OBITU ARY
In Memoriam Robin Holliday (November 6, 1932–April 9, 2014)
T
HE death of Robin Holliday in Sydney, Australia, on April 9, 2014 defines a solemn landmark in the story of genetic recombination that began 50 years ago with his publication in Genetical Research of the proposed structure of the DNA recombination intermediate, which has come to be known as the “Holliday junction.” Unsatisfied with existing efforts to explain meiotic gene conversion through copy choice, Robin looked for a fresh approach through the nascent field of DNA repair. Using the complementarity of the two strands of DNA to explain specific chromatid pairing at the molecular level, Holliday delineated his proposal for the successive stages of effective pairing and recombination of homologous chromatids. So familiar today are the principles embodied in the Holliday model of recombination that they are easily taken for granted. Yet the innovative nature of his work led to a breakthrough that has prompted reflective events at its 30th, 40th, and now 50th anniversaries. Sadly, Robin, who had been planning to attend this year’s event, died before it took place. Robin was born in Palestine in 1932 where his mother and architect father had been living since 1921 during the years of the British Mandate. The family returned to England three years later but stayed only a few years before sailing to Sri Lanka (then Ceylon) in April 1939 for what was expected to be a six-month assignment. This planning was overturned by the outbreak of World War II, which meant that the family did not return for eight years, spending a year of this time in South Africa, before returning to Sri Lanka. From Sri Lanka, they sailed to Gibraltar in 1944 where they remained until their eventual return to England in 1947. Robin seems to have thrived on the rich experiences of his early life, which left him with a profound love of travel. In 1952, Robin won a scholarship to Emmanuel College, Cambridge, where he would later write that it “proved hard to find congenial company.” He was still an undergraduate at Cambridge when in 1953 Francis Crick and James Watson made their momentous discovery of the double helix structure Copyright © 2014 by the Genetics Society of America doi: 10.1534/genetics.114.167684 1 Address for correspondence: [email protected]
of DNA. In the last year of his degree, Robin heard a lecture about this discovery and its genetic implications and he decided immediately that genetics would be the focus of his future career. Robin began postgraduate research in Cambridge where a galaxy of stars in the new science of molecular biology was assembled and where visitors from around the world would drop by. With his PhD research done, but not fully written up, Robin took a position at the John Innes Horticultural Institution in Hertfordshire. Here he found himself thinking hard about how two DNA molecules—the chromosome pairs—with almost the same genetic information could get together to produce a crossover. It was during a sabbatical visit in 1963 to the University of Washington, Seattle, that he elaborated the idea of the Holliday junction that he published first in 1964. An elegant account of the origins and subsequent development of this concept was provided by Franklin Stahl in a Perspectives review published in GENETICS to mark its 30th anniversary (Stahl 1994). In 1965, Robin was appointed to a position at the National Institute for Medical Research (NIMR) in Mill Hill, where he would form and lead the Genetics Division. Robin continued his work on recombination and during the 1970s he, together with John Pugh, suggested that chemical modifications to the DNA helix—methylation— might have important effects on how the genome plays out its instructions. Their ground-breaking article in Science in 1975 prepared the way for the burgeoning field of epigenetics. When Robin joined the NIMR, the director was Sir Peter Medawar, a Nobel Prize-winning immunologist also known for his scientific interest in ageing. Robin was fascinated by the question of how cells might age and rapidly developed a strong interest in this topic, contributing in important ways to understanding how human cells grown in culture have limited division capacity (replicative senescence, also known as the “Hayflick limit”) and to testing ideas about the role of damage to proteins within the ageing process, building upon a hypothesis proposed by his friend Leslie Orgel. In much of this work, Robin evinced the same intuitive fascination with
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quantitative theoretical study that underpinned his early work on recombination. Robin was a scientist who delighted in the interplay between theory and experiments. In 1988, Robin left London to take a position with the Commonwealth Scientific and Industrial Research Organization in Sydney, where he worked until retirement, although he continued scientific writing until the time of his death. Robin had a natural gift for writing. His book Understanding Ageing (1995) provides a superb introduction for the general scientist. He was also a talented sculptor and two of his bronze works are at the Royal Society in London where
424
Obituary
he was elected fellow in 1976 and honored with a Royal Medal in 2011. Robin is survived by his wife Lily and their daughter Mira, by his first wife Diana, their children David, Caroline, Rebecca, and Emma, and by ten grandchildren and a great grandchild.
Literature Cited Stahl, F. W., 1994 The Holliday junction on its thirtieth anniversary. Genetics 138: 241–246. Holliday, R., 1995 Understanding Ageing. Cambridge University Press, Cambridge.
Tom Kirkwood1
In Memoriam Robin Holliday (November 6, 1932–April 9, 2014)
T
HE death of Robin Holliday in Sydney, Australia, on April 9, 2014 defines a solemn landmark in the story of genetic recombination that began 50 years ago with his publication in Genetical Research of the proposed structure of the DNA recombination intermediate, which has come to be known as the “Holliday junction.” Unsatisfied with existing efforts to explain meiotic gene conversion through copy choice, Robin looked for a fresh approach through the nascent field of DNA repair. Using the complementarity of the two strands of DNA to explain specific chromatid pairing at the molecular level, Holliday delineated his proposal for the successive stages of effective pairing and recombination of homologous chromatids. So familiar today are the principles embodied in the Holliday model of recombination that they are easily taken for granted. Yet the innovative nature of his work led to a breakthrough that has prompted reflective events at its 30th, 40th, and now 50th anniversaries. Sadly, Robin, who had been planning to attend this year’s event, died before it took place. Robin was born in Palestine in 1932 where his mother and architect father had been living since 1921 during the years of the British Mandate. The family returned to England three years later but stayed only a few years before sailing to Sri Lanka (then Ceylon) in April 1939 for what was expected to be a six-month assignment. This planning was overturned by the outbreak of World War II, which meant that the family did not return for eight years, spending a year of this time in South Africa, before returning to Sri Lanka. From Sri Lanka, they sailed to Gibraltar in 1944 where they remained until their eventual return to England in 1947. Robin seems to have thrived on the rich experiences of his early life, which left him with a profound love of travel. In 1952, Robin won a scholarship to Emmanuel College, Cambridge, where he would later write that it “proved hard to find congenial company.” He was still an undergraduate at Cambridge when in 1953 Francis Crick and James Watson made their momentous discovery of the double helix structure Copyright © 2014 by the Genetics Society of America doi: 10.1534/genetics.114.167684 1 Address for correspondence: [email protected]
of DNA. In the last year of his degree, Robin heard a lecture about this discovery and its genetic implications and he decided immediately that genetics would be the focus of his future career. Robin began postgraduate research in Cambridge where a galaxy of stars in the new science of molecular biology was assembled and where visitors from around the world would drop by. With his PhD research done, but not fully written up, Robin took a position at the John Innes Horticultural Institution in Hertfordshire. Here he found himself thinking hard about how two DNA molecules—the chromosome pairs—with almost the same genetic information could get together to produce a crossover. It was during a sabbatical visit in 1963 to the University of Washington, Seattle, that he elaborated the idea of the Holliday junction that he published first in 1964. An elegant account of the origins and subsequent development of this concept was provided by Franklin Stahl in a Perspectives review published in GENETICS to mark its 30th anniversary (Stahl 1994). In 1965, Robin was appointed to a position at the National Institute for Medical Research (NIMR) in Mill Hill, where he would form and lead the Genetics Division. Robin continued his work on recombination and during the 1970s he, together with John Pugh, suggested that chemical modifications to the DNA helix—methylation— might have important effects on how the genome plays out its instructions. Their ground-breaking article in Science in 1975 prepared the way for the burgeoning field of epigenetics. When Robin joined the NIMR, the director was Sir Peter Medawar, a Nobel Prize-winning immunologist also known for his scientific interest in ageing. Robin was fascinated by the question of how cells might age and rapidly developed a strong interest in this topic, contributing in important ways to understanding how human cells grown in culture have limited division capacity (replicative senescence, also known as the “Hayflick limit”) and to testing ideas about the role of damage to proteins within the ageing process, building upon a hypothesis proposed by his friend Leslie Orgel. In much of this work, Robin evinced the same intuitive fascination with
Genetics, Vol. 198, 423–424 September 2014
423
quantitative theoretical study that underpinned his early work on recombination. Robin was a scientist who delighted in the interplay between theory and experiments. In 1988, Robin left London to take a position with the Commonwealth Scientific and Industrial Research Organization in Sydney, where he worked until retirement, although he continued scientific writing until the time of his death. Robin had a natural gift for writing. His book Understanding Ageing (1995) provides a superb introduction for the general scientist. He was also a talented sculptor and two of his bronze works are at the Royal Society in London where
424
Obituary
he was elected fellow in 1976 and honored with a Royal Medal in 2011. Robin is survived by his wife Lily and their daughter Mira, by his first wife Diana, their children David, Caroline, Rebecca, and Emma, and by ten grandchildren and a great grandchild.
Literature Cited Stahl, F. W., 1994 The Holliday junction on its thirtieth anniversary. Genetics 138: 241–246. Holliday, R., 1995 Understanding Ageing. Cambridge University Press, Cambridge.
Tom Kirkwood1
