Chapter 23 : Transposable Elements of Maize

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The two-component system is the most aggressive DNA transposon yet characterized. Even PCR strategies based on the most highly conserved sequences of the terminal inverted repeats (TIRs) may not recover all the Mu elements in maize or other species. The two-element system was recovered independently as Cy/rcy; this element system yielded the same high frequency of late somatic excisions as Mutator, but the regulator exhibited near- Mendelian segregation. Autoregulation of the TIR promoters by MuDR encoded products has been assessed by examining TIRB-GUS and TIRB-luciferase expression in Mutator and non-Mutator plants. Effects are weak and may be best explained by the ability of MURA to interfere with host methylation of the promoter regions. Transgenes are also subject to epigenetic silencing, but independently of Mutator silencing. In one’s view, the key to understanding Mutator activities requires elucidating which MuDR-encoded products are necessary and sufficient for each component of somatic and germinal activities. Clarification of which MURA and MURB products are required for somatic insertion and the insertional events in germinal cells will be the foundation for testing models proposed for the control of transposition outcome. If different MUR proteins are required for specific activities, then regulation of protein production and/or localization will be the key research area. On the other hand, if the same MUR proteins conduct all the biochemistry, then their posttranslational modification or the changing contributions of host proteins will be the focus of future analysis.

Citation: Walbot V, Rudenko G. 2002. Transposable Elements of Maize, p 533-564. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch23

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Image of Figure 1.
Figure 1.

Structure of and the nonautonomous element. The shared TIRs are shown in black, and the unrelated internal sequences are crosshatched in and open in Arrows above the elements indicate the orientation, length, and position of short, conserved repeats found in some elements. element is composed of two genes, and separated by a repeat-rich intergenic region (hatched box).

Citation: Walbot V, Rudenko G. 2002. Transposable Elements of Maize, p 533-564. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch23
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Image of Figure 2.
Figure 2.

Alignment of the TIRs of the mobile elements. Terminal inverted repeat sequences of the autonomous element are shown on the top. MuDRA is the TIR next to and MuDRB is the TIR next to ; the few mismatches between these TIRs are shown as shadowed letters. The positions of the presumptive CCAAT and TATA boxes and the transcription initiation start sites for both and are indicated with white letters on a black field. In the alignment of TIRs from nonautonomous elements, dots represent matching bases and dashes represent missing bases. Regions with assigned biological function and sequence motifs discussed in the text are boxed.

Citation: Walbot V, Rudenko G. 2002. Transposable Elements of Maize, p 533-564. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch23
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Image of Figure 3.
Figure 3.

Phylogenetic analysis of elements. Unrooted phylograms were generated by using PAUP to estimate relatedness between the virtual translation of hMUR proteins and the -encoded proteins. (Left) N-terminal part of hMURA, including exon 2. (Right) Full-length hMURB proteins and their wild-type counterparts.

Citation: Walbot V, Rudenko G. 2002. Transposable Elements of Maize, p 533-564. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch23
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Image of Figure 4.
Figure 4.

Major transcripts and predicted proteins encoded by (A) Transcriptional units in Heavy arrows depict the exon (filled) and intron (open) organization and direction of transcription of the and genes. Transcription initiates in the terminal inverted repeats of the element,as indicated by bent arrows above the element diagram, and terminates in the intergenic region. The ATG initiation codons discussed in the text are indicated above the diagram for and below the diagram for The position of the stop codon for the longest transcript from each gene is similarly indicated. (B) Major transcripts. Alternative transcription initiation start sites for and alternative splicing of the introns in pre-mRNA for both genes generate a variety of transcripts. (C) Major predicted proteins. Alternative splicing and two possible translation initiation start sites in result in multiple predicted proteins encoded by the known transcripts. MURA proteins contain a region of similarity to bacterial transposases and retroviral integrases,several functional nuclear localization sequences,and a zinc knuckle domain. Alternative splicing results in four predicted MURB proteins encoded by the characterized transcripts of

Citation: Walbot V, Rudenko G. 2002. Transposable Elements of Maize, p 533-564. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch23
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Image of Figure 5.
Figure 5.

Models for developmental timing and outcome of excision. (A) Double-stranded gap repair masks early and germinal excision. Excision of a element results in a double-stranded gap (1). In germinal and early somatic cells (2), which express abundant MURB protein,the gaps that were generated postreplicatively can be repaired by using the copy of the original locus from the sister chromatid. After insertion of the excised element, there is a net increase in the element copy number and restoration of the original insertion site. Evidence of excision is effectively removed. In late somatic cells (3), which express a low amount of MURB, gap repair does not occur, resulting in a measurable excision event. This is true even when the homologous chromosome contains the identical insertion allele. (B) Competition for binding model to explain developmental timing. / elements (1) are accessible for MURA transposase (2). In dividing cells, however, abundant transcriptional factors might assemble on specific motifs in the TIRs (3),potentially displacing transposase or affecting the geometry of the transpososome complex (4). Late in development, these transcriptional factors are depleted (5),allowing assembly of the transpositionally active complex, which results in late excision events (6).

Citation: Walbot V, Rudenko G. 2002. Transposable Elements of Maize, p 533-564. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch23
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Table 1.

Characteristics of elements

Citation: Walbot V, Rudenko G. 2002. Transposable Elements of Maize, p 533-564. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch23
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Table 2.

Summary of sequence differences between and -encoded genes

Citation: Walbot V, Rudenko G. 2002. Transposable Elements of Maize, p 533-564. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch23
Generic image for table
Table 3.

Large predicted protein products

Citation: Walbot V, Rudenko G. 2002. Transposable Elements of Maize, p 533-564. In Craig N, Craigie R, Gellert M, Lambowitz A (ed), Mobile DNA II. ASM Press, Washington, DC. doi: 10.1128/9781555817954.ch23

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