Maize Transposable Elements
Barbara McClintock originally proposed that transposable elements explained
the results of genetic analysis of variegated sectors
of maize aleurone. The aleurone layer is part of the
endosperm and is 3n due to double fertilization. Dosages
affects exist in this 3n tissue because the female parent
can contribute zero to four copies of an allele in 3n
tissue. The female parent contributes 2n and the male
parent contributes 1n.
Initially McClintock studied the bridge-break-fusion
cycle in an abnormal chromosome 9 of maize. This bridge-break-fusion
cycle was due to a chromosome that was dicentric. Dicentric
chromosomes have two centromeres. The dicentric chromosome
resulted in variegated sectors of purple and yellow
aleurone tissue in the same kernel of maize. Later she
discovered that the Ds allele, located on chromosome
9 could cause chromosome breakage. The Ds allele can
initiate the bridge-break-fusion cycle on a chromosome
that initially had only one centromere. As McClintock
did a more in-depth genetic analysis, she discovered
that a second element was required to cause chromosome
breakage.
The Ac or Activator element was necessary to activate the Ds or Dissociation
element. When the Ac element was absent from the nucleus, the Ds element
was not activated. The Ds element induces a breakage at a point on the
chromosome adjacent to its location. McClintock later discovered that
both the Ds and Ac elements can relocate in the genome to either a different
site on the same chromosome or to a non-homologous chromosome. Ds moves
to a different site only when Ac is present in the genome. Ac is an
autonomous element because it can move on its own initiative. Either
the Ds or the Ac element can be transposed to a location within a gene,
the effect is the same as a gene mutation to a different allelic form.