Lynn Margulis, noted biologist and professor at UMass-Amherst’s Department of Geosciences, died Nov. 22 at the age of 73.

A widely respected pioneer in the field of microbiology, Margulis was a fierce proponent of symbiotic theories of evolution that often went against the grain of established neo-Darwinian theory. After she published The Origin of Eukaryotic Cells in 1970, Margulis’ formerly marginalized work on organelles (structures within cells that perform specific functions) became a central and pivotal body of research, and a foundation for much of the field of endosymbiotic theory. She helped conceive the Gaia Theory (a biological variant of systems theory), which holds that the Earth is a living entity that seeks its own wellbeing, with English chemist James Lovelock, and was instrumental in gaining widening acceptance for it in the scientific community beginning in the 1970s.

Margulis held numerous honors and awards, including the Linnean Society of London’s Darwin-Wallace Medal (2008), the National Medal Medal of Science (1999) and the William Proctor Prize for Scientific Achievement. She was inducted to several scientific societies and academies, including the American Academy of Arts and Sciences, the Russian Academy of Natural Sciences, the National Academy of Sciences, the Leonardo DaVinci Society of Thinking and the World Academy of Art and Science. Margulis married astronomer Carl Sagan in 1957, and is survived by her four children, Jennifer Margulis, Zachary Margulis-Ohnuma, Jeremy Sagan and Dorion Sagan. With the latter she co-authored several recent books, including What is Sex?, Microcosmos: Four Billion Years of Evolution from Our Microbial Ancestors and Slanted Truths: Essays on Gaia, Symbiosis, and Evolution.