OBITUARY

In Memoriam Robin Holliday (November 6, 1932–April 9, 2014)

HE death of Robin Holliday in Sydney, Australia, on April 9, of DNA. In the last year of his degree, Robin heard a lecture T2014 defines a solemn landmark in the story of genetic about this discovery and its genetic implications and he de- recombination that began 50 years ago with his publication cided immediately that genetics would be the focus of his in Genetical Research of the proposed structure of the DNA future career. recombination intermediate, which has come to be known Robin began postgraduate research in Cambridge where a as the “Holliday junction.” Unsatisfied with existing efforts galaxy of stars in the new science of molecular biology was to explain meiotic gene conversion through copy choice, assembled and where visitors from around the world would Robin looked for a fresh approach through the nascent field drop by. With his PhD research done, but not fully written up, of DNA repair. Using the complementarity of the two strands Robin took a position at the John Innes Horticultural In- of DNA to explain specific chromatid pairing at the molecu- stitution in Hertfordshire. Here he found himself thinking hard lar level, Holliday delineated his proposal for the successive about how two DNA molecules—the chromosome pairs—with stages of effective pairing and recombination of homologous almost the same genetic information could get together to chromatids. So familiar today are the principles embodied in produce a crossover. It was during a sabbatical visit in 1963 the Holliday model of recombination that they are easily to the University of Washington, Seattle, that he elaborated taken for granted. Yet the innovative nature of his work the idea of the Holliday junction that he published first in led to a breakthrough that has prompted reflective events 1964. An elegant account of the origins and subsequent de- at its 30th, 40th, and now 50th anniversaries. Sadly, velopment of this concept was provided by Franklin Stahl in Robin, who had been planning to attend this year’s event, a Perspectives review published in GENETICS to mark its 30th died before it took place. anniversary (Stahl 1994). Robin was born in Palestine in 1932 where his mother and In 1965, Robin was appointed to a position at the Na- architect father had been living since 1921 during the years of tional Institute for Medical Research (NIMR) in Mill Hill, the British Mandate. The family returned to England three where he would form and lead the Genetics Division. years later but stayed only a few years before sailing to Sri Robin continued his work on recombination and during Lanka (then Ceylon) in April 1939 for what was expected to the 1970s he, together with John Pugh, suggested that be a six-month assignment. This planning was overturned by chemical modifications to the DNA helix—methylation— the outbreak of World War II, which meant that the family might have important effects on how the genome plays did not return for eight years, spending a year of this time in out its instructions. Their ground-breaking article in Sci- South Africa, before returning to Sri Lanka. From Sri Lanka, ence in 1975 prepared the way for the burgeoning field of they sailed to Gibraltar in 1944 where they remained until epigenetics. their eventual return to England in 1947. Robin seems to When Robin joined the NIMR, the director was Sir Peter have thrived on the rich experiences of his early life, which Medawar, a Nobel Prize-winning immunologist also known left him with a profound love of travel. for his scientific interest in ageing. Robin was fascinated by In 1952, Robin won a scholarship to Emmanuel College, the question of how cells might age and rapidly developed Cambridge, where he would later write that it “proved hard a strong interest in this topic, contributing in important ways to find congenial company.” He was still an undergraduate to understanding how human cells grown in culture have at Cambridge when in 1953 Francis Crick and James Watson limited division capacity (replicative senescence, also known made their momentous discovery of the double helix structure as the “Hayflick limit”) and to testing ideas about the role of damage to proteins within the ageing process, building upon Copyright © 2014 by the Genetics Society of America doi: 10.1534/genetics.114.167684 a hypothesis proposed by his friend Leslie Orgel. In much of 1Address for correspondence: [email protected] this work, Robin evinced the same intuitive fascination with

Genetics, Vol. 198, 423–424 September 2014 423 quantitative theoretical study that underpinned his early he was elected fellow in 1976 and honored with a Royal work on recombination. Robin was a scientist who delighted Medal in 2011. in the interplay between theory and experiments. Robin is survived by his wife Lily and their daughter Mira, by In 1988, Robin left London to take a position with the his first wife Diana, their children David, Caroline, Rebecca, and Commonwealth Scientific and Industrial Research Orga- Emma, and by ten grandchildren and a great grandchild. nization in Sydney, where he worked until retirement, although he continued scientific writing until the time of Literature Cited his death. Stahl, F. W., 1994 The Holliday junction on its thirtieth anniver- Robin had a natural gift for writing. His book Understand- sary. Genetics 138: 241–246. ing Ageing (1995) provides a superb introduction for the Holliday, R., 1995 Understanding Ageing. Cambridge University general scientist. He was also a talented sculptor and two Press, Cambridge. of his bronze works are at the Royal Society in London where Tom Kirkwood1

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