+ = equal author contribution/co‑first author; * = corresponding/co‑corresponding author
Submitted
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Aierken, D.; Aland, S.; Bo, S.; Boeynaems, S.; Cai, D.; Carra, S.; Case, L.; Chan, H.; Espinosa, J.; GrandPre, T.; . Roadmap for Condensates in Cell Biology. arXiv preprint arXiv:2601.03677 2026.
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Hess, N.; Joseph, J. A. Biomolecular Condensates Dictate the Folding Landscape of Proteins. bioRxiv 2026.
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Wani, Y.; Joseph, J. A. Non-Equilibrium Modeling of Directed Flux through Biomolecular Condensates. bioRxiv 2026.
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Pathak, S.; Chakravarti, A.; Bersche, C.; Joseph, J. A.; Strader, L. Molecular Basis for Thermoresponsive Protein Condensation in Plants. bioRxiv 2025.
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Wadsworth+, G.; Aierken+, D.; Thurston, G.; Joseph*, J. A.; Banerjee*, P. The Critical Role of the 2’-OH Group in Phase Separation and Percolation Transitions of RNA. bioRxiv 2025.
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Aierken, D.; Zhang, V.; Sealfon, R.; Marecki, J.; Raney, K.; Gladfelter, A. S.; Joseph*, J. A.; Roden*, C. A. Biomolecular Condensates Control and Are Defined by RNA-RNA Interactions That Arise in Viral Replication. bioRxiv 2024.
2026
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Garcia, P. L.; Joseph*, J. A. Molecular Origins of Viscoelasticity in Biomolecular Condensates. The Journal of Chemical Physics (Editor’s Pick) 2026.
2025
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Alberti, S.; Arosio, P.; Best, R. B.; Boeynaems, S.; Cai, D.; Collepardo-Guevara, R.; Dignon, G. L.; Dimova, R.; Garfinkle, S. E.-; Fawzi, N. L.; Fuxreiter, M.; Gladfelter, A. S.; Honigmann, A.; Jain, A.; Joseph, J. A.; Knowles, T. P. J.; Lasker, K.; Lemke, E. A.; Lindorff-Larsen, K.; Lipowsky, R.; Mittal, J.; Mukhopadhyay, S.; Vecchiarelli, A. G.; Wegmann, S.; Zhang, H.; Zhang, M.; Zubieta, C.; Zweckstetter, M.; Dormann, D.; Mittag, T. Current Practices in the Study of Biomolecular Condensates: A Community Comment. Nature Communications 2025, 16, 7730. https://doi.org/10.1038/s41467-025-62055-8.
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Chakravarti, A.; Joseph*, J. A. Accurate Prediction of Thermoresponsive Phase Behavior of Disordered Proteins. Protein Science 2025, 34, e70284. https://doi.org/10.1002/pro.70284.
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Emelianova, A.; Garcia, P. L.; Tan, D.; Joseph*, J. A. Prediction of Small-Molecule Partitioning into Biomolecular Condensates from Simulation. JACS Au (ACS Editors’ Choice) 2025.
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Hess, N.; Joseph*, J. A. Structured Protein Domains Enter the Spotlight: Modulators of Biomolecular Condensate Form and Function. Trends in Biochemical Sciences 2025, 50, 206–223.
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Luo+, C.; Hess+, N.; Aierken, D.; Qiang, Y.; Joseph*, J. A.; Zwicker*, D. Theory of Condensate Size Control by Molecular Charge Asymmetry. ACS Macro Letters 2025, 14, 1484–1491.
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Maristany, J.; Gonzalez, A.; Espinosa, J.; Huertas, J.; Collepardo-Guevara, R.; Joseph*, J. A. Decoding Phase Separation of Prion-Like Domains through Data-Driven Scaling Laws. eLife 2025, 13, RP99068.
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Maristany, J.; Emelianova, A.; Chew, P. Y.; Aguirre, A.; Collepardo-Guevara, R.; Joseph, J. A. Modeling Biomolecular Condensates across Scales: Atomistic, Coarse-Grained, and Data-Driven Approaches. Advances in Physics: X 2025, 10, 2592547.
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Ng, T.; Hoare, M.; Maristany, J.; Wilde, E.; Sneideris, T.; Huertas, J.; Agbetiameh, B.; Furukawa, M.; Joseph, J. A.; Knowles, T.; Collepardo-Guevara, R.; Itzhaki, L. S.; Kumita, J. R. Tandem-Repeat Proteins Introduce Tuneable Properties to Engineered Biomolecular Condensates. Chemical Science 2025, 16, 10532–10548.
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Tan, D.; Aierken, D.; Garcia, P. L.; Joseph*, J. A. Biomolecular Condensate Microstructure Is Invariant to Sequence-Encoded Molecular and Macroscopic Properties. Soft Matter 2025, 21, 8635–8654.
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Tjo, H.; Jiang, V.; Joseph, J. A.; Conway, J. Maltodextrin Transport in the Extremely Thermophilic, Lignocellulose Degrading Bacterium Anaerocellum Bescii (f. Caldicellulosiruptor Bescii). Journal of Bacteriology 2025, 207, e00401–24.
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Tjo, H.; Jiang, V.; Calvo, A.; Joseph, J. A.; Conway, J. A Highly Conserved ABC Transporter Mediates Cello-Oligosaccharide Uptake in the Extremely Thermophilic, Lignocellulolytic Bacterium Anaerocellum Bescii (f. Caldicellulosiruptor Bescii). Applied and Environmental Microbiology 2025, e01284–25.
2024
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Aierken, D.; Joseph*, J. A. Accelerated Simulations Reveal Physicochemical Factors Governing Stability and Composition of Rna Clusters. Journal of Chemical Theory and Computation 2024, 20, 10209–10222.
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Chew, P. Y.; Joseph, J. A.; Collepardo-Guevara, R.; Reinhardt, A. Aromatic and Arginine Content Drives Multiphasic Condensation of Protein-RNA Mixtures. Biophysical Journal 2024, 123, 1342–1355.
2023
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Chew, P. Y.; Joseph, J. A.; Collepardo-Guevara, R.; Reinhardt, A. Thermodynamic Origins of Two-Component Multiphase Condensates of Proteins. Chemical Science 2023, 14, 1820–1836.
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Nicy, N.; Collepardo-Guevara, R.; Joseph*, J. A.; Wales*, D. J. Energy Landscapes and Heat Capacity Signatures for Peptides Correlate With Phase Separation Propensity. QRB discovery 2023, 4, e7.
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Zhu+, H.; Narita+, M.; Joseph+, J.; Krainer+, G.; Arter, W.; Olan, I.; Saar, K.; Ermann, N.; Espinosa, J.; Shen, Y.; . The Chromatin Regulator HMGA1a Undergoes Phase Separation in the Nucleus. ChemBioChem 2023, 24, e202200450.
2022
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Garaizar, A.; Espinosa, J.; Joseph, J. A.; Collepardo-Guevara, R. Kinetic Interplay Between Droplet Maturation and Coalescence Modulates Shape of Aged Protein Condensates. Scientific Reports 2022, 12 (1), 4390. https://doi.org/10.1038/s41598-022-08130-2.
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Garaizar, A.; Espinosa, J.; Joseph, J. A.; Krainer, G.; Shen, Y.; Knowles, T.; Collepardo-Guevara, R. Aging Can Transform Single-Component Protein Condensates into Multiphase Architectures. Proceedings of the National Academy of Sciences of the United States of America 2022, 119 (26), e2119800119. https://doi.org/10.1073/pnas.2119800119.
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Sanchez-Burgos, I.; Espinosa, J.; Joseph, J. A.; Collepardo-Guevara, R. RNA Length Has a Non-Trivial Effect in the Stability of Biomolecular Condensates Formed by RNA-Binding Proteins. PLoS Computational Biology 2022, 18 (2), e1009810. https://doi.org/10.1371/journal.pcbi.1009810.
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Welsh, T.; Krainer, G.; Espinosa, J.; Joseph, J. A.; Sridhar, A.; Jahnel, M.; Arter, W.; Saar, K.; Alberti, S.; Collepardo-Guevara, R.; Knowles, T. Surface Electrostatics Govern the Emulsion Stability of Biomolecular Condensates. Nano Letters 2022, 22 (2), 612-621. https://doi.org/10.1021/acs.nanolett.1c03138.
2021
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Farr, S.; Woods, E.; Joseph, J. A.; Garaizar, A.; Collepardo-Guevara, R. Nucleosome Plasticity Is a Critical Element of Chromatin Liquid-Liquid Phase Separation and Multivalent Nucleosome Interactions. Nature Communications 2021, 12 (1), 2883. https://doi.org/10.1038/s41467-021-23090-3.
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Joseph+, J. A.; Espinosa+, J.; Sanchez-Burgos, I.; Garaizar, A.; Frenkel, D.; Collepardo-Guevara, R. Thermodynamics and Kinetics of Phase Separation of Protein-RNA Mixtures by a Minimal Model. Biophysical Journal 2021, 120 (7), 1219-1230. https://doi.org/10.1016/j.bpj.2021.01.031.
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Joseph+*, J. A.; Reinhardt+*, A.; Aguirre, A.; Chew, P.; Russell, K.; Espinosa, J.; Garaizar, A.; Collepardo-Guevara*, R. Physics-Driven Coarse-Grained Model for Biomolecular Phase Separation With Near-Quantitative Accuracy. Nature Computational Science 2021, 1 (11), 732-743. https://doi.org/10.1038/s43588-021-00155-3.
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Krainer+, G.; Welsh+, T.; Joseph+, J. A.; Espinosa, J.; Wittmann, S.; de Csilléry, E.; Sridhar, A.; Toprakcioglu, Z.; Gudiškytė, G.; Czekalska, M.; Arter, W.; Guillén-Boixet, J.; Franzmann, T.; Qamar, S.; St George-Hyslop, P.; Hyman, A.; Collepardo-Guevara, R.; Alberti, S.; Knowles, T. Reentrant Liquid Condensate Phase of Proteins Is Stabilized by Hydrophobic and Non-Ionic Interactions. Nature Communications 2021, 12 (1), 1085. https://doi.org/10.1038/s41467-021-21181-9.
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Sanchez-Burgos, I.; Espinosa, J.; Joseph, J. A.; Collepardo-Guevara, R. Valency and Binding Affinity Variations Can Regulate the Multilayered Organization of Protein Condensates With Many Components. Biomolecules 2021, 11 (2). https://doi.org/10.3390/biom11020278.
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Sanchez-Burgos, I.; Joseph, J. A.; Collepardo-Guevara, R.; Espinosa, J. Size Conservation Emerges Spontaneously in Biomolecular Condensates Formed by Scaffolds and Surfactant Clients. Scientific Reports 2021, 11 (1), 15241. https://doi.org/10.1038/s41598-021-94309-y.
2020
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Espinosa, J.; Joseph, J.; Sanchez-Burgos, I.; Garaizar, A.; Frenkel, D.; Collepardo-Guevara, R. Liquid Network Connectivity Regulates the Stability and Composition of Biomolecular Condensates With Many Components. Proceedings of the National Academy of Sciences of the United States of America 2020, 117 (24), 13238-13247. https://doi.org/10.1073/pnas.1917569117.
2019
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Joseph, J. A.; Chakraborty, D.; Wales, D. J. Energy Landscape for Fold-Switching in Regulatory Protein RfaH. Journal of Chemical Theory and Computation 2019, 15 (1), 731-742. https://doi.org/10.1021/acs.jctc.8b00912.
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Röder+, K.; Joseph+, J.; Husic, B. E.; Wales, D. J. Energy Landscapes for Proteins: From Single Funnels to Multifunctional Systems. Advanced Theory and Simulations 2019, 2, 1800175.
2018
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Joseph, J. A.; Wales, D. Intrinsically Disordered Landscapes for Human CD4 Receptor Peptide. The Journal of Physical Chemistry B 2018, 122 (50), 11906-11921. https://doi.org/10.1021/acs.jpcb.8b08371.
2017
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Joseph, J. A.; Röder, K.; Chakraborty, D.; Mantell, R.; Wales, D. Exploring Biomolecular Energy Landscapes. Chemical Communications 2017, 53 (52), 6974-6988. https://doi.org/10.1039/c7cc02413d.
2016
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Joseph, J. A.; Whittleston, C.; Wales, D. Structure, Thermodynamics, and Folding Pathways for a Tryptophan Zipper As a Function of Local Rigidification. Journal of Chemical Theory and Computation 2016, 12 (12), 6109-6117.
2015
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Joseph, J. A.; McDowell, S. A. C. Comparative Computational Study of Model Halogen-Bonded Complexes of FKrCl. The journal of Physical Chemistry A 2015, 119 (11), 2568-77. https://doi.org/10.1021/jp5091262.
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McDowell, S. A. C.; Joseph, J. A. A Comparative Study of Model Halogen-Bonded, π-Hole-Bonded and Cationic Complexes Involving NCX AND H2O (X = F, Cl, Br). Molecular Physics 2015, 113, 16-21. https://doi.org/10.1080/00268976.2014.939116.
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McDowell, S. A. C.; Joseph, J. A. Interplay of Hydrogen, Halogen, Lithium and Beryllium Bonds in Complexes of Thiirane; Scheiner, S., Ed.; Springer International Publishing: Cham, 2015; pp. 357–389. https://doi.org/10.1007/978-3-319-14163-3_12.
2014
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McDowell, S. A. C.; Joseph, J. A. Variation of Sigma-Hole Magnitude With M Valence Electron Population in MX(n)Y(4-N) Molecules (n = 1-4; M = C, Si, Ge; X, Y = F, Cl, Br). Physical Chemistry Chemical Physics 2014, 16 (2), 669-71. https://doi.org/10.1039/c3cp53738b.
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McDowell, S. A. C.; Joseph, J. A. Destabilization of the Halogen Bond in Complexes of Protonated NCX (X=F, Cl, Br) Molecules. Chemical Physics Letters 2014, 603, 37-40.
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McDowell, S.; Joseph, J. The Effect of Atomic Ions on Model σ-Hole Bonded Complexes of AH3Y (A = C, Si, Ge; Y = F, Cl, Br). Physical chemistry chemical physics : PCCP 2014, 16 (22), 10854-60. https://doi.org/10.1039/c4cp01074d.
2013
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McDowell, S. A. C.; Joseph, J. A. Cooperative Effects in Novel LiF∕HF⋯LiF⋯XF (X = F, Cl, Br) Clusters. The Journal of Chemical Physics 2013, 138 (16), 164313. https://doi.org/10.1063/1.4801863.
2012
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McDowell, S. A. C.; Joseph, J. A. Communication: An Unusual Halogen-Bonding Motif: The LiBr···BrF Dimer As a Model System. The Journal of Chemical Physics 2012, 137 (17), 171103. https://doi.org/10.1063/1.4766932.
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McDowell, S. A. C.; Joseph, J. A. Cooperative Effects of Noncovalent Bonds to the Br Atom of Halogen-Bonded H3N. BrZ and HCN. BrZ (Z = F, Br) Complexes. The Journal of Chemical Physics 2012, 137 (7), 074310. https://doi.org/10.1063/1.4745838.