Abstract
H. B. D. Kettlewell's (1955, 1956) classic field experiments on industrial melanism in polluted and unpolluted settings using the peppered moth, Biston betularia, are routinely cited as establishing that the melanic (dark) form of the moth rose in frequency downwind of industrial centers because of the cryptic advantage dark coloration provides against visual predators in soot-darkened environments. This paper critiques three common myths surrounding these investigations: (1) that Kettlewell used a model that identified crypsis as the only selective force responsible for the spread of the melanic gene, (2) that Kettlewell's field experiments alone established that selection for crypsis was the most important factor in the spread of melanic forms, and (3) that Kettlewell's investigations in an unpolluted wood near Dorset constituted a control for his earlier Birmingham studies (contra Hagen 1993, 1996). This analysis further identifies two features that distinguish manipulative experiments in evolutionary biology from experiments in other contexts. First, experiments in evolutionary biology rest on a wealth of information provided by strictly observational ecological studies; in the absence of such information experiments in evolutionary biology make no sense. Second, there is a trade-off between how much control investigators have over the conditions being studied and how informative the results of the experiment will be with regard to natural populations.
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References
Achinstein, P. and Hannaway, O. (eds.): 1985, Observation, Experiment and Hypothesis in Modern Physical Science, MIT Press, Cambridge, MA.
Allen, P.M.B.: 1955, ‘Review of E. B. Ford's Moths’, The Entomologist's Record and Journal of Variation 67, 104.
Batens, D. and van Bendegem, J.P. (eds.): 1988, Theory and Experiment: Recent Insights and New Perspectives on Their Relation, D. Reidel Publishing Company, Dordrecht.
Berry, R.J.: 1990, ‘Industrial Melanism and Peppered Moths (Biston betularia (L))’, Biological Journal of the Linnean Society 39, 301–322.
Bishop, J.A. and Cook, L.M.: 1980, ‘Industrial Melanism and the Urban Environment’, Advances in Ecological Research 11, 373–404.
Bowater, W.: 1914, ‘Heredity of Melanism in the Lepidoptera’, Journal of Genetics 3, 209–315.
Brandon, R.N.: 1990, Adaptation and Environment, Princeton University Press, Princeton.
Brandon, R.N.: [1994] 1996, ‘Theory and Experiment in Evolutionary Biology’, reprinted in R.N. Brandon (ed.), Concepts and Methods in Evolutionary Biology, Cambridge University Press, Cambridge, England.
Chapman, T.A.: 1888, ‘On Melanism in Lepidoptera’, The Entomologist's Monthly Magazine 25, 40.
Clarke C.A. and Sheppard, P.M.: 1966, ‘A Local Survey of the Distribution of the Industrial Melanic Forms of the moth Biston betularia and Estimates of the Selective Values of These in an Industrial Environment’, Proceedings of the Royal Society (B) 165, 424–439.
Cooke, N.: 1887, ‘On Melanism in Lepidoptera’, The Entomologist's Monthly Magazine 10, 92–96, 151–153.
Culp, S.: 1995, ‘Objectivity in Experimental Inquiry: Breaking Data Technique Circles’, Philosophy of Science 62, 438–458.
Demuth, R.P.: 1955, ‘Birds and Moths’, The Entomologist's Record and Journal of Variation 67, 157–158.
Diamond, J.: 1986, ‘Laboratory Experiments, Field Experiments and Natural Experiments’, in Diamond, J. and Case, T. (eds.), Community Ecology, Harper & Row, New York.
Dietrich, M.R.: 1991, Theory and Experiment in Molecular Population Genetics, Ph.D.Thesis, University of California, San Diego.
Fisher, R.A.: 1930, The Genetical Theory of Natural Selection, Dover Publications Inc., New York.
Ford, E.B.: 1937, ‘Problems of Heredity in the Lepidoptera’, Biological Reviews 12, 461–503.
Ford, E.B.: 1940, ‘Genetic Research on the Lepidoptera’, Annals of Eugenics 10, 227–252.
Ford, E.B.:1955, Moths, Collins Press, London.
Franklin, A.: 1986, The Neglect of Experiment, Cambridge University Press, Cambridge.
Franklin, A.: 1990, Experiment, Right or Wrong, Cambridge University Press, Cambridge.
Galison, P.: 1987, How Experiments End, University of Chicago Press, Chicago.
Gooding, D., Pinch, T. and Schaffer, S. (eds.): 1989, The Uses of Experiment, Cambridge University Press, New York.
Grant, B.S., Owen, D.F. and Clarke, C.A.: 1996, ‘Parallel Rise and Fall of Melanic Peppered Moths in America and Britain’, Journal of Heredity 87, 351–357.
Hacking, I.: 1983, Representing and Intervening, Cambridge University Press, New York.
Hagen, J.: 1993, ‘Kettlewell and the Peppered Moths Reconsidered’, Bioscene 19, 3–9.
Hagen, J.: 1996, ‘H. B. D. Kettlewell and the Peppered Moths’, in Hagen, J., Allchin, D. and Singer, F. (eds.), Doing Biology, Harper Collins, New York.
Haldane, J.B.S.: 1924, ‘A Mathematical Theory of Natural and Artificial Selection’, Transactions of the Cambridge Philosophical Society 23, 3–41.
Harrison, J.W.H.: 1920a, ‘Genetical Studies in the Moths of the Geometrid Genus Oporabia (Oporinia) with a Special Consideration of Melanism in the Lepidoptera’, Journal of Genetics 9, 195–280.
Harrison, J.W.H.: 1920b, ‘The Inheritance of Melanism in the Genus Tephorsia (Ectropis), with Some Consideration of the Inconstancy of Unit Characters’, Journal of Genetics 10, 61–85.
Harrison, J.W.H.: 1926, ‘The Inheritance of Wing Colour and Pattern in the Leptidopterous Genus Tephrosia (Ectropis) II. Experiments Involving Melanic T. bistortata and Typical T. crepuscularia’, Journal of Genetics 17, 1–19.
Harrison, J.W.H.: 1927, ‘The Induction of Melanism in the Lepidoptera, and its Evolutionary Significance’, Nature 119, 127–129.
Hughes, A.W.: 1932, ‘Induced Melanism in the Lepidoptera’, Proceedings Royal Society B 110, 378–402.
Kettlewell, H.B.D.: 1955, ‘Selection Experiments on Industrial Melanism in the Lepidoptera’, Heredity 9, 323–342.
Kettlewell, H.B.D.: 1956, ‘Further Selection Experiments on Industrial Melanism in the Lepidoptera’, Heredity 10, 287–301.
Kettlewell, H.B.D.: 1958, ‘A Survey of the Frequencies of Biston betularia (L.) (Lep.) and its Melanic Forms in Great Britain’, Heredity 12, 51–72.
Kettlewell, H.B.D.: 1973, The Evolution of Melanism: The Study of a Recurring Necessity, Clarendon Press, Oxford.
Kimler, W.C.: 1983a, ‘Mimicry: Views of Naturalists and Ecologists Before the Modern Synthesis’, in Greene, M. (ed.), Dimensions of Darwinism, Cambridge University Press, Cambridge, pp. 97–127.
Kimler, W.C.: 1983b, One Hundred Years of Mimicry: History of an Evolutionary Exemplar, Ph.D. Thesis, Cornell University, Ithaca NY.
Knorr Cetina, K.: 1990, ‘The Couch, the Cathedral and the Lab: On the Relationship between Experiment and Laboratory in Science’, in Pickering, A. (ed.), Science as Practice and Culture, University of Chicago Press, Chicago.
Lambert, D.M., Miller, C.D. and Hughes, T.J.: 1986, ‘On the Classic Case of Natural Selection’, Biology Forum 79, 11–49.
Le Grand, H.E. (ed.): 1990, Experimental Inquires: Historical, Philosophical and Social Studies of Experimentation in Science, Kluwer Academic Publishers, Dordrecht.
Mader, Sylvia S.: 1988, Inquiry into Life (5th edn.), Wm. C. Brown Publishers, Dubuque, Iowa.
Mani, G.: 1990, ‘Theoretical Models of Melanism in Biston betularia — A Review’, Biological Journal of the Linnean Society 39, 355–371.
Merrifield, F.: 1890, ‘Systematic Temperature Experiments on Some Lepidoptera, in All Their Stages’, Transactions of the Royal Entomological Society of London 1890, 131–159.
Mikkola, K.: 1984, ‘On the Selective Forces Acting in the Industrial Melanism of Biston and Oligia Moths (Lepidoptera, Geometridae and Noctuidae)’, Heredity 52, 9–16.
Onslow, H.: 1920, Journal of Genetics 9, 339–346 (as cited in Ford 1940).
Rheinberger, H.: 1992a, ‘Experiment, Difference and Writing: I. Tracing Protein Synthesis’, Studies in the History and Philosophy of Science 23, 305–331.
Rheinberger, H.: 1992b, ‘Experiment, Difference and Writing: II. The Laboratory Production of Transfer RNA’, Studies in the History and Philosophy of Science 23, 389–422.
Rudge, D.W.: 1996, A Philosophical Analysis of the Role of Selection Experiments in Evolutionary Biology, Ph.D. Thesis, University of Pittsburgh, Pittsburgh.
Thomsen, M. and Lemeche, H.: 1933, ‘Experimente zur Erzielung eines Erblichen Melanismus bei dem Spanner Selenia bilunaria Esp.’, Biologisches Zentralblatt 53, 541–560.
Timoféeff-Ressovsky, N.W.: 1933, Archiv für Naturgeschicht N. F. 2, 285–290 (as cited in Ford 1940).
Turner, J.R.G.: 1983, ‘“The Hypothesis that Explains Mimetic Resemblance Explains Evolution”: The Gradualist-Saltationist Schism’, in Greene, M. (ed.), Dimensions of Darwinism, Cambridge University Press, Cambridge, pp. 129–169.
Turner, J.R.G.: 1985a, ‘Fisher's Evolutionary Faith and the Challenge of Mimicry’, in Dawkins, R. and Ridley, M. (eds.), Oxford Surveys in Evolutionary Biology, Vol. 2, Oxford University Press, New York, pp. 159–196.
Turner, J.R.G.: 1985b, ‘Random Genetic Drift, R. A. Fisher, and the Oxford School of Ecological Genetics’, in Kruger, L., Gigerenzer, G. and Morgan, M. (eds.), The Probabilistic Revolution Volume 2: Ideas in the Sciences, MIT Press, Cambridge, MA, pp. 312–354.
Turner, J.R.G.: 1987, ‘Random Genetic Drift, R. A. Fisher, and the Oxford School of Ecological Genetics’, in Kruger, L., Gigerenzer, G. and Morgan, M. (eds.), The Probabilistic Revolution Volume 2: Ideas in the Sciences, MIT Press, Cambridge, MA, pp. 313–354.
Turner, J.R.G.: 1990, ‘Henry Bernard Davis Kettlewell’, in Holmes, F.L. (ed.), The Dictionary of Scientific Biography, Vol. 17,Supplement 2, Charles Scribner's Sons, New York, pp. 469–471.
Tutt J.W.: 1890, ‘Melanism and Melanochroism in British Lepidoptera’, The Entomologist's Record and Journal of Variation 1, 5–7, 49–56, 84–90, 121–125, 169–172, 228–234, 293–300, 317–325.
Tutt J.W.: 1891, ‘Melanism and Melanochroism in British Lepidoptera’, The Entomologist's Record and Journal of Variation 2, 3–7, 31–35, 49–53, 77–80, 97–98, 145–149.
Weismann, A.: 1882, Studies in the Theory of Descent, Part I., Sampson, Low, Marston, Searle & Rivington, London.
Williams, H.B.: 1932–1933, ‘Notes on Boarmia repandata and B. rhomboidaria’, Proceedings of the South London Entomology and Natural History Society 1932–1933, 1–10.
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Rudge, D.W. Taking the Peppered Moth with a Grain of Salt. Biology & Philosophy 14, 9–37 (1999). https://doi.org/10.1023/A:1006524501723
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DOI: https://doi.org/10.1023/A:1006524501723