Viktor Hamburger

Forum TRENDS in Neurosciences Vol.24 No.11 November 2001 673

to a given sensory input (e.g. hunger to 10 Frasure-Smith, N. and Prince, R. (1989) Long 24 Hasselmo, M.E. et al. (1997) Noradrenergic feeding, stomach movements to nausea term follow-up of the Ischemic Heart Disease Life suppression of synaptic transmission may and stuffy air to respiration) but visceral Stress Monitoring Program. Psychosom. Med. 51, influence cortical signal-to-noise ratio. 485–513 J. Neurophysiol. 77, 3326–3339 sensory inputs have now been linked to 11 Talbot, F. and Nouwen, A. (2000) A review of the 25 Zagon, A. and Kemeny, A.A. (2000) Slow long-term, and sometimes pathological relationship between depression and diabetes in hyperpolarization in cortical neurones – a possible modulation of behavioural patterns. adults: is there a link? Diabetes Care 23, mechanism behind vagus nerve stimulation Any further analysis of the generation 1556–1562 therapy for refractory epilepsy? Epilepsia 41, of physiological and pathological 12 Frasure-Smith, N. (1991) In-hospital symptoms of 1382–1389 psychological stress as predictors of long-term 26 Valentino, R.J. et al. (1993) The locus coeruleus as behavioural responses need to consider the outcome after acute myocardial infarction in men. a site for integrating corticotropin-releasing factor delicate and constant interaction between Am. J. Cardiol. 67, 121–127 and noradrenergic mediation of stress responses. the CNS and the body to adjust the ability 13 George, M.S. (2000) Vagus nerve stimulation: a Ann. New York Acad. Sci. 697, 171–187 of the CNS to interpret and initiate a new tool for brain research and therapy. Biol. 27 van Bockstaele, E.J. et al. (1998) Amygdaloid corticotropin-releasing factor targets locus behavioural response appropriate to the Psychiatry 47, 287–295 14 Schachter, S.C. and Saper, C.B. (1998) Vagus coeruleus dendrites: substrate for the body’s ability to execute this response. nerve stimulation. Epilepsia 39, 677–686 coordination of emotional and cognitive limbs of When the body’s ability and the feedback 15 Zagon, A. et al. (1999) Late vagal inhibition in the stress response. J. Neuroendocrinol. 10, provided to the CNS are not in balance, the neurones of the ventrolateral medulla oblongata 743–757 discrepancy between the interpretation of in the rat. Neuroscience 92, 877–888 28 Ziegler, D.R. et al. (1999) Excitatory influence of the locus coeruleus in hypothalamic–pituitary– a sensory input and the execution of the 16 Zagon, A. (2001) Sciatic and vagal sensory inputs converge onto non-baroreceptive neurones of the adrenocortical axis responses to stress. response is expected to manifest in rostral ventrolateral medulla. Brain Res. 896, J. Neuroendocrinol. 11, 361–369 maladaptive behavioural responses. 64–68 29 Arborelius, L. et al. (1999) The role of 17 Sun, M.K. and Guyenet, P.G. (1987) Arterial corticotropin-releasing factor in depression and References baroreceptor and vagal inputs to anxiety disorders. J. Endocrinol. 160, 1–12 1 Onions, C.T., ed. (1965) The Shorter Oxford sympathoexcitatory neurons in rat medulla. Am. 30 Timpl, P. et al. (1998) Impaired stress response English Dictionary, Oxford University Press J. Physiol. 252, R699–R709 and reduced anxiety in mice lacking a functional 2 Crucia, G.P. et al. (2000) Emotional and 18 Aston-Jones, G. et al. (1986) The brain nucleus corticotropin-releasing hormone receptor. Nat. physiological responses to false feedback. Cortex locus coeruleus: restricted afferent control of a Genet. 19, 162–166 36, 623–647 broad efferent network. Science 234, 734–736 31 Maier, S.F. et al. (1998) The role of the vagus nerve 3 Valins, S. (1967) Emotionality and information 19 vanBockstaele, A.J. and Aston-Jones, G. (1995) in cytokine-to-brain communication. Ann. New concerning internal reactions. J. Pers. Soc. Integration in the ventral medulla and York Acad. Sci. 840, 289–300 Psychol. 6, 458–463 32 Bartorelli, C. et al. (1960) Inhibitory control of 4 Myerburg, R.J. and Castellanos, A. (1997) Cardiac coordination of sympathetic pain and arousal functions. Clin. Exp. Hypertens. 17, 153–165 sinocarotid pressoceptive afferents on arrest and sudden cardiac death. In Heart hypothalamic autonomic activity and sham rage Disease; A Textbook of Cardiovascular Medicine 20 Zagon, A. and Hughes, D.I. (2001) Somatic inhibition of vagal inputs is mediated pre- behavior. Arch. Ital. Biol. 98, 308–326 (Braunwald, E., ed.), Saunders Co. 33 Bonvallet, M. and Bloch, V. (1961) Bulbar control 5 Green, A.I. and Austin, C.P. (1993) synaptically in the rostral ventrolateral medulla oblongata. Eur. J. Neurosci. 13, 1–15 of cortical arousal. Science 133, 1133–1134 Psychopathology of pancreatic cancer: a 34 Juhasz, G. et al. (1985) Effects of hypnogenic psychobiologic probe. Psychosomatics 34, 208–221 21 Zagon, A. (2000) Activation of cardiac vagal afferents facilitates late vagal inhibition in vagal stimulation on thalamic neuronal activity 6 Hall, R.C. (2000) Anxiety and endocrine disease. in cats. Brain Res. Bull. 15, 437–441 Semin. Clin. Neuropsychiatry 4, 72–83 neurones of the rostral ventrolateral medulla 7 ten Thoren, C. and Petermann, F. (2000) Reviewing oblongata (RVLM) bilaterally. Brain Res. 854, asthma and anxiety. Respir. Med. 94, 409–415 172–177 Aniko Zagon 8 van Dixhoorn, J. et al. (1989) Physical training 22 Aston-Jones, G. et al. (1996) Role of the locus Dept Anatomy and Developmental Biology, coeruleus in emotional activation. Prog. Brain and relaxation therapy in cardiac rehabilitation Royal Free and University College Medical Res. 107, 379–402 assessed through a composite criterion for School, Rowland Hill Street, London, training outcome. Am. Heart J. 118, 545–552 23 Bremner, J.D. et al. (1996) Noradrenergic UK NW3 2PF. 9 Carr, R.E. (1999) Panic disorder and asthma. mechanisms in stress and anxiety. I. Preclinical J. Asthma 44, 143–152 studies. Synapse 23, 28–38 e-mail: [email protected]

Obituary

passion for outdoor activities, especially an instructorship back at Freiburg from Viktor Hamburger observing and collecting insects and larvae 1927–1932, Viktor accepted a Rockefeller in the fields and ponds near the family Fellowship to study in Frank Lillie’s (1900–2001) home. Following undergraduate training laboratory at Chicago, bringing the in Zoology at Breslau, Heidelberg, and precepts and methods of experimental ‘Our real teacher has been and still is the Munich, in 1921 Viktor became a graduate embryology to a laboratory in which the embryo, who is, incidentally, the only teacher student in the laboratory of Hans Spemann chick embryo was atop the pecking order. who is always right.’ V. Hamburger at Freiburg, during the period when the Because of political changes in his organizer was discovered and concepts of homeland, Viktor remained in the United Viktor Hamburger died 12 June 2001, one tissue interaction and cell determination States and in 1935 he began a nearly month shy of his 101st birthday. Born in were having their initial bloom. 50-year tenure at Washington University Landeshut, Silesia (then part of Germany; Following post-doctorates in in St Louis, including 25 years as now within Poland), Viktor grew up with a Göttingham and Berlin-Dahlem, and then chairman of the Zoology Department. He

inspiration during Viktor’s student days. (in the 1930s) to analyses of the influence Shortly after World War II, Viktor invited of vitalism on early embryologists. The Rita Levi-Montalcini to join him at Heritage of Experimental Embryology: Washington University to investigate Hans Spemann and the Organizer3 in together what seemed to be substantial 1988 brings to the current generations a differences in trophic effects on embryonic sense of the excitement and uncertainties sensory and motor neurons. This that were present during this early period collaboration led to the first identification of discovery, illuminating the personalities of a developmental growth factor, nerve and conceptual perspectives of key growth factor (NGF), which is essential for researchers involved. the differentiation and survival of many sensory and sympathetic (but not motor) ‘…Viktor’s interests and life perspectives neurons. Viktor certainly appreciated the were very much a melding of natural irony that decades of unsuccessful sciences with philosophy, literature and attempts to isolate the chemical nature of the arts.’ Spemann’s Organizer were eclipsed by this discovery of a neuron-specific trophic Viktor touched all who knew him with Portrait photo courtesy of Washington University factor. Today, many growth factors, often his wisdom, dignity, generosity and archives. acting in concert, are known to modulate kindness. He had little patience for cell phenotype in embryonic and adult imprecision of thought or language, in also participated for many years in the tissues, and several of Viktor’s early NGF science and also other endeavors, but Embryology Course at the Marine collaborators shared in receiving the taught always by example rather than Biological Laboratories in Woods Hole, Nobel Prize in 1986. criticism. In his presence discussions continuing the fondness for comparative seemed more purposeful and disciplined, embryology developed earlier. His Manual ‘[Viktor’s] Manual of Experimental yet his conversations betrayed a of Experimental Embryology1, first Embryology, first published in 1942, wonderful, often playful, sense of humor. published in 1942, demystified accessing demystified accessing and manipulating Viktor knew and worked with many of the and manipulating early embryos for early embryos for several generations of founders of experimental embryology several generations of students. students.’ during the first half of the 20th century, While embracing the methods of and of neuroembryology in the second half. experimental embryology, which Viktor, however, turned his attention to However, he did not dwell upon this past combined an analytical approach with the functional aspects of neurogenesis, with nostalgia or sentimentality. Rather, aesthetic appeal of directly engaging the investigating the interactions necessary he taught that the ideas enabling great living embryo, it was applications of these for the orderly development of embryonic accomplishments in science are timeless, methods to problems of the developing motility and patterns of limb movement. and we who practice – or are learning to nervous system that most captivated His goal was to dissect those network practice – developmental biology today Viktor and set the directions he would components that come into play benefit by understanding not only what actively pursue throughout his career. independent of inputs from those that our predecessors accomplished but also Viktor’s early studies on redirected axon require functional feedback. He and his why and how they did so. outgrowth led him to postulate that colleagues isolated parts of the embryonic A full description of the life, trophic factors released by nerves and nervous system and found that most publications, and many awards of Viktor target tissues are necessary for the proper behavior-generating neuronal networks Hamburger is available through the development of both. His subsequent develop normally. As always it was the Washington University Library web site studies showing that motor neuron chick embryo that served as a model (library.wustl.edu/~biology/vh/). survival was dependent upon the amount system. Indeed, it is because of the stage of target tissue available furthered this series of chick development that he and Drew M. Noden belief. During these studies involving Howard Hamilton devised in the early Professor of Anatomy and Animal chick wing transplantations or 1950s that Hamburger’s name is most Development, College of Veterinary extirpations, Viktor discovered that up to widely known among students of Medicine, Cornell University, Ithaca, 60% of developing motor neurons die in vertebrate development2. NY 14853-6401, USA. normal embryos. This discovery of Though best known for his work in e-mail: [email protected] programmed cell death, which is a crucial experimental embryology and component of most developing tissues, has neuroembryology, Viktor’s interests and References 1 Hamburger, V. (1942) Manual of Experimental subsequently emerged as a major focus in life perspectives were very much a Embryology, University of Chicago Press biomedical research. melding of natural sciences with 2 Hamburger, V. (1951) Stage 21 (4-day) chick Viktor’s interests in nerve-target philosophy, literature and the arts. His embryo from ‘A series of normal stages in the interactions led to, among others, his deep publication topics range from color development of the chick embryo.’ J. Morph. 88, 49–92 admiration for and friendship with changes in fish during their mating season 3 Hamburger, V. (1988) The Heritage of Ross Harrison, whose frequent visits to to the geology of Silesia, from mechanistic Experimental Embryology: Hans Spemann and Freiburg had been a source of much explanations of human birth defects the Organizer, Oxford University Press

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