Recombination between Components of a Mutable Gene System in Maize.

An unstable component (I-R) of the R-stippled allele interacts with a linked modifier which enhances stippled's expression ( M-st) to delete the intervening segment. The precision of deletion formation suggests a recombinational basis, specifically unequal crossing over between I-R and M-st. Four deletions were selected as losses of R function from plants homozygous for R-st and M-st. Also lost are stippled's near-colorless seed phenotype, its paramutagenicity and a closely linked gene, Inhibitor of striate. The deletion chromosomes, missing a 6-cM segment, are transmitted normally by ovules but in reduced frequency by pollen. Homozygous and heteroallelic combinations of the deletions confer defective seed lethality. The four did not differ detectably in transmissibility or breakpoint termini. The recurrence of deletions that have the same termini is explained by recombination between I-R and M-st. The homology between M-st and I-R, and their presence in the same chromosome arm, favors the view that M-st originated by I-R transposition.