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The Wonderful Form of Cosmic Order: Bringing Statistics to Evolution

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The introduction of statistical reasoning into evolutionary theory – and the corresponding changes in our concepts of chance and evolution that made it possible – did not happen overnight. In particular, many of the relevant biologists between 1900 and 1930 (such as W. F. R. Weldon, Karl Pearson, and R. A. Fisher) pointed back to Francis Galton as a vitally important precursor for their work. When we read Galton, however, it is far from immediately obvious why he would be taken to be so crucial by so many important successors. In this talk, I’ll explore some of Galton’s ideas about the relationship between micro-level variation and population change, and try to reconstruct what within his thought might have been seen to be so groundbreaking by biologists in the first part of the twentieth century. I will close by drawing some morals for our contemporary conception of evolutionary theory.

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The Wonderful Form of Cosmic Order: Bringing Statistics to Evolution

  1. 1. The Wonderful Form of Cosmic Order Bringing Statistics to Evolution cefises wip, 27.2.2019 Charles H. Pence @pencechp
  2. 2. Outline 1. A historical paradox 2. Galton’s early work 3. Natural Inheritance 3.1 Particulate inheritance 3.2 Chances and causes 3.3 Natural selection (or supposed to be) 4. A research program for biometry 5. Two perennial questions in evolution The take-home: Galton builds a school around him in no small part because he identifies two central, recurring philosophical issues regarding statistical theories. Charles H. Pence 2 / 36
  3. 3. A Historical Paradox Charles H. Pence A Historical Paradox 3 / 36
  4. 4. Charles H. Pence A Historical Paradox 4 / 36
  5. 5. Charles H. Pence A Historical Paradox 5 / 36
  6. 6. The next year was to place in Weldon’s hands a book – Francis Galton’s Natural Inheritance, by which one avenue to the solution of such problems [in variation, correlation, and evolution], one quantitative method of attacking organic correlation, was opened out to Weldon; and from this book as source spring two of the friendships and the whole of the biometric movement, which so changed the course of his life and work. (Pearson 1906, 13–14) Charles H. Pence A Historical Paradox 6 / 36
  7. 7. no one who studied it on its appearance and had a receptive and sufficiently trained mathematical mind could deny its great suggestiveness, or be other than grateful for all the new ideas and possible problems which it provided. The methods of Natural Inheritance may be antiquated now, but in the history of science it will be ever memorable as marking a new epoch... (Pearson 1930, 57–58) Charles H. Pence A Historical Paradox 7 / 36
  8. 8. Charles H. Pence A Historical Paradox 8 / 36
  9. 9. the argumentation concerning this concept [the idea of “natural ability”] was, as frequently with Galton, very bad, but the concept was powerful if vague.... (Norton 1978, 43) Charles H. Pence A Historical Paradox 9 / 36
  10. 10. [The biologist and textbook author W. K. Brooks] included a whole chapter on Galton in his Foundations of Zoology and explicitly addressed the ambiguities in his writings by quoting passages that seem both to support and to deny a role for selection in determining the fate of the new characters produced by saltation. In the end, however, it seems that Galton was simply ambiguous on the question of the role played by selection... (Bowler 2014, 276, 274) Charles H. Pence A Historical Paradox 10 / 36
  11. 11. Galton’s Early Work Charles H. Pence Galton’s Early Work 11 / 36
  12. 12. Charles H. Pence Galton’s Early Work 12 / 36
  13. 13. L’homme que je considère ici est, dans la société, l’analogue du centre de gravité dans les corps ; il est la moyenne autour de laquelle oscillent les élémens sociaux : ce sera, si l’on veut, un être fictif pour qui toutes les choses se passeront conformément aux résultats moyens obtenus pour la société. Si l’on cherche à établir, en quelque sorte, les bases d’une physique sociale, c’est lui qu’on doit considérer, sans s’arrêter aux cas particuliers ni aux anomalies, et sans rechercher si tel individu peut prendre un développement plus ou moins grand dans l’une de ses facultés. (Quetelet 1835, 1:21) Charles H. Pence Galton’s Early Work 13 / 36
  14. 14. Triste condition de l’espèce humaine ! Nous pouvons énumérer d’avance combien d’individus souilleront leur mains du sang de leurs semblables, combien seront faussaires, combien empoisonneurs, à peu près comme on peut énumérer d’avance les naissances et les décès qui doivent avoir lieu. (Quetelet 1835, 1:10) Charles H. Pence Galton’s Early Work 14 / 36
  15. 15. I know of scarcely anything so apt to impress the imagination as the wonderful form of cosmic order expressed by the “law of error.” A savage, if he could understand it, would worship it as a god. It reigns with serenity in complete self-effacement amidst the wildest confusion. The huger the mob and the greater the anarchy the more perfect its sway. (Galton 1886, 494–5) Charles H. Pence Galton’s Early Work 15 / 36
  16. 16. Charles H. Pence Galton’s Early Work 16 / 36
  17. 17. The outline of my problem of this evening is, that since the characteristics of all plants and animals tend to conform to the law of deviation, let us suppose a typical case, in which the conformity shall be exact, and which shall admit of discussion as a mathematical problem, and find what the laws of heredity must then be to enable successive generations to maintain statistical identity. (Galton 1877a, 493) Charles H. Pence Galton’s Early Work 17 / 36
  18. 18. Charles H. Pence Galton’s Early Work 18 / 36
  19. 19. [I]n addition to [the constant effects of the mean and gravity] there were a host of petty disturbing influences, represented by the spikes among which the pellets tumbled in all sorts of ways. The theory of combination shows that the commonest case is that where a pellet falls equally often to the right of a spike as to the left of it.... It also shows that the cases are very rare of runs of luck carrying the pellet much oftener to one side than the other. (Galton 1877a, 495) Charles H. Pence Galton’s Early Work 19 / 36
  20. 20. Charles H. Pence Galton’s Early Work 20 / 36
  21. 21. [A]lthough characteristics of plants and animals conform to the law, the reason of their doing so is as yet totally unexplained. The essence of the law is that differences should be wholly due to the collective actions of a host of independent petty influences in various combinations.... Now the processes of heredity...are not petty influences, but very important ones. (Galton 1877b, 512) Charles H. Pence Galton’s Early Work 21 / 36
  22. 22. Natural Inheritance Charles H. Pence Natural Inheritance 22 / 36
  23. 23. It would seem that while the embryo is developing itself, the particles more or less qualified for each new post wait as it were in competition, to obtain it. Also that the particle that succeeds, must owe its success partly to accident of position and partly to being better qualified than any equally well placed competitor to gain a lodgement. (Galton 1889, 9) Charles H. Pence Natural Inheritance 23 / 36
  24. 24. The child inherits partly from his parents, partly from his ancestry. Speaking generally, the further his genealogy goes back, the more numerous and varied will his ancestry become, until they cease to differ from any equally numerous sample taken at haphazard from the race at large. Their mean stature will then be the same as that of the race; in other words, it will be mediocre. Or, to put the same fact into another form, the most probable value of the mid-ancestral deviates in any remote generation is zero. (Galton 1885, 1209) Charles H. Pence Natural Inheritance 24 / 36
  25. 25. [The first chapter’s] intention has been to show the large part that is always played by chance in the course of hereditary transmission, and to establish the importance of an intelligent use of the laws of chance and of the statistical methods that are based upon them, in expressing the conditions under which heredity acts. (Galton 1889, 17) Charles H. Pence Natural Inheritance 25 / 36
  26. 26. A parade of great accuracy is foolish, because precision is unattainable in biological and social statistics; their results being never strictly constant. [...] We require no more than a fairly just and comprehensive method of expressing the way in which each measurable quality is distributed among the members of any group.... (Galton 1889, 36) Charles H. Pence Natural Inheritance 26 / 36
  27. 27. Evolution may produce an altogether new type of vessel that shall be more efficient than the old one, but when a particular type of vessel has become adapted to its functions through long experience it is not possible to produce a mere variety of its type that shall have increased efficiency in some one particular without the rest. So it is with animals. (Galton 1889, 124) Charles H. Pence Natural Inheritance 27 / 36
  28. 28. A New Research Program Charles H. Pence A Research Program for Biometry 28 / 36
  29. 29. Charles H. Pence A Research Program for Biometry 29 / 36
  30. 30. Charles H. Pence A Research Program for Biometry 30 / 36
  31. 31. Charles H. Pence A Research Program for Biometry 31 / 36
  32. 32. heredity and deep ancestry Charles H. Pence A Research Program for Biometry 32 / 36
  33. 33. heredity and deep ancestry evolutionary dynamics Charles H. Pence A Research Program for Biometry 32 / 36
  34. 34. Two Perennial Questions Charles H. Pence Two Perennial Questions 33 / 36
  35. 35. What is the relationship between the statistical trends described by biological theory and the processes in the world those trends are intended to describe? Charles H. Pence Two Perennial Questions 34 / 36
  36. 36. How can we express evolutionary dynamics in statistical terms? Charles H. Pence Two Perennial Questions 35 / 36
  37. 37. Questions? charles@charlespence.net https://charlespence.net @pencechp

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