TIMELINE

1833 Robert Brown describes the nucleus in cells of orchids
1866 Gregor Mendel publishes ‘Studies of plant hybridisation’
1868 Friedrich Miescher discovers ‘nuclein’ (DNA) in pus cells
1878 Albrecht Kossel isolates ‘yeast nuclein’ (later shown to be RNA)
1880 Walther Flemming describes nuclear ‘threads’ made of ‘chromatin’ during cell division (‘mitosis’) in the salamander
1882 Flemming suggests that chromatin and nuclein are identical
1885 Kossel extracts two bases, guanine and adenine, from thymus nuclein, followed by thymine (1893), cytosine (1894) and uracil (1900)
1888 Wilhelm Waldeyer renames Flemming’s threads ‘chromosomes’
1889 Richard Altmann renames nuclein ‘nucleic acid’
1900 Mendel’s work is ‘rediscovered’ by Carl Correns, Hugo de Vries and Erich von Tschermak
1903 Walter Sutton formulates the ‘chromosome theory of inheritance’
1904 William Bateson begins a pro-Mendel crusade and coins the word ‘genetics’
1909 Wilhelm Johannsen invents the words ‘gene’, ‘genotype’ and ‘phenotype’ Phoebus Levene identifies the sugar in yeast nucleic acid (RNA) as ribose
1912 Levene proposes that nucleic acids are a small ‘tetranucleotide’, containing one of each of the four bases Max von Laue takes the first X-ray photograph of a crystal
1914 Lawrence Bragg formulates Bragg’s Law of X-ray crystallography; with his father William, develops ‘a new crystallography’
1915 Thomas Hunt Morgan publishes The Mechanism of Mendelian Inheritance, based on mutations in the fruit fly
1927 Fred Griffith shows that dead pneumococci bacteria can ‘transform’ (change the genetic characteristics of) live pneumococci, when injected into living mice
1928 Levene and Kossel both claim that genes are made of protein, not nucleic acid
1929 Levene identifies the sugar in thymus nucleic acid (DNA) as deoxyribose

Martin Dawson, in Oswald Avery’s lab at the Rockefeller, confirms Griffith’s finding of transformation of pneumococci, also in living mice

1931 Dawson and Richard Sia achieve transformation in vitro
1932 Lionel Alloway in Avery’s lab extracts the ‘transforming principle’ responsible for transformation but cannot identify it chemically
1937 Torbjörn Caspersson deduces that DNA molecules are very long, thin cylinders, and much bigger than a ‘tetranucleotide’
1938 Florence Bell takes X-ray photographs of DNA; she and Bill Astbury suggest that the bases in the DNA molecule are stacked ‘like a pile of pennies’
1940 Colin MacLeod in Avery’s lab detects DNA in extracts of ‘transforming principle’ but does not follow up the observation
1941 Alfred Mirsky extracts ‘chromosin’ (DNA with associated protein) from cell nuclei
1942 Maclyn McCarty and Avery show that the ‘transforming principle’ consists of DNA, with tiny amounts of contaminating protein
1944 Erwin Schrödinger suggests in his book What is Life? that genes are ‘aperiodic crystals’

Avery, MacLeod and McCarty publish their landmark paper showing that DNA is the ‘transforming principle’ and the genetic material in pneumococci

Mirsky insists that protein, not DNA, mediates transformation and is the genetic material

1947 Rollin Hotchkiss shows that DNA contains unequal amounts of the four bases, thus ruling out the hypothetical ‘tetranucleotide’

André Boivin proves that DNA also transforms other bacteria (E. coli)

Masson Gulland proposes that the DNA molecule is held together by hydrogen bonding between bases

Gulland’s PhD student Michael Creeth proposes that DNA consists of two straight strands of DNA, linked by hydrogen bonding between bases on opposing strands

1948 Erwin Chargaff reports that amounts of adenine and thymine are equal, as are those of cytosine and guanine, in different sources of DNA

Linus Pauling discovers the alpha-helix, crucial in shaping protein molecules

1949 Sven Furberg works out that the bases lie perpendicular to the backbone of DNA, and proposes a single-stranded, helical structure for DNA
1950 Ray Gosling at King’s takes an X-ray photograph showing a regular ‘crystalline’ appearance of DNA (the A form)
1951 January: Rosalind Franklin joins the Biophysics Unit at King’s, to work on the X-ray structure of DNA

May: Wilkins presents the crystalline DNA structure at a meeting in Naples and inspires Jim Watson to solve its structure

Elwyn Beighton in Leeds takes an X-ray photograph that shows the helical features of DNA (B form). The photograph is ignored

July: Wilkins presents DNA structures at a meeting in Cambridge and is told by Franklin to stop working on DNA

Alec Stokes at King’s predicts the X-ray pattern of a helical molecule

October: Jim Watson joins Francis Crick at the Cavendish Laboratory in Cambridge and persuades him to pursue the structure of DNA

November: Wilkins meets Watson and Crick and tells them that the most likely structure contains three helical strands of DNA

Watson attends a colloquium at King’s where Wilkins and Franklin present their work on DNA

Bruce Fraser at King’s builds a model of DNA containing three helical strands, which Wilkins rejects

December: Using data from King’s, Crick and Watson build a three-stranded model of DNA, which is fatally flawed; Wilkins breaks off their collaboration

1952 January: Franklin and Gosling characterise the A and B forms of DNA

April: John Griffith in Cambridge calculates that hydrogen bonding will attract adenine to thymine, and cytosine to guanine

May: Gosling takes Photograph 51, showing the helical features of DNA (B form)

July: Franklin decides that ‘crystalline’ DNA (A form) cannot be a helix, causing Wilkins to have doubts about the helical nature of DNA in general

December: Pauling proposes a DNA model with three helical strands, also fatally flawed

1953 February: Watson visits King’s; Wilkins shows him Photograph 51, in which Watson sees the diagnostic features of a helical structure

March: Franklin leaves King’s to study viral structure at Birkbeck College, London

Watson realises that the pairing of bases on opposing strands is the key to the structure of DNA. Using Franklin’s data and without her knowledge, he and Crick construct the double helix

April: Nature publishes three papers on the double helix, by Watson and Crick; Wilkins et al; and Franklin and Gosling

July: Watson and Crick publish a follow-up paper in Nature on the self-replication of DNA

1958 16 April: Rosalind Franklin dies of ovarian cancer, aged 38
1962 Watson, Crick and Wilkins share the Nobel Prize for Physiology or Medicine
1968 Watson publishes The Double Helix
2001 Independent scientific tribunal clears Gregor Mendel of having falsified his data