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In Pursuit of the Gene

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CELL BIOLOGY © 153<br />

whose lifetime ambition was to establish <strong>the</strong> facts <strong>of</strong> heredity experimentally.<br />

The 22-year-old Boveri began work in Richard Hertwig’s Zoological<br />

<strong>In</strong>stitute in Munich in 1885, just at <strong>the</strong> moment when <strong>the</strong> idea that <strong>the</strong> hereditary<br />

material was located in <strong>the</strong> as-yet-unnamed chromosomes began<br />

to gain currency. Through Hertwig, Boveri received <strong>the</strong> ideal introduction<br />

to cytology, and in particular to <strong>the</strong> study <strong>of</strong> <strong>the</strong> nucleus, which was rapidly<br />

becoming a subfield <strong>of</strong> its own. Deeply impressed by van Beneden’s results,<br />

Boveri spent his first several years in Hertwig’s institute repeating van<br />

Beneden’s work on egg cell formation in Ascaris. 25 <strong>In</strong> January 1888, Boveri<br />

visited Naples, where as fate would have it he was a guest in <strong>the</strong> same<br />

pensione where Weismann was staying.<br />

Although few men were better prepared than Boveri to accept Weismann’s<br />

views on <strong>the</strong> role <strong>of</strong> <strong>the</strong> reduction division in <strong>the</strong> formation <strong>of</strong> sex<br />

cells, Boveri and Weismann had a fundamental disagreement over <strong>the</strong> nature<br />

<strong>of</strong> <strong>the</strong> chromosomes. Whereas Weismann believed that <strong>the</strong> chromosomes<br />

were all essentially identical, each containing multiple copies <strong>of</strong> “ancestral<br />

germ-plasms,” Boveri’s work on <strong>the</strong> chromosomes in <strong>the</strong> egg cells <strong>of</strong><br />

Ascari had convinced him that <strong>the</strong> individual chromosomes differed in form<br />

and function, and each evening after returning from <strong>the</strong> Zoological Station<br />

<strong>the</strong> two men argued. 26 That spring, Boveri began to collect a variety <strong>of</strong> marine<br />

forms from <strong>the</strong> Sea <strong>of</strong> Naples and made detailed renderings <strong>of</strong> carefully<br />

prepared chromosome arrays isolated from <strong>the</strong>ir sex cells, documenting<br />

<strong>the</strong> differences in <strong>the</strong> sizes and shapes <strong>of</strong> <strong>the</strong> individual chromosomes, as<br />

well as <strong>the</strong> way <strong>the</strong>y changed from one generation to <strong>the</strong> next. He soon<br />

found that in each case <strong>the</strong> sperm and egg cells contained identical sets <strong>of</strong><br />

chromosomes; 27 this was consistent with <strong>the</strong> idea that <strong>the</strong> chromosomes are<br />

<strong>the</strong> repository <strong>of</strong> <strong>the</strong> hereditary material, but it did not yet constitute a<br />

pro<strong>of</strong>. 28<br />

Boveri continued <strong>the</strong> work on chromosomes when he returned to Munich<br />

in April 1888, and later <strong>the</strong> same year he presented <strong>the</strong> fruits <strong>of</strong> this<br />

new line <strong>of</strong> inquiry in <strong>the</strong> second <strong>of</strong> six “Cell Studies,” a unique amalgam <strong>of</strong><br />

genetics, cytology, and embryology that immediately established him as a<br />

world leader in <strong>the</strong> study <strong>of</strong> heredity. <strong>In</strong> this paper Boveri clearly showed<br />

that <strong>the</strong> chromosomes, which disappeared during <strong>the</strong> resting period be-

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