Harry Rubin, who helped establish an understanding of how normal cells are transformed into cancer cells and won (along with Rento Dulbecco) a 1964 Albert Lasker Basic Medical Research Award, has died at age 93. Rubin, whose work with the Rous sarcoma virus (RSV) brought him to the Virus Laboratory at the University of California, Berkeley, demonstrated how normal in vitro cells could turn into cancer cells when infected with RSV.

Rubin showed that cells in RSV-induced tumors are capable of releasing the virus, implying that RSV was permanently associated with the host cell and suggested that it plays a direct and continuing role in perpetuating the cell in its malignant state. Rubin’s investigations laid the groundwork for further research that established the existence of oncogenes.