemistry in Biology and Medicine, ed. T. Shed- lovsky, pp. 167-86. New York: {ohn Wiley & Sons. With I. B. Wilson. Choline acetylase. In: Methods in Enzymology, vol. 1, ed. S. P. Colowick and N. O. Kaplan, pp. 619-24. New York: Academic Press. With I. B. Wilson. Acetylcholinesterase. In: Methods in Enzymology,

DAVID NACHMANSOHN 385 vol. 1, ed. S. P. Colowick and N. O. Kaplan, pp. 642-51. New York: Academic Press. With I. B. Wilson. Reactivation of human serum esterase inhibited by alkylphosphates. J. Am. Chem. Soc., 77:2383-86. Metabolism and function of the nerve cell. In: Neurochemistry, ed. K. A. C. Elliott, I. H. Page, and I. H. Quastel, pp. 399-425. Springfield, Ill.: Charles C Thomas. With I. B. Wilson and E. K. Meislich. The reactivation of acetyl- cholinesterase inhibited by tetraethyl pyrophosphate and di- isopropyl fluorophosphate. l. Am. Chem. Soc., 77:4286-91. With I. B. Wilson and S. Ginsburg. A powerful reactivator of al- kylphosphate-inhibited acervlcholinesterase Biochim Hionhvs Acta,18:168-70. —--or-- -- The generation of bioelectric potentials. Circ. Res., 3:429-33. Principles for testing drug effects during growth. In: Biochemistry of the Developing Nervous System; Proceedings of the First Inter- national Neurochemical Symposium, Oxford, England, July 1954, pp. 479 - 99. New York: Academic Press. Die Role des Acetylcholins in den Elementarvogaengen der Ner- venleitung. In: Ergebnasse der Physiology, vol. 48, pp. 575-683. Heidelberg: Springer-Verlag. With M. Altamirano. Properties of the innervated membrane of the electroplax of electric eel. I. Cell. Comp. Physiol., 46: 249-78. With I. B. Wilson. The interaction of tensilon and neostigmine with acetylcholinesterase. Arch. Int. Pharmacodyn. Ther., 104: 204-13. With I. B. Wilson. Promotion of acetylcholinesterase activity by the anionic site. Faraday Discuss. Chem. Soc., 20: 119-25. . . 1956 With H. Kewitz and I. B. Wilson. A specific antidote against lethal alkylphosphate intoxication. Arch. Biochem. Biophys., 60: 261-63. With I. B. Wilson and E. Cabib. Acetylcholinesterase: enthalpies and entropies of activation. I. Am. Chem. Soc., 78:202-7. With M. Cohen. Concentration of choline acetylase in conducting tissue. Arch. Biochem. Biophys., 60:284-96. With A. Weber. The ultracentrifugal separation of L-myosin and

386 BIOGRAPHICAL MEMOIRS actin in an actomyosin sol under the influence of AT P. Biochim. Biophys. Acta, 19:345-51. With I. B. Wilson. Chemical control of ion movements during nerve activity. In: Proceedings of the Third International Congress of Biochemistry, Brussels, 1955, ed. C. Liebecq, pp.440-44. New York: Academic Press. With I. B. Wilson and M. Altamirano. Action of tertiary and qua- ternary nitrogen derivatives upon the acetylcholine receptor. In: Neurochemistry, ed. S. R. Korey, pp. 155-68. New York: Hoeber-Harper. With I. B. Wilson. Trends in the biochemistry of nerve activity. In: Currents in Biochemical Research 1956, ed. D. E. Green, pp. 628— 52. New York: Interscience. With M. Altamirano. Effect of acetylcholine in the electroplax of electric eel. Biochim. Biophys. Acta, 20:323-36. With H. Kewitz and I. B. Wilson. A specific antidote against lethal alkyl phosphate intoxication. II. Antidotal properties. Arch. Biochem. Biophys., 64:456-65. 1957 With M. A. Eisenberg. The acetate-activating mechanism of Rho- dospirillum rubrum. Biochim. Biophys. Acta, 23 :327-32. With S. Ginsburg and I. B. Wilson. Oximes of the pyridine series. I. Am. Chem. Soc., 79:481-85. With H. Kewitz. A. specific antidote against lethal alkyl phosphate intoxication. III. Repair of chemical lesion. Arch. Biochem. Biophys., 66:263 - 70. With H. Kewitz. A specific antidote against lethal alkyl phosphate intoxication. IV. Effects in brain. Arch. Biochem. Biophys., 66:271-83. With R. Berman-Reisberg. Properties and biological significance of choline acetylase. Yale }. Biol. Med., 29:403-35. With M. Altamirano and C. W. Coates. Effect of potassium on elec- troplax of Electrophorus electricus. ]. Cell. Comp. Physiol., 49: 69-102. With I. B. Wilson and F. Sondheimer. A specific antidote against lethal alkyl phosphate intoxication. V. Antidotal properties. Arch. Biochem. Biophys., 69:468-74. Etudes sur la conduction de ['influx nerveux au niveau molecu- laire. Bull. Soc. Chim. Biol., 39:1021-35.

DAVID NACHMANSOHN 387 Carl Neuberg, 1877-1956. Proc. Rudolf Virchow Med. Soc. City N.Y., 15:75 -82. With E. Schoffeniels. An isolated single electroplax preparation. I. New data on the effect of acetylcholine and related compounds. Biochim. Biophys. Acta, 26:1-15. With E. Schoffeniels. An isolated single electroplax preparation. II. Improved preparation for studying ion flux. Biochim. Bio- phys. Acta, 26:585-96. 1958 With I. B. Wilson and C. Quan. Acetylcholinesterase studies on molecular complementariness. Arch. Biochem. Biophys., 73: 131-43. With M. A. Eisenberg. Intermediate metabolism of electric tissue in relation to function. I. Glycolytic enzymes and succinic oxi- dase. Arch. Biochem. Biophys., 74:372-89. With I. B. Wilson. Designing of a new drug with antidotal prop- erties against the nerve gas Sarin. Biochem. Biophys. Acta. 27: 196-99. With E. Schoffeniels Potassium concentration and potential differ- ence in the single isolated electroplax of the electric eel. Biochim. Biophys. Acta, 27:660. With E. Schoffeniels. Electrical activity of isolated single electroplax of electric eel as affected by temperature. Science, 127: 1117-18. With E. Schoffeniels. A method for studying separately the prop- erties of the innervated and non-innervated membrane of an isolated single electroplax of the Skate. Nature (London), 181: 287-88. With E. Schoffeniels and I. B. Wilson. Overshoot and block of con- duction by lipid soluble acetylcholine analogues. Biochim. Bio- phys. Acta, 27:629-33. With I. B. Wilson, S. Ginsburg, and C. Quan. Molecular comple- mentariness as basis for reactivation of alkylphosphate inhib- ited enzyme. Arch. Biochem. Biophys., 77:286-96. With I. B. Wilson. A specific antidote for nerve gas and insecticide (alkylphosphate) intoxication. Neurology (Suppl. 1), 8:41-43. Molecular forces controlling ion movements during nerve activity. In: Proceedings of the Fourth International Congress of Biochemistry. Vol. 3, Biochemistry of the Central Nervous System, pp. 26-35. Lon- don: Pergamon Press.

388 BIOGRAPHICAL MEMOIRS With W.-D. Dettbarn and I. B. Wilson. Action of lipid soluble qua- ternary ammonium ions on conduction membrane. Science, 128: 1275-76. Acetylcholine, nerve gases and an antidote. (Loewi Festschrift.) Med. Circle Bull., 5~7~:6-8. 1959 With F. C. G. Hoskin. Intermediate metabolism of electric tissue in relation to function. II. Comparison of glycolysis rates in organs of Electrophor~s electr~cus. Arch. Biochem. Biophys., 81 :330-39. With L. P. Hinterbuchner and I. B. Wilson. Muscle response to long chain quaternary ammonium ions. II. Biochim. Biophys. Acta, 32:375-80. With W.-D. Dettbarn. Action of lipid soluble quaternary ammo- nium ions on the resting potential nerve fibers of the frog. Biochim. Biophys. Acta, 32:381-86. With I. B. Wilson and S. Ginsburg. Reactivation of alkylphosphate inhibited acetylcholinesterase by his quaternary derivatives of 2-PAM and 4-PAM. i. Biochem. Pharmacol., 1:200-206. With L. P. Hinterbuchner and I. B. Wilson. Muscle response to long chain quaternary ammonium ions. I. Biochim. Biophys. Acta, 31 :323-27. Chemical factors controlling ion movements during nerve activity. In: The Method of Isotopic Tracers Applied to the Study of Active Ion Transport (Premier Colloque de Biologie de Saclay), pp. 63-87. London: Pergamon Press. Role of acetylcholine in axonal conduction and neuromuscular transmission. (Utrecht Symposium.) Am. I. Phys. Med., 38: 190-206. With H. C. Lawler. A simplified procedure for the partial purifi- cation of acetylcholinesterase electric tissue. I. Biol. Chem., 234:799-801. With E. Schoffeniels. Ion movements studied with single isolated electroplax. Ann. N.Y. Acad. Sci., 81:285-306. With W.-D. Dettbarn. Distinction between sodium and potassium in change in permeability effected by lipid-soluble analogues of acetylcholine. Nature (London), 183 :465-66. With S. Ehrenpreis. Interaction of curare and related substances with acetylcholine receptor-like protein. Science, 1 ?9: 1613 -14.

DAVID NACHMANSOHN 389 With A. Weber. On the role of calcium in the activity of adenosine 5-triphosphate hydrolysis by actomyosin. J. Biol. Chem., 234: 2764-69. With I. B. Wilson. Molecular complementarily in antidotes for nerve gases. Ann. N.Y. Acad. Sci., 81:307-16. Basic problems of drug action on the myoneural junction. Anes- thesiology, no. 4, 20:421-38. With F. C. G. Hoskin. Intermediate metabolism of electric tissue in relation to function. III. Oxidation of substrates by tissues of Electrophorus electr~cus as compared to other vertebrates. Arch. Biochem. Biophys., 85:141-48. With I. B. Wilson. Molecular complementarily and antidotes for alkylphosphate poisoning. Fed. Proc. Fed. Am. Soc. Exp. Biol., no. 2, part 1, 18: 752-58. Chemical and Molecular Basis of Nerve Activity. New York: Academic Press. 1960 The neuromuscular junction. B. The role of the acetylcholine sys- tem. In: The Structure and Function of Muscle, vol. 2, ed. G. H. Bourne, pp. 199-302. New York: Academic Press. With I. B. Wilson. Aspects of the molecular basis of nervous activ- ity. In: Molecular Biology. Elementary processes of nerve conduction and muscle contraction, pp. 163-71. New York: Academic Press. With W.-D. Dettbarn. New evidence for the role of acetylcholine in conduction. Biochim. Biophys. Acta, 41:337-86. With P. Rosenberg. In viva reactivation by PAM of brain cholines- terase inhibited by Paraoxon. Biochem. Pharmacol., 3:312-19. With F. C. G. Hoskin. Effect of inhibitors on the metabolism of specifically labelled glucose by brain. Biochim. Biophys. Acta, 40:309-13. With R. Whittam and M. Guinnebault. The efflux of potassium from electroplax of electric eels. I. Gen. Physiol., 43: 1171-91. With R. Whittam and M. Guinnebault. The effect of blocking elec- trical activity on the efflux of potassium from electroplax. Biochim. Biophys. Acta, 45:336-47. With W.-D. Dettbarn. The effect of curare on conduction in mye- linated, isolated nerve fibers of the frog. Nature (London), 186:891-92.

390 BIOGRAPHICAL MEMOIRS With F. C. G. Hoskin. A source of error in the use of radioactive substrates for metabolic studies. Arch. Biochem. Biophys., 87: 151-52. The aims of the symposium. In: Molecular Biology. Elementary pro- cesses of nerve conduction and muscle contraction, pp. 13-16. New York: Academic Press. With S. Ehrenpreis. Isolation and identification of the acetylcho- line receptor protein of electric tissue. Biochim. Biophys. Acta, 44:56 1-77. With P. Rosenberg and H. Higman. An improved isolated single electroplax preparation. I. Effect of compounds acting primar- ily at the synapses. Biochim. Biophys. Acta, 44:151-60. With S. Ehrenpreis and M. M. Fishman. The interaction of quaternary ammonium compounds with chondroitin sulfate. Biochim. Biophys. Acta, 44:577-85. With V. G. Longo and D. Bovet. Aspects electronoencephalogra- phiques de l'antagonisme entre le iodomethylate de 2-pyridine aldoxime (PAM) et le methylfluorophosphate dtisopropyle (Sarin). Arch. Int. Pharmacodyn. Ther., 123:282-90. With L. P. Hinterbuchner. Electrical activity evoked by a specific chemical reaction. Biochim. Biophys. Acta, 44:554-60. Chemical and molecular forces underlying nerve activity. (In me- moriam: Peter Rona.) Arzneim.-Forsch., 10:387-90. With P. Rosenberg and H. B. Higman. An improved isolated single electroplax preparation. II. Compounds acting on the con- ducting membrane. Biochim. Biophys. Acta, 45:348-54. With W.-D. Dettbarn, H. B. Higman, and P. Rosenberg. Rapid and reversible block of electrical activity by powerful marine biotox- ins. Science, 132:300-301. With S. Ehrenpreis and M. G. Kellock. Acetylcholine receptor pro- tein and nerve activity. I. Specific reaction of local anesthetics with the protein. Biochem. Biophys. Res. Commun., 2:31 1-15. With E. Battels, W.-D. Dettbarn, H. B. Higman, and P. Rosenberg. Acetylcholine receptor protein and nerve activity. II. Cationic group in local anesthetics and electrical response. Biochem. Biophys. Res. Commun., 2:316-19. With S. Ehrenpreis and M. G. Kellock. The interaction of quater- nary ammonium compounds with hyaluronic acid. Biochim. Biophys. Acta, 45 :525-28.

DAVID NACHMANSOHN 391 With S. Ochoa and F. A. Lipman. Otto Meyerhof, 1894-1951. In: Biographical Memoirs, vol. 34, pp. 153-82. New York: Columbia University for the National Academy of Sciences. With F. C. G. Hoskin. Chemical stimulation and modifications of glucose metabolism by brain. Arch. Biochem. Biophys., 91: 43-46. 1961 Biochemical basis of nerve activity. In: Radioactive Isotopes in Physi- ology, Diagnostics and Therapy, vol. 2, 2d ea., ed. H. Schwiegk and F. Turba, pp. 229-51. Heidelberg: Springer-Verlag. Chemical and molecular aspects of bioelectrogenesis. In: Bioelectro- genes~s: Proceedings of the Symposium on Comparative Bioelectrogen- s~s, ed. C. Chagas and A. Paes de Carvalho, pp. 237-61. New York: Elsevier Publ. Co. With W.-D. Dettbarn. New evidence for the role of acetylcholine in bioelectrogenesis. In: Bioelectrogenes~s: Proceedings of the Sympo- sium on Comparative Bioelectrogenesis, ed. C. Chagas and A. Paes de Carvalho, pp. 262-87. New York: Elsevier Publ. Co. With S. Ehrenpreis. The isolation and identification of the acetyl- choline receptor protein from electric tissue of Electrophorus electr~cus. In: Bioelectrogenes~s: Proceedings of the Symposium on Comparative Bioelectrogenesis, ed. C. Chagas and A. Paes de Car- valho, pp. 379-96. New York: Elsevier Publ. Co. Chemical and molecular forces controlling ion movements. In: Problems of the Evolution and Enzymochem~stry of Excitation Processes (Koshtoyants Memorial Volume), ed. T. M. Turpajew, pp. 215- 28. Moscow: USSR Academy of Sciences. The role of acetylcholine in nerve activity. In: Glaucoma: Transac- tions of the Fifth Conference, ed. W. Newell, pp. 137-91. New York: iosiah Macy ~r. Foundation. With R. Whittam. Some effects of electrical activity and depolariz- ing agents on the efflux of potassium from electroplax of elec- tric eels. In Bioelectrogenes~s: Proceedings of the Symposium on Com- parative Bioelectrogenesas, ed. C. Chagas and A. Paes de Carvalho, pp. 166-68. New York: Elsevier Publ. Co. With P. Rosenberg and S. Ehrenpreis. Reversible block of axonal conduction by curare after treatment with cobra venom and a detergent. Nature (London), 190: 728-29.

392 BIOGRAPHICAL MEMOIRS Le probleme de role l'acetylcholine dans l'activite nerveuse a l'etat actuel. In: Actualites Neurophysiologiques, vol. 3, ed. A. M. Mon- nier, pp. 299-337. Paris: Masson. With H. C. Lawler. Turnover time of acetylcholinesterase. l. Biol. Chem., 236:2296-301. With P. Rosenberg and S. Ehrenpreis. Reversible block of axonal conduction by curare after treatment with cobra venom. Bio- chem. Pharmacol., 8:192-206. Chemical factors controlling ion movements during nerve activity. Koshtoyants volume, English edition, ed. }. W. S. Pringle. Ox- ford: Pergamon Press. With W.-D. Dettbarn and F. C. G. Hoskin. Changes of glucose me- tabolism during lobster nerve activity. Biochim. Biophys. Acta, 50:568-70. With W. H. Harrison. Enzymic reactions competing with noradren- aline N-methyl transferase. Biochim. Biophys. Acta, 50:202-4. With A. M. Gold. Synthesis of a series of organophosphorus esters containing alkylating groups. J. Org. Chem., 26:3991-94. Chemical factors controlling nerve activity. Science, 134: 1962-68. With H. B. Higman and E. Bartels. The competitive nature of the action of acetylcholine and local anesthetics. Biochim. Biophys. Acta, 54:543 -54. 1962 Chemical and molecular basis of nerve activity. In Neurochem?stry, ed. K. A. C. Elliot, I. H. Page, and J. H. Quastel, pp. 522-57. Springfield, Ill.: Charles C Thomas. With W.-D. Dettbarn. The active form of local anesthetics. Biochim. Biophys. Acta, 57:73-76. With H. B. Higman and E. Bartels. New method for record- ing electrical characteristics of the monocellular electroplax. Biochim. Biophys. Acta, 57:77-82. Nerve activity, chemical basis of. In: McGraw-Hill Yearbook of Science and Technology, pp. 352-56. New York: McGraw-Hill. With W.-D. Dettbarn. Acetylcholinesterase actibity in Nitella. Na- ture (London), 1 94: 1 175 - 76. With W.-D. Dettbarn and F. A. Davis. Effect of acetylcholine on the electrical activity of somatic nerves of the lobster. Science, 136:716-17.

DAVID NACHMANSOHN 393 The propagation of nerve impulses. Nature's mechanism of mes- sage transmission. Yale Sci., 36~5) :20 -26. Basic aspects of nerve activity explained by biochemical analysis. I. Am. Med. Assoc., 179~8) :639-43. With F. C. G. Hoskin. Specificity of the stimulation by quinones of direct oxidation of glucose by brain slices. Biochim. Biophys. Acta, 62: 11-16. With P. Rosenberg and T. R. Podleski. Block of axonal conduction by acetylcholine and d-tubocurarine after treatment with cot- tonmouth moccasin venom. I. Pharmacol. Exp. Ther., 137:249- 62. Answer to Ehrenpreis. Science, 136: 177-81. With W.-D. Dettbarn and F. C. G. Hoskin. Electrical and esterase activity in axons. Biochim. Biophys. Acta, 62:566-73. With H. Greenberg. Isolation of serine phosphate from the active site of human prostatic acid phosphorase; inhibition of the en- zyme by DFP. Biochem. Biophys. Res. Commun., 7:186-89. With F. A. Davis and W.-D. Dettbarn. Depolarizing action of cal- cium-ion depletion on frog nerve and its inhibition by com- pounds acting on the acetylcholine system. Biochim. Biophys. Acta, 63:349-57. With W.-D. Dettbarn and F. A. Davis. "Sucrose gap" technique ap- plied to single-nerve-fiber preparation. Biochim. Biophys. Acta, 60:648-50. With T. R. Podleski. Revised distinction direct and indirect re- sponse of electroplax. Biochim. Biophys. Acta, 63:358-64. With E. Bartels. Structure-activity relationship studied on the iso- lated single electroplax. Biochim. Biophys. Acta, 63:365-73. With W.-D. Dettbarn and P. Rosenberg. Sources of error in relating electrical and acetylcholinesterase activity. Biochem. Pharma- col.,ll:1025-30. With W.-D. Dettbarn and P. Rosenberg. Acetylcholinesterase in aplysia. Biochim. Biophys. Acta, 65:362. 1963 Choline acetylase. In: Handbuch der experimentellen Pharmakologie, Erg. Bd. 15, ed. G. Koelle, pp. 40-54. Heidelberg: Springer- Verlag. Discussion remarks. In: Proceedings of the First International Phar-

394 BIOGRAPHICAL MEMOIRS macolog~cal Meeting, Stockholm, 1961, vol. 7, pp. 134-43. Ox- ford: Pergamon Press. Actions on axons and the evidence for the role of acetylcholine in axonal conduction. In: Handbuch der experimentellen Pharmako- log7e, Erg. Bd.15, ed. G. Koelle, pp.701-40. Heidelberg: Sprin- ger-Verlag. With H. C. Lawler. Purification and properties of an acetylcholin- esterase polymer. l. Biol. Chem., 238:132-37. Facteurs chimiques controlant les mouvements ioniques pendant l'activite nerveuse. Bull. Soc. Chim. Biol., 45:29-54. With P. Rosenberg and W.-D. Dettbarn. Ester splitting activity of the electroplax. Biochim. Biophys. Acta, 69:103-14. With F. C. G. Hoskin and C. von Eschen. Action of arylsulfatase on vitamin K3 disulfate. Biochim. Biophys. Acta, 67:669-71. With D. E. Fahrney and A. M. Gold. Sulfonyl fluorides as inhibi- tors of esterases. I. Rates of reaction with acetylcholinesterase, c~-chymotrypsin, and trypsin. l. Am. Chem. Soc., 85:997-1000. With P. Rosenberg and F. C. G. Hoskin. Demonstration of in- creased permeability as a factor responsible for the effect of acetylcholine on the electrical activity of venom treated axons. J. Gen. Physiol., 46: 1065-73. With W. H. Harrison. Detection of intermediate oxidation of ad- renaline and noradrenaline by fluorescence spectrometric anal- ysis. Arch. Biochem. Biophys., 101:116-23. With D. E. Fahrney and A. M. Gold. On the problem of the serine- histidine hydrogen bond in the active site of a-chymotrypsin. J. Am. Chem. Soc., 85:349. With W.-D. Dettbarn and F. A. Davis. Effects of acetylcholine on axonal conduction of lobster nerve. Biochim. Biophys. Acta, 66:397-405. With P. Rosenberg, H. B. Higman, and E. Bartels. The active struc- ture of local anesthetics. Effects on electrical and cholinesterase activity. Biochim. Biophys. Acta, 66:406-14. With A. M. Gold and D. E. Fahrney. The mechanism of reactiva- tion of phenylmethanesulfonyl cx-chymotrypsin. Biochem. Bio- phys. Res. Commun., 10:55-59. With P. Rosenberg and T. R. Podleski. Ability of venoms to render squid axons sensitive to curare and acetylcholine. Biochim. Bio- phys. Acta, 75: 104-15. With P. Rosenberg and K. Y. Ng. Factors in venoms leading to block

DAVID NACHMANSOHN 395 of axonal conduction by curare. Biochim. Biophys. Acta, 75:116-28. With W. H. Harrison. Ascorbic acid-induced fluorescence of a noradrenaline oxidation product. Biochim. Biophys. Acta, 78: 705-10. With H. B. Higman, T. R. Podleski, and E. Bartels. Apparent dis- sociation constants between carbamylcholine, d-tubocurarine and the receptor. Biochim. Biophys. Acta, 75:187-93. With T. R. Podleski and E. Bartels. Difference between tetracaine and d-tubocurarine in the competition with carbamylcholine. Biochim. Biophys. Acta, 75:387. With W.-D. Dettbarn. Hydrolysis of choline esters by invertebrate nerve fibers. Biochim. Biophys. Acta, 77:430-35. With F. C. G. Hoskin. Stereospecificity in the reactions of acetyl- cholinesterase. Proc. Soc. Exp. Biol. Med., 113:320-21. The chemical basis of Claude Bernard's observations on curare. Biochem. Z., 338:454-73. With F. C. G. Hoskin and C. von Eschen. Stimulation by quinones of initial pentose phosphate pathway steps in soluble brain preparations. Arch. Biochem. Biophys., 103: 111-16. 1964 With P. Rosenberg, E. A. Machey, H. B. Higman, and W.-D. Dett- barn. Choline acetylase and cholinesterase activity in dener- vated electroplax. Biochim. Biophys. Acta, 82:266-75. With H. B. Higman, R. R. Podlewski, and E. Bartels. Correlation of membrane potential and K flux in the electroplax of Electro- phorus. Biochim. Biophys. Acta, 79:138-50. With H. C. Lawler. The preparation of a soluble acetylcholinester- ase from brain. Biochim. Biophys. Acta, 81:280-88. Chemical control of ion movements across conducting membranes. In: Symposium on New Perspectives in Biology, BBA Library vol. 4, ed. M. Sela, pp. 176-204. Amsterdam: Elsevier. With F. C. G. Hoskin and P. Rosenberg. Alteration of acetylcholine penetration into, and effects on, venom-treated squid axons by physostigmine and related compounds. ~. Gen. Physiol., 47: 1117-27. With W.-D. Dettbarn. Action of acetylcholine and curare on lobster axons. Life Sci., 12 :910 -16. With E. Bartels and T. R. Podleski. Action of nicotine on the elec-

396 BIOGRAPHICAL MEMOIRS troplax and difference of potency between ionized and union- ized forms. Biochim. Biophys. Acta, 79:511-20. With W.-D. Dettbarn and P. Rosenberg. Restoration by a specific chemical reaction of "irreversibly" blocked axonal electrical ac- tivity. Life Sci., 3:55-60. With W.-D. Dettbarn. Distinction between action on acetylcholin- esterase and on acetylcholine receptor in axons. Biochim. Bio- phys. Acta, 79:629-30. With F. A. Davis. Acetylcholine formation in lobster sensory axons. Biochim. Biophys. Acta, 88:384-89. With H. D. Markman, P. Rosenberg, and W.-D. Dettbarn. Eye drops and diarrhea: Diarrhea as a first symptom of phospholine iodide toxicity. New Engl. I. Med., 271:197-99. With P. Rosenberg and W.-D Dettbarn. Increased acetylcholines- terase activity of intact cells produced by venoms. Biochem. Pharmacol., 13: 1157-65. Perspectives in research on the molecular basis of nerve activity. In: Tribute to ~ A. Engelhardt. Molecular Biology: Problems and Per- spectives, pp. 282-303. Moscow: Academy of Sciences of the USSR. Chemical control of bioelectric currents in membranes of conduct- ing cells. I. M. Sinai Hosp. N.Y., 31 :549-83. With A. Karlin. The association of acetylcholinesterase and of membrane in subcellular fractionations of the electric tissue of Electrophorus. ]. Cell Biol., 25: 159-69. With F. C. G. Hoskin. Stimulation of respiration and inhibition of glycolysis in lobster axons by quinones. Arch. Biochem. Bio- phys., 108:506-9. With A. M. Gold and D. Fahrney. Sulfonyl fluorides as inhibitors of esterase. II. Formation and reactions of phenylmethanesul- fonyl cx-chymotrypsin. Biochemistry, 3:783 -91. With I. Steinhardt and S. Beychok. Interaction of proteins with hydrogen ions and other small ions and molecules. In: Proteins, vol. 2, ed. H. Neurath, pp. 139-304. New York: Academic Press. With S. Beychok. Effect of ligands on the optical rotatory disper- sion of hemoglobin. Biopolymers, 3 :575-84. With S. Beychok and G. D. Fasman. Circular dichroism of poly-L- tyrisine. Biochemistry, 3: 1675 -78. . , . .. ~ . .

DAVID NACHMANSOHN 1965 397 With W.-D. Dettbarn, H. B. Higman, E. Bartels, and T. R. Podleski. Effects of marine toxins on electrical activity and K ion efflux of excitable membranes. Biochim. Biophys. Acta, 94:472-78. With S. Beychok. On the problem of isolation of the specific ace- tylcholine receptor. Biochem. Pharmacol., 14:1249-55. With G. D. Webb. Affinity of benzoguinonium and ambenonium derivatives for the acetylcholine receptor, tested on the electro- plax, and for acetylcholinesterase in solution. Biochim. Bio- phys. Acta, 102: 172-84. With E. Breslow, S. Beychok, K. Hardman, and F. R. N. Gurd. Rel- ative conformations of sperm whale metmyoglobin and apo- myoglobin in solution. }. Biol. Chem., 240:340-49. With H. Greenberg. Studies of acid phosphomonoesterase and their inhibition by diisopropylphosphorofluoridate. J. Biol. Chem., 240:1639-46. With M. Brzin, W.-D. Dettbarn, and P. Rosenberg. Acetylcholines- terase activity per unit surface of conducting membranes. I. Cell Biol., 26:353-64. With A. de Roetth, Jr., W.-D. Dettbarn, P. Rosenberg, i. G. Wilen- sky, and A. Wong. Effect of phospholine iodide on blood cholinesterase levels of normal and glaucoma subjects. Am. I. Ophthalmol., 59:586 -91. With M. Brzin, W.-D. Dettbarn, and P. Rosenberg. Penetration of enostigmine, physostigmine and paraxon into the squid giant axon. Biochem. Pharmacol., 14:919-24. With P. Rosenberg and W.-D. Dettbarn. Cholinesterase activity of rabbit aorta. Life Sci., 4:567-72. With A. Karlin and N. I. A. Overweg. An inhibitor of oxytocin from the urinary bladder of the toad, Bufo marinus. Nature (London), 207: 1401-2. With E. Bartels. Relationship between acetylcholine and local anes- thetics. Biochim. Biophys. Acta, 109:194-203. With F. C. G. Hoskin and P. Rosenberg. Penetration of sugars, ste- roids, amino acids and other organic compounds into the in- terior of the squid giant axon. }. Gen. Physiol., 49:47-56. Chemische Kontrolle des Permeabilitaetszyklus. Erregbarer Mem- branen Wahrend Elektrischer Aktivitat. Nova Acta Leopold., 30:207-33.

398 BIOGRAPHICAL MEMOIRS With P. Rosenberg and W.-D. Dettbarn. Use of venoms in testing for essentiality of cholinesterase in conduction. In: Animal Toxin. Oxford: Pergamon Press. With E. Bartels. Molecular structure determining the action of lo- cal anesthetics on the acetylcholine receptor. (Ochoa Anniver- sary Volume.) Biochem. Z., 342:359-74. With P. Rosenberg and F. C. G. Hoskin. Penetration of acetylcho- line into squid giant axons. Biochem. Pharmacol., 14: 1765-72. With P. Rosenberg. Effects of venoms on the squid giant axon. Toxicon,3:125-31. Chemical control of the permeability cycle in excitable membranes during activity. Isr. J. Med. Sci., 1: 1201-19. 1966 Sechs deutschjuedische Wissenschaftler: Haber, Willstatter, Neu- berg, Meyerhof, Bergmann and Schonheimer. Das Neue Israel (Zurich), 18:826 -33. Chemical forces controlling permeability changes of excitable membranes during electrical activity. In: Nerve As A Tissue, ed. K. Rodahl, pp. 141-61. New York: McGraw-Hill. Role of acetylcholine in neuromuscular transmission. (Presented at a symposium on myasthenia gravis.) Ann. N.Y. Acad. Sci., 135: 136-49. With H. G. Mautner, E. Bartels, and G. D. Webb. Sulfur and selen- ium isologs related to acetylcholine and choline. IV. Activity in the electroplax preparation. Biochem. Pharmacol., 15: 187-93. With A. K. Prince. Spectrophotometric study of the acetylcholin- esterase-catalyzed hydrolysis of 1-methyl-acetoxyquinolinium iodides. Arch. Biochem. Biophys., 113:195-204. With A. K. Prince. A sensitive fluorometric procedure for the de- termination of small quantities of acetylcholinesterase. Bio- chem. Pharmacol., 15:411-17. With F. C. G. Hoskin. Anaerobic glycolysis in parts of the giant axon of squid. Nature (London), 210:856 -59. With S. H. Bryant and M. Brzin. Cholinesterase activity of isolated giant synapses. J. Cell. Physiol., 68: 107-8. With P. Rosenberg, W.-D. Dettbarn, and M. Brzin. Acetylcholine and choline acetylase in squid axon, ganglia, and retina. Nature (London), 210:858 - 59. With F. C. G. Hoskin, P. Rosenberg, and M. Brzin. Reexamination

DAVID NACHMANSOHN 399 of the effect of DFP on electrical and cholinesterase activity of squid giant axon. Proc. Natl. Acad. Sci. USA, 55:1231-35. Properties of the acetylcholine receptor protein analyzed on the excitable membrane of the monocellular electroplax prepara- tion. In: Current Aspects of Biochemical Energetics: Lipmann Dedi- catory Volume, ed. N. O. Kaplan and E. P. Kennedy, pp. 145-72. New York: Academic Press. With A. de Roetth, ir., A. Wong, W.-D. Dettbarn, P. Rosenberg, and i. G. Wilensky. Blood cholinesterase activity in glaucoma patients treated with phospholine iodide. Am. J. Ophthaln~ol., 62:834-38. Chemical control of the permeability cycle in excitable membranes during electrical activity. Isr. I. Med. Sci., 1:201-19. With W.-D. Dettbarn and P. Rosenberg. Effect of ions on the efflux of acetylcholine from peripheral nerve. l. Gen. Physiol., 50: 447-60. With P. Rosenberg and H. G. Mautner. Similarity of effects of ox- ygen, sulfur, and selenium isologs on the acetylcholine receptor in excitable membranes on junctions and axons. Proc. Natl. Acad. Sci. USA, 55:835-38. The biochemical basis of cholinergic drugs. In: Biochemistry and Pharmacology of the Basal Ganglia, ed. E. Costa, L. J. Cote, and M. D. Yahr, pp. 1-15. Hewlett, N.Y.: Raven Press. With M. Brzin, V. M. Tennyson, and P. E. Duffy. Acetylcholinester- ase in frog sympathetic and dorsal root ganglia: A study by electron microscope cytochemistry and microgassometric anal- ysis with the magnetic diver. J. Cell Biol., 31:215-42. With G. D. Webb, W.-D. Dettbarn, and M. Brzin. Biochemical and pharmacological aspects of the synapses of the squid stellate ganglion. Biochem. Pharmacol., 15: 1813-19. With A. Karlin and E. Bartels. Effects of blocking sulfhydryl groups and of reducing disulfide bonds on the acetylcholine-activated permeability system of the electroplax. Biochim. Biophys. Acta, 126:525-35. With T. R. Podleski. Similarities between active sites of acetylcho- line-receptor and acetylcholinesterase with quinolinium ions. Proc. Natl. Acad. Sci. USA, 56:1034-39. With M. Brzin. The localization of acetylcholinesterase in axonal membranes of frog nerve fibers. Proc. Natl. Acad. Sci. USA, 56: 1560-63.

400 BIOGRAPHICAL MEMOIRS 1967 With P. Rosenberg and E. Bartels. Drug effects on the spontaneous electrical activity of the squid giant axon. I. Pharmacol. Exp. Ther., 155:532. With W.-D. Dettbarn. The acetylcholine system in peripheral nerve. (Presented at a symposium on cholinergic mechanism.) Ann. N.Y. Acad. Sci., 144:483. With M. Brzin and W.-D. Dettbarn. Cholinesterase activity of nodal and internodal regions of myelinated nerve fibers of frog. I. Cell Biol., 32:577. With P. Rosenberg and H. G. Mautner. Acetylcholine receptor: Similarity in axons and junctions. Science, 155: 1569. With W. Leuzinger and A. L. Baker. Acetylcholinesterase. I. Large scale purification, homogeneity, amino acid analysis. Proc. Natl. Acad. Sci. USA, 57:446. 1968 With }.-P. Changeux, W. Leuzinger, and M. Huchet. Specific bind- ing of acetylcholine to acetylcholinesterase in the presence of eserine. FEBS (Fed. Eur. Biochem Soc.) Lett., 2:77. 1969 With W. Leuzinger. Structure and function of acetylcholinesterase. In: Progress in Brain Research, ed. K. Ackert and P. G. Waser, vol. 31, pp. 241-45. Amsterdam: Elsevier. With E. Bartels. Organophosphate inhibitors of acetylcholine- receptor and -esterase tested on the electroplax. Arch. Bio- chem. Biophys., 133: 1-10. With W. l. Deal and B. F. Erlanger. Photoregulation of biological activity by photochromic reagents. III. Photoregulation of bioe- lectricity by acetylcholine receptor inhibitors. Proc. Natl. Acad. Sci. USA, 64:1230 - 34. 1970 Proteins in bioelectricity. In: Protein Metabolism of the Nervous System, ed. A. Lajtha, pp. 313-33. New York: Plenum Press. Proteins in bioelectricity. In: Colloquium Macromolecules, Biosyn- thesis and Function, vol. 21, ed. S. Ochoa, C. F. Heredia, and C.

DAVID NACHMANSOHN 40 Asensio, pp. 321-28. FEES Proceedings of the Sixth Meeting, Madrid, April 7-11, 1969. London and New York: Academic Press. With E. Bartels, W. Deal, A. Karlin, and H. G. Mautner. Affinity oxidation of the reduced acetylcholine receptor. Biochim. Bio- phys. Acta, 203 :568-71. Proteins in excitable membranes. Their properties and function in bioelectricity. Science, 168: 1059-66. With W.-D. Dettbarn, E. Bartels, F. C. G. Hoskin, and F. Welsch. Spontaneous reactivation of organophosphorus inhibited elec- troplax cholinesterase in relation to acetylcholine induced de- polarization. Biochem. Pharmacol., 19:2949-55. With H. G. Mautner and E. Bartels. Interactions of p-nitrobenzene diazonium fluoroborate and analogs with the active sites of acetylcholine-receptor and -esterase. Proc. Natl. Acad. Sci. USA, 67:74-78. 1971 With E. Bartels. Depolarization of electroplax membrane in cal- cium-free Ringer's solution. I. Membr. Biol., 5:121-32. With E. Bartels, N. H. Wassermann, and B. F. Erlanger. Photo- chromic activators of the acetylcholine receptor. Proc. Natl. Acad. Sci. USA, 68:1820-23. With E. Bartels and T. L. Rosenberry. Snake neurotoxins; effects of disulfide reduction on interaction with electroplax. Science, 174: 1236-37. Similarity of chemical events in conducting and synaptic mem- branes during electrical activity. Proc. Natl. Acad. Sci. USA, 68:3170-72. 1972 Bioenergetics and properties and function of proteins in excitable membranes associated with bioelectrogenesis. In: Molecular Bioenergetics and Macromolecular Biochemistry (Meyerhof Sympo- sium, Heidelberg, July 1970), ed. H. H. Weber, pp. 172-93. Heidelberg: Springer-Verlag. Biochemistry as part of my life. (Prefatory chapter.) In: Annual Review of Biochemastry, pp. 1-28. Stanford: Annual Reviews. With T. L. Rosenberry, H. W. Chang, and Y. T. Chen. Purification of acetylcholinesterase by a~nity chromatography and deter-

402 BIOGRAPHICAL MEMOIRS mination of active site stoichiometry. I. Biol. Chem., 247: 1555- 65. With E. Bartels and P. Rosenberg. Correlation between electrical activity and phospholipid splitting by snake venom in the single electroplax. J. Neurochem., 19: 1251-65. With I. Del Castillo, E. Bartels, and }. A. Sobrino. Microelectro- phoretic application of cholinergic compounds, protein oxidiz- ing agents and mercurials to the chemically excitable mem- brane of the electroplax. Proc. Natl. Acad. Sci. USA, 69:2081- 85. 1973 The neuromuscular junction. The role of acetylcholine in excitable membranes. In: The Structure and Function of Muscle, vol.3, Phys- iology and Biochemistry, ed. G. H. Bourne, pp.32-117. New York: Academic Press. With E. Bartels and T. L. Rosenberry. Modification of electroplax excitability by veratridine. Biochim. Biophys. Acta, 298:973- 85. With E. Neumann and A. Katchalsky. An attempt at an interpre- tation of nerve excitability. Proc. Natl. Acad. Sci. USA, 70:727- 31. Proprietes et fonction des proteines dans les membranes excitables. Un modele integrate de l'excitabilite nerveuse. Biochimie, 55: 365-76. 1974 Importance of structure and organization for the chemical reac- tions in excitable membranes. In: Central Nervous System: Studies on Metabolic Regulation and Function, ed. E. Genazzini and H. Herken, pp. 121-37. Heidelberg: Springer-Verlag. Organophosphate insecticides. A challenging problem of environ- ment control. Rehovot, 7:4-6. With E. Neumann. Properties and function of proteins in excitable membranes. An integral model of nerve excitability. (Presented at the New York Academy of Sciences Conference on the Mech- anism of Energy Transduction of Biological Systems. Ann. N.Y. Acad. Sci., 227:275-84. With Y. T. Chen, T. L. Rosenberry, and H. W. Chang. Subunit het-

DAVID NACHMANSOHN 403 erogeneity of acetylcholinesterase. Arch. Biochem. Biophys., 161 :479-87. With H. W. Chang. Purification and characterization of acetylcho- line receptor-I from Electrophorus electricus. Proc. Natl. Acad. Sci. USA, 71:2113-17. With T. L. Rosenberry, Y. T. Chen, and E. Bock. Structure of 11 S acetylcholinesterase. Subunit composition. Biochemistry, 13: 3068-79. With E. Neumann. Nerve excitability- towards an integrating con- cept. In: Biomembranes, ed. L. A. Manson. New York: Plenum Press. Biochemical foundation of an integral model of nerve excitability. (Presented at 25. Mosbacher Colloquium der Gesellschaft fuer Biologische Chemie, April 25 - 27.) In: Biochemistry of Sensory Functions, ed. L. Jaenicke, pp. 431-64. Berlin/Heidelberg/New York: Springer-Verlag. Chemical and Molecular Basis of Nerve Activity, 2d. rev. ed. including: Suppl. 1, "Properties and Function of the Proteins of the Ace- tylcholine Cycle in Excitable Membranes," and suppl. 2 (by E. Neumann), "Towards a Molecular Model of Bioelectricity." New York: Academic Press. 1976 Highlights of a friendship. In: Repections on Biochemistry, pp. 405- 11. London: Pergamon Press. The transduction of chemical into electrical energy. Proc. Natl. Acad. Sci. USA, 73:82-85. 50 years ago: Acetylcholine its role in nerve excitability. Trends Biochem. Sci., 1:237-38. 1977 Nerve excitability: Transition from descriptive phenomenology to chemical analysis of mechanisms. (Herken Festschrift.) Klin. Wochenschr., 55:715-23. Nerve excitability: From descriptive phenomenology to molecular interpretation. In: P. ~ S. Biomedical Sciences Symposia, Arden House Conference on Neuronal Information Transfer, ed. H. Vogel.

404 BIOGRAPHICAL MEMOIRS FURTHER READINGS First Conference of Physicochemical Mechanism of Nerve Activity. New York: Academy of Sciences, 1946. Metabolism and Function: Anniversary Volume in Honor of Otto Meyer- hof. Biochim. et Biophys. Acta. Amsterdam: Elsevier Publishing Co., 1950. First Conference on Nerve Impulse. New York: Josiah Macy, fir., Foun- dation, 1950. Second Conference on Nerve Impulse. New York: Josiah Macy, fir., Foundation, 1951. Fourth Conference on Nerve Impulse. New York: Josiah Macy, Jr., Foundation, 1953. Fifth Conference on Nerve Impulse. New York: Josiah Macy, fir., Foun- dation, 1954. Ion Transport Across Membranes. (Symposium at Columbia Univer- sity.) New York: Academic Press, 1954. Chemical and Molecular Basis of Nerve Activity. (Monograph.) New York: Academic Press, 1959. Second Conference on Physicochemical Mechanism of Nerve Activity. New York: New York Academy of Sciences, 1959. Molecular Biology. Elementary Process of Nerve Conduction and Muscle Contraction. New York: Academic Press, 1960.

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Biographic Memoirs: Volume 58 contains short biographies of deceased members of the National Academy of Sciences.

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