Roadmap
Open Access

Roadmap for Condensates in Cell Biology

Dilimulati Aierken ^1,2, Sebastian Aland ^3,4, Stefano Bo ⁵, Steven Boeynaems ^6,7,8,9,10, Danfeng Cai ^11,12,13, Serena Carra ¹⁴, Lindsay B. Case ¹⁵, Hue Sun Chan ¹⁶, Jorge R. Espinosa ^17,18,19 et al.

Trevor K. GrandPre^20,21, Alexander Y. Grosberg ²², Ivar S. Haugerud ²³, William M. Jacobs ²⁴, Jerelle A. Joseph ^1,2,*, Frank Jülicher ^25,†, Kurt Kremer²⁶, Guido Kusters ²⁷, Liedewij Laan ²⁸, Keren Lasker ²⁹, Kathrin S. Laxhuber ²⁵, Hyun O. Lee ^16,‡, Kathy F. Liu^{30,31,32,33,34}, Dimple Notani³⁵, Yicheng Qiang²⁷, Paul Robustelli ³⁶, Leonor Saiz ³⁷, Omar A. Saleh ³⁸, Helmut Schiessel ³⁹, Jeremy D. Schmit ⁴⁰, Meng Shen ⁴¹, Krishna Shrinivas ^42,43, Antonia Statt ⁴⁴, Andres R. Tejedor ^17,18,19, Tatjana Trcek ⁴⁵, Christoph A. Weber ²³, Stephanie C. Weber ^46,47,§, Ned S. Wingreen ^48,49,∥, Huaiying Zhang ⁵⁰, Yaojun Zhang ^51,52, Huan-Xiang Zhou ⁵³, and David Zwicker ^27,¶

¹Department of Chemical and Biological Engineering, Princeton University, New Jersey 08544, USA
²Omenn-Darling Bioengineering Institute, Princeton University, New Jersey 08544, USA
³Institute of Numerical Mathematics and Optimization, TU Bergakademie Freiberg, 09599 Freiberg, Germany
⁴Faculty of Informatics/Mathematics, HTW Dresden, 01069 Dresden, Germany
⁵Department of Physics, King's College London, London WC2R 2LS, United Kingdom
⁶Department of Molecular and Human Genetics, Baylor College of Medicine, Houston, Texas 77030, USA
⁷Jan and Dan Duncan Neurological Research Institute, Texas Children's Hospital, Houston, Texas 77030, USA
⁸Therapeutic Innovation Center (THINC), Baylor College of Medicine, Houston, Texas 77030, USA
⁹Center for Alzheimer's and Neurodegenerative Diseases (CAND), Baylor College of Medicine, Houston, Texas 77030, USA
¹⁰Dan L Duncan Comprehensive Cancer Center (DLDCCC), Baylor College of Medicine, Houston, Texas 77030, USA
¹¹Department of Biochemistry and Molecular Biology, Johns Hopkins Bloomberg School of Public Health, Baltimore, Maryland 21205, USA
¹²Department of Biophysics and Biophysical Chemistry, Johns Hopkins School of Medicine, Baltimore, Maryland 21205, USA
¹³Department of Oncology, Johns Hopkins School of Medicine, Baltimore, Maryland 21205, USA
¹⁴Department of Biomedical, Metabolic and Neural Sciences, University of Modena and Reggio Emilia, Modena 41125, Italy
¹⁵Department of Biology, Massachusetts Institute of Technology, Cambridge, Massachusetts 02139, USA
¹⁶Department of Biochemistry, University of Toronto, Toronto, Ontario M5S 1A8, Canada
¹⁷Department of Physical Chemistry, University Complutense of Madrid, Madrid 28040, Spain
¹⁸Instituto Pluridisciplinar, Universidad Complutense de Madrid, P.º de Juan XXIII, 1, Moncloa - Aravaca, 28040 Madrid, Spain
¹⁹Yusuf Hamied Department of Chemistry, University of Cambridge, Lensfield Road, Cambridge CB2 1EW, United Kingdom
²⁰Department of Physics and Center for Biomolecular Condensates, Washington University in St. Louis, St. Louis, Missouri 63130, USA
²¹National Institute for Theory and Mathematics in Biology, Northwestern University and The University of Chicago, Chicago, Illinois 60611, USA
²²Department of Physics and Center for Soft Matter Research, New York University, 726 Broadway, New York, New York 10003, USA
²³Faculty of Mathematics, Natural Sciences, and Engineering, Institute of Physics, University of Augsburg, Universitätsstrasse 1, 86159 Augsburg, Germany
²⁴Department of Chemistry, Princeton University, Princeton, New Jersey 08544, USA
²⁵Max Planck Institute for the Physics of Complex Systems, Nöthnitzer Str. 38, 01187 Dresden, Germany
²⁶Max Planck Institute for Polymer Research, Ackermannweg 10, 55128 Mainz, Germany
²⁷Max Planck Institute for Dynamics and Self-Organization, Am Faßberg 17, 37077 Göttingen, Germany
²⁸Department of Bionanoscience, Kavli Institute of Nanoscience Delft, Delft University of Technology, Delft, The Netherlands
²⁹Department of Integrative Structural and Computational Biology, The Scripps Research Institute, La Jolla, California 92037, USA
³⁰Department of Biochemistry and Biophysics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA
³¹Graduate Group in Biochemistry and Molecular Biophysics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA
³²Abramson Family Cancer Research Institute, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA
³³Penn Institute for RNA Innovation, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA
³⁴Penn Center for Genome Integrity, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA
³⁵National Centre for Biological Sciences-TIFR, GKVK Campus, Bellary Road, Bangalore 560065, India
³⁶Department of Chemistry, Dartmouth College, Hanover, New Hampshire 03755, USA
³⁷Department of Biomedical Engineering, University of California, Davis, California 95616, USA
³⁸Materials and Physics Departments, University of California, Santa Barbara, California 93110, USA
³⁹Cluster of Excellence Physics of Life, TU Dresden, 01062 Dresden, Germany
⁴⁰Department of Physics, Kansas State University, Manhattan, Kansas 66506, USA
⁴¹Department of Physics, California State University, Fullerton, California 92831, USA
⁴²Department of Chemical and Biological Engineering, Northwestern University, Evanston, Illinois 60208, USA
⁴³Center for Synthetic Biology, Northwestern University, Evanston, Illinois 60208, USA
⁴⁴Department of Materials Science and Engineering, University of Illinois Urbana-Champaign, Urbana, Illinois 61801, USA
⁴⁵Department of Biology, Johns Hopkins University, Baltimore, Maryland 21218, USA
⁴⁶Department of Biology, McGill University, Montreal, H3A 1B1, Canada
⁴⁷Department of Physics, McGill University, Montreal H3A 2T8, Canada
⁴⁸Department of Molecular Biology, Princeton University, New Jersey 08544, USA
⁴⁹Lewis-Sigler Institute for Integrative Genomics, Princeton University, New Jersey 08544, USA
⁵⁰Department of Biological Sciences, Carnegie Mellon University, Pittsburgh, Pennsylvania, 15213, USA
⁵¹Department of Physics & Astronomy, Johns Hopkins University, Baltimore, Maryland 21218, USA
⁵²Department of Biophysics, Johns Hopkins University, Baltimore, Maryland 21218, USA
⁵³Department of Chemistry and Department of Physics, University of Illinois Chicago, Chicago, Illinois 60607, USA

^*Contact author: jerellejoseph@princeton.edu
^†Contact author: julicher@pks.mpg.de
^‡Contact author: hyunokate.lee@utoronto.ca
^§Contact author: steph.weber@mcgill.ca
^∥Contact author: wingreen@princeton.edu
^¶Contact author: david.zwicker@ds.mpg.de

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PRX Life 4, 032501 – Published 20 August, 2026

DOI: https://doi.org/10.1103/3b2k-ftwx

Abstract

Biomolecular condensates govern essential cellular processes yet elude description by traditional equilibrium models. This roadmap, distilled from structured discussions at a workshop and reflecting the consensus of its participants, clarifies key concepts for researchers, funding bodies, and journals. After unifying terminology that often separates disciplines, we outline the core physics of condensate formation, review their biological roles, and identify outstanding challenges in nonequilibrium theory, multiscale simulation, and quantitative in-cell measurements. We close with a forward-looking outlook to guide coordinated efforts toward predictive, experimentally anchored understanding and control of biomolecular condensates.

Physics Subject Headings (PhySH)

Article Text

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PRX Life 4, 032501– Published 20 August, 2026

Vol. 4, Iss. 3 — August - October 2026

Received 8 January 2026
Accepted 10 July 2026

DOI: https://doi.org/10.1103/3b2k-ftwx

Published by the American Physical Society under the terms of the Creative Commons Attribution 4.0 International license. Further distribution of this work must maintain attribution to the author(s) and the published article's title, journal citation, and DOI. Open access publication funded by Max Planck Society.

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Roadmap for Condensates in Cell Biology

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