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The passing of Marilyn Farquhar on Nov. 23 brought to a close a long and inspiring career in cell biology. She rightly ranks among the pioneers in her field and especially as a role model for women in science.

Rockefeller University Press

Marilyn Farquhar was an electron microscopy pioneer and a treasured mentor to generations of cell biologists.

We met Marilyn more than 50 years ago at the Rockefeller University when she was a visiting professor in George Palade’s lab. Marilyn had worked previously with Palade, and their early and classic insights into the organization of epithelial junctions, published in the period 1963-65, remain a staple in medical and cell biology textbooks to this day. 

Here were characterized, in , along with basement membranes that Marilyn was later to . These are the defining feature of all epithelia in our bodies.
 
At Rockefeller in 1969, Marilyn was part of an amazing group of colleagues that included David Sabatini, G眉nter Blobel, Phil Siekevitz, Jim Jamieson and David Luck. Marilyn and George had close interactions with Christian De Duve and his group, located in the same building at Rockefeller. Collectively, the pioneers’ focus was on deciphering mechanistically the workings of the cell, especially its membrane-bounded organelles, which often were being seen and studied for the first time. 

Marilyn came for her sabbatical at Rockefeller from her lab in the department of experimental pathology at the University of California, San Francisco. Shortly after she arrived, a paper from her work at UCSF was published that conceptually unified and showed in intricate detail the structures that captivated the Palade and De Duve labs.

She was clearly among the standard setters in a department and institution loaded with scientific talent. Furthermore, because she was both friendly and easily approachable, she enriched the scientific environment of numerous students and colleagues and a host of collaborators. She married George Palade in 1970, and her sabbatical at Rockefeller turned into an extended transcontinental migration. Together, they moved to Yale School of Medicine in 1973, where she became the Sterling professor of cell biology. At Yale, she was a major force in shaping the new section of cell biology that ultimately became the department of cell biology. As she continued to make contributions in discovery research, she also developed a modern cell biology course that became a valued component of the medical school curriculum.

At Rockefeller and at Yale, Marilyn and George were close friends and colleagues of visiting Romanian scientists Nicolae and Maya Simionescu, and they strongly encouraged Nicolae and Maya, who were pioneers in bringing modern cell biology to Romania. Marilyn and George were honored guests at the inauguration of the Institute of Cell Biology and Pathology in Bucharest, celebrated in 1979 with an international symposium featuring giants in the field, all of whom were Marilyn’s close friends and colleagues.

Courtesy of Maya Simionescu

This photo taken in 1979 at the inaugural symposium of ICBP Bucharest shows, in the front row, from left to right, Werner Franke, Maya Simionescu, Gunter Blobel, Marilyn Farquhar, George Palade, Christian De Duve, Nicolae Simionescu and David Sabatini.

Marilyn was always proud that she was California born and raised. She earned her undergraduate and graduate degrees at the University of California, Berkeley, and then moved to the Rockefeller University in New York City for postdoctoral training with George Palade before joining the faculty at UCSF. In 1990, she returned to California as the founding chair of the department of cellular and molecular medicine of the University of California, San Diego, Medical School.

We continue to appreciate Marilyn’s early contributions to understanding the developmental regulation of leukocyte granulogenesis and granule turnover in endocrine cells by lysosomal degradation. She made numerous contributions to unraveling the intricacies of the Golgi apparatus, including biogenesis, sorting and recycling. Her discoveries on the biogenesis of glomerular basement membranes included the classic definition of the role of associated proteoglycans in regulating glomerular filtration. She and her colleagues clarified the organization, trafficking, signaling and pathological roles of podocalyxin and megalin in renal podocytes and proximal tubule cells.

Rockefeller University Press

Left: This image from a 1965 paper in the Journal of Cell Biology by Marilyn Farquhar and George Palade shows a desmosome linking the plasma membranes of cells located in the granular (SG) and cornified layers of amphibian skin. Magnification x 135,000. Right: This image from a 1970 paper in the Journal of Cell Biology by Dorothy Bainton and Marilyn Farquhar shows concentration of myelocyte peroxidase within the secretory pathway of a rabbit eosinophil beginning in the endoplasmic reticulum (labeled er) and progressing through the Golgi (Gc) and maturing secretory granules (ig). Parallel observations for acid phosphatase and arylsulfatase pointed to the lysosome-like nature of these granules. Magnification x 43,000.

As time progressed, Marilyn became a major force in exploring the roles of G protein-coupled receptors, growth factor receptors and associated regulatory complexes in intracellular membrane trafficking. As with the work of all great scientists, the approaches employed by Marilyn’s laboratory evolved with the times. She maintained her creative use of electron microscopy as a signature experimental tool throughout her career.

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Marilyn Farquhar received the FASEB/Excellence in Science award at the 91亚色传媒's centennial meeting in San Francisco.

Marilyn’s achievements were recognized with many awards and honors including election to both the National Academy of Sciences and the American Academy of Arts and Sciences. She received the American Society for Cell Biology's E.B. Wilson Medal and the American Society of Nephrology’s Homer W. Smith Award. She was particularly gratified by the Federation of American Societies for Experimental Biology that she received in 2006 at the 91亚色传媒 Annual Meeting in San Francisco.

Marilyn followed the accomplishments of her trainees and colleagues, including us, with eager interest and always with encouragement. We remember her ready smile and cheerful disposition. Her memory provides enduring inspiration to all of us who had the good fortune to follow the beauty and impact of her discoveries.

The authors thank Maya Simionescu (ICBP, Bucharest); Susan Ferro-Novick (University of California, San Diego); Stuart Kornfeld (Washington University, St. Louis); Colin Hopkins (Imperial College, London); Bill Brown (Cornell University); Pietro De Camilli (Yale University); David Sabatini (New York University) and Kathleen Dickson (McGill University) for their comments and suggestions.