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Theodore H. Bullock: Pioneer of Integrative and Comparative Neurobiology

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Theodore H. Bullock: Pioneer of Integrative and Comparative Neurobiology J Comp Physiol A (2008) 194:119–134 DOI 10.1007/s00359-007-0286-y REVIEW Theodore H. Bullock: pioneer of integrative and comparative neurobiology G. K. H. Zupanc · M. M. Zupanc Received: 15 August 2007 / Revised: 24 October 2007 / Accepted: 27 October 2007 © Springer-Verlag 2007 Abstract Theodore H. Bullock (1905–2005) was a Keywords Theodore H. Bullock · pioneer of integrative and comparative neurobiology and Comparative neurobiology · Integrative neurobiology · one of the founders of neuroethology. His work—distin- Systems neurobiology · History of neuroscience guished by the tremendous number of diVerent research themes and animal taxa studied—provided the basis for a comprehensive analysis of brain evolution. Among his Introduction major achievements are: one of the Wrst physiological W analyses of rhythmic central pattern generators; the rst Truth is bound to prevail in spite of us simultaneous recording from both the presynaptic and Theodore H. Bullock postsynaptic region of a chemical synapse; the demonstra- tion of intercellular communication through graded poten- On 20 December 2005, Theodore (“Ted”) Holmes Bull- tials; and the discovery of two novel sensory organs formed ock passed away at the age of 90. Like no one else, he had by infrared receptors in pit vipers and electroreceptors in inXuenced the development of comparative and integrative electric Wsh. He was also one of the Wrst who applied com- neurobiology in the twentieth century. There is hardly any putational tools to the analysis of complex neural signals major taxonomic group of animals he did not study during and to perform a comparative analysis of cognitive events. his lifetime. Similarly, his research interests were remark- His two-volume treatise “Structure and function in the ner- ably diverse, including, among others, the study of physio- vous system of invertebrates” (with G. Adrian Horridge) logical processes at chemical and electrical synapses, the remains the most comprehensive, authoritative review of anatomy and physiology of sense organs, the analysis of this topic ever written. In addition to his research merits, his electrical brain activity associated with cognitive events, legacy is particularly based on his cosmopolitan way of and the mathematical modeling of neurophysiological thinking and acting, his large, worldwide school of activity. Despite this diversity, he pursued a rigorous scien- students, and his committed advocacy for comparative and tiWc approach, aiming at a detailed and comprehensive systems-oriented neurobiology. analysis of a phenomenon under investigation. Possibly even more important, he inspired numerous students and co-workers all over the world to identify new research directions and to establish their own Weld of study—an atti- G. K. H. Zupanc (&) · M. M. Zupanc tude that will leave its stamp on generations of scientists to School of Engineering and Science, Jacobs University Bremen, come. P.O. Box 750 561, 28725 Bremen, Germany e-mail: [email protected] The following review of Ted Bullock’s life and major scientiWc achievements is based on analysis of his own pub- G. K. H. Zupanc · M. M. Zupanc lications and the comprehensive collection of documents Section of Neurobiology, Division of Biological Sciences, from his laboratory, deposited in the Archives of the and Department of Neurosciences, School of Medicine, University of California, San Diego, 9500 Gilman Drive, Scripps Institution of Oceanography, University of Califor- La Jolla, CA 92093, USA nia at San Diego; interviews with his children and several 13 120 J Comp Physiol A (2008) 194:119–134 of his former friends, colleagues, and students; as well as our own recollection of numerous meetings and conversa- tions with him. In addition to a discussion of some of his major accomplishments, we have placed particular empha- sis on a reXection and illustration of his unique approach— to both science and people. Childhood, education, and early career Ted was born in Nanking, China, on 16 May 1915, the sec- ond of four children of his Presbyterian missionary parents, Amasa Archibald Bullock and Ruth Beckwith Bullock (Fig. 1). His father had studied at the University of Califor- Fig. 2 Ted Bullock in 1940. (Photographer unknown; courtesy nia, Berkeley, The University of Chicago, and Columbia Scripps Institution of Oceanography Archives) University. Among other activities, Archibald Bullock established in China University programs in chemistry and Perhaps not only the research associated with the thesis agriculture, and directed schools. The family returned to the topic, but also the person behind his mentor, drew Ted United States in 1928, one year after the outbreak of the Bullock to Light. Like Bullock in his early years, Light Chinese Civil War. These Wrst 13 years of Ted’s life, with (1886–1947) had spent many years abroad, teaching in the immersion in Chinese society, seemed to have had a Japan, China, and on the Philippines. Although, in contrast lasting inXuence on his respect towards other cultures and to Bullock, Light seemed to be a rather reserved man, his his lifelong feelings of being “a citizen of the world Wrst, of attitude towards people included a number of traits that the United States second” (Bullock 1996, p. 112). later could easily have been attributed to Ted. In a letter Back in the States, Ted attended GarWeld Junior High included in a tribute of Park College, Parkville (Missouri), School in Berkeley, and Pasadena Junior College. After from which Light had graduated in 1908, Ted Bullock (then graduating from the latter with an associate-in-art degree in representing the University of California at Los Angeles) 1932, he transferred to the University of California, Berke- and Richard M. Eakin (representing the University of ley for his junior and senior years, and majored in zoology California at Berkeley) wrote on 4 April 1958: in 1936. He stayed at Berkeley and did his Ph.D. thesis research under the guidance of Sol Felty Light (known to In all his friends and students, whether intimates or most people as S.F. Light) on the anatomy and physiology not, he inspired something more than respect—a of the nervous system of enteropneusts, one of the two clas- personal conWdence and an attachment to the man. ses of acorn worms (phylum Hemichordata). Figure 2 He will be remembered for his modesty, extending to shows Ted Bullock during that time in Berkeley. an underestimation of self, for exacting criticism in Fig. 1 Left Baby Ted Bullock at the age of three months, with his of the twentieth century. Right Ted Bullock (second from right) with father Amasa Archibald Bullock, in Kuling (China). This town, a his mother, Ruth Beckwith, and his siblings (from left to right) Bur- mountain resort on the Yangtze river in Kiangsi province, was partic- lingane, Donald, and Beatrice. (Photographers unknown; courtesy ularly popular among foreigners living in China in the Wrst few decades Scripps Institution of Oceanography Archives.) 13 J Comp Physiol A (2008) 194:119–134 121 the use of words and ideas, which drove him now to Before commencing his Wrst postdoc position at Yale caution and again to very forward positions, for a University, Ted Bullock began a tradition to spend the sum- sincere interest in human relations, and for a strong mer months at the Marine Biology Laboratory at Woods appreciation of natural beauty. Hole on Cape Cod. Often, he was accompanied by his wife Martha Runquist (whom he married in 1937) and later by After receiving his Ph.D. in 1940, Bullock was awarded their children Christine (born 1946) and Stephen (born a Sterling Fellowship to work in zoology as a postdoc under 1949). These visits enabled him to meet numerous other John Spangler Nicholas at Yale University. One year later, scientists (Fig. 3) and to initiate a number of collaborations. now as recipient of a Rockefeller Fellowship, he joined the The summers at Woods Hole appear also to have been for- laboratory of Harold Saxton Burr in the Medical School of mative and inXuential on his lifelong interest in applying a Yale. Shortly thereafter, in early 1942, he was recruited wide range of techniques to a large, albeit carefully into a research project commissioned by the OYce of Sci- selected, number of species to address biologically impor- entiWc Research and Development (an agency of the US tant questions. As he put it (Bullock 1996, p. 119): federal government created to coordinate scientiWc research for military purposes during World War II) and led by Besides meeting a wide cross section of people in Louis Goodman and Alfred Gilman in Yale’s new Depart- zoology, physiology, anatomy, and related Welds, the ment of Pharmacology. The group was assigned the task to opportunities to learn new techniques, especially study the eVects of nitrogen mustard gas and to develop electrophysiological ones, and to apply them to simple antidotes against this chemical warfare agent (ironically, invertebrate preparations, were golden. I became although nitrogen mustard has never been used in the bat- imprinted on comparative physiology and on the tleWeld, this work led to a major scientiWc breakthrough— importance of combining anatomy and physiology, on the development of chemotherapy, as nitrogen mustard was the value of simple systems, and on the diversity of the Wrst agent found to be eVective in producing remission integrative mechanisms in the nervous system. of some forms of cancer). In 1942, after two postdoctoral years, Ted Bullock was appointed Instructor in Neuroanatomy in the Yale School Controversy about the role of acetylcholine of Medicine. In 1944, he left Yale to accept a position as in the conduction of nerve impulses Assistant Professor of Anatomy at the University of Mis- souri School of Medicine, Columbia. During the latter ten- During the summers at Woods Hole, Ted Bullock initiated ure, he taught courses in neuroanatomy, topographic and an extensive collaboration with David Nachmansohn (then applied anatomy, and elementary anatomy.
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