The culture media and meiotic time course were essentially performed as previously described (Kim et al., 2010 (link)). Cells were patched to YPG plates (1% yeast extract, 2% peptone, 3% glycerol, and 2% bactoagar) for 24 h at 30°C. To select single colonies, cells from the YPG plate were streaked onto YPD plates (1% yeast extract, 2% peptone, 2% glucose, and 2% bactoagar) and grown at 30°C for 2 days. A single diploid colony resulting from this streaking was inoculated into 2 ml liquid YPD medium (1% yeast extract, 2% peptone, 2% glucose) and incubated at 30°C for 24 h. For synchronous meiosis, YPD cultures were inoculated in SPS medium (1% potassium acetate, 1% bactopeptone, 0.5% yeast extract, 0.17% yeast nitrogen without amino acids, 0.5% ammonium sulfate, 0.05 M potassium biphthalate, and 2 drops/L antifoam [Sigma, USA], pH 5.5) at a 1:500 dilution and cultured for 18 h. Meiosis was initiated in SPM medium (0.2% potassium acetate, 0.02% raffi-nose, and 2 drops/L antifoam). Meiotic cells were harvested and resuspended in 50 mM Tris-HCl and 50 mM EDTA. Cross-linking of cells was performed with psoralen under UV light for 10 min.