Abstract
Chapter 6 turns to the investigation of the mechanism of carbon reduction in photosynthesis, which, by then, was known to be the “dark” (that is, light-independent) reaction of the process. The turning point of this project came with the advent of radioisotopes, as it was found that they could be used to trace the metabolic processes of plants and animals. The path-breaking work done by Martin Kamen and Samuel Ruben is described, that is, their experimental design to use radioactive carbon in the study of photosynthesis and their discovery of carbon-14 in 1940. However, the main part of the chapter reconstructs the subsequent work (after World War II) of the research team headed by Melvin Calvin and Andrew A. Benson, which eventually succeeded in elucidating the complex reaction cycle of photosynthetic carbon reduction. This team—generously supported by the AEC—was one of the first non-physical, large and interdisciplinary research groups at the time. It is analysed how the labour was divided within this group and which strategies guided the course of actions. Widespread heuristic moves were, for example, the transfer of knowledge from respiration to photosynthesis, the assumption that all biochemical reactions also run in reverse and the recombination of structural formula on paper.
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Notes
- 1.
The story of how the path of carbon in photosynthesis was elucidated has been told before, mostly in the form of autobiographical reports. See, e.g., Bassham (2003); Benson (2002a, b); Calvin (1964, 1989, 1992); Florkin (1979) (Chapter 56, pp. 81–108), Kamen (1974, 1985, 1989); Lehmann (1968) and Morton (2007). The book by Rabinowitch (1956) also has a section on the “Evolution of the CO\(_2\) Reduction Mechanism”, in which the different stages of the work carried out at Berkeley are summarised (pp. 1688–1698).
- 2.
On Lawrence, the Berkeley Radiation Laboratory and the early history of the cyclotron, see Heilbron and Seidel (1989); Heilbron et al. (1981) and Herken (2002). For a more general view that enlarges on the links with the Manhattan Project, see also Boorse et al. (1989) and Rhodes (1986). See Creager (2013) for an illuminating account of how profoundly the availability of radioisotopes through the cyclotron (and later the nuclear power stations) changed the life sciences after 1945.
- 3.
This is not to be confused with the equally famous MIT Radiation Laboratory, although both were also abbreviated to the same nickname “Rad Lab”.
- 4.
Cf. Creager (2013, pp. 24–41) and Creager (2006).
- 5.
He was honoured “for the invention and development of the cyclotron and for results obtained with it, especially with regard to artificial radioactive elements”. See www.nobelprize.org.
- 6.
His father was born in Belarus, his mother in “Lithuania or Latvia”, as he wrote in his autobiography, Kamen (1985, p. 2). Kamen (1989) provides a shorter account of his memories. See also Gest (2005), who was Kamen’s first doctoral student at Washington University, St. Louis.
- 7.
Cf. Creager (2013, pp. 41–49).
- 8.
On the life and work of Ruben, see, e.g., Gest (2004) and Johnston (2003), Chapter 3.
- 9.
Kamen (1985, pp. 81–82).
- 10.
In his autobiography, Kamen stressed that Ruben’s reputation was boosted when, after the latter had achieved some visibility in the department through the isotope studies, some graduate students chose to work with him on strictly physico-chemical problems, so that it became apparent that Ruben was not exclusively committed to biochemical work. See Kamen (1985, p. 114).
- 11.
See also Creager (2013, pp. 224–227).
- 12.
Urey was awarded the 1934 Nobel Prize in chemistry for these achievements.
- 13.
These studies’ results were later summarised in Schoenheimer (1940). See also Simoni et al. (2002) for a review of the work by Schoenheimer and Rittenberg. The classic papers are Schoenheimer and Rittenberg (1935, 1937). Kohler (1977) provides a thorough historical study of the episode.
- 14.
See Kamen (1985, p. 82).
- 15.
On Hassid, see Ballou and Barker (1979).
- 16.
This observation is supported by the case studies described in Holmes (2004).
- 17.
Kamen later was brought, by the same line of reasoning (although under completely different circumstances), to the study of bacterial metabolism—a topic to which he had never dreamed to contribute, but one that would greatly benefit from his specific skills and interests.
- 18.
See Kamen (1985, p. 83), for a description of the early stages of the project.
- 19.
In retrospect, one can understand the nature of these two stumbling blocks: first, as was later demonstrated, plants prefer “normal” carbon to its isotopes, so that radioactively labelled carbon dioxide tends to be absorbed at lower rates; second in contrast to standard (even very recently published) textbooks, Walker (2007, p. 182), has underlined the fact that “like sucrose, free glucose is not a major product of carbon assimilation by photosynthesising chloroplasts if, indeed, it is formed in the light at all”.
- 20.
Kamen (1985, p. 84).
- 21.
In the terminology suggested in Graßhoff et al. (2000), this implies turning the Herstellungsprozess into the actual Untersuchungsprozess. In the framework proposed by Hans-Jörg Rheinberger one may want to describe it as turning a “technical object” into an “epistemic object”; cf. Rheinberger (1997).
- 22.
Kamen (1985, p. 84). Additional background information on these early experiments can be found in Benson (1982) and Barker (1982).
- 23.
Kamen (1985, p. 86).
- 24.
Kamen (1985, p. 87).
- 25.
Kamen (1989, p. 140); Barker (1982, p. 68). After receiving his PhD in chemistry in 1933, Barker had been the first postdoctoral student of Cornelis B. van Niel at the Hopkins Marine Station. This was exactly the time when van Niel was elaborating his concept of a general equation of photosynthesis; cf. Chapter 4. Again, the closely interrelated network of actors and institutions in photosynthesis research during the 1930s is obvious.
- 26.
It was at the Carnegie Institution that Emerson and Lewis, during the former’s leave of absence from Caltech, started their work on the maximum quantum yield of photosynthesis; see Chapter 5. For a broader account of this institution’s history, see Craig (2005).
- 27.
Kamen (1985, p. 104).
- 28.
On the entanglement of cooperation and competition in the sciences, see Nickelsen (2014).
- 29.
Kamen (1985, p. 107).
- 30.
This was one of the findings that prompted the model developed by Franck and Herzfeld (1941); see Chapter 4. Ruben and Kamen’s research findings were summarised by Kamen in 1949 in a contribution to the volume edited by Franck and Loomis; see Kamen (1949). For the original papers, see: Ruben et al. (1939a, b, 1940a, b) and Ruben and Kamen (1940a).
- 31.
Kamen (1985, p. 109).
- 32.
Lipmann (1941) and Kalckar (1941). See also Chapter 7.
- 33.
Ruben (1943, p. 280).
- 34.
Kamen (1985, p. 162). However, Ruben’s reaction sequence did make Kamen think intensely about the general relationship between phosphate metabolism and biochemical energy storage, which later became a central theme of Kamen’s research work.
- 35.
Ruben (1943, p. 281).
- 36.
The seminal papers based on the study of propionic bacteria were Wood and Werkman (1935, 1936, 1938). See also Singleton (1997) and Krebs (1974) for historical accounts of this discovery.
- 37.
Ruben and Kamen (1940b).
- 38.
Cf. Kamen (1985, pp. 110–112).
- 39.
Kamen (1985), Chapter 7, is just one example; other references are given therein.
- 40.
The discovery was published in Ruben and Kamen (1941).
- 41.
The disadvantage of long half-lives is, of course, that the number of disintegrations per minute becomes very small—sometimes too small to be detected. Fortunately, though, carbon-14 turned out to be still within the biochemically useful range.
- 42.
This was a study in the metabolism of propionic acid bacteria, published as Carson et al. (1941).
- 43.
Kamen (1985, p. 140).
- 44.
See Ruben et al. (1941); confirmation was provided by Dole and Jenks (1944) and Holt and French (1948). Ruben et al. (1941) did not refer to the important work carried out by Robin Hill at the University of Cambridge (UK), which was described in Chapter 4; nor did Hill refer to Ruben et al. (1941), although they had arrived at similar conclusions at roughly the same time.
- 45.
See Vinogradov and Teiss (1941, 1947).
- 46.
For more background information, see Kamen (1985, 1989).
- 47.
See Seidel (1983), as well as Calvin (1992, p. 52); Seaborg and Benson (1997, p. 8).
- 48.
Laurence (1945, p. 6, 1946a, pp. 267–268, b, p. 41).
- 49.
On Calvin’s life and work, see, e.g., Loach (1997), Bassham (1997) and Seaborg and Benson (1997). See also Calvin (1989, 1992) for autobiographical accounts.
- 50.
On Benson’s life and work, see, e.g., Buchanan et al. (2007) and the autobiographical perspectives Benson (2002a, b, 2010).
- 51.
It was James A. Bassham, in particular, who emphasised in retrospect the importance of collaborating with such highly skilled glassblowers, machinists and carpenters; see Bassham (2003, p. 38).
- 52.
See, e.g., Benson (2002a, p. 34). There is a discrepancy between Benson’s and Calvin’s accounts of who was given the vial. Calvin claims to have inherited this vial from Ruben himself, see Calvin (1992, p. 53). However, this version seems highly unlikely in view of the fact that Benson, not Calvin, was, at the time, collaborating with Ruben.
- 53.
In a large-scale Oral History Project, carried out by Vivian Moses, in collaboration with his wife Sheila, a substantial number of scientists were asked about their recollections of working in Calvin’s group. The interviews were published online as Moses and Moses (2000). This quote is taken from the interview with Moses himself, at page 17/6 (i.e. from page 6 of the interview number 17 of the collection; all the interviews were numbered according to this pattern).
- 54.
Moses and Moses (2000), interview with Bassham, p. 7/2. On this episode, see also Bassham (2003, pp. 37–38).
- 55.
It is clear from the interviews carried out in the course of a Oral History Project by Vivian Moses, together with his wife Sheila, that only a few of the people who came to work with Calvin were aware of his status in science. The younger scientists mostly came across his laboratory accidentally.
- 56.
Moses and Moses (2000), interview with Bassham, p. 7/4.
- 57.
On Aronoff’s life and work, see Govindjee (2010).
- 58.
See Moses and Moses (2000), interview with Bassham, p. 7/3.
- 59.
Moses and Moses (2000), interview with Moses, p. 17/8.
- 60.
Moses and Moses (2000), interview with Goodman, p. 14/4.
- 61.
Moses and Moses (2000), interview with Quayle p. 3/9.
- 62.
On the seminar’s policy, see, e.g., Moses and Moses (2000), interview with Calvin, pp. 1/22–23.
- 63.
Moses and Moses (2000), interview with Quayle, p. 3/13.
- 64.
Moses and Moses (2000), interview with Taylor, p. 1/60.
- 65.
See for shorter treatments of the following episode, although with differing foci, Nickelsen (2012b) and Schüring (2006).
- 66.
Thimann (1938, p. 506).
- 67.
Thimann (1938, p. 506).
- 68.
Thimann (1938, p. 507).
- 69.
Krebs and Johnson (1937).
- 70.
Krebs and Henseleit (1932). On the discovery of the urea cycle, see Nickelsen and Graßhoff (2008); Holmes (1991).
- 71.
See Calvin (1989, p. 9).
- 72.
According to Benson, this came about through Calvin’s involvement as a consultant to Dow Chemicals, where new resins were being developed at the time. See Moses and Moses (2000), interview with Benson, p. 12/18.
- 73.
See Benson and Calvin (1947) and Calvin and Benson (1948) for the first publications from Berkeley. The approach was criticised in, e.g., Brown et al. (1949); Fager (1949); Gaffron and Fager (1951).
- 74.
Benson and Calvin (1947, p. 648), Table 1.
- 75.
Calvin and Benson (1948).
- 76.
Calvin (1962, pp. 880–881, 1989, p. 9).
- 77.
Calvin and Benson (1948, pp. 478–479).
- 78.
The same model hypothesis was still published in Benson et al. (1949, p. 399).
- 79.
See Bechtel and Richardson (1993); Bechtel and Abrahamsen (2005). The concepts of “functional” and “structural” decomposition also looms large in Bechtel (2006).
- 80.
“Using some of the reactions already established in animal tissue and bacteria, it is possible to account for the above results as well as the observed distribution of radiocarbon in short photosynthesis.” Benson and Calvin (1947, pp. 648–649).
- 81.
See Consden et al. (1944) for the publication of the method; their work was based on the seminal suggestion published in Martin and Synge (1941).
- 82.
The plant biochemist Albert Frenkel emphasised that it was Charles Dent who had brought paper chromatography from the UK to the USA in the first place and who was instrumental in first attempting to identify \(^{14}\)C-labelled amino acids; see Frenkel (1993, p. 106). Among the early relevant publications were Dent et al. (1947a, b) and Fink and Fink (1948). On the advent of paper chromatography in Berkeley see also Kamen (1985, p. 193), and several of the interviews in the Oral History Collection, Moses and Calvin (1958).
- 83.
See Stepka et al. (1948) for the first publication from the Berkeley Group based on paper chromatography (on amino acid separation, which was Stepka’s field of expertise). Benson et al. (1950) then demonstrated how successfully paper chromatography could be applied to identify carboxylic acids and phosphate esters.
- 84.
For a detailed description of this procedure, see also Calvin (1962, p. 881, 1989, p. 9); and Bassham and Calvin (1960, pp. 890–895).
- 85.
Moses and Moses (2000); Interview with Bassham, p. 7/10. See also Benson (2002a) and Bassham (2003).
- 86.
Bassham (2003, p. 41).
- 87.
Moses and Moses (2000), interview with Moses, pp. 17/12 ff.
- 88.
See Gaffron and Fager (1951, p. 91).
- 89.
Bassham (2003, p. 39).
- 90.
Emerson to Gaffron, 4 April 1950. Robert Emerson Papers, 1923-61, Record Series 15/4/28, Box 1, Folder: Gaffron, Hans, University of Illinois Archives.
- 91.
Benson and Calvin (1950).
- 92.
See, e.g., Badin and Calvin (1950); Calvin and Massini (1952).
- 93.
See Bassham et al. (1950). The idea that the “malic” enzyme should somehow also be involved in photosynthetic carbon dioxide fixation was, nevertheless, rather persistent. In Ochoa and Vishniac (1952), which was a very influential paper (see Chapter 7), the idea was not only revived but also expanded on, despite evidence to the contrary supplied by the inhibition studies.
- 94.
The first report on ribulose diphosphate (RDP) was given in Benson (1951); the first report on sedoheptulose phosphate in Benson et al. (1951).
- 95.
Benson (2002a, p. 39).
- 96.
Benson (2002a, p. 39). See also Bassham (2003, p. 42), for this episode.
- 97.
See Benson et al. (1951) for the pertinent publication.
- 98.
Benson (1951, p. 2972).
- 99.
Benson and Calvin (1947, p. 713).
- 100.
Benson et al. (1952, p. 4481).
- 101.
Moses and Moses (2000); interview with Kay, p. 20/5.
- 102.
Moses and Moses (2000); interview with Kay, p. 20/6.
- 103.
Moses and Moses (2000); interview with Kay, p. 20/5.
- 104.
Moses and Moses (2000); interview with Kay, p. 20/6.
- 105.
Moses and Moses (2000); interview with Tolbert, pp. 28/2–3.
- 106.
Moses and Moses (2000); Interview with Tolbert, p. 28/3.
- 107.
See Moses and Moses (2000), interview with Tolbert, p. 28/7. In a later section of the interview (p. 28/9), Tolbert described how she took a long career break to raise a family and then eventually became an accountant.
- 108.
See Calvin and Massini (1952, p. 451).
- 109.
Moses and Moses (2000); interview with Massini, p. 48/2. In this oral interview with Massini, his use of “switching out” the light has been corrected to “switching off” the light.
- 110.
Calvin and Massini (1952, p. 454).
- 111.
Wilson and Calvin (1955, p. 5949).
- 112.
Moses and Moses (2000); interview with Wilson, p. 13/7.
- 113.
See Moses and Moses (2000); interview with Wilson, p. 13/7.
- 114.
Wilson and Calvin (1955, p. 5952).
- 115.
Cf. Bassham et al. (1954, p. 1766).
- 116.
Quayle et al. (1954, p. 1766).
- 117.
See Wilson and Calvin (1955).
- 118.
Fuller (1999, p. 8).
- 119.
See Moses and Moses (2000); interview with Holtham, p. 23/8.
- 120.
The story is well remembered by many members of the laboratory. See Fuller (1999, pp. 8–9), and Moses and Moses (2000), e.g., the interviews with Calvin, Holtham, Kay, Moses, Wilson, etc.
- 121.
See Bassham et al. (1954). Bassham thought that, although the group had had a “pretty good handle on the cycle” before 1954, he always regarded “Path XXI” as being the definitive publication. See Moses and Moses (2000), interview with Bassham, p. 7/10.
- 122.
Bassham et al. (1954, p. 1767).
- 123.
Bassham et al. (1954, p. 1767).
- 124.
On the assumption of reverse reactions of known processes as a typical strategy within the biomedical sciences of the 1950s, see the argument in Scholl and Nickelsen (2015).
- 125.
Bassham et al. (1954, p. 1767).
- 126.
Benson (2002a, p. 45).
- 127.
Quayle et al. (1954, p. 3611). On Quayle, see Kornberg (2006); see also Fuller (1999).
- 128.
See Moses and Moses (2000), interviews with Mayaudon and Benson. See also Benson (2002a).
- 129.
Ning Pon recalled that there were endless discussions about what to call the enzyme. See Moses and Moses (2000, p. 9/3). In the same vein, Benson wrote on a Christmas card to Warburg: “An enthusiastic belgian [sic], Mayaudon, and I have finally succeeded in purifying the carboxylation enzyme. Should we call it ‘Photosynthase’ or ‘ribulose diphosphate carboxylase’? It seems to be a major leaf protein.” Archive of the BBAW, NL Warburg 114. Card undated.
- 130.
See Wildman (1992, 1998, 2002) for autobiographical accounts.
- 131.
Arne Tiselius, the inventor of this apparatus, received the 1948 Nobel Prize in chemistry for analytical work done with the help of this instrument.
- 132.
Wildman (2002, p. 245). The results were published in Wildman and Bonner (1947).
- 133.
Benson (2002a, p. 46). The paper referred to is Mayaudon et al. (1957); see also Mayaudon (1957).
- 134.
See Benson (2010) for an account of his “Last days in the old radiation lab”. In 2012, an interview with Benson on these events was recorded and posted to the web. For a description of the interview as well as the link to the video, see Buchanan and Wong (2013).
- 135.
Fuller (1999, pp. 9–10), pointedly summarised the disconcerting feeling with which one is left after having read Calvin’s autobiography, Calvin (1992), in which there is no single reference to Benson. Also thereafter, Calvin consistently rejected the fact that Benson played a crucial role in the photosynthesis project; cf., e.g., Moses and Moses (2000), interview with Calvin, p. 1/30. This, of course, is a cruel distortion of history.
- 136.
See Horecker et al. (1954) and Weissbach et al. (1956).
- 137.
See Dorner et al. (1957).
- 138.
On the history of Rubisco, see also Portis and Parry (2007).
- 139.
Moses and Moses (2000), interview with N. Tolbert, p. 29/20.
- 140.
Moses and Moses (2000), interview with Bassham, p. 7/17.
- 141.
See Moses and Moses (2000), Introduction by Moses, p. Intro-4. This point was emphasised also by others, see, e.g., the interviews with Park, p. 25/11, and Benson, pp. 12/29–30.
- 142.
See on these heuristics Scholl and Nickelsen (2015).
- 143.
See Kortschak et al. (1965).
- 144.
Cf. Hatch and Slack (1966); Hatch et al. (1967); retrospective accounts are given in Hatch (1992), Hatch (2002). Around the same time the occurrence of a “reductive carboxylic acid cycle” in some bacteria was reported, which in fact is a tricarboxylic acid cycle run in reverse in order to produce organic compounds from carbon dioxide in water; see Evans et al. (1966).
- 145.
The following references were cited in Bassham et al. (1954): Axelrod et al. (1953), Horecker and Smyrniotis (1952, 1953) and Racker et al. (1953).
- 146.
Perhaps the most prominent of the red herrings that were followed up in the laboratory was Calvin’s “thioctic acid theory” on which he worked intensely for more than two years. See Barltrop et al. (1954) and Calvin (1954) for central papers on this theme.
- 147.
Moses and Moses (2000), interview with Kay, p. 20/6.
- 148.
For some time, the University of Chicago group, which included Hans Gaffron, Edmund Fager, Jerome Rosenberg and Allan Brown, had been competing with Berkeley; but as they made much slower progress than their highly endowed Berkeley colleagues, they eventually dropped the project.
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Nickelsen, K. (2015). The Path of Carbon in Photosynthesis (1937–1954). In: Explaining Photosynthesis. History, Philosophy and Theory of the Life Sciences, vol 8. Springer, Dordrecht. https://doi.org/10.1007/978-94-017-9582-1_6
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