How was it concluded that genes are located on chromosomes ?

Morgan confirmed Mendelian laws of inheritance and the hypothesis that genes are located on chromosomes. Morgan had discovered that eye colour in Drosophila expressed a sex-linked trait. All first-generation offspring of a mutant white-eyed male and a normal red-eyed female would have red eyes because every chromosome pair would contain at least one copy of the $ X$ chromosome with the dominant trait.