The transposition of Mu elements underlying Mutator activity in maize requires a transcriptionally active MuDR element. Despite variation in MuDR copy number and RNA levels in Mutator lines, transposition events are consistently late in plant development, and Mu excision frequencies are similar. Here, we report previously unsuspected and ubiquitous MuDR homologs that produce both RNA and protein. MuDR transcript levels are proportional to MuDR copy number, and homolog transcript levels increase in active Mutator lines. A subset of homologs exhibits constitutive transcription in MuDR Ϫ and epigenetically silenced MuDR lines, suggesting independent transcriptional regulation. Surprisingly, immunodetection demonstrated nearly invariant levels of MuDR and homolog protein products in all tested Mutator and non-Mutator stocks. These results suggest a strict control over protein production, which might explain the uniform excision frequency of Mu elements. Moreover, the nonfunctional proteins encoded by homologs may negatively regulate Mutator activity and represent part of the host defense against this transposon family.
INTRODUCTIONOptimized transmission and restricted transposition activity are characteristics of many transposable elements. These features combine to allow efficient transposon proliferation with a minimal cost to the host genome. A variety of regulatory mechanisms operating at both the transcriptional and post-transcriptional levels can result in specific temporal and spatial patterns of transposon activity. Some mechanisms are epigenetic and act through several generations. In plants, increased DNA methylation of promoter regions of autonomous elements such as Ac , En / Spm , and MuDR correlates with decreased production of transposase transcripts and loss of transposition reactions (Fedoroff and Chandler, 1994; Martienssen and Baron, 1994). In some cases, host methylation acts in concert with transposon autoregulation. For example, the TnpA transposase protein encoded by Spm binds to its own unmethylated promoter, an interaction that decreases transcription; in the absence of Spm -encoded products, the promoter becomes methylated, and both transcription and transposition cease (Schläppi et al., 1994). Introduction of a functional Spm element results in activation of the silenced promoter and resumption of transposon activity.More dynamic regulation occurs during a single life cycle to limit the production or function of active transposase proteins.For example, tissue-specific RNA splicing restricts expression of the functional transposase of the Drosophila melanogaster P element to the germ line (Laski et al., 1986;Siebel et al., 1992). In bacteria, the inhibition of bacterial Tn10 transposition occurs through production of element-encoded antisense RNA (Simons and Kleckner, 1988). The maize transposable element Ac exhibits a negative dosage phenomenon: with increasing copy number and RNA levels, there is a lower excision frequency, and transposition events occur later in development (McClintock, 1...