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Transcriptional regulation of meiosis in budding yeast

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Fig. 9. A schematic structure <strong>of</strong> IME1 5’ untranslated region show<strong>in</strong>g the positive and<br />

negative transcription factors which regulate its expression.<br />

The regulated transcription <strong>of</strong> IME1 is mediated by the comb<strong>in</strong>atorial effect <strong>of</strong> dist<strong>in</strong>ct elements.<br />

The MAT signal is transmitted to UCS4 by Rme1. The activity <strong>of</strong> Rme1 as a repressor requires<br />

Rgr1 and S<strong>in</strong>4. The cAMP/PKA signal transduction pathway transmits the glucose signal to IREu<br />

through two transcription factors, Sok2 and Msn2. In glucose growth media phosphorylation <strong>of</strong><br />

Sok2 by PKA promotes its repression activity, whereas phosphorylation <strong>of</strong> Msn2 sequesters the<br />

prote<strong>in</strong> <strong>in</strong> the cytoplasm. In acetate growth media Sok2 repression is relieved by a decrease <strong>in</strong> the<br />

activity <strong>of</strong> PKA, as well as by the function <strong>of</strong> Ime1. The <strong>in</strong>creased b<strong>in</strong>d<strong>in</strong>g <strong>of</strong> Msn2 to IREu leads<br />

to transcriptional activation. Yhp1 b<strong>in</strong>ds to UASv, however, its activity is not required for the<br />

transcription <strong>of</strong> IME1. Nitrogen, via Cdc25, regulates the URS activity <strong>of</strong> UCS1. It is not known<br />

whether the effect <strong>of</strong> Cdc25 is mediated through the cAMP/PKA and/or MAPK pathways.<br />

Filled box - elements required for transcriptional activation. Open box – elements that whose<br />

function is only to repress transcription. A positive role is marked with an arrow, a negative role<br />

<strong>in</strong> l<strong>in</strong>e. Dashed l<strong>in</strong>es and a question mark – prelim<strong>in</strong>ary data.<br />

Fig. 10. The expression and activity <strong>of</strong> Ime1 are regulated by nutrients.<br />

Schematic representation on how glucose and nitrogen regulates the availability and activity <strong>of</strong><br />

Ime1. The nucleus is depicted between the two circular l<strong>in</strong>es, and cell membrane by a heavy l<strong>in</strong>e.<br />

A positive role is marked with an arrow, a negative role by a straight l<strong>in</strong>e. The transcription <strong>of</strong><br />

IME1 is <strong>in</strong>hibited <strong>in</strong> the presence <strong>of</strong> glucose and nitrogen. Translation <strong>of</strong> IME1 mRNA is<br />

<strong>in</strong>hibited <strong>in</strong> the presence <strong>of</strong> a nitrogen source. In the presence <strong>of</strong> nitrogen, Ime1is sequestered<br />

from the nuclei due to phosphorylation by Cdc28/Cln. Interaction <strong>of</strong> Ime1 with Ume6, and<br />

consequently transcriptional activation <strong>of</strong> early <strong>meiosis</strong>-specific genes (EMG) is <strong>in</strong>hibited by<br />

glucose and nitrogen.<br />

Fig. 11. Conditions lead<strong>in</strong>g to the choice between silenc<strong>in</strong>g and expression <strong>of</strong> early <strong>meiosis</strong>specific<br />

genes.<br />

A. Vegetative growth conditions B. Meiotic conditions.<br />

Ume6 b<strong>in</strong>ds to the URS1 element <strong>in</strong> early-<strong>meiosis</strong> specific genes (EMG) under all growth<br />

conditions. In vegetative growth conditions Ume6 recruits to repression complexes, S<strong>in</strong>3/Rpd3<br />

and Isw2, lead<strong>in</strong>g to histone H4 deacetylase and chromat<strong>in</strong> remodel<strong>in</strong>g, respectively, and<br />

silenc<strong>in</strong>g. Under these conditions Rim15 and most probably Rim11 are non-active. The activity<br />

<strong>of</strong> Rim15 is repressed by PKA (Tpk1,2,3) phosphorylation. Under meiotic conditions (acetate<br />

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