摘要
The nucleolus, a well-known membraneless organelle and the largest substructure with the highest transcriptional activity in the nucleus, was discovered over 200 years ago and is best known for its critical role in ribosome biogenesis(Kresoja-Rakic and Santoro, 2019;Navarro, 2019). Previous studies have reported that the nucleolus comprises three radially distributed layers known as the fibrillar center (FC).