by Matthew Cobb
17.Pardee et al. (1959).
18.Szilárd (1960), Schaffner (1974).
19.Monod (1972b), p. 199.
20.Monod (1972b), p. 199.
21.Summarised in Szilárd (1960). For Maas’s only published speculations on the matter, made in a conference discussion in September 1957 and published in 1958, see Schaffner (1974), p. 361. See also Maas (2004).
22.Schaffner (1974), p. 360; Maas (2004).
23.Vogel (1957).
24.Pardee et al. (1958); Schaffner (1974), p. 374.
25.Jacob (1988), p. 298. The fiftieth anniversary of the operon led to a flurry of papers putting the discovery into historical perspective, for example Beckwith (2011), Gann (2010), Lewis (2011) and Yaniv (2011). For Jacob’s work on phage, see Peyrieras and Morange (2002).
26.Grmek and Fantini (1982), p. 209.
27.Jacob (1988), p. 308.
28.Kay (2000), p. 217.
29.Jacob (1988), p. 304. As Lily Kay has noted, in his 1965 Nobel Lecture, Jacob used the more neutral analogy of doors in a house, each controlled ‘by a little radio receiver’ (Jacob, 1972, p. 154).
30.Monod (1959).
31.Pardee et al. (1959).
32.Pontecorvo (1952).
33.Lederberg (1957), p. 753.
34.Benzer (1957), p. 70. For the impact of Benzer’s work on a young researcher, see Holliday (2006).
35.Benzer (1966).
36.Benzer (1957), p. 90.
37.Benzer (1959, 1961).
38.Beadle (1957), p. 129. For a full discussion of Benzer’s work on the rII region and its implications, see Holmes (2006).
39.Meselson and Stahl (1958). See also: Holmes (2001), Davis (2004) and Hanawalt (2004).
40.Delbrück and Stent (1957).
41.Judson (1996), p. 416.
42.Crick (1988), p. 119. See also Brenner (2001), pp. 73–87.
43.Jacob (1988), p. 312.
44.Judson (1996), p. 419.
45.Jacob (1988), pp. 313–14.
46.Yčas and Vincent (1960).
47.Brenner et al. (1961), Gros et al. (1961). For Gros’s view of the race to identify the messenger, see Gros (1979). Watson’s telegram to Brenner asking him to delay publication until his group’s paper was ready can be seen at http://libgallery.cshl.edu/items/show/66514.
48.Jacob and Monod (1961a, b), Monod and Jacob (1961).
49.Jacob and Monod (1961a), p. 318.
50.Jacob and Monod (1961a), p. 334.
51.Jacob and Monod (1961a), p. 344. They had first used the term operon in the previous year, in a French publication (Jacob et al., 1960). The precise functioning of the Lac operon would not be fully understood for some years. See Müller-Hill (1996) for a detailed insider’s account. For an overview of the operon and its legacy, see Morange (2005a).
52.Jacob and Monod (1961a), p. 354.
53.Jacob and Monod (1961a), p. 354. Jacob could not recall any conscious reference to Schrödinger (Morange, 1998, p. 295, note 36).
54.Monod and Jacob (1961), p. 401.
55.Jacob (2011).
56.Ptashne (2013), p. 1181.
57.Brenner (1961); Monod and Jacob (1961), p. 393.
Chapter 10
1.Crick (1959).
2.Belozersky and Spirin (1958), Sueoka et al. (1959).
3.Golomb (1962a), p. 100.
4.Rheinberger (1997), p. 213.
5.There is a large amount of material covering Nirenberg’s career in the Modern Manuscripts Collection, History of Medicine Division, National Library of Medicine at Bethesda. This includes over 40 volumes of diaries and notebooks – a future researcher’s goldmine. The diary covering this period is D9 IXA, 1960 Sep–[1961] May, to be found in Box 22, Folder 44, Marshall W. Nirenberg Papers, 1937–2003. The Nirenberg diary entries quoted here are all taken from Kay (2000). A tiny proportion of the papers can be found online at http://profiles.nlm.nih.gov/JJ/.
6.Kay (2000), p. 240.
7.Hoagland et al. (1957, 1958). For a thorough exploration of Zamecnik’s work and its conceptual implications, see Rheinberger (1997).
8.Grunberg-Manago et al. (1955). Ochoa shared the 1959 Nobel Prize with his ex-student Arthur Kornberg, who in 1956 had isolated the enzyme that enables DNA molecules to copy themselves. There was no prize for the co-discoverer of polynucleotide phosphorylase, the French biochemist Marianne Grunberg-Manago.
9.Singer (2003).
10.Kay (2000), p. 241.
11.Nirenberg (1960).
12.Lamborg and Zamecnik (1960); Rheinberger (1997), pp. 208–21.
13.Kay (2000), p. 246.
14.Tissières et al. (1960).
15.Kay (2000), p. 246.
16.Nirenberg (1963), p. 84.
17.Kay (2000), p. 247.
18.Rheinberger (1997), p. 213.
19.Crick et al. (1957), p. 420.
20.Crick (1958), p. 160.
21.Kay (2000), p. 248.
22.Matthaei and Nirenberg (1960).
23.Matthaei and Nirenberg (1961a).
24.Matthaei and Nirenberg (1961a), pp. 405–6.
25.Matthaei and Nirenberg (1961a), p. 407.
26.Kay (2000), p. 249.
27.Judson (1996), pp. 458–9.
28.Judson (1996), p. 460.
29.Judson (1996), p. 462.
30.Dr Jerry Hurwitz, e-mail to the author, 9 April 2014.
31.Lengyel (2012).
32.Hargittai (2002), p. 140.
33.Matthaei and Nirenberg (1961b), p. 1587. One of the controls later caused much confusion: to show that acidity was not involved in the DNase effect, they added several compounds, including polyadenylic acid, none of which affected protein synthesis. Polyadenylic acid is better known as poly(A); this ‘negative control’ later led some competitors to unfairly cast doubt on whether they had intended to get an effect with poly(U). See Kay (2000), pp. 249–50; Rheinberger (1997), p. 210.
34.Nirenberg and Matthaei (1961), p. 1601.
35.When interviewed by Judson in the 1970s, Nirenberg seemed unaware of these key papers (Judson, 1996, p. 462).
36.Anonymous (1961).
37.Anonymous (1961), Morgan (1961), Slater (2003). See also the informal photos of the Congress in the collections of Watson and Brenner, held at Cold Spring Harbor: http://libgallery.cshl.edu/items/show/51693 and http://libgallery.cshl.edu/items/show/52212.
38.The manuscript version of this talk, with Nirenberg’s handwritten edits, can be found at http://profiles.nlm.nih.gov/ps/access/JJBBKB.pdf. For the published version see Nirenberg and Matthaei (1963).
39.Nirenberg (2004) recalled the size of the audience as ‘~35’ (p. 49).
40.Hargittai (2002), p. 137.
41.Judson (1996), pp. 463–4.
42.Watson (2001), p. 265.
43.Watson and Berry (2003), p. 76; Crick et al. (1961), p. 1232; Nirenberg (2004), p. 49.
44.Judson (1996), p. 464.
45.Interview with Nirenberg by Ruth Harris, 1995–1996. http://history.nih.gov/archives/downloads/Nirenberg%200ral%20history%20Chap%203a-%20%20Recognition%20Moscow,%20MIT.pdf
46.Judson (1996), p. 464.
47.Dr Jerry Hurwitz, e-mail to the author, 9 April 2014.
48.Varmus (2009), p. 24.
49.Letter from Lengyel to Ochoa, 19 August 1961, in Supplemental Material, Lengyel (2012).
50.Hurwitz recalled, ‘I was impressed with the data presented by Nirenberg in Moscow but puzzled by the properties of the product (which were due to my own lack of information)’ (e-mail to the author, 9 April 2014).
51.Judson (1996), p. 464.
52.Lipmann to Crick, 27 November 1961. http://profiles.nlm.nih.gov/ps/retrieve/ResourceMetadata/SCBBBV.
53.Kay (2000), p. 255.
54.Judson (1996), p. 465.
55.Judson (1996), pp. 464–5.
56.Stent (1971) described the experiment as follows: ‘One day, Nirenberg added artificially synthesised polyuridylic acid to this reaction mixture instead of natural mRNA and obtained a most surprising result’ (p. 528). See al
so Brenner (2001), p. 99.
57.Woese (1967), p. 53, note 1; Nirenberg (2004), p. 50. According to Woese, Beljanski’s results were ‘uninterpretable’. Nirenberg claims that Tissière had also tried and failed to get poly(A) to work, but Tissière said that although poly(A) was sitting in a freezer in the lab next door, he never thought to use it. He described his lack of initiative as ‘idiotic’ (Judson, 1996, p. 465).
58.Zamecnik (2005).
59.Nirenberg (2004), p. 50.
60.Nirenberg (2004), p. 49.
61.Judson (1996), p. 465.
62.Ochoa (1980), p. 20. Lengyel (2012), p. 32, uses very similar words.
63.Judson (1996), p. 469.
64.Nirenberg (2004), p. 49.
65.Martin (1984), p. 293.
66.Nirenberg (2004), p. 50.
67.Transcript of BBC talk ‘Cracking the genetic code’ by Crick, 22 January 1962. http://profiles.nlm.nih.gov/ps/access/SCBBFX.pdf.
68.Crick et al. (1961), p. 1229. For an appreciation of this paper, see Yanofsky (2007).
69.Judson (1996), p. 467.
70.Crick et al. (1961), p. 1227.
71.Crick et al. (1961), p. 1231.
72.Crick et al. (1961), p. 1232.
73.Anonymous (1962), p. 19.
74.Transcript of BBC talk ‘Cracking the genetic code’ by Crick, 22 January 1962. http://profiles.nlm.nih.gov/ps/access/SCBBFX.pdf.
75.Crick (1962), p. 16.
Chapter 11
1.In chronological order: Lengyel et al. (1961, 1962), Basilio et al. (1962), Gardner et al. (1962), Speyer et al. (1962a, b), Wahba et al. (1962, 1963a, 1963b).
2.Letter from Tomkins to Nirenberg, 25 October 1961. http://profiles.nlm.nih.gov/ps/access/JJBCBB.pdf.
3.Lengyel et al. (1961), p. 1941.
4.Martin et al. (1961).
5.Lengyel (2012), p. 35.
6.The Sunday Times, 31 December 1961. According to the paper, Crick was ‘leader of the Cambridge team which discovered code’, while Ochoa’s colleague Speyer was ‘a British biologist’.
7.‘I have stressed that it is your discovery which was the real breakthrough.’ Crick to Nirenberg, 4 January 1962. http://profiles.nlm.nih.gov/ps/access/JJBBFL.pdf.
8.Nirenberg to Crick, 15 January 1962. http://profiles.nlm.nih.gov/ps/access/JJBBFJ.pdf.
9.Speyer et al. (1962a, b).
10.When each batch of these synthetic RNAs was created, the nucleotides were assembled in a different, random way, producing slightly different molecular sequences, and making it difficult to compare studies that claimed to be looking at the same nucleotide ratios (Matthaei et al., 1962, p. 671).
11.Martin et al. (1961), Speyer et al. (1962a, b).
12.Crick to Ochoa, 21 September 1962. http://profiles.nlm.nih.gov/ps/access/SCBBSY.pdf.
13.Speyer et al. (1962b), p. 443.
14.Speyer et al. (1962b), p. 445.
15.Matthaei et al. (1962), p. 674.
16.Matthaei et al. (1962). See also Crick to Nirenberg, 29 January 1962. http://profiles.nlm.nih.gov/ps/access/JJBBGN.pdf.
17.Crick (1966a), p. 6.
18.Eck (1961). See also Jukes (1962, 1963), Lanni (1962), Wall (1962) and Woese (1962).
19.Tsugita (1962).
20.Crick (1963a), p. 170.
21.Ageno (1962).
22.Woese (1962).
23.Roberts (1962).
24.Eck (1963).
25.Zubay and Quastler (1962).
26.Golomb (1962b).
27.Bretscher and Grunberg-Manago (1962), Gardner et al. (1962).
28.Gardner et al. (1962).
29.Chantrenne (1963), p. 30.
30.Couffignal (1965), p. 182.
31.Couffignal (1965), p. 78.
32.Chantrenne (1963), p. 27. The term was also used at the meeting by André Lwoff (Couffignal, 1965, p. 176).
33.Ochoa (1964), pp. 4, 3.
34.Tatum (1963), p. 175.
35.Vogel et al. (1963), p. 517.
36.Nirenberg and Jones (1963), p. 461.
37.Vogel et al. (1963), p. 503.
38.Vogel et al. (1963), pp. 517–18.
39.Crick (1963a), pp. 177, 180.
40.Crick (1963a), p. 182.
41.Crick (1963a), p. 212. The original draft of the article contains some even sharper formulations (http://libgallery.cshl.edu/items/show/52223). Crick wrote to Ochoa apologising for criticising his work ‘in certain ways’ (Crick to Ochoa, 21 September 1962; http://profiles.nlm.nih.gov/ps/access/SCBBSY.pdf). A few months later Crick published a less combative version of these arguments in Science, but repeated his crushing statement that the experimental evidence for establishing a codon ‘falls short of proof in almost all cases’ (Crick, 1963b, p. 463).
42.Crick (1963a), p. 198.
43.Crick (1963a), p. 202.
44.Crick (1966a), p. 5.
45.Crick (1963a), pp. 213–14.
46.A number of Watson’s colleagues, including the joker Seymour Benzer, sent Watson a congratulatory telegram that concluded ‘PLEASE DONT REFUSE’ http://libgallery.cshl.edu/items/show/46113.
47.Nirenberg et al. (1963).
48.Nirenberg et al. (1963), p. 557.
49.Speyer et al. (1963).
50.Ochoa (1964).
51.Nirenberg and Leder (1964).
52.Heaton (2010), pp. 26, 31.
53.Heaton (2010), pp. 29–30.
54.Crick (1966b), p. 554.
55.Söll et al. (1965).
56.Salas et al. (1965). For a contemporaneous review, see Stent (1966).
57.Clark and Marcker (1966).
58.Friedberg (2010), p. 150.
59.Yanofsky et al. (1964).
60.Brenner et al. (1967).
61.Subak-Sharpe et al. (1966), Woese et al. (1966).
62.Crick (1966a), p. 3.
63.Jacob (1977).
64.http://profiles.nlm.nih.gov/ps/access/JJBCCQ_.jpg.
65.Stent (1968a).
66.Yčas (1969), p. 284.
67.Jacob (2011).
68.Cairns (1966).
Chapter 12
1.Monod and Jacob (1961), p. 393.
2.Crick (1970), p. 561.
3.Lewin (1974).
4.Chow et al. (1977), Klessig (1977), Dunn and Hassell (1977), Lewis et al. (1977), Berk and Sharp (1977), Berget et al. (1977).
5.Gilbert (1978), Crick (1979), Witkowski (1988).
6.Gilbert (1978).
7.Boyce et al. (1991), Fedorov et al. (1992), Hong et al. (2006).
8.Henikoff et al. (1986).
9.Crick (1979).
10.Burnet (1956), p. 22.
11.Crick (1959).
12.Rogozin et al. (2012).
13.Schmucker et al. (2000), Neves et al. (2004).
14.Lah et al. (2014).
15.Barrell et al. (1979).
16.Sapp (2009).
17.Archibald (2014).
18.Lane and Martin (2010), McInerney et al. (2014), Yong (2014).
19.Sánchez-Silva et al. (2003).
20.Hatfield and Gladyshev (2002), Srinivasan et al. (2002), Hao et al. (2002), Lobanov et al. (2006).
21.Kryukov et al. (2003).
22.Berry et al. (1993).
23.Rinke et al. (2013), Ivanova et al. (2014).
24.Starck et al. (2012).
25.Cavalcanti and Landweber (2004).
26.Lozupone et al. (2001), Lekomtsev et al. (2007).
27.Ring and Cavalcanti (2007).
28.Lajoie et al. (2013a, b).
29.Lane (2009).
30.Theobald (2010).
31.Holley et al. (1965).
32.Sanger (1988), p. 22. See García-Sancho (2010) for how Sanger’s sequencing strategy changed as he moved from proteins to DNA.
33.Sanger et al. (1977), van Noorden et al. (2014).
34.Sanger et al. (1978).
35.Rabinow (1996). For Mullis’s account of how he came up with the method see http://www.nobelprize.org/nobel_prizes/chemistry/laureates/1993/mullis-lecture.html.
36.Chien et al. (1976).
&
nbsp; 37.Zagorski (2006).
38.Botstein et al. (1980).
39.García-Sancho (2012).
40.Davies (2002), Sulston and Ferry (2002), Shreeve (2004), Ashburner (2006), Venter (2007).
41.The transcript of the ceremony, which includes some creepy banter between Clinton and Blair, can be found here: http://transcripts.cnn.com/TRANSCRIPTS/0006/26/bn.01.html.
42.Lander et al. (2001), Venter et al. (2001).
43.The Guardian, 13 June 2013.
44.The Guardian, 5 September 2014. Bizarrely, the court allowed patenting of genes only in material that has been removed from the human body. How else would the sequence be determined?
45.Mardis (2008), Schuster (2008).
46.Li et al. (2010), Worley and Gibbs (2010).
47.For microbes there is Genome Announcements, http://genomea.asm.org/.
48.Lim et al. (2014).
49.Hayden (2014).
50.MIT Technological Review, September 2014. http://www.technologyreview.com/news/531091/emtech-illumina-says-228000-human-genomes-will-be-sequenced-this-year/.
51.Genome of the Netherlands consortium (2014).
52.http://cancergenome.nih.gov.
53.Callaway (2014b).
54.Philippe et al. (2011).
55.http://www.genomesonline.org. For how to sequence a genome, see http://sciblogs.co.nz/tuataragenome/2013/06/25/first-find-your-tuatara-or-how-to-sequence-a-genome/.
56.Neale et al. (2014).
57.Bennett and Moran (2013).
58.McCutcheon and Moran (2011).
59.Woese and Fox (1977a, b). For an analysis of Woese’s work and its implications, see Sapp (2009).
60.Williams et al. (2013), McInerney et al. (2014).
61.Axelsson et al. (2013), Freedman et al. (2014).
62.Moroz et al. (2014).
63.Regier et al. (2010).
64.Crick (1958), p. 142. For historical analyses of the significance of this approach, see Stevens (2013) and Suárez-Díaz (2014).
65.http://timetree.org. The app is also called Timetree.
66.Orlando et al. (2013).
67.Penney et al. (2013).
68.Shapiro and Hofreiter (2014).
69.Pääbo (2014).
70.Krings et al. (1997), Green et al. (2010).
71.Fu et al. (2014).
72.Higham et al. (2014).
73.Sankararaman et al. (2014), Vernot and Akey (2014).
74.Prüfer et al. (2014).
75.Reich et al. (2010).
76.Jeong et al. (2014).
77.Huerta-Sánchez et al. (2014).
78.Hammer et al. (2011), Callaway (2014a), Prüfer et al. (2014).