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In  Saccharomyces cerevisiae , meiosis and sporulation are highly regulated responses that are driven in part by changes in RNA expression. Alternative mRNA forms with extended 5′ UTRs are atypical in  S. cerevisiae , and 5′ extensions with upstream open reading frames (uORFs) are even more unusual. Here we characterize the gene  YPR036W-A , now renamed  SPO24 , which encodes a very small (67-amino-acid) protein. This gene gives rise to two mRNA forms: a shorter form throughout meiosis and a longer, 5′-extended form in mid-late meiosis. The latter form includes a uORF for a 14-amino-acid peptide (Spo24 u14 ). Deletion of the downstream ORF (dORF) leads to sporulation defects and the appearance of pseudohyphae-like projections. Experiments with luciferase reporters indicate that the uORF does not downregulate dORF translation. The protein encoded by the dORF (Spo24 d67 ) localizes to the prospore membrane and is differentially phosphorylated during meiosis. Transcription of the 5′-extended mRNA in mid-meiosis depends upon the presence of two middle sporulation elements (MSEs). Removal of the MSEs severely inhibits the mid-meiotic appearance of the 5′-extended mRNA and limits the ability of plasmid-borne  SPO24  to rescue the sporulation defect of a  spo24 Δ mutant, suggesting that the 5′-extended mRNA is functionally important. These results reveal Spo24 d67  as a sporulation-related factor that is encoded by a transcriptionally dynamic, uORF-containing locus.

SPO24 Is a Transcriptionally Dynamic, Small ORF-Encoding Locus Required for Efficient Sporulation in Saccharomyces cerevisiae