The Ac-st2 element of maize exhibits a positive dosage effect and epigenetic regulation

A novel derivative of the maize transposable element Ac, termed Ac-st2, that displays a positive dosage effect in maize has been identified. Although identical in sequence to other Ac elements, increasing the copy number of the element in the endosperm results in earlier and more frequent Ds excision. Ac-st2 autonomously transposes and catalyzes somatic excision of Ds elements. Germinal transpositions of either Ac-st2 or Ds, however, were not observed. The Ac-st2 phenotype includes a reduction in Ac transcript accumulation that is associated with increased methylation at specific sites in the promoter region of the major transcriptional start site within Ac (ORFa). This element differs from metastable (cycling) Ac derivatives in that Ac-st2 conditions a uniform transposition pattern throughout endosperm and plant development. Ac-st2 undergoes frequent increases in activity after its association with an active Ac element. This change in activity correlates with reduced levels of methylation in the ORFa promoter region. Using a competitive PCR assay, Ac transcript accumulation was followed through endosperm development. From these data, a model is proposed to explain the patterns of variegation associated with both "wild type" active Ac and Ac-st2 elements.