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J. Biol. Chem., Vol. 283, Issue 21, 13, May 23, 2008
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Classics
Properties of the Specific Binding of 125I-Nerve Growth Factor to Responsive Peripheral Neurons
(Frazier, W. A., Boyd, L. F., and Bradshaw, R. A. (1974) J. Biol. Chem. 249, 5513–5519)
Properties and Specificity of Binding Sites for 125I-Nerve Growth Factor in Embryonic Heart and Brain
(Frazier, W. A., Boyd, L. F., Pulliam, M. W., Szutowicz, A., and Bradshaw, R. A. (1974) J. Biol. Chem. 249, 5918–5923)
Ralph A. Bradshaw was born in Dorchester, Massachusetts in 1941. He attended Colby College in Waterville, Maine, where he majored in chemistry. During his senior year at Colby, he took a course in biochemistry taught by Wilmon B. Chipman, a recent graduate of the University of Illinois and a new addition to the Colby faculty. Chipman was young and enthusiastic about biochemistry, and less than 2 weeks into the course Bradshaw decided that he would go to graduate school to pursue studies in biochemistry. He applied to several schools but settled on Duke University because of the prominent faculty in the biochemistry department and because Duke offered the most financial support.
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Bradshaw received his Ph.D. in 1966 and then spent a year doing a postdoctoral fellowship with Frank Gurd at Indiana University where he sequenced two aquatic mammal myoglobins. After that he joined JBC Classic author Hans Neurath (3) at the University of Washington where he completed the sequence of bovine carboxypeptidase A.
In 1969, Bradshaw was offered a position on the faculty of the Department of Biological Chemistry at Washington University School of Medicine in St. Louis. He accepted the offer knowing that the position would provide ample opportunity for productive collaborations. Sure enough, soon after arriving in St. Louis, Bradshaw, in collaboration with Rita Levi-Montalcini, determined the sequence of mouse nerve growth factor (NGF) with Ruth Angeletti (4). This initial collaboration with Levi-Montalcini, who was awarded the Nobel Prize in 1986 for her pioneering work in discovering NGF, led Bradshaw to spend more than 35 years studying growth factors, their receptors, and their signaling mechanisms. Some of his NGF receptor studies are the subject of the two JBC Classics reprinted here.
Both papers are binding studies that showed that iodinated NGF bound to NGF-responsive cells (peripheral neurons) with high affinity and specificity, thus establishing that they had cell surface receptors and were hormonal in nature, an idea that directly arose from the amino acid sequence determination. In the first paper Bradshaw and his colleagues study the interaction of NGF with its receptors on the surface membranes of responsive chick embryo sympathetic and sensory neurons and report that the binding is a complex process with properties similar to the binding of insulin to fat and liver cell membranes and human lymphocytes. In the second Classic, Bradshaw and his colleagues continue their binding studies, this time using embryonic heart and brain tissues from chickens, and report that binding in these tissues is very similar to that in the sympathetic and sensory nervous systems. This latter study presaged the much wider range of responsive tissues for NGF, particularly the central nervous system, that was eventually established several years later.
The results of these two papers shook up the field as, up until then, NGF and epidermal growth factor, which had been discovered in the mouse submandibular gland by JBC Classic author Stanley Cohen (5) while studying NGF, were not even considered to be related to hormones. Immediately, many other substances such as platelet-derived growth factor, the somatomedins (now insulin-like growth factors), the transforming growth factors, and the fibroblast growth factors shot to center stage, and the growth factor field, as an expansion of endocrinology, was born with a vengeance. Bradshaw and colleagues eventually characterized and isolated the NGF receptor, TrkA (6–8), substantially before it was cloned.
In 1982, Bradshaw moved to California where he became Professor and Chair of the Department of Biological Chemistry at the University of California, Irvine. He is currently Professor Emeritus in the Department of Physiology & Biophysics at the University of California, Irvine as well as Professor in the Department of Pharmaceutical Chemistry and Deputy Director of the Mass Spectrometry Facility at the University of California, San Francisco.
Bradshaw has been very active in the American Society for Biochemistry and Molecular Biology, serving on the Committee on Equal Opportunities for Minority Groups, the Nominating Committee, and the Membership Committee and also acting as Society Treasurer and Councilor. He is currently Deputy Chair of the ASBMB Public Affairs Advisory Committee and Society Historian. Bradshaw was also the President of the Federation of American Societies for Experimental Biology, Treasurer of the Keystone Symposia, and the founding President of the Protein Society. He has served on many editorial boards and was an Associate Editor of the Journal of Biological Chemistry. He is currently Co-editor of Molecular and Cellular Proteomics.
In recognition of his contributions to science, Bradshaw has received many awards and honors including the Passano Foundation Young Scientist Award (1976), the American Chemical Society St. Louis Section Award (1979), the Italian National Research Council Gold Medal (1987), and the Australia Society of Medical Research Gold Medal (1999).1,2
The first author of both the Bradshaw Classics, William A. Frazier, has also gone on to have a very productive career in science. At the time the papers were published, Frazier was a predoctoral student in Bradshaw's laboratory. A few years later, he became an Assistant Professor in the Department of Biological Chemistry at Washington University School of Medicine. He remains there today and is currently a Professor of Cell Biology and Physiology and Professor of Biomedical Engineering. His research focuses on the function, structure, and mechanism of thrombospondin-1 (TSP1) and its receptors, CD36 and CD47.
FOOTNOTES
1 Biographical information on Ralph Bradshaw was taken from Ref. 2. 
2 We would like to thank Ralph Bradshaw for providing background information for this introduction.
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