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Encyclopedia of Evolution.pdf - Online Reading Center

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Fisher, R. A. (1890–1962) British Mathematician, Geneticist<br />

Sir R. A. Fisher paved the way for the acceptance <strong>of</strong><br />

natural selection as the mechanism <strong>of</strong> evolution and helped<br />

to make the modern synthesis possible.<br />

Born February 17, 1890, Ronald Aylmer Fisher was a<br />

mathematical prodigy. He was accustomed to solitude, both<br />

because <strong>of</strong> his upbringing and because he was extremely<br />

nearsighted. Because his doctor had forbidden him to read by<br />

electric light, he had to receive his mathematics tutoring at<br />

Harrow School without the aid <strong>of</strong> pencil, paper, and blackboard.<br />

He developed the ability to work out complex mathematical<br />

solutions in his head.<br />

He began to attend Cambridge University in 1909, the<br />

centennial <strong>of</strong> Darwin’s birth and the fiftieth anniversary<br />

<strong>of</strong> the publication <strong>of</strong> his most famous work (see Darwin,<br />

Charles; origin <strong>of</strong> species [book]). Despite the ceremonies<br />

that attended the anniversary, Darwin’s theories did not<br />

meet with widespread acceptance at that time. Nevertheless,<br />

young Fisher was strongly influenced by them.<br />

Fisher was even more strongly influenced by the recent<br />

rediscovery <strong>of</strong> Mendelian genetics, which were being<br />

investigated by William Bateson, geneticist at Cambridge.<br />

Both from his studies <strong>of</strong> Mendelian inheritance patterns, and<br />

from his association with the statistician Karl Pearson, Fisher<br />

became well-grounded in the study <strong>of</strong> statistics. His 1927 book<br />

Statistical Methods for Research Workers was a landmark in<br />

the history <strong>of</strong> scientific research methods, and he eventually<br />

became one <strong>of</strong> the giants <strong>of</strong> statistical theory. He developed<br />

the variance ratio, one <strong>of</strong> the most widely used statistical<br />

parameters in the world today. This ratio in effect expressed<br />

the variation in a sample that could be explained relative to<br />

the proportion that could not be explained. He developed a<br />

probability table that showed how likely the results were to<br />

be significant, depending on the sample size and the number<br />

<strong>of</strong> categories. For example, a variance ratio <strong>of</strong> 3 (three times<br />

as much variation explained as unexplained) might be credible<br />

for a large sample size but not for a small one. As a result <strong>of</strong><br />

F<br />

Fisher’s table, scientists no longer need to say that “the results<br />

appear to not be due to chance” but can say something like<br />

“the probability that the results occurred by chance are less<br />

than one percent.” Modern scientific research, and its application<br />

in fields as diverse as space travel, industrial quality control,<br />

and the control <strong>of</strong> diseases, would scarcely be possible<br />

without the statistical tools pioneered by Fisher. The variance<br />

ratio is called the F ratio, after Fisher.<br />

Fisher was one <strong>of</strong> the founding fathers <strong>of</strong> population<br />

genetics (see also Haldane, J. B. S.; Wright, Sewall).<br />

Fisher demonstrated mathematically how Mendelian genetics,<br />

which had previously been considered anti-Darwinian,<br />

fit together with Darwinian natural selection, as proclaimed<br />

in the title <strong>of</strong> his 1930 book The Genetical Theory <strong>of</strong> Natural<br />

Selection. The concept <strong>of</strong> “fitness,” which had previously<br />

been vague, was given a precise mathematical definition. His<br />

equations made some oversimplifications—for example, he<br />

assumed that each gene had an independent effect on the phenotype,<br />

rather than interacting with other genes—but were<br />

more advanced than anything that had gone before. His equations<br />

also showed that even the smallest fitness advantages,<br />

over enough time, allowed natural selection to produce big<br />

effects. At the same time he demonstrated that it was small<br />

mutations, not large ones, that caused evolutionary change in<br />

populations.<br />

Fisher’s interest in genetics spilled over into a passion<br />

for eugenics, especially from his association with Charles<br />

Darwin’s son, Major Leonard Darwin. Fisher came to believe<br />

that natural selection was weakened as a result <strong>of</strong> the comforts<br />

<strong>of</strong> civilization, that the genes <strong>of</strong> Englishmen were<br />

becoming weaker, which he believed was the reason that<br />

the British Empire was in decline. One <strong>of</strong> the founders <strong>of</strong><br />

eugenics was Darwin’s cousin (see Galton, Francis), who<br />

was also an expert in statistics, which made it appeal even<br />

more to Fisher. Fisher noted with alarm that the upper class<br />

in British society had fewer <strong>of</strong>fspring than the lower classes.<br />

He put his eugenic theories (with himself at the top, despite

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