Oliver Smithies. 23 June 1925—10 January 2017

Oliver Smithies was born in Copley, near Halifax in Yorkshire, UK. He received his doctorate from Oxford in 1951, then began working at the Connaught Laboratories in Toronto, where he developed starch gel electrophoresis. This technology allowed identification of genetic variants in human serum proteins and revolutionized protein analysis. After moving to the University of Wisconsin, he studied the genetics of antibody variability, then turned to nucleic acid methods, developing safe cloning vectors, driving production of software for genetic analysis, sequencing several human genes and finally creating genetically engineered animals, for which he later received the Nobel Prize. He then moved to the University of North Carolina, where he developed methods for altering gene dosage in mice, which he used to develop ways to attack complex physiological questions, including blood pressure regulation. Finally, he formulated a new hypothesis to explain kidney glomerular filtration, then devised methods that confirmed the hypothesis. Oliver collaborated with his wife, Nobuyo Maeda, during a long and happy marriage. While maintaining separate laboratories, they stimulated each other's scientific understanding and frequently published together. During a 70-year life in science, he mentored many students, postdoctoral fellows and collaborators, nearly all remaining his friends for life.