However, it was later shown that they were engaged without an enclosing membrane ( Yanagawa et al., 1995 Schwarz-Romond et al., 2005 Kim W. For example, the Dvl protein family was initially assumed to transmit the Wnt signals in response to the binding of extracellular Wnt depending on membrane-bound vesicles. Moreover, the membrane-less compartments in the cytoplasm are involved in cellular signal transduction. Additionally, these structures and organelles are usually composed of macromolecules such as proteins and RNA and are in many cases known as ribonucleoprotein (RNP) granules ( Anderson and Kedersha, 2009). Notably, membrane-less organelles such as the nucleolus, have been known since the 1830s when the structure of the cell nucleus was first described ( Wagner and Barry, 1836 David, 1964). Such organelles include nuclear structures like nuclear paraspeckles, the nucleolus as well as Cajal bodies and cytoplasmic organelles such as the P-bodies, stress granules and centrosomes. Apart from the traditional membrane-enclosed organelles, such as the mitochondria and endoplasmic reticulum, many organelles do not have membranous structures yet remain compartmentalized and can concentrate certain types of molecules ( Handwerger and Gall, 2006 Mao et al., 2011a Pederson, 2011). Intracellular components are compartmentalized, respectively, in living cells to facilitate the regulation of cellular activities in time and space. Additionally, the review briefly introduces the basic rules of LLPS, and cellular signaling that potentially plays a role in cancer, including pathways relevant to immune responses and autophagy. Therefore, the present review summarizes the recent progress made on the existing roles of LLPS in cancer, including cancer-related signaling pathways, transcription regulation and maintenance of genome stability. While the basic rules driving LLPS are continuously being uncovered, their function in cellular activities is still unclear, especially within a pathological context. The studies also showed how these principles determine the molecular properties, LLPS behavior, and composition of liquid condensates. Previous studies highlighted the biophysical and biochemical principles underlying the formation of these liquid condensates. In addition, typical membrane-less condensates such as nuclear speckles, stress granules and cell signaling clusters play important roles in various cellular activities, including regulation of transcription, cellular stress response and signal transduction. It drives the formation of liquid-like condensates containing biomolecules in the absence of membrane structures in living cells. Liquid–liquid Phase Separation (LLPS) of proteins and nucleic acids has emerged as a new paradigm in the study of cellular activities. 6Shulan (Hangzhou) Hospital Affiliated to Shulan International Medical College, Zhejiang Shuren University, Hangzhou, China.5Organ Transplantation Institute, Zhejiang University, Hangzhou, China.4Key Laboratory of Organ Transplantation, Research Center for Diagnosis and Treatment of Hepatobiliary Diseases, Hangzhou, China.3Key Laboratory of the Diagnosis and Treatment of Organ Transplantation, Research Unit of Collaborative Diagnosis and Treatment For Hepatobiliary and Pancreatic Cancer, Chinese Academy of Medical Sciences, Hangzhou, China.2NHC Key Laboratory of Combined Multi-Organ Transplantation, Hangzhou, China.1Division of Hepatobiliary and Pancreatic Surgery, Department of Surgery, First Affiliated Hospital, School of Medicine, Zhejiang University, Hangzhou, China.
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