JOSEPH E. VARNER

October 7, 1921–July 4, 1995

BY MAARTEN J. CHRISPEELS

JOSEPH E. VARNER'S fifty-year career (1945–95) spanned the emergence and development of plant biochemistry, and he was one of the major contributors to this field. His most notable research achievements were the definition of cell death as an active process; discovery that the hormone gibberellin regulates the expression of α-amylase in barley aleurone cells at the level of the gene; and cloning of the cDNA for the cell wall protein extensin, which laid the foundation for the study of the role of cell wall proteins in plants. Together with James Bonner, Varner edited Plant Biochemistry, which remained the standard single-volume textbook in the field for fifteen years. During the last ten years of his life he was probably the most widely admired and loved plant biologist in the country, the elder statesman of his discipline. He was extremely knowledgeable about biochemistry and whenever he talked to colleagues or students he generously shared his many ideas. He was a tireless promoter of the study of plants and talked about experiments until the final days of his life. In addition, Varner was a sought-after advisor to government, universities, and industry. He was a major supporter of the American Society of Plant Physiologists, which he served as president in 1970–71



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