Banani SF, Lee HO, Hyman AA, Rosen MK. Biomolecular condensates: organizers of cellular biochemistry. Nat Rev Mol Cell Biol. 2017;18(5):285–98.

CAS  PubMed  PubMed Central  Google Scholar 

Lafontaine DLJ, Riback JA, Bascetin R, Brangwynne CP. The nucleolus as a multiphase liquid condensate. Nat Rev Mol Cell Biol. 2021;22(3):165–82.

CAS  PubMed  Google Scholar 

Xiao Q, McAtee CK, Su X. Phase separation in immune signalling. Nat Rev Immunol. 2022;22(3):188–99.

CAS  PubMed  Google Scholar 

Holehouse AS, Kragelund BB. The molecular basis for cellular function of intrinsically disordered protein regions. Nat Rev Mol Cell Biol. 2024;25(3):187–211.

CAS  PubMed  Google Scholar 

Henninger JE, Oksuz O, Shrinivas K, Sagi I, LeRoy G, Zheng MM, Andrews JO, Zamudio AV, Lazaris C, Hannett NM, et al. RNA-Mediated feedback control of transcriptional condensates. Cell. 2021;184(1):207–e225224.

CAS  PubMed  Google Scholar 

Roden C, Gladfelter AS. RNA contributions to the form and function of biomolecular condensates. Nat Rev Mol Cell Biol. 2021;22(3):183–95.

CAS  PubMed  Google Scholar 

Hess N, Joseph JA. Structured protein domains enter the spotlight: modulators of biomolecular condensate form and function. Trends Biochem Sci. 2025;50(3):206–23.

CAS  PubMed  Google Scholar 

Hyman AA, Weber CA, Jülicher F. Liquid-liquid phase separation in biology. Annu Rev Cell Dev Biol. 2014;30:39–58.

CAS  PubMed  Google Scholar 

Mittag T, Pappu RV. A conceptual framework for understanding phase separation and addressing open questions and challenges. Mol Cell. 2022;82(12):2201–14.

CAS  PubMed  PubMed Central  Google Scholar 

Galvanetto N, Ivanović MT, Chowdhury A, Sottini A, Nüesch MF, Nettels D, Best RB, Schuler B. Extreme dynamics in a biomolecular condensate. Nature. 2023;619(7971):876–83.

CAS  PubMed  PubMed Central  Google Scholar 

Boija A, Klein IA, Sabari BR, Dall’Agnese A, Coffey EL, Zamudio AV, Li CH, Shrinivas K, Manteiga JC, Hannett NM, et al. Transcription factors activate genes through the Phase-Separation capacity of their activation domains. Cell. 2018;175(7):1842–e18551816.

CAS  PubMed  Google Scholar 

Strzyz P. Phase separation tunes signal transduction. Nat Rev Mol Cell Biol. 2019;20(5):263.

CAS  PubMed  Google Scholar 

Su X, Ditlev JA, Hui E, Xing W, Banjade S, Okrut J, King DS, Taunton J, Rosen MK, Vale RD. Phase separation of signaling molecules promotes T cell receptor signal transduction. Science. 2016;352(6285):595–9.

CAS  PubMed  PubMed Central  Google Scholar 

Strom AR, Emelyanov AV, Mir M, Fyodorov DV, Darzacq X, Karpen GH. Phase separation drives heterochromatin domain formation. Nature. 2017;547(7662):241–5.

CAS  PubMed  PubMed Central  Google Scholar 

Povarova OI, Antifeeva IA, Fonin AV, Turoverov KK, Kuznetsova IM. The role of liquid-liquid phase separation in actin polymerization. Int J Mol Sci. 2023. https://doi.org/10.3390/ijms24043281.

PubMed  PubMed Central  Google Scholar 

Yang S, Liu C, Guo Y, Li G, Li D, Yan X, Zhu X. Self-construction of actin networks through phase separation-induced abLIM1 condensates. Proc Natl Acad Sci U S A. 2022;119(29): e2122420119.

CAS  PubMed  PubMed Central  Google Scholar 

Savastano A, Flores D, Kadavath H, Biernat J, Mandelkow E, Zweckstetter M. Disease-associated tau phosphorylation hinders tubulin assembly within tau condensates. Angew Chem Int Ed Engl. 2021;60(2):726–30.

CAS  PubMed  Google Scholar 

Sahu S, Chauhan P, Lumen E, Moody K, Peddireddy K, Mani N, Subramanian R, Robertson-Anderson R, Wolfe AJ, Ross JL. Interplay of self-organization of microtubule asters and crosslinking protein condensates. PNAS Nexus. 2023;2(7):pgad231.

PubMed  PubMed Central  Google Scholar 

Basu A, Krug T, du Pont B, Huang Q, Sun S, Adam SA, Goldman RD, Weitz DA. Vimentin undergoes liquid-liquid phase separation to form droplets which wet and stabilize actin fibers. Proc Natl Acad Sci U S A. 2025;122(10): e2418624122.

CAS  PubMed  Google Scholar 

Quiroz FG, Fiore VF, Levorse J, Polak L, Wong E, Pasolli HA, Fuchs E. Liquid-liquid phase separation drives skin barrier formation. Science. 2020. https://doi.org/10.1126/science.aax9554.

PubMed  PubMed Central  Google Scholar 

Shen M, Astapov D, Fedorov D, Välisalmi T, Linder MB, Aranko AS. Phase separation drives the folding of recombinant collagen. Int J Biol Macromol. 2024;282(Pt 5):137170.

CAS  PubMed  Google Scholar 

Rangachari V. Biomolecular condensates - extant relics or evolving microcompartments? Commun Biol. 2023;6(1):656.

CAS  PubMed  PubMed Central  Google Scholar 

Dasmeh P, Wagner A. Natural selection on the phase-separation properties of FUS during 160 my of mammalian evolution. Mol Biol Evol. 2021;38(3):940–51.

CAS  PubMed  Google Scholar 

Feric M, Misteli T. Phase separation in genome organization across evolution. Trends Cell Biol. 2021;31(8):671–85.

CAS  PubMed  PubMed Central  Google Scholar 

Portz B, Lee BL, Shorter J. FUS and TDP-43 phases in health and disease. Trends Biochem Sci. 2021;46(7):550–63.

CAS  PubMed  PubMed Central  Google Scholar 

Dang M, Wu L, Zhang X. Structural insights and milestones in TDP-43 research: a comprehensive review of its pathological and therapeutic advances. Int J Biol Macromol. 2025;306(Pt 3):141677.

CAS  PubMed  Google Scholar 

Yu Y, Liu Q, Zeng J, Tan Y, Tang Y, Wei G. Multiscale simulations reveal the driving forces of p53C phase separation accelerated by oncogenic mutations. Chem Sci. 2024;15(32):12806–18.

CAS  PubMed  PubMed Central  Google Scholar 

Brangwynne CP, Eckmann CR, Courson DS, Rybarska A, Hoege C, Gharakhani J, Jülicher F, Hyman AA. Germline P granules are liquid droplets that localize by controlled dissolution/condensation. Science. 2009;324(5935):1729–32.

CAS  PubMed  Google Scholar 

Li P, Banjade S, Cheng HC, Kim S, Chen B, Guo L, Llaguno M, Hollingsworth JV, King DS, Banani SF, et al. Phase transitions in the assembly of multivalent signalling proteins. Nature. 2012;483(7389):336–40.

CAS  PubMed  PubMed Central  Google Scholar 

Kato M, Han TW, Xie S, Shi K, Du X, Wu LC, Mirzaei H, Goldsmith EJ, Longgood J, Pei J, et al. Cell-free formation of RNA granules: low complexity sequence domains form dynamic fibers within hydrogels. Cell. 2012;149(4):753–67.

CAS  PubMed  PubMed Central  Google Scholar 

Galganski L, Urbanek MO, Krzyzosiak WJ. Nuclear speckles: molecular organization, biological function and role in disease. Nucleic Acids Res. 2017;45(18):10350–68.

CAS  PubMed  PubMed Central  Google Scholar 

Alexander KA, Coté A, Nguyen SC, Zhang L, Gholamalamdari O, Agudelo-Garcia P, Lin-Shiao E, Tanim KMA, Lim J, Biddle N, et al. p53 mediates target gene association with nuclear speckles for amplified RNA expression. Mol Cell. 2021;81(8):1666–e16811666.

CAS  PubMed  PubMed Central  Google Scholar 

Wang Y, Chen LL. Organization and function of paraspeckles. Essays Biochem. 2020;64(6):875–82.

CAS  PubMed  Google Scholar 

Li Y, Ma X, Wu W, Chen Z, Meng G. PML nuclear body biogenesis, carcinogenesis, and targeted therapy. Trends Cancer. 2020;6(10):889–906.

CAS  PubMed  Google Scholar 

Nsengimana B, Khan FA, Ngowi EE, Zhou X, Jin Y, Jia Y, Wei W, Ji S. Processing body (P-body) and its mediators in cancer. Mol Cell Biochem. 2022;477(4):1217–38.

CAS  PubMed