Keith Roberts Porter: 1912–1997

K eith Roberts Porter died on May 2, 1997, just over a month short of his 85th birthday. He had the perspicacity, good fortune, and patience to take advantage of the fast moving frontier of analytical biology after the Second World War to provide many of the techniques and experimental approaches that established the new field of biomedical research now known as cell biology. He was renowned for taking the first electron micro-graph of an intact cell, but his contributions went far beyond that seminal instance. They ranged from technical developments, such as the roller flask for cell culture and the Porter-Blum ultramicrotome, to experimental and observational achievements, such as studies on the synthesis and assembly of collagen, on the role of coated vesicles in endocytosis, on lipid digestion in the intestine, and on the universality of the 9 ϩ 2 axoneme in cilia. The initial ultra-structure descriptions of the endoplasmic reticulum and the sarcoplasmic reticulum, identification of the role of T-tubules in excitation–contraction coupling in muscle and the role of the cytoskeleton in cell transformation and shape change, were his, as were many other contributions, described in some detail elsewhere (Peachey and Brinkley, 1983; Moberg, 1996). Absent from this list are his early pioneering work establishing the androgenetic haploid in frogs, an exercise in nuclear transplantation with consequences for the recent cloning of mammals, and his later adventures with pigment migration in fish chromatophores. In addition to his specific scientific contributions, Keith Porter also made more important philosophical contributions to the field that he helped to shape. These principles include the understanding that the cell is not a " bag of enzymes " ; that organelle structure is consistent from cell to cell throughout a wide range of protists, animals, and plants; that this means that cell structure and function have a macromolecular basis; that self-assembly is a critical mor-phogenetic principle; and, most presciently, that the cell is structurally integrated down to molecular resolutions by an intricate network of cytoplasmic proteins, this integration having consequences for signal transduction and function. These contributions sometimes involved experienced collaborators and, in later years, young disciples with whom Porter developed particular empathy. Keith Porter was a boy from Yarmouth, Nova Scotia, who grew up far from the centers of science in the United States and Europe. He went to college at Acadia University in Nova Scotia and then did graduate work at Harvard …