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In Pursuit of the Gene

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192 ¨ THE FLY ROOM<br />

near one ano<strong>the</strong>r in a linear series along <strong>the</strong> chromosome, adding that “<strong>the</strong><br />

proportions that result are not so much <strong>the</strong> expression <strong>of</strong> a numerical system<br />

[as Bateson had suggested] as <strong>of</strong> <strong>the</strong> relative location <strong>of</strong> <strong>the</strong> factors in<br />

<strong>the</strong> chromosomes.” 21<br />

Sturtevant took <strong>the</strong> next big step later that year. 22 Half a century later,<br />

he still vividly recalled <strong>the</strong> afternoon that he hit upon what was without<br />

doubt <strong>the</strong> greatest idea <strong>of</strong> his life. He and Morgan had been discussing a<br />

1909 paper by <strong>the</strong> American geneticist W. E. Castle in which Castle seemed<br />

to be suggesting, contrary to <strong>the</strong> inspired deductions <strong>of</strong> <strong>the</strong> great European<br />

cytologist Wilhelm Roux in 1883 and Morgan’s recent elaboration on <strong>the</strong><br />

same <strong>the</strong>me, that genes were not arranged in a line along <strong>the</strong> chromosomes<br />

after all. Morgan and Sturtevant agreed that Castle’s idea was not<br />

worth fur<strong>the</strong>r consideration—<strong>the</strong> genes were almost certainly arranged in<br />

a line along <strong>the</strong> chromosomes, and <strong>the</strong> association between <strong>the</strong>m, as Morgan<br />

had recently pointed out, was a function <strong>of</strong> <strong>the</strong>ir distance apart. It was<br />

<strong>the</strong>n that Sturtevant had his insight, suddenly seeing that <strong>the</strong> ratio <strong>of</strong> crossover<br />

to noncrossover types among <strong>the</strong> progeny could be used as a measure<br />

<strong>of</strong> <strong>the</strong> distances between two genes on a chromosome and that <strong>the</strong> use <strong>of</strong><br />

this measure made possible <strong>the</strong> construction <strong>of</strong> a linear map <strong>of</strong> <strong>the</strong> genes.<br />

While Morgan had been vague about <strong>the</strong> relationship between association<br />

<strong>of</strong> factors and <strong>the</strong>ir placement on <strong>the</strong> chromosome, Sturtevant proposed<br />

a concrete way to quantify <strong>the</strong> distance between genes. Fur<strong>the</strong>rmore,<br />

he claimed that if one knew AB (<strong>the</strong> distance between genes A and B) and<br />

BC (<strong>the</strong> distance between B and C), <strong>the</strong>n AC (<strong>the</strong> distance between A and<br />

C) would equal <strong>the</strong>ir difference or <strong>the</strong>ir sum.<br />

<strong>Gene</strong> mapping. From Hermann J. Muller, “The Mechanism <strong>of</strong> Crossing Over,” American Naturalist 50 (1916): 200.<br />

By applying <strong>the</strong> same tests to strategically chosen combinations <strong>of</strong><br />

genes, he could make a map <strong>of</strong> <strong>the</strong> genes showing <strong>the</strong>ir linear order and

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