Abstract
Many physicists and philosophers coming to the philosophy of science through the inanimate sciences find biology an untidy subject1 in certain respects. It does not seem to have conceptual boundaries that can be made self-contained as in physics. This paper examines the problem of functional explanation, together with the role, if any, of theoretical entities (or ‘units’) in biology. To begin with I shall examine, fairly precisely, the physicist’s idea of a typical ‘theoretical entity’, and then look to see whether we in fact find theoretical entities of a similar type appearing in biological theory. I shall be considering four examples: the ‘gene’, the ‘organiser’, ‘messenger R.N.A.’ and the ‘repressor’. Then I shall apply my conclusions to the problems of functional explanation.
An earlier version of this paper was presented to the British Society for the Philosophy of Science in September, 1967. Much of the work involved in this paper has been supported by the National Science Foundation (Grant No. GS-1269). I thank them for their support.
I wish to acknowledge with gratitude the help I have received through discussions with Professor Gerald Edelmann, Professor John Maynard-Smith, Sir Peter Medawar, Dr. Stephen Toulmin, and Dr. Paul Weiss. Dr. Yehuda Elkana has given me extensive and perceptive help with the German translations.
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References
I had a delightful exchange on this point with Russ Hanson, in the spring of 1967 — a few weeks before he was killed. We had been discussing the whole problem of the ‘organiser’ in embryological theory, and Hanson revealed that he once wrote a paper about this while he was in Cambridge during the 1950’s. When I asked him what he said about the organiser, he was his characteristic, forceful self. “I wrote that it was just yet another of those damned theoretical entities that scientists are always thinking up.” But when I asked him what he had done with the paper, he gave me an answer that was completely out of character, for he admitted that he had thrown it in the wastepaper basket. When I pressed him as to why he had done this, he said, “Because it was messy.” Further discussion revealed that it was not so much the content of the paper that was messy, but the fact that as his study progressed he found that the ‘organiser’ did not present him with a nice, neat, tidy problem which could be wrapped up in a nice tidy solution. It was from this discussion, however, that the germ of this paper emerged so that I thought it worthwhile to look at the question of theoretical entities and functional explanation.
G. Mendel, ‘Experiments in Plant-Hybridization’, 1865. Translated by William Bateson, and reprinted in Classic Papers in Genetics (ed. by J. A. Peters), 1959, pp. 1-20.
W. Johannsen, Elemente der exakten Erblichkeitslehre, Jena 1909. And The American Naturalist 45 (1911) 149-159.
W. Johannsen, Über Erblichkeit in Populationen und in reinen Linien, Jena 1903. Translated by James A. Peters in Classic Papers in Genetics, London 1959.
W. Johannsen, Elemente, 1909, p. 124.
W. Johannsen, Elemente, p. 143, 2nd ed., 1913. Translated for me by Dr. Yehuda Elkana.
W. Johannsen, The American Naturalist 45 (1911) 132–133.
W. Johannsen, The American Naturalist 45 (1911) 154.
T. H. Morgan, The Theory of the Gene, 1st ed., 1926; revised 1928; 2nd ed., 1929. Reprinted New York, 1965, pp. 26-27.
See, for instance, P. Medawar, ‘A Biological Retrospect’. Presidential Address to the British Association, 1965. Reprinted in The Art of the Soluble, 1967, pp. 99-110.
P. Medawar, The Art of the Soluble, 1967, p. 106.
C. H. Waddington, New Patterns in Genetics and Development, 1962, p. 241.
V. Hamburger, ‘Some Thoughts on the Organiser’, Presidential Address to the American Society of Zoologists, East Lansing, Michigan, September, 1965: “The term ‘organiser’ is closely associated with some key notions in biology such as organisation, organism, organ. It suggests that we are at the heart of a basic problem: the coming into being of organised structure or form.… The organiser concept seems to tackle the problem of morphgenesis boldly on the highest level; it focusses on the coming into being of the organism as a whole. Invertebrates, wholeness is represented by the axial organs, their polarity, bilateral symmetry and the pattern of their arrangement. It is certainly one of the foremost and most challenging tasks of embryology to explain how this organisation, this spatial pattern comes into being, and to search for the causal factors which are instrumental in its manifestation.”
For a survey of the problem of the time-control in development, and an attempt to produce a ‘dynamic theory of cellular-control processes’, see B. C. Goodwin, Temporal Organisation in Cells, London 1963.
Many elementary texts describe this process. For instance, an excellent simple account is in C. H. Waddington, Principles of Development and Differentiation, 1966.
Triton species.
Amblystoma species.
A. Weismann, Über Vererbung, Jena 1883. The Germ Plasm, translated, 1902.
H. Spemann, Embryonic Development and Induction, 1938. For Spemann’s discussion of Weismann’s theory, and its influence on Roux, see pp. 14-39.
H. Spemann, ‘Entwicklungsphysiologische Studien am Tritonei, I’, Arch. f. Entw. Mech. 12 (1901b) 224–264. — Idem II, Arch.f. Entw. Mech., 1902. — Idem III, Arch.f. Entw. Mech. 16 (1903a) 551-631.
J. Loeb, ‘Über eine einfache Methode, zwei oder mehr zusammengewachsene Embryonen aus einem Ei hervorzubringen’, Pflügers Arch. 55 (1894) 525–530.
H. Spemann, ‘Die Entwicklung seitlicher und dorso-ventraler Keimhälften bei verzögerter Kernversorgung’, Ztschr.f. wiss. Zool. 132 (1928). 105–134. This paper of Spemann’s written in 1928 has been abridged and translated by M. L. Gabriel. It will be found in Great Experiments in Biology (ed. by Gabriel and Fogel) as ‘The Development of Lateral and Dorso-Ventral Embryo Halves with Delayed Nuclear Supply’, 1955, pp. 215-219.
A whole range of articles on the transplantation of cell nuclei exists, but see for example: T. J. King and R. Briggs, Proceedings National Academy of Sciences, Washington 41 (1955) 321. — T. J. King and R. Briggs, Cold Spring Harbor Symposium of Quantitative Biology 21, 271. — J. B. Gurdon, ‘Transplantation of Cell Nuclei’, in Advances in Morphogenesis (ed. by M. Abercrombie and J. Brachet), Vol. IV, 1964, pp. 1-43.
See notes 20, 21, and 22.
H. Spemann, ‘Über die Determination der ersten Organanlagen des Amphibienembryo, I–VI’, Arch.f. Entw. Mech. 43 (1918) 448–555.
H. Spemann, ibid., 1918. See especially the last two sections of the discussion. Section D: ‘The Point of Origin and the Spatial Spread of the Determination’, pp. 529-540; and Section E: ‘The Theory of Double Formations’, pp. 540-543.
H. Spemann, ‘Die Erzeugung tierischer Chimaeren durch hetero-plastische embryonale Transplantation zwischen Triton cristatus u. Taeniatus’, Arch.f. Entw. Mech. 48 (1921a) 533–570.
H. Spemann, ibid. (1921) 568. (Translated by Dr. Yehuda Elkana.)
H. Spemann and Hilde Mangold, ‘Über Induktion von Embryonalanlagen durch Implantation artfremder Organisatoren’, Arch. f. Mikr., Anat. u Entw. Mech. 100 (1924) 599–638. This article was translated by Professor Viktor Hamburger and reprinted in Foundations of Experimental Embryology (ed. by Willier and Oppenheimer), 1964. I have used the translation of this paper throughout, and where in the footnotes a page reference is given, it refers to the English translation, not the German original.
H. Spemann and H. Mangold, ibid., 1924, 147.
B. H. Willier and J. M. Oppenheimer, Foundations of Experimental Embryology, 1964, p. 146.
H. Spemann and H. Mangold, ibid., 1924, 153.
H. Spemann and H. Mangold, ibid., 1924, 180.
H. Spemann and H. Mangold, ibid., 1924, 182.
My italics.
H. Spemann and H. Mangold, ibid., 1924, 182.
An essential part of the research which I am conducting — whose end result will be, a book upon the concept of the organiser in the history of embryology — is a series of extensive interviews with many distinguished embryologists who worked either directly with Spemann or who were active in the field during the 20 years between 1920 and 1945 when the ‘organiser’ was very much in the news. In addition, a questionnaire has been sent out to some 75 embryologists about the whole history of the episode and their views on the current status of the concept. These interviews and questionnaires will be fully documented subsequently.
In analyzing Spemann’s attitude to the organiser during the years 1920 and 1924, and up to approximately 1927, it must be clearly understood that what I am here describing is — insofar as I am able to judge from my researches up to this point — his attitude during these years. It is to be in no way taken for granted that this remained static. On the contrary, there is a great deal of evidence that over the years his beliefs in the nature of the ‘organiser’ changed to a great extent. It seems — though this awaits further documentation — that he began by regarding the ‘organiser’ somewhat vitalistically and ended up, by the time of the publication of his Silliman Lectures in 1938, treating it heuristically, indeed, operationally.
J. S. Huxley and G. R. DeBeer, Elements of Experimental Embryology, 1934, p. 12. (This book is dedicated to Ross Harrison and Hans Spemann.)
This anecdote has been reported to me by several people. It is also supposed to have suggested the’ sandwich’ technique for further induction experiments.
C. H. Waddington, New Patterns in Genetics and Development, 1962, p. 14.
Personal communication.
H. Spemann, ‘The Organiser Effect in Embryonic Development’, Nobel Prize Lecture, December 12, 1935.
For another discussion of this see, for instance, Sydney Smith, ‘The Use of Models in the Teaching of Embryology’, in Models and Analogues in Biology (Symposia of the Society for Experimental Biology), Academic Press, New York, 1960, p. 228.
F. Jacob and J. Monod, ‘Genetic Regulatory Mechanisms in the Synthesis of Proteins’, Journal of Molecular Biology 3 (1961) 318–56.
D. Gillispie and S. Spiegelman, ‘A Quantitative Assay for DNA-RNA Hybrids with DNA immobilized on a Membrane’, Journal Molecular Biology 12 (1965) 839–842.
F. Jacob and J. Monod, ibid., Journal of Molecular Biology 3 (1961) 318-56.
W. Gilbert and B. Miller-Hill, ‘The Isolation of the Lac Repressor’, Proceedings of the National Academy of Sciences 56, No. 6 (1966) 1891–1898.
E. W. Sinnott, L. C. Dunn, and Th. Dobzhansky, Principles of Genetics, 4th ed., 1950, p. 461.
This statement caused considerable difficulty in the discussion at the Boston Colloquium. Certain people disputed it, preferring to interpret the phrase ‘theoretical entity’ as referring to anything in a theoretical explanation which can be said to have ‘material existence’; and arguing therefore that many examples could be found of explanations of this form in biology. Some of the examples cited (e.g., ‘cells’) were well-chosen; others (e.g.,’ species’) were, I think, certainly not. In my own argument, however (and in my initial discussion with Russ Hanson), I was referring to explanation in terms of hypothetical units, and in this sense, my point stands. In this regard it will be interesting to see what becomes of J. Z. Young’s ‘units of memory’. These hypothetical units do not manifest themselves like genes, in any gross behavioural patterns, but are utilised in constructing a theoretical model for the memory-system of the brain. (J. Z. Young, ‘The Organisation of a Memory System’, The Croonian Lecture, Proceedings of the Royal Society, B. 163 (1965) 285–320, especially 285-288). Though Young, too, is aware of how the problem of biological complexity makes explanations of this unit kind hazardous: “It may be that in nervous systems, with their immensely rich web of connexions, the very conception of unit-changes is irrelevant and indeed misleading” (p. 285). See also the following notes, 50 and 51, for further discussion of this problem.
For a full discussion of this whole problem see: G. J. Goodfield, The Growth of Scientific Physiology, London 1960, pp. 60–75 and 135-155.
Considerations of these sorts may have been at the root of Bernard’s well-known reservations about the applicability of statistical methods to the study of organisms. Bernard is the man who could — for the reason of the complexity of the interacting phenomena — endorse the statement of Regnault and Reiset: “It was therefore by a fortuitous circumstance in the experiments of Lavoisier, Dulong and Despretz, that the quantity of heat liberated by an animal was found to be about equal to what the carbon (contained in the carbonic acid produced) and the hydrogen would fire off in burning.” Regnault and Reiset, ‘Recherches chimiques sur la respiration des animaux des diverses classes’, Annales de chimie et de physique, Third series, 26, p. 573. Quoted in C. Bernard, An Introduction to the Study of Experimental Medicine, translated by Copley Greene, 1927, p. 133.
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Goodfield, J. (1969). Theories and Hypotheses in Biology. In: Cohen, R.S., Wartofsky, M.W. (eds) Boston Studies in the Philosophy of Science. Boston Studies in the Philosophy of Science, vol 5. Springer, Dordrecht. https://doi.org/10.1007/978-94-010-3381-7_14
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