David Lack | |
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![]() Lack in 1966, photo byEric Hosking | |
Born | David Lambert Lack (1910-07-16)16 July 1910 London, England |
Died | 12 March 1973(1973-03-12) (aged 62) |
Alma mater | University of Cambridge |
Known for |
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Spouse | Elizabeth Lack |
Children | 4, includingAndrew Lack |
Awards |
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Scientific career | |
Fields | Ornithology |
Institutions | |
Doctoral students | |
Other notable students | Robert H. MacArthur |
David Lambert LackFRS[1] (16 July 1910 – 12 March 1973) was a Britishevolutionary biologist who made contributions toornithology, ecology, andethology.[4] His 1947 book,Darwin's Finches, on thefinches of the Galapagos Islands was a landmark work as were his other popular science books onLife of the Robin andSwifts in a Tower.[5] He developed what is now known asLack's Principle which explained the evolution of avian clutch sizes in terms of individual selection as opposed to the competing contemporary idea that they had evolved for the benefit of species (also known asgroup selection). His pioneering life-history studies of the living bird helped in changing the nature of ornithology from what was then a collection-oriented field. He was a longtime director of theEdward Grey Institute of Field Ornithology at theUniversity of Oxford.
Lack was born in London, the oldest of four children of Harry Lambert LackMDFRCS, who later became President of theBritish Medical Association.[6] The name 'Lack' is derived from 'Lock'. His father grew up in a farming family from Norfolk and became a leading ear, nose and throat surgeon at theLondon Hospital. Although his father had some interest in birds as a boy it does not appear that he influenced David's interest. His mother Kathleen was the daughter of Lt. Col. McNeil Rind of theIndian army. Kathleen's father was Scottish and on her mother's side was part Irish, Greek and Georgian.[1] Kathleen had been an actor and was a supporter of women's suffrage. At home they organized meetings of the poetry society. David was schooled at home until seven and then went to the Open Air School in Regent's Park before going to The Hall, Hampstead followed by Foster's School, Stubbington andGresham's School,Holt, Norfolk. Lack was taught biology at Gresham's by W.H. Foy and G.H. Lockett.[7] He went toMagdalene College, Cambridge and received a BA second class in 1933 after studying botany, zoology and geology for part I of the Tripos and zoology for part II.[8][9]
Until the age of fifteen, Lack lived in a large house in Devonshire Place, London. The family spent their summers in New Romney Kent where Lack became familiar with the local birds especially on Romney Marsh.[10] By the age of nine, he had learnt the names of most birds and had written out an alphabetically arranged life-list.[1] In 1926, Lack won the Holland-Martin Natural History Prize for an essay on "Three Birds of Kelling Heath".[11] In 1928, with an essay on 'My favourite birds' he was the national winner of the senior prize (a silver medal) in the Public School Essay Competition, organised by theRoyal Society for the Protection of Birds.[12] David did not wish to follow his father's profession in medicine and took an interest in zoology. His father then considered entomology which was then the only professional field in zoology and found work for David at the Frankfurt museum in the summer of 1929. He spent four months pinning insects in theSenckenberg Museum and found it “extremely” boring.[13] He joined the Cambridge Ornithological Club whose members includedPeter Scott,Arthur Duncan,Dominic Serventy, andTom Harrisson. His first scientific paper was on the display of nightjars, published in the Ibis in 1932. He joined on several expeditions with Cambridge researchers including two to the Arctic.[14] Lack wroteThe Birds of Cambridgeshire (1934) which was published by the Cambridge Bird Club. In this work, he departed from the contemporary style with a distinct de-emphasis on rare and accidental birds.[15]
Lack received an Sc.D. from Cambridge University in 1948.[16]
After Cambridge, Lack, on the recommendation ofJulian Huxley took up a position as a science mentor atDartington Hall School,[17]Devonshire from 1934 until Summer 1938 when he took a year off to study bird behaviour on theGalapagos Islands. In 1935 he made his first trip to the United States as a chaperone for a Dartington Hall student returning to California. Here he metJoseph Grinnell andRobert McCabe and gave a talk at theCooper Ornithological Club. In New York, he metErnst Mayr at the American Museum of Natural History. He returned via theSS Bremen, only one of about four English speakers on the German ship.[18] He was only in the Galapagos for part of that year, starting August 1938. With the data that he collected in the Galapagos, especially on the finches he went to the United States. April to August 1939 was spent at theCalifornia Academy of Sciences which held a large collection of the finches of Galapagos that had been studied earlier byHarry Swarth and atErnst Mayr's home inNew Jersey. While in the US he made a study of the tricoloured blackbird with John T. Emlen.[19] He returned home in September 1939, after the outbreak of war.[20] Lack publishedThe Galapagos Finches (Geospizinae), A Study in Variation in which he examined variations within species across islands and considered that many of them were non-adaptive and due to founder effect and genetic drift.[21] Lack's first major work wasThe Life of the Robin, which was based on four years of field work that he conducted while teaching atDartington Hall School. He examined robin behaviour, song, territory, pairing and breeding using ringing to mark and track individual birds.[22] The manuscript was completed in 1942 and it went through five editions from 1943 to 1970. One of Lack's students at Dartington Hall wasEva Ibbotson. A colleague who helped in filming some of the robins for Lack was the geography teacher Bill Hunter. In 1934 Lack went to Tanganyika on an invitation fromR.E. Moreau.[23]
Lack was committed to pacificism and debated the philosophy even during his Dartington days with the founder of the college,Leonard Knight Elmhirst. DuringWorld War II Lack however served with aBritish Army unit called the Army Operational Research Group on the Orkney Islands working onradar use. During this work he met other biologists who had been inducted into the war includingGeorge Varley, an entomologist who introduced him to the idea of density-dependent regulation of animal populations.[24] Lack's observations on spurious echoes produced by birds would later allow him to establish the field ofradar ornithology to studybird migration.[25][26] Lack was released from wartime duty in August 1945 so as to take a position to as Director (succeedingW.B. Alexander) of theEdward Grey Institute of Field Ornithology atOxford University, a position that he held until his death in 1973.
Lack's work in ornithology was almost entirely based on studies of theliving bird. He was one of the pioneers of life-history studies in Britain, especially those based on quantitative approaches, when some traditional ornithologists of the time were focussing their studies on morphology and geographic distribution.[27] Lack's major scientific research included work onpopulation biology and density dependent regulation. His work suggested that natural selection favouredclutch sizes that ensured the greatest number of surviving young. This interpretation was however debated byV.C. Wynne-Edwards who suggested that clutch size was density-independent. This was one of the earliest debates ongroup selection. Lack's studies were based onnidicolous birds and some recent studies have suggested that his findings may not hold for other groups such as seabirds.[28]
As a mentor for numerous doctoral students, Lack followed a hands-off method, letting students decide their own research topics. He encouraged students to sort out their ideas and find the "simplest explanation, which was probably best." He would make students work on their papers and give only one reading to their thesis asking them to choose either a draft or a final version to submit.[29]
He wrote numerous papers in ornithological journals, and had a knack of choosing memorable titles: he once claimed to have single-handedly caused the renaming of a group of birds through the submission of a scientific paper with the title "Territory and Polygamy in a Bishop". This 1935 publication was subsequently titled "Territory and polygamy in a bishop bird,Euplectes hordeacea hordeacea (Linn.)" in the journalIbis as the journal editor felt that the title might cause misunderstanding.[30]
Lack's most famous work isDarwin's Finches, a landmark study whose title linkedDarwin's name with theGalapagos group of species and popularised the term "Darwin's finches" in 1947, though the term had been introduced byPercy Lowe in 1936.[31] There are two versions of this work, differing significantly in their conclusions. The first is a book-length monograph, written after his visit to the Galapagos, but not published until 1945.[32] In it Lack interprets the differences in bill size as species recognition signals, that is, asisolating mechanisms.
The second is the later book in which the differences in bill size are interpreted as adaptations to specific food niches, an interpretation that has since been abundantly confirmed.[3] This change of mind, according to Lack's Preface, came about as a result of his reflections on his own data whilst he was doing war work. The effect of this change in interpretation is to put the emphasis for speciation onto natural selection for appropriate food handling instead of seeing it primarily as a by-product of an isolating mechanism. In this way his work contributed to themodern evolutionary synthesis, in whichnatural selection came to be seen as the prime mover in evolution, and not random or mutational events. Lack's work laid the foundations for the much more extensive work ofPeter and Rosemary Grant and their colleagues.[33] Lack's work feeds into studies of island biogeography which continue the same range of issues presented by the Galapagos fauna on a more varied canvas.[34] According toErnst Mayr,
In 1943 Lack took an interest in clutch size after reading Moreau's manuscript sent to theIbis. Lack was then an assistant to the editor of theIbis.[36] Lack postulated what is now known as Lack's Principle, which states that "the clutch size of each species of bird has been adapted by natural selection to correspond with the largest number of young for which the parents can, on average, provide enough food".[37][38][39]
Lack took a keen interest in the mechanisms involved in regulating populations in nature.The Natural Regulation of Animal Numbers is one of Lack's most frequently cited works. Here he gave primacy to natural selection in determining the rate of reproduction and he especially countered the idea that it was adjusted with mortality rates so that constant populations are maintained. It was critiqued byJ.B.S. Haldane who found it lacking mathematical precision and biased to bird studies. The other major critic wasV.C. Wynne-Edwards with whom he clashed for nearly a decade.[40] Lack followed up on the criticisms in his later books including Population Studies of Birds (1966).
The centenary of Lack's birth, 16 July 2010, was marked by a 'David Lack Centenary Symposium', hosted by theEdward Grey Institute. A programme of talks focused on and celebrated the scientific contributions of Lack to ornithology, and the broader fields of ecology and evolution, and assessed the development of these fields in the 21st century.[42]
David Lack marriedElizabeth Lack (née Silva) who was also an ornithologist. Elizabeth Silva was born in Hertfordshire in 1916 and took an early interest in music. She wished to join the Royal Academy of Music in London but the war led to her serving in the Auxiliary Territorial Service as an ambulance driver in Europe. After the war she applied for work and due to her interest in birds she sent her resume toRichard Fitter who passed it on to David Lack with a note "Here's another for your reject file." Lack however interviewed her and appointed her as a secretary. Noting her interest in birds, he also invited her to serve as a field assistant for studies in theWytham Woods. She also helped in the study of swifts. One day Elizabeth did not return to her office after her observations of the swifts and David, worried that she might have fallen off a ladder, found her engrossed in observation. They became engaged in 1948 and were married on July 9, 1949. The best man was George Varley.[43] They had four children: Peter Lack (born 1952, a biologist),Andrew Lack (born 1953, also a biologist and academic), Paul Lack (born 1957, a freelance teacher), and Catherine Lack (born 1959, a university chaplain).[4] In Oxford, the Lacks initially lived in a flat inPark Town, Oxford, and later onBoars Hill, just south of Oxford. Lack enjoyed music and was also a fan of field hockey and tennis in which he also participated.[44] Lack died fromNon-Hodgkin lymphoma despite radiation treatments.[45]
Lack's parents belonged to the Church of England, and he was an agnostic as an early adult but became a convert toAnglicanism in 1948, possibly influenced by Dan and Mary Neylan, friends at Dartington Hall.[46] He sought to find a compromise between science and religion and wrote, in 1957,Evolutionary theory and Christian belief, on the relationship between Christian faith and evolutionary theory. Lack believed that evolution could not account for morality, truth, beauty, free will, self-awareness and individual responsibility.[47] This book foreshadows, in some ways, thenon-overlapping magisteria conception of the relationship betweenreligion and science later popularised byStephen Jay Gould.
Arthur Cain remarked of him, "David Lack was the only religious man I knew at that period who did not allow his religion to dictate his view of natural selection."[48]