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What specific trait in fly studies "demonstrate without doubt" that genes are on chromosomes? How did...

What specific trait in fly studies "demonstrate without doubt" that genes are on chromosomes? How did it show this?

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Sutton and Boveri have given the chromosomal theory of inheritance. This theory states that the hereditary units are present on the chromosomes of the individual as the behavior of the heredity units and chromosomes is similar.

In 1911, Thomas Hunt Morgan observed sex linkage of the eye color in fruit fly. Based on the experimental studies on the inheritance of the eye color, Morgan argued that the genes are on the chromosomes. His student Sturtevant used the phenomena of genetic linkage and showed that genes are linearly arranged on the chromosomes. They confirmed the chromosomal theory of inheritance. the inheritance of the eye color was depending on the sex of the parent. Because the sex is determined by the chromosomes, the gene for eye color must be there on the sex chromosomes. That is how the genes are confirmed to be present on the chromosomes.

His student Alfred Sturtevant has developed method to construct gene map on the chromosomes based on the recombination frequency.

When fruit flies were crossed they found that

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