Abstract
In the first decades of the twentieth century, the process of photosynthesis was still a mystery: Plant scientists were able to measure what entered and left a plant, but little was known about the intermediate biochemical and biophysical processes that took place. This state of affairs started to change between the two world wars, when a number of young scientists in Europe and the United States, all of whom identified with the methods and goals of physicochemical biology, selected photosynthesis as a topic of research. The protagonists had much in common: They had studied physics and chemistry (although not necessarily plant physiology) to a high level; they used physicochemical methods to study the basic processes of life; they believed these processes were the same, or very similar, in all life forms; and they were affiliated with institutions that fostered this kind of study. This set of cognitive, methodological, and material resources enabled these protagonists to transfer their knowledge of the concepts and techniques from microbiology and human biochemistry, for example, to the study of plant metabolism. These transfers of knowledge had a great influence on the way in which the biochemistry and biophysics of photosynthesis would be studied over the following decades. Through the use of four historical cases, this paper analyzes these knowledge transfers, as well as the investigative pathways that made them possible.
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Notes
In this paper, the term “mechanism” is used roughly in the sense of the actors’ categories at the time, namely the pathways and the sequence of steps in the light-dependent reactions underlying the physiological phenomena. I believe that there exist constructive links between this contemporary understanding of the term and the “philosophy of mechanisms,” proposed by Bechtel and Richardson (1993) and Machamer, Darden, and Craver (2000), but they need to be explored in another paper.
There was, of course, a longer tradition of photosynthesis research in plant physiology, both in the laboratory and in the field, notably that undertaken by scientists in the Desert Laboratory in Tuscon, Arizona. On the latter tradition, see Craig (2005) and Kingsland (2005). For an overview of the American tradition from the perspective of the actors, see Went (1958). However, most of this research did not yet include the investigation of biochemical and biophysical mechanisms on which this paper is focused.
This term was introduced by Holmes (2004).
See also Warburg (1914b, p. 320), in which he discusses the advantage of working with cells rather than tissues.
Kok (1960) provides an overview of how the manometric methods that Warburg used in his photosynthesis studies developed over time.
These figures, however, turned out to be inaccurate by a factor of two to three, a mistake that Warburg never acknowledged. In the years after 1945, however, the discrepancy between Warburg’s measurements and those of almost everybody else became the subject of acrimonious controversy; see Nickelsen and Govindjee (2011).
His father had tried to persuade Warburg to study photosynthesis and, in particular, its quantum efficiency, back in 1912. Nothing came of it, though, as Otto Warburg was then too deeply involved in the study of respiration; cf. Nickelsen (2009).
For a philosophical perspective on analogical reasoning, see Bartha (2016).
See the interview with Delbrück in 1978, carried out as part of the Caltech Archives Oral History Project; for his years in Berlin, see Delbrück (1978, pp. 41–60). Delbrück claimed that he was particularly inspired by Bohr’s lecture on “Light and Life”; see Bohr (1933). On the question of how Bohr and Delbrück regarded the relationship between biology and physics, see Roll-Hansen (2011).
The Delbrück club and the paper on the nature of the gene, in particular, became famous primarily through its reception by Erwin Schrödinger and the (not always accurate) reference to this paper in Schrödinger’s lectures of 1944, entitled “What is Life?” From what we know, though, it is highly probable that photosynthesis was equally important as a topic of debate at the club’s gatherings. At least two other photosynthesis experts are reported to have also joined Delbrück’s club (e.g. French 1979, p. 7): the plant physiologist C. Stacy French (1907–1995), who in 1935 was a postdoctoral student in Warburg’s laboratory and later led the Department of Plant Biology at the Carnegie Institution, Stanford (see Craig 2005; Govindjee and Fork 2006), and the physical chemist Eugene Rabinowitch (1901–1973), who was then at the KWI for Physical Chemistry and in 1946 became co-director, with Emerson, of the Photosynthesis Project at the University of Illinois at Urbana–Champaign (see Bannister 1972; Govindjee 2004).
He was soon proven wrong by the development of the theory of resonance energy transfer; see Förster (1950).
For more detail, see Sloan (2011, p. 79). Sloan, however, erroneously assumed that Gaffron and Wohl (1936) had already accepted Cornelis B. van Niel’s generalized equation for photosynthesis (see “Isolated Chloroplasts, Oxygen Generation, and General Biochemistry” section of this paper), including the hypothesis that photosynthetic oxygen came from water. Gaffron himself explained that he delayed accepting the equation primarily out of enormous loyalty to (and admiration for) Warburg at the time; see Gaffron (1969).
Hill succeeded Muriel Wheldale Onslow (1880–1932), a pioneer of biochemistry from 1926 until her untimely death in 1932; see Richmond (2007).
On Keilin’s life and work, see Mann (1964).
Among those who had tried to investigate chlorophyll was the German scientist Alexander Tschirch (1856–1939). In his memoir, Tschirch wrote how he had discovered that it was impossible to study the chemistry of this pigment without clashing fiercely with one’s colleagues: “It is as if there were a goblin sitting in the chlorophyll, who takes great pleasure in setting all those researchers attempting to unravel its mysteries against each other” (1921, p. 182). See Höxtermann (1991).
This assumption only changed in 1941, after experiments with “heavy” water that incorporated the isotope oxygen-18 were conducted. The finding that the isotope ratio of photosynthetically evolved oxygen was identical to that of the water finally provided conclusive evidence that oxygen was produced by the splitting of water; see Ruben et al. (1941).
The “agent” is introduced in the paper on pp. 215–216. See also Friedmann (2004, p. 54).
Kluyver drew attention to this concept in a series of lectures in London, published as Kluyver (1931).
For a timeline of research in anoxygenic photosynthesis, see Gest and Blankenship (2004).
In the same paper, van Niel interpreted the metabolism of the Athiorhodaceae as being analogous to the metabolism of the Thiorhodaceae, although the former used “primarily simple organic substances as hydrogen donors for the photoreduction of carbon dioxide.” Van Niel (1941, p. 281).
See van Niel (1967, p. 10).
These “general” branches of biology were not directly related to the contemporary debates on “general biology”; see Laubichler (2006) and Sucker (2002). These debates were concerned with the role of the organism and its environment (see also Baedke 2018), while general physiologists and others were mostly interested in the proximate causes of cellular processes, which they investigated using the simplest of organisms possible (such as unicellular algae).
The Cold Spring Harbor Laboratory conferences were instigated with the purpose of “fostering a closer relationship between biology and the basic sciences” (Harris 1935, preface). Biologists, physicists, chemists, mathematicians, and other scientists were invited to the symposia held each summer to discuss different areas of biological research.
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Acknowledgements
I would first like to thank the editors, Jan Baedke and Christina Brandt, for proposing this thematic special issue and for including my contribution in this compilation, as well as the editors of the JHB for giving me the opportunity to do so. I am grateful to Robert Meunier, David Munns, Raphael Scholl, and Caterina Schürch for their helpful comments on earlier drafts of this paper, and to Margareta Simons, who carefully edited the final version. Finally, I would like to thank the two anonymous referees for their detailed reviews and valuable comments and suggestions. Although their widely divergent opinions made the task of incorporating their recommendations rather challenging, their observations certainly prompted me to rethink and rework the paper thoroughly—hopefully for the better.
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Nickelsen, K. Physicochemical Biology and Knowledge Transfer: The Study of the Mechanism of Photosynthesis Between the Two World Wars. J Hist Biol 55, 349–377 (2022). https://doi.org/10.1007/s10739-019-9559-x
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DOI: https://doi.org/10.1007/s10739-019-9559-x