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A cell biologist who uses the combined tools of biochemistry and fine structural analysis to illuminate functional and biogenetic relationships between subcellular organells. Since the early sixties, his work on the role of the endoplasmic reticulum (ER) in the synthesis of secretory, membrane and organelle proteins has provided many new insights into the processes that generate the organizational complexity of the eukaryotic cell. These include the formulation, with Gunter Blobel in 1971, of the signal hypothesis to explain the synthesis of secretory and membrane proteins on ribosomes of the rough endoplasmic, and the development of a system of virally infected cultured cells to study the generation of epithelial cell polarity and the intracellular sorting of proteins targeted to the cell surface.