Title:
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Evolutionary and Functional Analysis of TMAC: A Drosophila Transcriptional Regulatory Complex
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The testis meiotic arrest genes regulate transcription of genes required for
entry into meiosis and spermatid differentiation. This is achieved by the
formation of the multisubunit testis meiotic arrest complex, TMAC. Two
components of this complex Mip40 and Caf1 are also found in the Drosophila
Rb, E2F and Myb (dREAM) complex which acts to regulate transcription of
differentiation genes in the soma and female germline.
I have shown that two TMAC proteins have paralogues in the dREAM
complex; aly/mip130 and tomb/mip120. A further dREAM complex
component, Iin52 is a paralogue of the testis specific gene wuc/CG12442. I
have investigated the timings of these three duplications in diptera evolution
and propose a model for the stepwise evolution of the complexes, followed by
the repeated subfunctionalisation from the ancestral ubiquitous expression to
one testis specific complex and one complex functioning in the soma and
female germline.
In addition to wuc being paralogous to Iin52 it has also been found to bind the
TMAC protein Aly by yeast-two-hybrid. Wuc protein localises to chromatin in
primary spermatocytes but in a tomb (TMAC, DNA-binding protein) mutant
background this specific localisation. is lost.
I have investigated wuc's RNAi phenotype in the testis and flies are male
sterile. Some meiosis occurred, but multiple chromosome segregation defects
and no signs of differentiation/elongation were observed. Primary
spermatocytes also accumulated an unidentified phase dark granular material
in the cytoplasm and homologous chromosome pairing was disrupted. The
expression of multiple TMAC target genes are also decreased in wuc RNAi
testes which with all the above evidence suggests wuc is a potential new
meiotic arrest gene.
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