The Chromosomal Theory of Inheritance
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The chromosomal theory of inheritance, proposed by Walter Sutton and Theodor Boveri in 1902, holds that Mendel's hereditary factors are located on chromosomes.
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Sutton observed that chromosomes separate into gametes exactly as Mendel's factors must, one from each homologous pair.
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The theory predates any direct proof a gene sits on a chromosome — Morgan's crosses supplied that evidence a decade later.