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Physics of active polymers: Scaling analysis via a compounding formula

Takahiro Sakaue*

Enrico Carlon

  • Soft Matter and Biophysics, KU Leuven, Celestijnenlaan 200D, B-3001 Leuven, Belgium

  • *Contact author: sakaue@phys.aoyamna.ac.jp

Phys. Rev. E 114, 015424 – Published 31 July, 2026

DOI: https://doi.org/10.1103/fhd9-l9l4

Abstract

Active polymeric systems exhibit a rich spectrum of nonequilibrium phenomena arising from stochastic forces that explicitly break detailed balance. Despite the rapid growth of experimental and numerical studies, analytical progress remains limited. To date, theoretical understanding relies largely on variants of the active Rouse model, whose formal solutions, though exact, are often obscured by summations over Rouse modes and therefore provide limited direct physical insight. In this work, we develop a transparent scaling theory that captures the tagged-monomer mean-squared displacement (MSD) in active polymers through a compounding formula: The MSD of a monomer in the chain is expressed as that of an isolated active particle, modulated by a connectivity factor encoding tension propagation along the polymer backbone. This approach isolates the role of activity from that of polymer connectivity and reveals the emergent dynamical regimes in a physically intuitive manner. We test the scaling predictions against exact calculations for a broad class of generalized active polymer models driven by diverse noise statistics. The agreement demonstrates the robustness of the scaling framework across microscopic details. Our results provide a simple and extensible theoretical structure that can be applied to complex and analytically intractable active polymer systems, thereby offering a unifying perspective on nonequilibrium polymer dynamics.

Physics Subject Headings (PhySH)

See Also

Compounding Formula Approach to Chromatin and Active Polymer Dynamics

Takahiro Sakaue and Enrico Carlon
Phys. Rev. Lett. 137, 058101 (2026)

Article Text

References (55)

  1. D. Mizuno, C. Tardin, C. F. Schmidt, and F. C. MacKintosh, Nonequilibrium mechanics of active cytoskeletal networks, Science 315, 370 (2007).
  2. M. C. Marchetti, J.-F. Joanny, S. Ramaswamy, T. B. Liverpool, J. Prost, M. Rao, and R. A. Simha, Hydrodynamics of soft active matter, Rev. Mod. Phys. 85, 1143 (2013).
  3. G. De Magistris and D. Marenduzzo, An introduction to the physics of active matter, Physica A 418, 65 (2015).
  4. F. Jülicher, S. W. Grill, and G. Salbreux, Hydrodynamic theory of active matter, Rep. Prog. Phys. 81, 076601 (2018).
  5. S. Banerjee, M. L. Gardel, and U. S. Schwarz, The actin cytoskeleton as an active adaptive material, Annu. Rev. Condens. Matter Phys. 11, 421 (2020).
  6. T. Eisenstecken, G. Gompper, and R. G. Winkler, Internal dynamics of semiflexible polymers with active noise, J. Chem. Phys. 146, 154903 (2017).
  7. R. G. Winkler and G. Gompper, The physics of active polymers and filaments, J. Chem. Phys. 153, 040901 (2020).
  8. A. Kaiser and H. Löwen, Unusual swelling of a polymer in a bacterial bath, J. Chem. Phys. 141, 044903 (2014).
  9. J. Shin, A. G. Cherstvy, W. K. Kim, and R. Metzler, Facilitation of polymer looping and giant polymer diffusivity in crowded solutions of active particles, New J. Phys. 17, 113008 (2015).
  10. H. Vandebroek and C. Vanderzande, Dynamics of a polymer in an active and viscoelastic bath, Phys. Rev. E 92, 060601(R) (2015).
  11. T. Sakaue and T. Saito, Active diffusion of model chromosomal loci driven by athermal noise, Soft Matter 13, 81 (2017).
  12. H. Vandebroek and C. Vanderzande, On the generalized Langevin equation for a Rouse bead in a nonequilibrium bath, J. Stat. Phys. 167, 14 (2017).
  13. D. Osmanović and Y. Rabin, Dynamics of active Rouse chains, Soft Matter 13, 963 (2017).
  14. J. Grimm and M. Dolgushev, Dynamics of networks in a viscoelastic and active environment, Soft Matter 14, 1171 (2018).
  15. S. Put, T. Sakaue, and C. Vanderzande, Active dynamics and spatially coherent motion in chromosomes subject to enzymatic force dipoles, Phys. Rev. E 99, 032421 (2019).
  16. V. Bianco, E. Locatelli, and P. Malgaretti, Globulelike conformation and enhanced diffusion of active polymers, Phys. Rev. Lett. 121, 217802 (2018).
  17. P. Malgaretti, E. Locatelli, and C. Valeriani, Coil-to-globule collapse of active polymers: A Rouse perspective, Mol. Phys. 123, e2384462 (2025).
  18. J. Smrek, I. Chubak, C. N. Likos, and K. Kremer, Active topological glass, Nat. Commun. 11, 26 (2020).
  19. H. Salari, M. Di Stefano, and D. Jost, Spatial organization of chromosomes leads to heterogeneous chromatin motion and drives the liquid- or gel-like dynamical behavior of chromatin, Genome Res. 32, 28 (2022).
  20. J.-X. Li, S. Wu, L.-L. Hao, Q.-L. Lei, and Y.-Q. Ma, Nonequilibrium structural and dynamic behaviors of polar active polymer controlled by head activity, Phys. Rev. Res. 5, 043064 (2023).
  21. A. Goychuk, D. Kannan, A. K. Chakraborty, and M. Kardar, Polymer folding through active processes recreates features of genome organization, Proc. Natl. Acad. Sci. USA 120, e2221726120 (2023).
  22. M. Kumar, A. Murali, A. G. Subramaniam, R. Singh, and S. Thutupalli, Emergent dynamics due to chemo-hydrodynamic self-interactions in active polymers, Nat. Commun. 15, 4903 (2024).
  23. L. Caprini, I. Abdoli, U. M. B. Marconi, and H. Löwen, Spontaneous self-wrapping in chiral active polymers, Newton 1, 100253 (2025).
  24. G. Forte, C. A. Brackley, N. Gilbert, and D. Marenduzzo, Nonequilibrium polymer models for chromatin, Curr. Opin. Genet. Dev. 96, 102426 (2026).
  25. S. C. Weber, A. J. Spakowitz, and J. A. Theriot, Nonthermal ATP-dependent fluctuations contribute to the in vivo motion of chromosomal loci, Proc. Natl. Acad. Sci. USA 109, 7338 (2012).
  26. M. M. Tortora, H. Salari, and D. Jost, Chromosome dynamics during interphase: A biophysical perspective, Curr. Op. Gen. Dev. 61, 37 (2020).
  27. N. Khanna, Y. Zhang, J. S. Lucas, O. K. Dudko, and C. Murre, Chromosome dynamics near the sol-gel phase transition dictate the timing of remote genomic interactions, Nat. Commun. 10, 2771 (2019).
  28. M. Hidalgo-Soria, Y. Haddad, E. Barkai, Y. Garini, and S. Burov, Directed motion and spatial coherence in the cell nucleus, Biophys. J. 125, 3404 (2026).
  29. T. Yuan, H. Yan, M. L. P. Bailey, J. F. Williams, I. Surovtsev, M. C. King, and S. G. J. Mochrie, Effect of loops on the mean-square displacement of Rouse-model chromatin, Phys. Rev. E 109, 044502 (2024).
  30. S. C. Weber, A. J. Spakowitz, and J. A. Theriot, Bacterial chromosomal loci move subdiffusively through a viscoelastic cytoplasm, Phys. Rev. Lett. 104, 238102 (2010).
  31. H. Hajjoul, J. Mathon, H. Ranchon, I. Goiffon, J. Mozziconacci, B. Albert, P. Carrivain, J.-M. Victor, O. Gadal, K. Bystricky, et al., High-throughput chromatin motion tracking in living yeast reveals the flexibility of the fiber throughout the genome, Genome Res. 23, 1829 (2013).
  32. M. P. Backlund, R. Joyner, and W. E. Moerner, Chromosomal locus tracking with proper accounting of static and dynamic errors, Phys. Rev. E 91, 062716 (2015).
  33. R. Wang, J. Mozziconacci, A. Bancaud, and O. Gadal, Principles of chromatin organization in yeast: Relevance of polymer models to describe nuclear organization and dynamics, Curr. Opin. Cell Biol. 34, 54 (2015).
  34. A. K. Yesbolatova, R. Arai, T. Sakaue, and A. Kimura, Formulation of chromatin mobility as a function of nuclear size during C. elegans embryogenesis using polymer physics theories, Phys. Rev. Lett. 128, 178101 (2022).
  35. S. S. Ashwin, K. Maeshima, and M. Sasai, Heterogeneous fluid-like movements of chromatin and their implications to transcription, Biophys. Rev. 12, 461 (2020).
  36. M. Socol et al., Rouse model with transient intramolecular contacts on a timescale of seconds recapitulates folding and fluctuation of yeast chromosomes, Nucleic Acids Res. 47, 6195 (2019).
  37. A. Javer, N. J. Kuwada, Z. Long, V. G. Benza, K. D. Dorfman, P. A. Wiggins, P. Cicuta, and M. Cosentino Lagomarsino, Persistent super-diffusive motion of Escherichia coli chromosomal loci, Nat. Commun. 5, 3854 (2014).
  38. T. Sakaue and E. Carlon, companion paper, Compounding formula approach to chromatin and active polymer dynamics, Phys. Rev. Lett. 137, 058101 (2026).
  39. M. Rubinstein and R. H. Colby, Polymer Physics (Oxford University Press, Oxford, 2003).
  40. T. Sakaue, Nonequilibrium dynamics of polymer translocation and straightening, Phys. Rev. E 76, 021803 (2007).
  41. P. Rowghanian and A. Y. Grosberg, Force-driven polymer translocation through a nanopore: An old problem revisited, J. Phys. Chem. B 115, 14127 (2011).
  42. T. Ikonen, A. Bhattacharya, T. Ala-Nissila, and W. Sung, Influence of pore friction on the universal aspects of driven polymer translocation, Europhys. Lett. 103, 38001 (2013).
  43. R. Frederickx, T. In't Veld, and E. Carlon, Anomalous dynamics of DNA hairpin folding, Phys. Rev. Lett. 112, 198102 (2014).
  44. T. Sakaue, Dynamics of polymer translocation: A short review with an introduction of weakly-driven regime, Polymers 8, 424 (2016).
  45. J. Sarabadani, S. Buyukdagli, and T. Ala-Nissila, Pulling a DNA molecule through a nanopore embedded in an anionic membrane: Tension propagation coupled to electrostatics, J. Phys.: Condens. Matter 32, 385101 (2020).
  46. A. Suma and C. Micheletti, Pore translocation of knotted DNA rings, Proc. Natl. Acad. Sci. USA 114, E2991 (2017).
  47. M. Doi and S. Edwards, The Theory of Polymer Dynamics (Oxford University Press, Oxford, 1988).
  48. J. Noolandi, G. W. Slater, and G. Allegra, Generalized Rouse model for polymer melt dynamics, Makromol. Chem., Rapid Commun. 8, 51 (1987).
  49. A. Amitai and D. Holcman, Polymer model with long-range interactions: Analysis and applications to the chromatin structure, Phys. Rev. E 88, 052604 (2013).
  50. K. Polovnikov, M. Gherardi, M. Cosentino-Lagomarsino, and M. Tamm, Fractal folding and medium viscoelasticity contribute jointly to chromosome dynamics, Phys. Rev. Lett. 120, 088101 (2018).
  51. T. Saito and T. Sakaue, Driven anomalous diffusion: An example from polymer stretching, Phys. Rev. E 92, 012601 (2015).
  52. P.-G. de Gennes, Scaling Concepts in Polymer Physics (Cornell University Press, Ithaca, NY, 1979).
  53. N. Katayama and T. Sakaue, Note on two-point mean square displacement, Soft Matter 21, 5871 (2025).
  54. A. Goychuk, D. Kannan, and M. Kardar, Delayed excitations induce polymer looping and coherent motion, Phys. Rev. Lett. 133, 078101 (2024).
  55. L. Remini, M. Segers, J. Palmeri, J.-C. Walter, A. Parmeggiani, and E. Carlon, Chromatin structure from high resolution microscopy: Scaling laws and microphase separation, Phys. Rev. E 109, 024408 (2024).

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