The CLIP–cohibin axis is required for survival during nitrogen starvation. (A) Strains BY4741 (WT), SCU3365 (atg1Δ), SCU3368 (nvj1Δ), SCU4325 (atg39Δ), SCU4419 (heh1Δ), SCU4400 (nur1Δ), SCU2761 (csm1Δ), and SCU4885 (lrs4Δ) were used. Exponentially growing cells were subjected to nitrogen starvation for 7 d. Dead cells were stained with 2 µg/ml phloxine B. Cells stained with phloxine B counted as dead cells in each experiment (>100 cells), and the mean value (± SD) obtained from three independent experiments is shown as a percentage. Significant differences (#, P < 0.001) between mutants and the WT were calculated using two-tailed Student's t test. (B) A cartoon for the movements of nucleolar proteins (nucleoli) and rDNA after TORC1 inactivation. CLIP, cohibin, and nucleophagy are required for this repositioning. (C) A hypothetical model for rDNA and nucleolar protein repositioning after TORC1 inactivation in the context of nucleophagy. The CLIP–cohibin axis is required for the repositioning. Micronucleophagy is necessary not only for nucleolar protein degradation but also for the preceding repositioning. See the text for details.