American Philosophical Society
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1Name:  Dr. Elizabeth H. Blackburn
 Institution:  University of California, San Francisco
 Year Elected:  2005
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Living
 Birth Date:  1948
   
 
Dr. Elizabeth Blackburn is a leader in the area of telomere and telomerase research and has made key discoveries in different aspects of telomere function and biology. In 1985, she discovered the ribonucleoprotein enzyme telomerase, and since that time, hers has become a lead laboratory in manipulating and studying telomerase activity in cells. Having amassed considerable knowledge and experience in the effects this has on cells, Dr. Blackburn and her research team at the University of California, San Francisco worked with a variety of organisms and human cells, especially cancer cells, with the goal of understanding telomerase and telomere biology. Her work on telomeres and telomerase has been published extensively in peer-reviewed journals. Dr. Blackburn earned her B.Sc. (1970) and M.Sc. (1972) degrees from the University of Melbourne in Australia, and her Ph.D. (1975) from the University of Cambridge in England. She did her postdoctoral work in molecular and cellular biology from 1975-77 at Yale University. In 1978, Dr. Blackburn joined the department of molecular biology at the University of California Berkeley. In 1990, she joined the departments of microbiology and immunology, and biochemistry and biophysics, at the University of California, San Francisco, and she was department chair of the department of microbiology and immunology from 1993-99, and the Morris Herzstein Professor of Biology and Physiology in the department of biochemistry and biophysics at UCSF as well as a non-resident fellow of the Salk Institute. In January 2016 Dr. Blackburn became professor emeritus at the University of California, San Francisco, and was President of the Salk Institute for Biological Studies until 2018. Throughout her career, Dr. Blackburn has been honored by her peers as the recipient of many prestigious awards. These include the Eli Lilly Research Award for Microbiology and Immunology (1988), the National Academy of Science Award in Molecular Biology (1990), and honorary doctorate degrees from Yale University (1991), the University of Pennsylvania (2004), Bard College (2004), Brandeis University (2004), and the University of Chicago (2004). She was a Harvey Society Lecturer at the Harvey Society in New York (1990) and recipient of the UCSF Women's Faculty Association Award (1995). Most recently, she was awarded the Australia Prize (1998), the Harvey Prize (1999), the Keio Prize (1999), the American Association for Cancer Research-G.H.A. Clowes Memorial Award (2000), the American Cancer Society Medal of Honor (2000), the AACR-Pezcoller Foundation International Award for Cancer Research (2001), the General Motors Cancer Research Foundation Alfred P. Sloan Award (2001), the Bristol Myers Squibb Award for Distinguished Cancer Research (2003), the Dr. A.H. Heineken Prize for Medicine (2004), the Kirk A. Landon-American Association for Cancer Research Prize for Basic Cancer Research (2005), the Benjamin Franklin Medal in Life Science (2005), and the Nobel Prize in Medicine (2009). She was named California Scientist of the Year in 1999 and was elected President of the American Society for Cell Biology for the year 1998. Dr. Blackburn is an elected Foreign Associate of the National Academy of Sciences (1993) and an elected Member of the Institute of Medicine (2000). She is an elected Fellow of the American Academy of Arts & Sciences (1991), the Royal Society of London (1992), and the American Association for the Advancement of Science (2000). She was elected Foreign Associate of the National Academy of Sciences in 1993 and was elected as a Member of the Institute of Medicine in 2000. She was awarded the Albert Lasker Medical Research Award in Basic Medical Research (2006). In 2007 she was named one of TIME Magazine's 100 Most Influential People, and she is the 2008 North American Laureate for L'Oreal-UNESCO for Women in Science.
 
2Name:  Dr. Aaron J. Ciechanover
 Institution:  Technion - Israel Institute of Technology
 Year Elected:  2005
 Class:  2. Biological Sciences
 Subdivision:  202. Cellular and Developmental Biology
 Residency:  International
 Living? :   Living
 Birth Date:  1947
   
 
Aaron Ciechanover was born in Haifa, Israel in 1947. He is currently on the academic staff of the Faculty of Medicine of the Technion in Haifa, Israel. He received his M.Sc. (1971) and M.D. (1975) from the Hebrew University in Jerusalem, Israel, and his D.Sc. (1982) from the Technion. There, as a graduate student with Dr. Avram Hershko and in collaboration with Dr. Irwin A. Rose from the Fox Chase Cancer Center in Philadelphia, they discovered that covalent attachment of ubiquitin to the target substrate signals it for degradation. They deciphered the mechanism of conjugation in a cell-free system, described the general proteolytic function of the system in cells, and proposed a model according to which this modification serves as a recognition signal for a specific downstream protease. As a post doctoral fellow with Dr. Harvey Lodish at the Massachusetts Institute of Technology, he collaborated with Drs. Alexander Varshavsky and Daniel Finley, and described the first mutant cell of the system, further corroborating the role of ubiquitin modification as a proteolytic signal in intact cells. Among the many prizes that Dr. Ciechanover received are the 2000 Albert Lasker Award for Basic Medical Research and the 2004 Nobel Prize in Chemistry. Dr. Ciechanover is a member of the Israeli National Academy of Sciences and Humanities, a foreign associate of the National Academy of Sciences of the USA and a member of the Pontifical Academy of Sciences of the Vatican.
 
3Name:  Dr. Elaine Fuchs
 Institution:  Rockefeller University; Howard Hughes Medical Institute
 Year Elected:  2005
 Class:  2. Biological Sciences
 Subdivision:  202. Cellular and Developmental Biology
 Residency:  Resident
 Living? :   Living
 Birth Date:  1950
   
 
Elaine Fuchs is a world leader in mammalian cell biology. She is internationally recognized for her outstanding and numerous contributions to skin biology and its human genetic disorders, including skin cancers and life-threatening genetic syndromes such as blistering skin disorders. For nearly three decades, Dr. Fuchs has focused on the molecular mechanisms that underlie development and differentiation of the epidermis and its appendages, and elucidating how perturbations of these mechanisms result in disease. She has systematically and brilliantly applied innovative approaches in biology, biochemistry and genetics. In doing so, Dr. Fuchs pioneered the use of "reverse genetics," an approach to start with a specific protein, study its biology and then use mice as a means to ultimately identify the genes responsible for inherited human disorders. A classical geneticist would start with a specific genetic disorder. Instead, Dr. Fuchs has employed this innovative cell biological approach to determine the genetic bases of numerous dermatological disorders in humans. The approach has since broadly benefited human medical genetics. Dr. Fuchs is widely recognized as having brought the field of dermatological research into modern day science. Her contributions are many, ranging from the identification of proteins and signal transduction pathways important in epidermal and hair functions to uncovering the molecular nature of skin diseases in humans. In addition, Dr. Fuchs and coworkers identified genetic defects in several disorders that arise from perturbations of cytoskeletal proteins related to those present in the skin, but whose expression resides outside the skin, particularly in the muscle and the nervous system. An elegant example of this is her use of reverse genetics to uncover the underlying genetic basis of blistering human skin disorder that arises from defects in epidermal keratin genes. Dr. Fuchs' 10 years of prior research set the groundwork for this discovery, which uncovered a key function of intermediate filament (IF) proteins as mechanical integrators of the cytoskeleton. The work also set the paradigm for more than 20 different human disorders of IF genes. Dr. Fuchs' ground-breaking research is often used in biology and medical textbooks as a landmark. Her science now focuses on understanding how tissues develop and dynamically respond to their environment. She has seamlessly transitioned from problems of signal transduction to transcriptional regulation and gene expression to the cytoskeleton and adhesion to stem cell lineage commitment. In the nineties, her team uncovered multiple roles for Wnt signaling in skin biology, discovering that sustained Wnt signaling can lead to stem cell activation and tumorigenesis. Their super-furry mice led them to identify stabilizing b-catenin mutations pilomatricomas, a human skin tumor. While b-catenin mutations had been previously linked to colon cancer, pilomatricomas represented the first example where b-catenin mutations are the leading cause of the tumor. Similarly, Dr. Fuchs' work on a-catenin provided insights into squamous cell carcinoma. The lab's transition from degenerative disorders to cancers has been a natural one, occurring concomitantly with their shift to tackling how growth and differentiation are balanced in stem cell lineage progression. Their recent work in isolating and characterizing the multipotent adult skin stem cells opens major new avenues for their future research in this area. Elaine Fuchs received her undergraduate degree with highest distinction in chemistry from the University of Illinois, Urbana-Champaign (1972). She received her Ph.D. in biochemistry from Princeton University (1977) and conducted her postdoctoral studies with Howard Green at the Massachusetts Institute of Technology, where she began her research in skin biology. She joined the faculty at the University of Chicago in 1980, where she progressed to become Amgen Professor of Basic Sciences prior to leaving for Rockefeller University in 2002, where she is Rebecca C. Lancefield Professor. She has been an Investigator of the Howard Hughes Medical Institute since 1988. Dr. Fuchs is a member of the National Academy of Sciences, the Institute of Medicine, and the American Academy of Arts & Sciences. She was President of the American Society of Cell Biology in 1991, and she holds an honorary doctorate from Mt. Sinai and New York University School of Medicine. Her scientific awards include the Richard Lounsbery Award (National Academy of Sciences), the Cartwright Award (Columbia University), the Novartis Award in Biomedical Research, the Dickson Prize in Medicine, the National Medal of Science, the 2010 L'Oreal-UNESCO prize, the 2012 March of Dimes Prize in Developmental Biology, the 2015 E. B. Wilson Medal, the 2016 Vanderbilt Prize in Biomedical Science, the 2019 AACR-G.H.A. Clowes Memorial Award, and the 2020 Canada Gairdner Award. She has trained more than 20 graduate students and has over 225 publications to her credit. Elaine Fuchs was elected a member of the American Philosophical Society in 2005.
 
4Name:  Dr. Avram Hershko
 Institution:  Technion-Israel Institute of Technology
 Year Elected:  2005
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  International
 Living? :   Living
 Birth Date:  1937
   
 
Avram Hershko was born in 1937 in Karcag, Hungary and emigrated with his family to Israel in 1950. He gained his M.D. (1965) and Ph.D. (1969) from the Hebrew University - Hadassah Medical School of Jerusalem, a period which included service as a physician in the Israel Defence Forces (1965-67). After a post-doctoral fellowship with Gordon Tomkins at the University of San Francisco (1969-72), he joined the faculty of the Haifa Technion, becoming professor in 1980. He is now Distinguished Professor in the Unit of Biochemistry in the B. Rappaport Faculty of Medicine of the Technion. His main research interests concern the mechanisms by which cellular proteins are degraded, a formerly neglected field of study. Dr. Hershko and his colleagues showed that cellular proteins are degraded by a highly selective proteolytic system. This system tags proteins for destruction by linkage to a protein called ubiquitin, which had previously been identified in many tissues, as the name suggests, but whose function was previously unknown. Subsequent work in Dr. Hershko's and many other laboratories has shown that the ubiquitin system has a vital role in controlling a wide range of cellular processes, such as the regulation of cell division, signal transduction and DNA repair. Abnormalities in the ubiquitin system result in diseases such as certain types of cancer. The full range of functions of the ubiquitin system in health and disease has still to be elucidated. Dr. Hershko was awarded the Nobel Prize in Chemistry (2004) jointly with his former Ph.D. student Aaron Ciechanover and their colleague Irwin Rose. His many honors include the Israel Prize for Biochemistry (1994), the Gardner Award (1999), the Lasker Prize for Basic Medical Research (2000), the Wolf Prize for Medicine (2001) and the Louisa Gross Horwitz Award (2001). Dr. Hershko is a member of the Israel Academy of Sciences (2000) and a Foreign Associate of the National Academy of Sciences (2003).
 
5Name:  Dr. Richard Losick
 Institution:  Harvard University
 Year Elected:  2005
 Class:  2. Biological Sciences
 Subdivision:  202. Cellular and Developmental Biology
 Residency:  Resident
 Living? :   Living
 Birth Date:  1943
   
 
Richard Losick received his B.A. from Princeton University in 1965 and his Ph.D. from the Massachusetts Institute of Technology in 1969. He was elected to the Harvard Society of Fellows as a Junior Fellow in 1969, and in 1972 he joined the faculty of Harvard University, where he is currently the Maria Moors Cabot Professor of Biology, a Harvard College Professor, and a Howard Hughes Medical Institute Professor. He is a past chairman of the Department of Molecular and Cellular Biology and the Department of Cellular and Developmental Biology. He teaches the introductory course on molecular biology at Harvard College, and as Head Tutor he is responsible for the undergraduate concentration in Biochemical Sciences. He is a member of the National Academy of Sciences, a Fellow of the American Academy of Arts & Sciences, a Fellow of the American Association for the Advancement of Science, a Fellow of the American Academy of Microbiology, and a former Visiting Scholar of the Phi Beta Kappa Society. His research interests include RNA polymerase, gene transcription and its control, and development in microorganisms. Recently, Dr. Losick was honored with the 2007 Selman A. Waksman Award in Microbiology for "discovering alternative bacterial sigma factors and his fundamnetal contributions to understanding the mechanicsm of bacterial sporulation" and the 2012 Louisa Gross Horwitz Prize for discovering the structure of bacteria.
 
6Name:  Dr. Roderick MacKinnon
 Institution:  Rockefeller University; Howard Hughes Medical Institute
 Year Elected:  2005
 Class:  2. Biological Sciences
 Subdivision:  206. Physiology, Biophysics, and Pharmacology
 Residency:  Resident
 Living? :   Living
 Birth Date:  1956
   
 
Roderick MacKinnon received an undergraduate degree from Brandeis University, a medical degree from Tufts University, and training in Internal Medicine at Beth Israel Hospital, Harvard Medical School. He then began his scientific career studying the biophysics of potassium channels at Brandeis University from 1986-89. He joined the faculty at Harvard Medical School as Assistant Professor of Physiology (1989), Associate Professor of Neurobiology (1992) and Professor of Neurobiology (1995). During this period he and his laboratory characterized potassium channels - their subunit stoichiometry, pore-lining amino acids, and components of their gates - through biochemical and functional analysis. He then moved to Rockefeller University in 1996 where his laboratory has provided the first atomic descriptions of ion selective membrane channels. He is currently a professor in the laboratory of Molecular Neurobiology and Biophysics at Rockefeller University and Investigator of the Howard Hughes Medical Institute. In 2003 he shared the Nobel Prize in Chemistry with Peter Agre for his structural and mechanistic studies of ion channels.
 
7Name:  Dr. Maynard V. Olson
 Institution:  University of Washington
 Year Elected:  2005
 Class:  2. Biological Sciences
 Subdivision:  207. Genetics
 Residency:  Resident
 Living? :   Living
 Birth Date:  1945
   
 
Maynard Olson graduated from Caltech with a Bachelor's degree in chemistry and received his Ph.D. in inorganic chemistry from Stanford University in 1970, where his thesis advisor was Henry Taube. After five years on the faculty of the Department of Chemistry at Dartmouth College, he changed his research emphasis to molecular genetics, working with Benjamin Hall in the Department of Genetics at the University of Washington. During that period, in the late 1970's, he participated in early applications of recombinant-DNA techniques to problems in yeast genetics; his research with Hall included the first sequencing of a mutant eukaryotic gene and one of the first applications of restriction-fragment length polymorphisms. In 1979, he moved to the Department of Genetics at Washington University in St. Louis, where he became a Professor of Genetics in 1986 and an Investigator of the Howard Hughes Medical Institute in 1989. At Washington University, he participated in the development of systematic approaches to the analysis of complex genomes, working both on the yeast and human genomes. This research included the development of new implementations of pulsed-field gel electrophoresis, including field-inversion gel electrophoresis, determination of the first complete electrophoretic karyotype of a eukaryotic organism, the development of computer-based methods for the construction of whole-genome physical maps based on clone fingerprints, the development of the yeast-artificial-chromosome cloning system, and the introduction of STS-content mapping as an approach to the low-resolution physical mapping of mammalian genomes. In 1992, he was awarded the Genetics Society of America Medal for outstanding contributions to genetics during the previous 15 years. Later that year, he moved back to the University of Washington where he is now Professor of Medicine and Genome Sciences and Director of the University of Washington Genome Center. In 1994, he was elected to the National Academy of Sciences. In 2000, he received the City of Medicine Award for exceptional contributions to medicine in the public interest, and in 2002, he received the Gairdner Foundation International Award for his scientific contributions to the Human Genome Project. Dr. Olson has also participated extensively in the formulation of policy for the Human Genome Project: in 1987, he served on the National Research Council Committee on Mapping and Sequencing of the Human Genome; from 1989-92, on the Program Advisory Committee on the Human Genome at the National Institutes of Health; from 1999-2003, and on the National Human Genome Research Institute Council. Dr. Olson's current research is focused on the analysis of natural genetic variation both in bacteria, particularly Pseudomonas aeruginosa, and humans.
 
8Name:  Dr. Louis Sokoloff
 Institution:  National Institutes of Health
 Year Elected:  2005
 Class:  2. Biological Sciences
 Subdivision:  209. Neurobiology
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1921
 Death Date:  July 30, 2015
   
 
Born in Philadelphia in 1921, the second of two sons of Eastern European immigrants, Louis Sokoloff early discovered the power of books in satisfying his curiosity and thirst for knowledge. His undergraduate education at the University of Pennsylvania was highlighted by exposure to the illustrious physiologist L. V. Heilbrunn. Contacts with this professor in and out of the classroom stimulated his love of science, which never faltered. It was Heilbrunn who first noted Sokoloff's potential and who steered him toward a professional career--he recommeneded medical studies. An interest in cell excitation on the one hand, stemming back to his undergraduate days, and assignments to psychiatric services in his medical internship as well as during his subsequent Army service on the other, intensified Sokoloff's interest in brain function. It was during this period that he met his future wife Betty. Following his return to civilian life in 1949, he sought to renew his interest in research, and was drawn to the laboratory of the then-35 year old Seymour Kety, who had just landed an NIH grant at Penn and was looking for a young associate. Soon thereafter, Dr. Sokoloff became immersed in learning the theoretical and practical aspects of the nitrous oxide technique for measuring the rate of cerebral blood flow (CBF) in humans. The method is based on Kety's mathematical model that derived the flow rate from measurement of brain uptake and release of diffusible substance. Low concentrations of the inert gas nitrous oxide was used for this purpose. During this period, Sokoloff made his first measurement of cerebral metabolism in hyperthyroidism, finding it not to be elevated even when the body's basal metabolic rate was nearly double. This led him to the hypothesis that thyroid hormones stimulate protein synthesis and to his interest in the thyroid hormone functions, a subject to which he subsequently made significant contributions. Many studies were performed on human subjects by Kety, Sokoloff, and co-workers, examining the rates of CBF and metabolism in mental activity, sleep, anesthesia, and under the influence of various pharmacological agents. In 1951, Kety moved to the National Institutes of Health (NIH), and offered Sokoloff the opportunity to join him. He accepted, and eventually was appointed Chief of the Laboratory of Cerebral Metabolism at the NIH, a title he retained for 35 years. At the NIH, Sokoloff collaborated with Kety, William Landau, Lewis Rowland and Walter Freygang, in developing a quantitative autoradiographic technique for measuring regional CBF in animals which he used to demonstrate a clear linkage between functional activity and regional blood flow in visual pathways of the brain. The autoradiographs from this study represented the first ever published demonstration of functional brain imaging. Sokoloff then used the quantitative autoradiographic technique to develop a method for the measurement of regional brain metabolism of glucose, the almost exclusive substrate for energy metabolism in the brain. There is little need to recount in detail Sokoloff's pioneering studies on regional cerebral glucose utilization for which he introduced the use of -2deoxy-D-[14C]glucose. The elegance of the deoxyglucose method itself and the great care taken in its quantification and in defining its limits are all reflections of Sokoloff's research style. Adaptation of the method to human studies was subsequently accomplished by means of single photon and positron emission tomography, in which [18F]fluorodeoxyglucose replaced the 14C=labeled compound, and was the result of a collaborative effort between Sokoloff, Martin Reivich, David Kuhl, Alfred Wolf, and Michael Phelps. The many tributes already paid Dr. Sokoloff attest to his accomplishments. Among his honors were membership in numerous societies and professional organizations including the National Academy of Sciences, the Institute of Medicine, and the American Academy of Arts & Sciences. He was elected a member of the American Philosophical Society in 2005. He served as president of the American Society for Neurochemistry, the International Society for Cerebral Blood Flow and Metabolism, and the Association for Research in Nervous and Mental Disease and on various editorial and advisory boards. He was the recipient of the Distinquished Service Award of the Department of Health, Education, and Welfare (1976); the F.O. Schmitt Medal in Neuroscience (1980); the Albert Lasker Clinical Medical Research Award (1981); the Karl Spencer Lashley Award of the American Philosophical Society (1987); the National Academy of Sciences Award in the Neurosciences (1988); the Georg Charles de Hevesy Nuclear Medicine Pioneer Award of the Society of Nuclear Medicine (1988); the Award of the Mihara Cerebrovascular Disorder Research Promotion Fund (1988); the Vicennial Medal, Georgetown University (1994); Lifetime Achievement Award of the Society of Biological Psychiatry (1996); and the Ralph Gerard Award of the Society of Neuroscience (1996). Dr. Sokoloff died July 30, 2015, at the age of 93, in Washington, DC.
 
9Name:  Sir David J. Weatherall
 Institution:  University of Oxford
 Year Elected:  2005
 Class:  2. Biological Sciences
 Subdivision:  204. Medicine, Surgery, Pathology and Immunology
 Residency:  International
 Living? :   Deceased
 Birth Date:  1933
 Death Date:  December 8, 2018
   
 
David Weatherall was a life-long student of the thalassemias. He was involved in identifying the general molecular nature of this group of hereditary anemias and in describing the genetic and clinical heterogeneity of both alpha- and beta-thalassemias. He also studied their influence on populations in many parts of the world and the role of malaria in determining their frequency. Both clinician and scientist, editor of the Oxford Textbook of Medicine and author of The New Genetics in Clinical Practice, Dr. Weatherall has played a significant role in bringing molecular genetics into the main stream of clinical medicine. He has been associated with the University of Oxford for more than thirty years as Nuffield Professor of Clinical Medicine (1974-92), Regius Professor of Medicine (1992-2000) and, after 2001, Regius Professor of Medicine Emeritus and Honorary Director of the Weatherall Institute of Molecular Medicine at the University of Oxford. In 2002 he was appointed Chancellor of Keele University. He was a member of the American Academy of Arts & Sciences (1988); the Royal Society (vice president, 1990-91); the National Academy of Sciences (1990); and the Institute of Medicine (1990). David Weatherall was elected a member of the American Philosophical society in 2005. He died on December 8, 2018 at the age of 85.
 
Election Year
2005[X]