Bar-Peled, L. & Kory, N. Principles and functions of metabolic compartmentalization. Nat. Metab. 4, 1232–1244 (2022).

Article  PubMed  PubMed Central  Google Scholar 

Zhao, Y. G. & Zhang, H. Phase separation in membrane biology: the interplay between membrane-bound organelles and membraneless condensates. Dev. Cell 55, 30–44 (2020).

Article  CAS  PubMed  Google Scholar 

Lyon, A. S., Peeples, W. B. & Rosen, M. K. A framework for understanding the functions of biomolecular condensates across scales. Nat. Rev. Mol. Cell Biol. 22, 215–235 (2021).

Article  CAS  PubMed  Google Scholar 

Shin, Y. & Brangwynne, C. P. Liquid phase condensation in cell physiology and disease. Science 357, eaaf4382 (2017).

Article  PubMed  Google Scholar 

Villegas, J. A., Heidenreich, M. & Levy, E. D. Molecular and environmental determinants of biomolecular condensate formation. Nat. Chem. Biol. 18, 1319–1329 (2022).

Article  CAS  PubMed  Google Scholar 

Samir, P. et al. DDX3X acts as a live-or-die checkpoint in stressed cells by regulating NLRP3 inflammasome. Nature 573, 590–594 (2019).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Hirose, T., Ninomiya, K., Nakagawa, S. & Yamazaki, T. A guide to membraneless organelles and their various roles in gene regulation. Nat. Rev. Mol. Cell Biol. 24, 288–304 (2023).

Article  CAS  PubMed  Google Scholar 

Welles, R. M. et al. Determinants that enable disordered protein assembly into discrete condensed phases. Nat. Chem. 16, 1062–1072 (2024).

Article  CAS  PubMed  Google Scholar 

Ye, S. et al. Micropolarity governs the structural organization of biomolecular condensates. Nat. Chem. Biol. 20, 443–451 (2024).

Article  CAS  PubMed  Google Scholar 

Choi, S., Meyer, M. O., Bevilacqua, P. C. & Keating, C. D. Phase-specific RNA accumulation and duplex thermodynamics in multiphase coacervate models for membraneless organelles. Nat. Chem. 14, 1110–1117 (2022).

Article  CAS  PubMed  Google Scholar 

Mu, W. et al. Superstructural ordering in self-sorting coacervate-based protocell networks. Nat. Chem. 16, 158–167 (2024).

Article  CAS  PubMed  Google Scholar 

Qi, C. et al. Multicompartmental coacervate-based protocell by spontaneous droplet evaporation. Nat. Commun. 15, 1107 (2024).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Kaur, T. et al. Sequence-encoded and composition-dependent protein–RNA interactions control multiphasic condensate morphologies. Nat. Commun. 12, 872 (2021).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Zhao, Q. H., Cao, F. H., Luo, Z. H., Huck, W. T. S. & Deng, N. N. Photoswitchable molecular communication between programmable DNA-based artificial membraneless organelles. Angew. Chem. Int. Ed. Engl. 61, e202117500 (2022).

Article  CAS  PubMed  Google Scholar 

Mashima, T. et al. DNA-mediated protein shuttling between coacervate-based artificial cells. Angew. Chem. Int. Ed. Engl. 61, e202115041 (2022).

Article  CAS  PubMed  PubMed Central  Google Scholar 

van Veldhuisen, T. W., Verwiel, M. A. M., Novosedlik, S., Brunsveld, L. & van Hest, J. C. M. Competitive protein recruitment in artificial cells. Commun. Chem. 7, 148 (2024).

Article  PubMed  PubMed Central  Google Scholar 

Silverman, A. D., Karim, A. S. & Jewett, M. C. Cell-free gene expression: an expanded repertoire of applications. Nat. Rev. Genet. 21, 151–170 (2020).

Article  CAS  PubMed  Google Scholar 

Garenne, D. & Noireaux, V. Cell-free transcription–translation: engineering biology from the nanometer to the millimeter scale. Curr. Opin. Biotechnol. 58, 19–27 (2019).

Article  CAS  PubMed  Google Scholar 

Holmes, J. A. et al. Caulobacter PopZ forms an intrinsically disordered hub in organizing bacterial cell poles. Proc. Natl Acad. Sci. USA 113, 12490–12495 (2016).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Lasker, K. et al. Selective sequestration of signalling proteins in a membraneless organelle reinforces the spatial regulation of asymmetry in Caulobacter crescentus. Nat. Microbiol. 5, 418–429 (2020).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Wei, S. P. et al. Formation and functionalization of membraneless compartments in Escherichia coli. Nat. Chem. Biol. 16, 1143–1148 (2020).

Article  CAS  PubMed  Google Scholar 

Farag, M., Borcherds, W. M., Bremer, A., Mittag, T. & Pappu, R. V. Phase separation of protein mixtures is driven by the interplay of homotypic and heterotypic interactions. Nat. Commun. 14, 5527 (2023).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Ebersbach, G., Briegel, A., Jensen, G. J. & Jacobs-Wagner, C. A self-associating protein critical for chromosome attachment, division, and polar organization in Caulobacter. Cell 134, 956–968 (2008).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Bowman, G. R. et al. A polymeric protein anchors the chromosomal origin/ParB complex at a bacterial cell pole. Cell 134, 945–955 (2008).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Lasker, K. et al. The material properties of a bacterial-derived biomolecular condensate tune biological function in natural and synthetic systems. Nat. Commun. 13, 5643 (2022).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Dzuricky, M., Rogers, B. A., Shahid, A., Cremer, P. S. & Chilkoti, A. De novo engineering of intracellular condensates using artificial disordered proteins. Nat. Chem. 12, 814–825 (2020).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Kedersha, N. et al. Stress granules and processing bodies are dynamically linked sites of mRNP remodeling. J. Cell Biol. 169, 871–884 (2005).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Courchaine, E. M. et al. DMA–tudor interaction modules control the specificity of in vivo condensates. Cell 184, 3612–3625 (2021).

Article  CAS  PubMed  PubMed Central  Google Scholar 

McNaughton, B. R., Cronican, J. J., Thompson, D. B. & Liu, D. R. Mammalian cell penetration, siRNA transfection, and DNA transfection by supercharged proteins. Proc. Natl Acad. Sci. USA 106, 6111–6116 (2009).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Thompson, D. B., Villaseñor, R., Dorr, B. M., Zerial, M. & Liu, D. R. Cellular uptake mechanisms and endosomal trafficking of supercharged proteins. Chem. Biol. 19, 831–843 (2012).

Article