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OTTO KRAYER October 22, 1899March 1S, 1982 BY AVRAM GOLDSTEIN Sie konnen eigentlich nur Soiche brauchen, die sick brauchen lassen. Schopenhauer. Neue Paralipomena 676, HandschriftlicherNachiass, Vol. 4 (Leipzig: P. Reclam, 1930~. For the style is the man, and where a man's treasure is there his heart, and his brain, and his writing, will be also. A. Quiller-Couch, On The Art Of Writing (London: G. P. Putnam's Sons, 1916~. IN your letter of 15 tune you state that you feel the barring of Jewish scientists is an injustice, and that your feelings about this injustice prevent you from accepting a position offered to you. You are of course personally free to feel any way you like about the way the government acts. It is not acceptable, however, for you to make the practice of your teaching profession dependent upon those feel- ings. You would in that case not be able in the future to hold any chair in a German university. Pending final decision on the basis of section 4 of the Law on the Restoration of the Professional Civil Service, I herewith forbid you, effective immediately, from entering any government academic insti- tution, and from using any State libraries or scientific facilities. 151

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152 BIOGRAPHICAL MEMOIRS THIS REMARKABLE LETTER, dated 20 June 1933, and here reprocluced in its entirety, was from the Prussian Minister for Science, Art, and National Education. The re- cipient, Otto Krayer, who cried IS March 1982, at the age of eighty-two, will be remembered for many thingshis out- standing research contributions to cardiovascular pharma- cology, his intensely enthusiastic teaching style, his very high stanclards of scientific publication and editorship, his guid- ance ant! support of the many young scientists who came under his influence and went on to significant careers in pharmacology or physiology. Krayer's unique contribution, however, was the example he set in ethical behavior behav- ior that in his thirty-fifth year anc! in the flowering of a prom- ising career brought upon him the full retribution of the Nazi hierarchy. Robert Jungk, in his book Brighter Than A Thousand Suns, A Personal History of the Atomic Scientists, writes about those clays in early 1933 in Gottingen: "Only a single one of Got- tingen's natural scientists had the courage to protest openly against the dismissal of the Jewish savants. This was the phys- iologist Krayer. He clid not allow himself to be intimidated either by his own dismissal, which was then orclered by the new Prussian Minister of Education, Stuckart, or by the threat of being clebarrecT from employment for the rest of his life." Yet rare though it was for a non-}ewish German intellec- tual to jeopardize his own future for the sake of a moral principle, "protest openly" is certainly not accurate. That was not Krayer's style. Never a political activist nor an organizer or preacher for causes Krayer wouic] have been the last to condemn his colleagues who, with various rationalizations, ' Robert Muck, Brighter Than A Thousand Suns, A Personal History of the Atomic Scientists (New York: Harcourt, Brace and Company, 1958), p. 36. .

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OTTO KRAYER 153 accepted the evil situation as beyond their control. Krayer believed, very simply, that a person tract to do what their con- science saict was right, that in such matters it was not a ques- tion of weighing consequences. His letter of 15 June 1933, which so infuriated the Nazi bureaucrat, is poignant testi- mony to this belief. He explains why he cannot accept the proffered appointment to the chair of pharmacology at Dus- seldorf the chair from which the Jewish incumbent Philipp Ellinger hack just been removed: . . . the primary reason tor my reluctance is that I feel the exclusion of Jewish scientists to be an injustice, the necessity of which I cannot under- stand, since it has been justified by reasons that lie outside the domain of science. This feeling of injustice is an ethical phenomenon. It is innate to the structure of my personality, and not something imposed from the outside. Under these circumstances, assuming such a position as the one in Dus- seldorf would impose a great mental burden on me a burden that would make it difficult to take up my duties as a teacher with joy and a sense of dedication, without which I cannot teach properly. I place a high value on the role of university teacher, and I myself would want the privilege of engaging in this activity to be given only to men who, apart from their research capabilities, also have special human qualities. Had I not expressed to you the misgivings that made me hesitate to accept your offer immediately, I would have compromised one of these essential human qualities, that of honesty. It seems to me, therefore, that the argument that in the interests of the task at hand I must defer my personal misgivings, is an empty one. I would not place even a lesser task in the hands of someone who cannot remain true to himself. Moreover, it is clear to me how great is the re- sponsibility that you have to carry a responsibility that gives you the right to expect honesty. The work to which I have heretofore dedicated all my strength, with the goal of applying my scientific knowledge and research expertise to effective university teaching, means so much to me that I could not com- promise it with the least bit of dishonesty. I therefore prefer to forego this appointment, though it is suited to my inclinations and capabilities, rather than having to betray my convic-

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54 BIOGRAPHICAL MEMOIRS tions; or that by remaining silent I would encourage an opinion about me that does not correspond with the facts. A moral dilemma arises when the policies of a legitimately constitutes] authority are morally unacceptable. Resistance to a tyranny that can make no claim to a popular mandate is difficult and risky enough. But HitIer's regime had all the trappings of legitimacy, it had come to power in a constitu- tional manner, and its support went creep anc! wide among the German people, not exclucting the university faculties and students. Noncompliance, under such circumstances, re- quires the courage of one's convictions to an extraordinary clegree. One's support has to come principally from one's own conscience, while one's peers, by and large, tend to distance themselves, in order to avert unpleasant repercussions and to avoic! confronting their own consciences. The events that facet! Krayer with a moral choice were unusual, from a historical perspective, but they were not unique. Fanaticism political, religious, tribal, racial, intel- lectual, nationalistic has perioclically infected one or an- other part of the earth's population since civilization began. No country and no time is immune, and so the moral di- lemma is an ever-recurring theme. During the agony of Viet- nam, American academics could witness the same cautious neutrality on the part of most of their colleagues, at least until it became acceptable anct popular to speak one's outrage. Ap- parently the simple ability to distinguish right from wrong and to act accordingly was incompatible with the scholarly temperament. "Not to clecicle is to decide," wrote the Amer- ican theologian Harvey Cox. Most fount! it easiest "not to clecicle." The surgeon Rudolf Nissen, writing of the German uni- versity faculties in 1933, has this to say: Another example of rare, almost isolated conduct amidst the crowd of opportunists was given by the Berlin Professor Extraordinarius, Otto

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OTTO KRAYER 155 Krayer. His pupil, M. Reiter, has these wonderful words for this conduct: "The world is not particularly rich with people who prefer to jeopardize their career rather than sanction it with alien injustice. Nothing is more characteristic of Krayer's personality than his repeated refusal in 1933 to take over the chair in Dusseldorf, whose former holder, Philipp Ellinger, was driven from it on account of his race. The Professor Extraordinarius in Berlin, who was 34 at the time, did something that those in power felt was an open revolt and that many of his colleagues felt was at least inop- portune and disturbing in the repercussions it had for them."2 Finally Nissen remarks: "It is unfortunate that such coura- geous and manly incliviclual actions in the universities were not collectecl ant! made available to the public by officials who occupied themselves with the history of the Nazi periocl." He concludes by quoting Shakespeare (The Winter's Tale, act I, scene 2) on the importance of publicly recognizing such ac- tions: "One good (leed, dying tongueless, slaughters a thou- sand waiting upon that." Krayer's own laconic account of this landmark event in his life is found in an autobiographical sketch he wrote after his retirement for the International Biographical Archives and Dic- tionaryofCentralEuropeanEmigres, 1933-45: In the Spring of 1933, while engaged in collaborative studies with Prof. H. Rein in the Department of Physiology, University of Gottingen, I was asked by the Department of Education of the State of Prussia to take over the Chair of Pharmacology in the Medical Academy of Dusseldorf. The vacancy had been created by the dismissal of the Jewish incumbent Prof. Philipp Ellinger. Refusal to fill the vacancy because of my stated disagree- ment with the unjust policies of the government led to my immediate suspension by the Prussian Minister of Education from my academic po- sitions. Moreover, I was forbidden to enter any university premises includ- ing University and State libraries. Returning from Gottingen to Berlin, where I could make use of private libraries, I was able to continue literary work in progress. I was especially anxious to complete and edit and to supervise the printing of Volume 2 of P. Trendelenburg: Die Hormone, a task 2 Rudolf Nissen, Helle Blatter dunkle Blatter: Erinnerungen eines Chirurgen (Stutt- gart: Deutsche Verlags-Anstalt, 1969), pp. 140-44.

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56 BIOGRAPHICAL MEMOIRS which had been entrusted to me by my teacher shortly before his death in 1931. Later in 1933 Krayer's academic privileges at the Univer- sity of Berlin were restored. However, he obtained a leave of absence and accepted an invitation to join the Department of Pharmacology at University College, London, with support from the Rockefeller Foundation, anct on the last day of 1933 he departed Germany. There followed an intense ant! pro- cluctive nine months of research in collaboration with E. B. Verney, who had been Starling's pupil. The substance of the investigations with Verney is recounted in a later section of this memoir. Krayer's former Berlin associate W. Felciberg, himself a recent refugee from the Nazis, was also in Lonclon. And dominating the scene was H. H. Dale, the foremost pharmacologist of the day. In the autumn of 1934 Krayer was callect to head the De- partment of Pharmacology at the American University of Beirut. His research and teaching accomplishments there are Ascribed later. Officially representing the American Univer- sity of Beirut at the Tercentenary Celebration of Harvard University in 1936, he was asked to stay on for a few months as a lecturer in pharmacology at the Harvard Meclical School. Then in 1937 an invitation was extencled for Krayer to join the faculty as associate professor of pharmacology. He ac- ceptecl and two years later became Reid Hunt's successor as heat] of the department, a position he held until his retire- ment in ~ 966. A littIe-known event of his early days in Boston sheds fur- ther light on the idealism that was a strong motivating force in Krayer's life. The Nobel peace prize had just been awar(le(1 to the German writer and journalist Car! van Ossietzky, a pacifist of international renown, who had exposed the secret rearming of Germany ant! who had been (and was until his

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OTTO KRAYER 157 death) incarcerated by the Nazi regime. Hitler's response to the award of the prize was a clecree forbidding Germans to accept any Nobel prize in the future. At the regular meeting of the German Chemical Society on May 8, 1937, the presi- clent of the Society, Professor Stock, acictressed himself to the honor bestowal upon van Ossietzky: "Every true German," he said, "must regard as a slap in the face this insulting abuse . . . an abuse dictated by political hatred. It is unclerstanclable that both the government and the people are indignant over this, and want nothing more to do with Nobel prizes . . . the crime of the Norwegian parliament's committee will be re- gretted cleeply by Science."3 Krayer's immediate reaction was the following brief note to the society's office: "The remarks of President A. Stock concerning the awarcl of the Nobel peace prize, which are printed on page 121 of the Proceedings of the German Chemical Society of 9 June 1937 oblige me to request that you strike my name from the list of members of the German Chemical Society." Professor Stock, in reply, couIct only imagine that he had been misunderstoocI. "I was only reflecting the feelings of every German scientist," he wrote, "in being upset by such a conscious provocation . . . by the honoring of a person who- even before the time of Hitler! hac] been branclect a traitor; and in cleploring that the scientific Nobel prizes had to suffer from this circumstance.... Perhaps you will be so kind as to write me a word of clarification." Krayer's response will ring a familiar note for all who, as students or colleagues, came under his influence. It recalls the curious blend of careful reasoning and objective presen- tation of facts on the one hancI, coupled with extraordinary 3 A. Stock, "Opening Remark," Berichte der Deutschen Chemischen Gesellschaft, 70(1937): 12 1.

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58 BIOGRAPHICAL MEMOIRS emotional intensity on the other, that colored many of his formal lectures and informal discourses. Dear Mr. President: I am happy to communicate to you the reason for my protest against your remarks. However, it is not my intention to enter into a discussion about the political expression "traitor". That this expres- sion does not necessarily have a precise ethical value must be obvious to everyone who has experienced how easily the meaning can be changed by various political trends that appear especially strongly and clearly at times of upheaval in the structure of a State. What made me write my letter of 3 September was the urge to express the view that not every German and- as I am convinced not every Ger- man scientist shares your feelings of being upset by the award of the recent Nobel peace prize. The reason for this conviction is what I have read over the last ten years of the writings of Carl van Ossietzky and have learned from other sources about him. I have had no occasion to meet this man personally. But whoever, over the past decade in Germany, has followed the course of his career in an unprejudiced way would even if he were a political op- ponentnot be able to ignore the fact of the man's extraordinary person- ality. Here is a man who, in a hard life full of work and an abundance of general human and political experience, has developed a world outlook and has deduced from it the principles of his life philosophy, who has made the profession of political writer his mission in life, and who is ready to dedicate to this profession not only all the strength of his spirit but also his whole personality. An unyielding character who, whenever the obli- gation of sincerity necessitates, openly uses his right of free speech to ex- press his opinion. A man who is not motivated by the lust for power and fame but who is forced to speak by the persuasion of the rightness of his beliefs, and who fights unafraid for that persuasion with the force of his arguments. Carl von Ossietzky has proven the sincerity of his mind and his selflessness by again refusing (he had already been amnestied once) to evade responsibility for his convictions. To back up his words with deeds was a necessity of life for him although he must have known that he could not expect any justice from his political enemies. The reason for such a judgment as you, Mr. President, have formu- lated, must be sought in an ethical evaluation of the man. I do not find

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OTTO KRAYER 159 sufficient basis for your interpretation, and I am not of the opinion that the scientific Nobel prizes have lost any of their value or significance by the honoring of Carl van Ossietzky. It is to the credit of the Nobel orga- nization that it honored the ethical qualities of this man; that is my con- viction. What can promote peace between nations if not the deeds of such men, who are motivated by a pure and deep consciousness of their re- sponsibility to a higher human order than is represented by the nation into which we [sic!] are born? A final incident is noteworthy, again for the light it sheets on the ethical stanciarcls by which Krayer consistently guiclect all his actions. In 1965 the Academic Council of the Medical Academy of Dusseldorf votect to confer honorary member- ship on Krayer. Writing about this decision, the rector of the University explainer] as follows: They would like thereby to show their appreciation of the stand you took when, on grounds of conscience, you refused the call to the chair of pharmacology and toxicology in Dusseldorf in 1933, which would have been your first opportunity to be head of your own institute. At the same time the Academic Council wishes to acknowledge the fact that even after your emigration, and despite the unpleasantness you experienced in Dus- seldorf, you nevertheless maintained and furthered your contacts with German science. Not the least, we would also like by our decision to ac- knowledge your scientific accomplishments, which relate to us in a special way through a traditional field of research at our Academy, namely, heart and circulation research. Krayer's immediate response was to accept the honorary membership with pleasure. But as time passed, he evidently became increasingly uneasy. Somehow a mutually suitable ciate for the presentation ceremony in Dusseldorf collie not be arranged. Finally, on January 26, 1966, Krayer sent what must have been a very difficult letter to write, as we can sur- mise from the three different preliminary handwritten drafts that are preserved, each full of cleletions ancT alternative

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160 BIOGRAPHICAL MEMOIRS worclings. Addressing the rector of the University of Dussel- clorf, Krayer wrote: In the course of the correspondence with you concerning the time of my visit to Dusseldorf, I have thought more deeply about the honor you are planning for me. I have come to the conclusion that the right thing for me to do is not to accept the honorary membership of the Medical Academy of Dusseldorf. Despite my happiness at your first letter, which reached me during my trip to Japan, I had certain reservations from the beginning. It is now clear to me that the original ethical position I took in 1933 does not permit of any external reward. I must ask you, therefore, to nullify the decision of the Scientific Council of the Medical Academy. I regret that I took so long to express my convictions clearly. Krayer closes with the hope that his decision will not cause bact feelings to mar his personal relationships with colleagues at Dusseldorf. The reference, in the rector's original letter, to Krayer's maintaining anct furthering contacts with German science will be cryptic to those unfamiliar with an episode that fol- lowed shortly on the close of World War Il. With Central Europe literally in ashes, its universities and research insti- tutes in ruins, and its people starving, the Unitarian Service Committee organized a medical mission to Czechoslovakia with Harvard carctiologist Paul DuctIey White as director and Krayer as an active participant. During that trip Krayer be- came fully aware of the (levastation of the German universi- ties through personal visits with university colleagues. It must have been then that he formulated a plan for rendering spe- cial material and moral assistance to the German academic communities. On his return to Harvard, he foundecI, and served as secretary-treasurer of, a Committee to Help Ger- man University Scientists. By lL948 a medical mission to Ger- many had been organized by the Unitarian Service Commit- tee, with Krayer as its chairman. This effort was supported

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OTTO KRAYER 215 f. O. Krayer. Der toxikologische Nachweis des Coniins. Arch. Exp. Pathol. Pharmakol., 162:342-72. g. O. Krayer and W. Koll. Coniinahnliche Eigenschaften einiger Aminbasen. Arch. Exp. Pathol. Pharmakol., 162:373-84. h. ~ Trendelenburg's Grundlagen der allgemeinen und speziellen Arznei- verordnung, 3d ed. rev. O. Krayer. Berlin: Springer. 1932 a. O. Krayer. Uber die Behandlung von Kreislaufstorungen mit Organ- und Muskelextrakten. Bemerkungen zur Pharmakolo- gie. Dtsch. Med. Wochenschr., 58:123-24. O. Krayer and E. Schutz. Mechanische Leistung und Aktions- strom des Warmbluterherzens. Verh. Dtsch. Pharmakol. Ges., XI. Tagung, 99 - 100. c. O. Krayer and E. Schutz. Mechanische Leistung und einphasi- sches Elektrogramm am Herz-Lungen-Praparat des Hundes. Z. Biol., 92:453-61. 1933 a. O. Krayer. Ist die Integritat der sympathischen Schilddrusenin- nervation notwendig fur die thyreotrope Wirkung des Hypo- physenvorderlappens? Arch. Exp. Pathol. Pharmakol., 171 :473-79. b. W. Feldberg and O. Krayer. Nachweis einer bei Vagusreiz frei- werdenden azetylcholinah nlich en Subs tan z am Warmbl u ter- herzen. Verh. Dtsch. Ges. Kreislaufforsch, VI. Tagung, 81-83. c. W. Feldberg and O. Krayer. Das Auftreten eines azetylcholinar- tigen Stoffes im Herzvenenblut von Warmblutern bei Reizung der Nervi vagi. Arch. Exp. Pathol. Pharmakol., 172: 170-93. d. O. Krayer. Zur Kreislaufwirkung der Leberpraparate des Han- dels. Dtsch. Med. Wochenschr., 59:576-78. e. O. Krayer. Zur Pharmakotherapie der Herzinsuffizienz. Er- krankungen des Herzmuskels und der Herzklappen, pp. 84- 94. Dresden and Leipzig: Steinkopff. 1934 a. F. Grabe, O. Krayer, and K. Seelkopf. Beitrag zur Aufklarung der kreislaufwirksamen (adrenalinahnlichen) Stoffe in Lebe- rextrakten. Klin. Wochenschr., 13: 1381-83.

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216 BIOGRAPHICAL MEMOIRS 1934 b. O. Krayer and E. B. Verney. Veranderung des Acetylcholinge- haltes im Blute der Coronarvenen unter dem Einfluss einer Blutdrucksteigerung durch Adrenalin. Klin. Wochenschr., 13: 1250-51. c. Die Hormone. Ihre Physiologte und Pharmakolog~e, by Paul Tren- delenburg, vol. 2, ed. O. Krayer. Berlin: Springer. 1935 a. O. Krayer. Beitrag zur Aufklarung der Natur der kreislauf- wirksamen Stoffe in als Heilmittel verwandten Leberextrakten. Institut de Recherches Physiologiques de Moscou. Problemes de Biologie et de Medecine Volume {ubilaire dedie au Prof. Lina Stern, pp. 179 - 83. b. O. Krayer and E. B. Verney. Reflektorische Beeinflussung des Gehaltes an Acetylcholin im Blute der Coronarvenen. Arch. Exp. Pathol. Pharmakol., 180:75-92. 1937 O. Krayer. Kurbissamen als Bandwurmmittel. Klin. Wochenschr., 16: 1651-52. 1938 Trendelenb?~rg's Grundlagen der allgemeinen und speziellen Arznei- verordnung, 4th ed. rev. O. Krayer. Berlin: Springer. 1941 a. D. G. Friend and O. Krayer. The estimation by a manometric method of the activity of cholinesterase in lymph. }. Pharmacol. Exp. Ther., 71:246-52. b. D. G. Friend and O. Krayer. The elimination of prostigmine. J. Pharmacol. Exp. Ther., 72:15. 1942 a. O. Krayer and R. Mendez. Studies on veratrum alkaloids. I. The action of veratrine upon the isolated mammalian heart. T. Pharmacol. Exp. Ther., 74:350-64. b. R. P. Linstead and O. Krayer. Effect of l-ascorbic acid on the isolated frog heart. Science, 95:332-33.

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OTTO KRAYER 217 c. O. Krayer, R. Mendez, E. Moisset de Espanes, and R. P. Lin- stead. Pharmacology and chemistry of substances with cardiac activity. I. Effect of unsaturated lactones on the isolated frog heart. I. Pharmacol. Exp. Ther., 74:372-80. d. O. H. Lowry, O. Krayer, A. B. Hastings, and R. P. Tucker. Effect of anoxemia on myocardium of the isolated heart of the dog. Proc. Soc. Exp. Biol. N.Y., 49:670-74. e. E. B. Astwood, l. M. Flynn, and O. Krayer. Effect of continuous intravenous infusion of glucose in normal dogs. }. Clin. Invest., 21:621. f. O. Krayer. The effect of veratrum alkaloids on circulatory reflexes. Fed. Proc. Fed. Am. Soc. Exp. Biol., 1:156. 1943 a. O. Krayer, R. P. Linstead, and D. Todd. Pharmacology and che- mistry of substances with cardiac activity. II. Effect of l-ascorbic acid and some related compounds and of hydrogen peroxide on the isolated frog heart. I. Pharmacol. Exp. Ther., 77:113- 22. b. G. K. Moe and O. Krayer. Studies on veratrum alkaloids. II. The action of veratridine and cevine upon the isolated mam- malian heart. I. Pharmacol. Exp. Ther., 77:220-28. c. O. Krayer. Action of l-ascorbic acid upon the isolated frog heart. Proc. Soc. Exp. Biol. N.Y., 53:51-52. d. O. Krayer, E. H. Wood, and G. Montes. Studies on veratrum alkaloids. IV. The sites of the heart rate lowering action of ver- atridine. J. Pharmacol. Exp. Ther., 79:215-24. e. S. Ellis, O. Krayer, and F. L. Plachte. Studies on physostigmine and related substances. III. Breakdown products of physostig- mine; their inhibitory effect on cholinesterase and their phar- macological action. J. Pharmacol. Exp. Ther., 79:309-19. 1944 a. O. Krayer, A. Goldstein, and F. L. Plachte. Studies on physo- stigmine and related substances. I. Quantitative relation be- tween dosage of physostigmine and inhibition of cholinesterase activity in the blood serum of dogs. J. Pharmacol. Exp. Ther., 80:8-30. b. G. K. Moe, D. L. Bassett, and O. Krayer. Studies on veratrum

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218 BIOGRAPHICAL MEMOIRS alkaloids. V. The effect of veratridine and cevine upon the cir- culation in anesthetized dogs, with particular reference to femoral arterial flow. i. Pharmacol. Exp. Ther., 80:272-84. c. O. Krayer, G. K. Moe, and R. Mendez. Studies on veratrum alkaloids. VI. Protoveratrine: Its comparative toxicity and its circulatory action. i. Pharmacol. Exp. Ther., 82: 167-86. d. O. Krayer. A difference in cardiodecelerator action between di- gitoxin and digitoxigenin. Proc. Soc. Exp. Biol. N.Y., 57:167- 69. 1946 a. H. M. Mating and O. Krayer. The action of erythrophleum al- kaloids upon the isolated mammalian heart. I. Pharmacol. Exp. Ther., 86:66 - 78. b. O. Krayer and G. H. Acheson. The pharmacology of the ver- atrum alkaloids. Physiol. Rev., 26:383-446. c. A. Farah and O. Krayer. The action of dimethylaminoethanol upon the heart-lung preparation of the dog. Fed. Proc. Fed. Am. Soc. Exp. Biol., 5:177-78. O. Krayer, A. Farah, and F. C. Uhle. Pharmacology and chem- istry of substances with cardiac activity. IV. Effect of methyl- aminoethanol, dimethylaminoethanol, and related substances on the isolated mammalian heart. I. Pharmacol. Exp. Ther., 88:277-86. d. 1947 O. Krayer, }. C. Aub, I. T. Nathanson, and P. C. Zamecnik. The influence of antitoxin upon the action of Clostr~dium oedematiens toxin in the heart-lung preparation of the dog. }. Clin. Invest., 26:411-15. 1948 a. O. Krayer and A. Farah. Action of cysteine and of dimercap- topropanol in heart failure caused by sodium bismuth tartrate. Fed. Proc. Fed. Am. Soc. Exp. Biol., 7:235. b. A. Wollenberger and O. Krayer. Experimental heart failure caused by central nervous system depressants and local anes- thetics. I. Pharmacol. Exp. Ther., 94:439-43.

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OTTO KRAYER 1949 219 a. O. Krayer, S. B. Wolbach, I. H. Mueller, and G. B. Wislocki. Reid Hunt. Harv. Med. Alumni Bull., 23:39 - 42. (Obituary.) b. A. Goldstein, O. Krayer, M. A. Root, G. H. Acheson, and M. E. Doherty. Plasma neostigmine levels and cholinesterase inhibi tion in dogs and myasthenic patients. I. Pharmacol. Exp. Ther. 96:56-85. c. O. Krayer. Veratramine, an antagonist to the cardioaccelerator action of epinephrine. Proc. Soc. Exp. Biol. N.Y., 70:631-32. d. O. Krayer. Studies on veratrum alkaloids. VIII. Veratramine, an antagonist to the cardioaccelerator action of epinephrine. {. Pharmacol. Exp. Ther., 96:422-37. e. E. Meilman and O. Krayer. Clinical studies on the pure ver- atrum alkaloids: Protoveratrine and veratridine. Forty-first An- nual Meeting, American Society for Clinical Investigation, At- lantic City, May 2. I. Clin. Invest., 28:798. f. O. Krayer. Studies on veratrum alkaloids. IX. The inhibition by veratrosine of the cardioaccelerator action of epinephrine and of norepinephrine. I. Pharmacol. Exp. Ther., 97:256-65. g. O. Krayer and E. F. Van Maanen. Studies on veratrum alka- loids. X. The inhibition by veratramine of the positive chrono- tropic effect of accelerans stimulation and of norepinephrine. I. Pharmacol. Exp. Ther., 97:301-7. h. O. Krayer. The pharmacological basis for the use of veratrum alkaloids in the treatment of hypertension. Proc. Rudolf Vir- chow Med. Soc. City N.Y., 8:126-27. 1950 a. Lectures Unitarian Service Committee Medical Mission to Germany, July2September3, 1948, ed. O. Krayer. Berlin: Springer. b. E. Meilman and O. Krayer. Clinical studies on veratrum alka- loids. I. The action of protoveratrine and veratridine in hyper- tension. Circulation, 1 :204-13. c. O. Krayer. A quantitative comparison of the antiaccelerator ac- tion of veratramine and jervine. I. Pharmacol. Exp. Ther., 98:19. d. K. Kramer, U. Luft, and O. Krayer. Action of epinephrine and veratramine upon heart rate and oxygen consumption in the

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220 BIOGRAPHICAL MEMOIRS heart-lung preparation of the dog. Fed. Proc. Fed. Am. Soc. Exp. Biol., 9:292. e. M. Reiter and O. Krayer. tervine and pseudojervine, antago- nists to the cardioaccelerator action of epinephrine and of ac- celerans stimulation. I. Pharmacol. Exp. Ther., 98:27. f. O. Krayer and M. Reiter. Studies on veratrum alkaloids. XI. Heroine and pseudojervine, antagonists to the cardioaccelerator action of epinephrine and of accelerans stimulation. Arch. Int. Pharmacodyn. Ther., 81:409-26. g. O. Krayer. Studies on veratrum alkaloids. XII. A quantitative comparison of the antiaccelerator cardiac action of veratra- mine, veratrosine, jervine and pseudojervine. I. Pharmacol. Exp. Ther. 98 :427-36. h. O. Krayer. Solanum alkaloids with antiaccelerator cardiac acti- vity. Fed. Proc. Fed. Am. Soc. Exp. Biol., 9:292. i. O. Krayer and L. H. Briggs. Studies on solanum alkaloids. I. The antiaccelerator cardiac action of ,13-dihydrosolasodine and tetrahydrosolasodine. Br. i. Pharmacol., 5:118-24. j. O. Krayer and L. H. Briggs. Studies on solanum alkaloids. II. The antiaccelerator cardiac action of solasodine and some of its derivatives. Br. J. Pharmacol., 5:517-25. k. O. Krayer. The antiaccelerator cardiac action of quinine and quinidine. I. Pharmacol. Exp. Ther., 100:146-50. O. Krayer. Untersuchungen uber die Kreislaufwirkung der Veratrumalkaloide. Arch. Exp. Pathol. Pharmakol., 209:405- 20. 1951 a. J. I. Mandoki, C. Mendez, R. R. Garcia, and O. Krayer. The action of veratramine and epinephrine on the functional re- fractory period of A-V conduction. I. Pharmacol. Exp. Ther., 101:25. b. O. Krayer. Quinine-like action of veratramine upon the single twitch and upon the "veratrine response" of the sartorius muscle of the frog. Fed. Proc. Fed. Am. Soc. Exp. Biol., 10:316. c. O. Krayer, F. C. Uhle, and P. Ourisson. Studies on veratrum alkaloids. XIV. The antiaccelerator cardiac action of derivatives of veratramine and jervine and of synthetic steroid secondary

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OTTO KRAYER 221 alkamines obtained from pregnenolone and from sapogenins. I. Pharmacol. Exp. Ther., 102:261 - 68. d. O. Krayer and H. W. George. Studies on veratrum alkaloids. XV. The quinine-like effect of veratramine upon the single twitch and upon the "veratrine response" of the sartorius muscle of the frog. I. Pharmacol. Exp. Ther., 103:249-58. e. O. Krayer, I. I. Mandoki, and C. Mendez. Studies on veratrum alkaloids. XVI. The action of epinephrine and of veratramine on the functional refractory period of the auriculo-ventricular transmission in the heart-lung preparation of the dog. J. Phar- macol. Exp. Ther., 103:412-19. 1952 a. O. Krayer, B. H. Rogers, S. M. Kupchan, and C. V. Deliwala. Pharmacological and chemical relation between the veratrum alkaloids and the zygadenus alkaloids. Fed. Proc. Fed. Am. Soc. Exp. Biol., 11:364. b. R. B. Arora, E. Meilman, and O. Krayer. Action of veratramine and of sympathomimetic amines upon the automaticity of the atrio-ventricular node. Fed. Proc. Fed. Am. Soc. Exp. Biol., 11:318. c. P. Ourisson and O. Krayer. Antagonistic action to the cardio- accelerator effect of ephedrine, synephrine? isuprel, tuamine and oenethyl. Fed. Proc. Fed. Am. Soc. Exp. Biol., 11:381. d. O. Krayer. Antiaccelerator cardiac agents. I. M. Sinai Hosp. N.Y., 19:53-69. e. E. Meilman and O. Krayer. Clinical studies on veratrum alka- loids. II. The dose-response relations of protoveratrine in hy- pertension. Circulation, 6:2 12 - 2 1 . f. R. B. Arora and O. Krayer. The antiveratrinic action of the cardiac glycosides and of bufotoxin. I. Pharmacol. Exp. Ther., 106:371-72. g. ~ Trendelenburg's Grundlagen der allgemeinen and speziellen Arznei- verordnung, 7th ea., rev. O. Krayer and M. Kiese. Berlin, Got- tingen, and Heidelberg: Springer. 1953 a. O. Krayer, S. M. Kupchan, C. V. Deliwala, and B. H. Rogers. Untersuchungen uber die Veratrumalkaloide. XVIII; Die

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222 BIOGRAPHICAL MEMOIRS chemischen und pharmakologischen Beziehungen zwischen den Zygadenusalkaloiden und den Veratrumalkaloiden. Arch. Exp. Pathol. Pharmakol., 219:371-85. b. O. Krayer. The history of the Bezold-tarisch effect. Presented at a symposium, Reflexes from the Cardiac and Pulmonary Areas. Nineteenth International Physiology Congress, Mon- treal. 1954 a. H. W. Kosterlitz, O. Krayer, and A. Matallana. The eject of moderately large doses of veratramine and veratrosine on the rhythm of the acutely denervated heart of the cat. }. Physiol. (London), 124:40P. b. O. Krayer and P. Ourisson. Studies on veratrum alkaloids. XIX. The action of veratramine upon cardioacceleration caused by ephedrine, tyramine, phenylephrine and isopropylarterenol. J. Pharmacol. Exp. Ther., 112:341 - 55. c. O. Krayer. Veratrum alkaloids. In: Pharmacology in Medicine, ed. V. A. Drill, pp. 1-10. New York: McGraw-Hill. 1955 c. a. O. Krayer and t. M. Benforado. Die Schlagfrequenz des akut denervierten Herzens im Herz-Lungen-Praparat des Hundes mit einem Hinweis auf die frequenzbeschleunigende Wirkung des Adrenalins. Pflug. Arch. Ges. Physiol., 260:177-87. b. O. Krayer, R. B. Arora, and E. Meilman. Studies on veratrum alkaloids. XXI. The action of veratramine upon impulse gene- ration in the dog heart. J. Pharmacol. Exp. Ther., 113:446-59. H. W. Kosterlitz, O. Krayer, and A. Matallana. Studies on ver- atrum alkaloids. XXII. Periodic activity of the sino-auricular node of the denervated cat heart caused by veratramine. i. Pharmacol. Exp. Ther., 113 :460-69. d. S. Ellis and O. Krayer. Properties of a toxin from the salivary gland of the shrew, Blarina brevicauda. ]. Pharmacol. Exp. Ther.,114:127 - 37. 1956 a. O. Krayer and }. Fuentes. Chronotropic cardiac action of reser- pine. Fed. Proc. Fed. Am. Soc. Exp. Biol., 15:1462.

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OTTO KRAYER 223 b. F. C. Uhle, B. A. Mitman, and O. Krayer. Synthetic esters of dimethylaminoethanol exhibiting positive inotropic cardiac ac- tivity. J. Pharmacol. Exp. Ther., 116:444 - 49. I. R. Innes, H. W. Kosterlitz, and O. Krayer. Studies on ver- atrum alkaloids. XXIV. The inhibition by veratramine and ver- atrosine of the cardioaccelerator effect of electrical stimulation of the accelerator nerves. l. Pharmacol. Exp. Ther., 117:317- 21. c. 1957 a. M. K. Paasonen and O. Krayer. Effect of reserpine upon the mammalian heart. Fed. Proc. Fed. Am. Soc. Exp. Biol., 16:326-27. b. O. Krayer and M. K. Paasonen. Direct cardiac action of reser- pine. Acta Physiol. Scand., 42:88-89. c. I. R. Innes and O. Krayer. Depletion of the cardiac catechol- amines by reserpine. I. Physiol. (London), 139: 18P. 1958 a. O. Krayer. Veratrum alkaloids. In: Pharmacology in Medicine, 2d ea., ed. V. A. Drill, pp. 515-24. New York: McGraw-Hill. b. O. Krayer and I. Fuentes. Changes in heart rate caused by di- rect cardiac action of reserpine. I. Pharmacol. Exp. Ther., 123: 145-52. M. K. Paasonen and O. Krayer. The release of norepinephrine from the mammalian heart by reserpine. I. Pharmacol. Exp. Ther., 123:153-60. d. I. R. Innes and O. Krayer. Studies on veratrum alkaloids. XXVII. The negative chronotropic action of veratramine and reserpine in the heart depleted of catecholamines. J. Pharma- col. Exp. Ther., 124:245-51. e. I. R. Innes, O. Krayer, and D. R. Waud. The action of Ranwolfia alkaloids on the heart rate and on the functional refractory pe- riod of atrio-ventricular transmission in the heart-lung prepa- ration of the dog. i. Pharmacol. Exp. Ther., 124:324-32. f. D. R. Wand, S. R. Kottegoda, and O. Krayer. Threshold dose and time course of norepinephrine depletion of the mamma- lian heart by reserpine. I. Pharmacol. Exp. Ther., 124:340-46. c.

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224 BIOGRAPHICAL MEMOIRS 1959 M. K. Paasonen and O. Krayer. The content of noradrenaline and adrenaline in the rat heart after administration of Ranwolfia alka- loids. Experientia, 15:75 -76. 1960 D. R. Wand and O. Krayer. The rate-increasing effect of epineph- rine and norepinephrine and its modification by experimental time in the isolated heart of normal and reserpine-pretreated dogs. J. Pharmacol. Exp. Ther., 128:352-57. 1961 a. O. Krayer. The history of the Bezold-larisch effect. Arch. Exp. Pathol. Pharmakol., 240:361-68. b. O. Krayer, E. B. Astwood, D. R. Wand, and M. H. Alper. Rate- increasing action of corticotropin and of a-intermedin in the isolated mammalian heart. Proc. Natl. Acad. Sci. USA, 47: 1227-36. 1962 a. O. Krayer, M. H. Alper, and M. K. Paasonen. Action of gua- nethidine and reserpine upon the isolated mammalian heart. }. Pharmacol. Exp. Ther., 135: 164-73. b. O. Krayer, B. D. Davis, and S. W. Kuffler. Obituary Otto Loewi. Pharmacologist, 4:47-49. c. O. Krayer. Accidents in the pursuit of knowledge. (Sollmann- Award oration.) Pharmacologist, 4:68-76. a. O. Krayer. 1963 Uber chronotrope Herzwirkung. Klin. Wo- chenschr., 41:272-76. b. M. H. Alper, W. Flacke, and O. Krayer. Pharmacology of reser- pine and its implications for anesthesia. Anesthesiology, 24:524-42. 1964 D. F. Hawkins, F. C. Uhle, and O. Krayer. Studies on veratrum alkaloids. XXXVII. Chronotropic cardiac action and toxicity of

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OTTO KRAYER 225 N-alkyl derivatives of veratramine. I. Pharmacol. Exp. Ther., 145:275-85. 1965 O. Krayer, W. Mosimann, and G. Schroder. Positive chronotropic cardiac effect of methyl guanidine. Fed. Proc. Fed. Am. Soc. Exp. Biol., 24:487. 1966 O. Krayer, W. Mosimann, and G. Silver. Rate-increasing action of methylguanidine upon the isolated mammalian heart. l. Phar- macol. Exp. Ther., 154:73-82. 1972 O. Krayer, N. Weiner, and W. Mosimann. Blood norepinephrine levels during responses of the heart-lung preparation to me- thylguanidine. J. Pharmacol. Exp. Ther., 181: 108-15. 1977 O. Krayer and E. Meilman. Veratrum alkaloids with antihyperten- sive activity. In: Handbook of Experimental Pharmacology, Hef~ter- Heubner, new ser., ed. G. V. R. Born, O. Eichler, A. Farah, H. Herken, and A. D. Welch, pp. 547-70. Berlin, Heidelberg, and New York: Springer-Verlag.