Richard Axel (born July 2, 1946) is an American molecular biologist and university professor in the Department of Neuroscience at Columbia University and investigator at the Howard Hughes Medical Institute. His work on the olfactory system won him and Linda Buck, a former postdoctoral research scientist in his group, the Nobel Prize in Physiology or Medicine in 2004. Education and early life Born in New York City to Polish Jewish immigrants, Axel grew up in Brooklyn. He graduated from Stuyvesant High School in 1963, (along with Bruce Bueno de Mesquita and Alexander Rosenberg), received his B.A. in 1967 from Columbia University, and his M.D. in 1971 from Johns Hopkins University. However, he was poorly suited to medicine and graduated on the promise to his department chairman that he would not practice clinically. He found his calling in research and returned to Columbia later that year, eventually becoming a full professor in 1978. Research and career During the late 1970s, Axel, along with microbiologist Saul J. Silverstein and geneticist Michael H. Wigler, discovered a technique of cotransformation via transfection, a process which allows foreign DNA to be inserted into a host cell to produce certain proteins. A family of patents, now colloquially referred to as the "Axel patents", covering this technique were filed for February 1980 and were issued in August 1983. As a fundamental process in recombinant DNA research as performed at pharmaceutical and biotech companies, this patent proved quite lucrative for Columbia University, earning it almost $100 million a year at one time, and a top spot on the list of top universities by licensing revenue. The Axel patents expired in August 2000. In their landmark paper published in 1991, Buck and Axel cloned olfactory receptors, showing that they belong to the family of G protein coupled receptors. By analyzing rat DNA, they estimated that there were approximately one thousand different genes for olfactory receptors in the mammalian genome. This research opened the door to the genetic and molecular analysis of the mechanisms of olfaction. In their later work, Buck and Axel have shown that each olfactory receptor neuron remarkably only expresses one kind of olfactory receptor protein and that the input from all neurons expressing the same receptor is collected by a single dedicated glomerulus of the olfactory bulb. Axel's primary research interest is on how the brain interprets the sense of smell, specifically mapping the parts of