On the origin of metacentric, attached-X (A-X) chromosomes in Drosophila melanogaster males

We describe here the isolation and cytogenetic characterization of a mutation inseparabile which generates in males a high frequency of A-X females. The mutation, segregating in low frequency in a laboratory stock, maps to cytological location 82F7–11 in the third chromosome. The mutation acts premeiotically in the male germ line. Disrupting the X chromosome centromeric heterochromatin suppresses the formation of A-X chromosome, implying that the mutation is involved in chromatid cohesion. The inseparabile mutation also affects disjunction of the chromosome 4 in males. We suspect that the mutation was responsible for the original A-X female found by L. V. Morgan in 1921.