The b locus of Zea mays encodes a transcriptional activator of the ant
hocyanin biosynthetic pathway. The B-Peru allele is expressed in the a
leurone layer of the seed, which results in dark purple pigmentation o
f this tissue. An unstable Mutator-induced B-Peru mutant allele, b-Per
um220, displays weak, variable pigment and a high germinal reversion r
ate not characteristic of other Mutator insertions. Characterization o
f relevant regions of b-Perum220 revealed a Mu2 element insertion in o
ne copy of a 534 bp sequence. This 534 bp sequence is tandemly triplic
ated in the progenitor B-Peru allele, upstream of the B-Peru transcrip
tion start site. In addition to the Mu2 insertion, the b-Perum220 alle
le contains a newly formed large tandem duplication of 4.0 kb, which i
ncludes the promoter region and the first three exons of the B-Peru ge
ne. The Mu2 element does not reside at any of the duplication breakpoi
nts. The molecular study of eleven independent germinal revertants rev
ealed five structural classes including structures in which the 4.0 kb
tandem duplication is partially or completely deleted, the Mu2 elemen
t is partially or completely deleted, or a combination of these events
has occurred. We hypothesize that most of the revertants arose by une
qual recombination between the duplicated regions. Based on these stru
ctural analyses, models are discussed to explain the reduced b gene ex
pression in b-Perum220.