- Open Access
Scaling Laws for Passive Polymer Dynamics in Active Turbulence
Phys. Rev. Lett. 135, 138301 – Published 24 September, 2025
DOI: https://doi.org/10.1103/w3gp-knnp
Abstract
Biological systems commonly combine intrinsically out-of-equilibrium active components with passive polymeric inclusions to produce unique material properties. To explore these composite systems, idealized models—such as polymers in active fluids—are essential to develop a predictive theoretical framework. We simulate a single, freely jointed passive chain in two-dimensional active turbulence. Active flows advect the polymer, producing a substantially enhanced diffusivity. Our results reveal that the dimensionless diffusivity obeys scaling laws governed by the Péclet, Weissenberg, and Ericksen numbers, which paves the way for designing active/polymeric hybrid materials with predictable properties that differ significantly from those of nondeformable inclusions.
Physics Subject Headings (PhySH)
- Biological fluid dynamics
- Biomolecular & subcellular processes
- Brownian motion
- Cellular organization, physiology & dynamics
- Complex fluids
- Mixing enhancement
- Non-Newtonian fluids
- Polymer behavior
- Protein dynamics, structure & function
- Transition to turbulence
- Turbulence
- Turbulent convection
- Active nematics
- Biological liquid crystals
- Biological materials
- Biomolecules
- Composite materials
- Liquid crystals
- Nematic liquid crystals
- Viscoelasticity
- Coarse graining
- Molecular dynamics
Article Text
Supplemental Material
References (139)
- Roland G. Winkler and Gerhard Gompper, The physics of active polymers and filaments, J. Chem. Phys. 153, 040901 (2020).
- Raymond Kapral and Alexander S. Mikhailov, Stirring a fluid at low Reynolds numbers: Hydrodynamic collective effects of active proteins in biological cells, Physica (Amsterdam) 318D, 100 (2016).
- Alexander S. Mikhailov and Raymond Kapral, Hydrodynamic collective effects of active protein machines in solution and lipid bilayers, Proc. Natl. Acad. Sci. U.S.A. 112, E3639 (2015).
- Wen Lu, Michael Winding, Margot Lakonishok, Jill Wildonger, and Vladimir I. Gelfand, Microtubule–microtubule sliding by kinesin-1 is essential for normal cytoplasmic streaming in drosophila oocytes, Proc. Natl. Acad. Sci. U.S.A. 113, E4995 (2016).
- Yara X. Mejia, Evgeny Nudler, and Carlos Bustamante, Trigger loop folding determines transcription rate of Escherichia coli’s RNA polymerase, Proc. Natl. Acad. Sci. U.S.A. 112, 743 (2015).
- Thomas Cremer, Marion Cremer, Barbara Hübner, Hilmar Strickfaden, Daniel Smeets, Jens Popken, Michael Sterr, Yolanda Markaki, Karsten Rippe, and Christoph Cremer, The 4D nucleome: Evidence for a dynamic nuclear landscape based on co-aligned active and inactive nuclear compartments, FEBS Lett. 589, 2931 (2015).
- Nirmalendu Ganai, Surajit Sengupta, and Gautam I. Menon, Chromosome positioning from activity-based segregation, Nucl. Acids Res. 42, 4145 (2014).
- Achal Mahajan, Wen Yan, Alexandra Zidovska, David Saintillan, and Michael J. Shelley, Euchromatin activity enhances segregation and compaction of heterochromatin in the cell nucleus, Phys. Rev. X 12, 041033 (2022).
- Iraj Eshghi, Alexandra Zidovska, and Alexander Y. Grosberg, Activity-driven phase transition causes coherent flows of chromatin, Phys. Rev. Lett. 131, 048401 (2023).
- Brian Chan and Michael Rubinstein, Activity-driven chromatin organization during interphase: Compaction, segregation, and entanglement suppression, Proc. Natl. Acad. Sci. U.S.A. 121, e2401494121 (2024).
- David Saintillan, Michael J. Shelley, and Alexandra Zidovska, Extensile motor activity drives coherent motions in a model of interphase chromatin, Proc. Natl. Acad. Sci. U.S.A. 115, 11442 (2018).
- Clemens Bechinger, Roberto Di Leonardo, Hartmut Löwen, Charles Reichhardt, Giorgio Volpe, and Giovanni Volpe, Active particles in complex and crowded environments, Rev. Mod. Phys. 88, 045006 (2016).
- Sumesh P. Thampi, Amin Doostmohammadi, Ramin Golestanian, and Julia M. Yeomans, Intrinsic free energy in active nematics, Europhys. Lett. 112, 28004 (2015).
- Alexander Morozov and Davide Marenduzzo, Enhanced diffusion of tracer particles in dilute bacterial suspensions, Soft Matter 10, 2748 (2014).
- Kyriacos C. Leptos, Jeffrey S. Guasto, Jerry P. Gollub, Adriana I. Pesci, and Raymond E. Goldstein, Dynamics of enhanced tracer diffusion in suspensions of swimming eukaryotic microorganisms, Phys. Rev. Lett. 103, 198103 (2009).
- Patrick T. Underhill, Juan P. Hernandez-Ortiz, and Michael D. Graham, Diffusion and spatial correlations in suspensions of swimming particles, Phys. Rev. Lett. 100, 248101 (2008).
- Xiao-Lun Wu and Albert Libchaber, Particle diffusion in a quasi-two-dimensional bacterial bath, Phys. Rev. Lett. 84, 3017 (2000).
- Joseph Harder, Chantal Valeriani, and Angelo Cacciuto, Activity-induced collapse and reexpansion of rigid polymers, Phys. Rev. E 90, 062312 (2014).
- Gastón Mino, Thomas E. Mallouk, Thierry Darnige, Mauricio Hoyos, Jeremi Dauchet, Jocelyn Dunstan, Rodrigo Soto, Yang Wang, Annie Rousselet, and Eric Clement, Enhanced diffusion due to active swimmers at a solid surface, Phys. Rev. Lett. 106, 048102 (2011).
- Guillaume Grégoire, Hugues Chaté, and Yuhai Tu, Active and passive particles: Modeling beads in a bacterial bath, Phys. Rev. E 64, 011902 (2001).
- Chantal Valeriani, Martin Li, John Novosel, Jochen Arlt, and Davide Marenduzzo, Colloids in a bacterial bath: Simulations and experiments, Soft Matter 7, 5228 (2011).
- Sattvic Ray, Jie Zhang, and Zvonimir Dogic, Rectified rotational dynamics of mobile inclusions in two-dimensional active nematics, Phys. Rev. Lett. 130, 238301 (2023).
- Luke Neville, Jens Eggers, and Tanniemola B. Liverpool, Controlling wall-particle interactions with activity, Soft Matter 20, 8395 (2024).
- Benjamin Loewe and Tyler N. Shendruk, Passive Janus particles are self-propelled in active nematics, New J. Phys. 24, 012001 (2022).
- Roberto Di Leonardo, Luca Angelani, Dario Dell’Arciprete, Giancarlo Ruocco, Valerio Iebba, Serena Schippa, Maria Pia Conte, Francesco Mecarini, Francesco De Angelis, and Enzo Di Fabrizio, Bacterial ratchet motors, Proc. Natl. Acad. Sci. U.S.A. 107, 9541 (2010).
- Sumesh P. Thampi, Amin Doostmohammadi, Tyler N. Shendruk, Ramin Golestanian, and Julia M. Yeomans, Active micromachines: Microfluidics powered by mesoscale turbulence, Sci. Adv. 2, e1501854 (2016).
- Daiki Nishiguchi, Igor S. Aranson, Alexey Snezhko, and Andrey Sokolov, Engineering bacterial vortex lattice via direct laser lithography, Nat. Commun. 9, 4486 (2018).
- Henning Reinken, Daiki Nishiguchi, Sebastian Heidenreich, Andrey Sokolov, Markus Bär, Sabine H. L. Klapp, and Igor S. Aranson, Organizing bacterial vortex lattices by periodic obstacle arrays, Commun. Phys. 3, 76 (2020).
- Ignasi Vélez-Ceron, Rodrigo C. V. Coelho, Pau Guillamat, Margarida Telo da Gama, Francesc Sagués, and Jordi Ignés-Mullol, Active nematic pumps, arXiv:2407.09960.
- Cody D. Schimming, C. J. O. Reichhardt, and Charles Reichhardt, Active nematic ratchet in asymmetric obstacle arrays, Phys. Rev. E 109, 064602 (2024).
- Kenji Kimura, Alexandre Mamane, Tohru Sasaki, Kohta Sato, Jun Takagi, Ritsuya Niwayama, Lars Hufnagel, Yuta Shimamoto, Jean-François Joanny, Seiichi Uchida, and Akatsuki Kimura, Endoplasmic-reticulum-mediated microtubule alignment governs cytoplasmic streaming, Nat. Cell Biol. 19, 399 (2017).
- Wen Lu and Vladimir I. Gelfand, Go with the flow–bulk transport by molecular motors, J. Cell Sci. 136, jcs260300 (2023).
- Rukshala Illukkumbura, Tom Bland, and Nathan W. Goehring, Patterning and polarization of cells by intracellular flows, Curr. Opin. Cell Biol. 62, 123 (2020).
- David B. Stein, Gabriele De Canio, Eric Lauga, Michael J. Shelley, and Raymond E. Goldstein, Swirling instability of the microtubule cytoskeleton, Phys. Rev. Lett. 126, 028103 (2021).
- Alicia G. Gubieda, John R. Packer, Iolo Squires, Jack Martin, and Josana Rodriguez, Going with the flow: Insights from caenorhabditis elegans zygote polarization, Phil. Trans. R. Soc. B 375, 20190555 (2020).
- Cynthia B. Whitchurch, Tim Tolker-Nielsen, Paula C. Ragas, and John S. Mattick, Extracellular DNA required for bacterial biofilm formation, Science 295, 1487 (2002).
- Dylan P. McCuskey, Raisa E. Achiriloaie, Claire Benjamin, Jemma Kushen, Isaac Blacklow, Omar Mnfy, Jennifer L. Ross, Rae M. Robertson-Anderson, and Janet Y. Sheung, DNA transport is topologically sculpted by active microtubule dynamics, PRX Life 3, 013015 (2025).
- Wentian Liao and Igor S. Aranson, Viscoelasticity enhances collective motion of bacteria, PNAS Nexus 2, pgad291 (2023).
- Scott Weady, David B. Stein, Alexandra Zidovska, and Michael J. Shelley, Conformations, correlations, and instabilities of a flexible fiber in an active fluid, Phys. Rev. Fluids 9, 013102 (2024).
- Andreas Kaiser and Hartmut Löwen, Unusual swelling of a polymer in a bacterial bath, J. Chem. Phys. 141, 044903 (2014).
- Jaeoh Shin, Andrey G. Cherstvy, Won Kyu Kim, and Ralf Metzler, Facilitation of polymer looping and giant polymer diffusivity in crowded solutions of active particles, New J. Phys. 17, 113008 (2015).
- S. Mahdiyeh Mousavi, Gerhard Gompper, and Roland G. Winkler, Active bath-induced localization and collapse of passive semiflexible polymers, J. Chem. Phys. 155, 044902 (2021).
- Thomas Eisenstecken, Ali Ghavami, Alexander Mair, Gerhard Gompper, and Roland G. Winkler, Conformational and dynamical properties of semiflexible polymers in the presence of active noise, AIP Conf. Proc. 1871, 050001 (2017).
- Thomas Eisenstecken, Gerhard Gompper, and Roland G. Winkler, Conformational properties of active semiflexible polymers, Polymers 8, 304 (2016).
- Thomas Eisenstecken, Gerhard Gompper, and Roland G. Winkler, Internal dynamics of semiflexible polymers with active noise, J. Chem. Phys. 146 (2017).
- Nairhita Samanta and Rajarshi Chakrabarti, Chain reconfiguration in active noise, J. Phys. A 49, 195601 (2016).
- Antina Ghosh and Nir S. Gov, Dynamics of active semiflexible polymers, Biophys. J. 107, 1065 (2014).
- Hans Vandebroek and Carlo Vanderzande, Dynamics of a polymer in an active and viscoelastic bath, Phys. Rev. E 92, 060601(R) (2015),
- Takahiro Sakaue and Takuya Saito, Active diffusion of model chromosomal loci driven by athermal noise, Soft Matter 13, 81 (2017).
- Brato Chakrabarti, Manas Rachh, Stanislav Y. Shvartsman, and Michael J. Shelley, Cytoplasmic stirring by active carpets, Proc. Natl. Acad. Sci. U.S.A. 121, e2405114121 (2024).
- Pyae Hein Htet and Eric Lauga, Cortex-driven cytoplasmic flows in elongated cells: Fluid mechanics and application to nuclear transport in Drosophila embryos, J. R. Soc. Interface 20, 20230428 (2023).
- Pyae Hein Htet and Eric Lauga, Analytical methods for cytoplasmic streaming in elongated cells, PNAS Nexus 4, pgaf057 (2025).
- Maik Drechsler, Lukas F. Lang, Layla Al-Khatib, Hendrik Dirks, Martin Burger, Carola-Bibiane Schönlieb, and Isabel M. Palacios, Optical flow analysis reveals that kinesin-mediated advection impacts the orientation of microtubules in the Drosophila oocyte, Mol. Biol. Cell 31, 1246 (2020).
- Luis H. Cisneros, John O. Kessler, Sujoy Ganguly, and Raymond E. Goldstein, Dynamics of swimming bacteria: Transition to directional order at high concentration, Phys. Rev. E 83, 061907 (2011).
- Henricus H. Wensink, Jörn Dunkel, Sebastian Heidenreich, Knut Drescher, Raymond E. Goldstein, Hartmut Löwen, and Julia M. Yeomans, Meso-scale turbulence in living fluids, Proc. Natl. Acad. Sci. U.S.A. 109, 14308 (2012).
- Jeanette D. Wheeler, Eleonora Secchi, Roberto Rusconi, and Roman Stocker, Not just going with the flow: The effects of fluid flow on bacteria and plankton, Annu. Rev. Cell Dev. Biol. 35, 213 (2019).
- Hüseyin Kurtuldu, Jeffrey S. Guasto, Karl A. Johnson, and Jerry P. Gollub, Enhancement of biomixing by swimming algal cells in two-dimensional films, Proc. Natl. Acad. Sci. U.S.A. 108, 10391 (2011).
- Omer Granek, Yariv Kafri, and Julien Tailleur, Anomalous transport of tracers in active baths, Phys. Rev. Lett. 129, 038001 (2022).
- Luca Angelani and Roberto Di Leonardo, Geometrically biased random walks in bacteria-driven micro-shuttles, New J. Phys. 12, 113017 (2010).
- A. Kaiser, H. H. Wensink, and H. Löwen, How to capture active particles, Phys. Rev. Lett. 108, 268307 (2012).
- A. Kaiser, K. Popowa, H. H. Wensink, and H. Löwen, Capturing self-propelled particles in a moving microwedge, Phys. Rev. E 88, 022311 (2013).
- Andreas Kaiser, Anton Peshkov, Andrey Sokolov, Borge Ten Hagen, Hartmut Löwen, and Igor S. Aranson, Transport powered by bacterial turbulence, Phys. Rev. Lett. 112, 158101 (2014).
- Stewart A. Mallory, Chantal Valeriani, and Angelo Cacciuto, Curvature-induced activation of a passive tracer in an active bath, Phys. Rev. E 90, 032309 (2014).
- Luca Angelani, Roberto Di Leonardo, and Giancarlo Ruocco, Self-starting micromotors in a bacterial bath, Phys. Rev. Lett. 102, 048104 (2009).
- Suraj Shankar, Anton Souslov, Mark J. Bowick, M. Cristina Marchetti, and Vincenzo Vitelli, Topological active matter, Nat. Rev. Phys. 4, 380 (2022).
- Ricard Alert, Jaume Casademunt, and Jean-François Joanny, Active turbulence, Annu. Rev. Condens. Matter Phys. 13, 143 (2022).
- See Supplemental Material at http://link.aps.org/supplemental/10.1103/w3gp-knnp for simulation details, and additional numerical results, which include Refs. [68–81].
- Tyler N. Shendruk and Julia M. Yeomans, Multi-particle collision dynamics algorithm for nematic fluids, Soft Matter 11, 5101 (2015).
- Hiroshi Noguchi, Norio Kikuchi, and Gerhard Gompper, Particle-based mesoscale hydrodynamic techniques, Europhys. Lett. 78, 10005 (2007).
- Elshad Allahyarov and Gerhard Gompper, Mesoscopic solvent simulations: Multiparticlecollision dynamics of three-dimensional flows, Phys. Rev. E 66, 036702 (2002).
- Ingo O. Gotze, Hiroshi Noguchi, and Gerhard Gompper, Relevance of angular momentum conservation in mesoscale hydrodynamics simulations, Phys. Rev. E 76, 046705 (2007).
- Arne W. Zantop and Holger Stark, Multi-particle collision dynamics with a nonideal equation of state. I, J. Chem. Phys. 154, 134904 (2021).
- Adam Hospital, Josep R. Goni, Modesto Orozco, and Josep L. Gelpl., Molecular dynamics simulations: Advances and applications, Adv. Appl. Bioinform. Chem. 37 (2015).
- Gary S. Grest and Kurt Kremer, Molecular dynamics simulation for polymers in the presence of a heat bath, Phys. Rev. A 33, 3628 (1986).
- Kurt Kremer and Gary S. Grest, Dynamics of entangled linear polymer melts: A molecular-dynamics simulation, J. Chem. Phys. 92, 5057 (1990).
- Gary W. Slater et al., Modeling the separation of macromolecules: A review of current computer simulation methods, Electrophoresis 30, 792 (2009).
- C. Patrick Royall, Patrick Charbonneau, Marjolein Dijkstra, John Russo, Frank Smallenburg, Thomas Speck, and Chantal Valeriani, Colloidal hard spheres: Triumphs, challenges, and mysteries, Rev. Mod. Phys. 96, 045003 (2024).
- John D. Weeks, David Chandler, and Hans C. Andersen, Role of repulsive forces in determining the equilibrium structure of simple liquids, J. Chem. Phys. 54, 5237 (1971).
- M. Ripoll, K. Mussawisade, R. G. Winkler, and G. Gompper, Dynamic regimes of fluids simulated by multiparticle-collision dynamics, Phys. Rev. E 72, 016701 (2005).
- Judit Clopés Llahí, Aitor Martín-Gómez, Gerhard Gompper, and Roland G. Winkler, Simulating wet active polymers by multiparticle collision dynamics, Phys. Rev. E 105, 015310 (2022).
- Andreas Zottl and Holger Stark, Emergent behavior in active colloids, J. Phys.: Condens. Matter 28, 253001 (2016).
- Timofey Kozhukhov and Tyler N. Shendruk, Mesoscopic simulations of active nematics, Sci. Adv. 8, eabo5788 (2022).
- Timofey Kozhukhov, Benjamin Loewe, and Tyler N. Shendruk, Mitigating density fluctuations in particle-based active nematic simulations, Commun. Phys. 7, 251 (2024).
- Pau Guillamat, Jordi Ignés-Mullol, and Francesc Sagués, Taming active turbulence with patterned soft interfaces, Nat. Commun. 8, 564 (2017).
- Ewan J. Hemingway, Prashant Mishra, M. Cristina Marchetti, and Suzanne M. Fielding, Correlation lengths in hydrodynamic models of active nematics, Soft Matter 12, 7943 (2016).
- Paarth Gulati, Suraj Shankar, and M. Cristina Marchetti, Boundaries control active channel flows, Front. Phys. 10, 948415 (2022).
- Luca Giomi, Geometry and topology of turbulence in active nematics, Phys. Rev. X 5, 031003 (2015).
- Malcolm Hillebrand and Ricard Alert, Discontinuous transition to active nematic turbulence, arXiv:2501.06085.
- Valentino Bianco, Emanuele Locatelli, and Paolo Malgaretti, Globulelike conformation and enhanced diffusion of active polymers, Phys. Rev. Lett. 121, 217802 (2018).
- Christian A. Philipps, Gerhard Gompper, and Roland G. Winkler, Dynamics of active polar ring polymers, Phys. Rev. E 105, L062501 (2022).
- S. R. Shannon and Tuck C. Choy, Dynamical scaling anomaly for a two dimensional polymer chain in solution, Phys. Rev. Lett. 79, 1455 (1997).
- Johannes M. Vianney A. Koelman, Cellular-automaton-based simulation of 2D polymer dynamics, Phys. Rev. Lett. 64, 1915 (1990).
- E. Falck, O. Punkkinen, Ilpo Vattulainen, and Tapio Ala-Nissila, Dynamics and scaling of two-dimensional polymers in a dilute solution, Phys. Rev. E 68, 050102(R) (2003).
- Olli Punkkinen, Emma Falck, Ilpo Vattulainen, and T. Ala-Nissila, Dynamics and scaling of polymers in a dilute solution: Analytical treatment in two and higher dimensions, J. Chem. Phys. 122, 094904 (2005).
- Robert J. Poole, The Deborah, and Weissenberg numbers, Rheol. Bull. 53, 32 (2012).
- John M. Dealy, Weissenberg and Deborah numbers—Their definition and use, Rheol. Bull. 79, 14 (2010).
- David P. Rivas, Tyler N. Shendruk, Robert R. Henry, Daniel H. Reich, and Robert L. Leheny, Driven topological transitions in active nematic films, Soft Matter 16, 9331 (2020).
- Alexander J. H. Houston and Nigel J. Mottram, Spontaneous flows and quantum analogies in heterogeneous active nematic films, Commun. Phys. 7, 375 (2024).
- Olga Bantysh, Berta Martínez-Prat, Jyothishraj Nambisan, Alberto Fernández-Nieves, Francesc Sagués, and Jordi Ignés-Mullol, First order alignment transition in an interfaced active nematic fluid, Phys. Rev. Lett. 132, 228302 (2024).
- Guillaume Duclos, Raymond Adkins, Debarghya Banerjee, Matthew S. E. Peterson, Minu Varghese, Itamar Kolvin, Arvind Baskaran, Robert A. Pelcovits, Thomas R. Powers, Aparna Baskaran et al., Topological structure and dynamics of three-dimensional active nematics, Science 367, 1120 (2020).
- Žiga Krajnik, Žiga Kos, and Miha Ravnik, Spectral energy analysis of bulk three-dimensional active nematic turbulence, Soft Matter 16, 9059 (2020).
- Pasquale Digregorio, Cecilia Rorai, Ignacio Pagonabarraga, and Federico Toschi, Coexistence of defect morphologies in three-dimensional active nematics, Phys. Rev. Lett. 132, 258301 (2024).
- Giuseppe Negro, Louise C. Head, Livio N. Carenza, Tyler N. Shendruk, Davide Marenduzzo, Giuseppe Gonnella, and Adriano Tiribocchi, Topology controls flow patterns in active double emulsions, Nat. Commun. 16, 1 (2025).
- Cecilia Rorai, Federico Toschi, and Ignacio Pagonabarraga, Active nematic flows confined in a two-dimensional channel with hybrid alignment at the walls: A unified picture, Phys. Rev. Fluids 6, 113302 (2021).
- Žiga Kos, Jure Aplinc, Urban Mur, and Miha Ravnik, Mesoscopic approach to nematic fluids, in Flowing Matter, edited by F. Toschi and M. Sega (Springer International Publishing, Cham, 2019), pp. 51–93.
- Santhan Chandragiri, Amin Doostmohammadi, Julia M. Yeomans, and Sumesh P. Thampi, Active transport in a channel: Stabilisation by flow or thermodynamics, Soft Matter 15, 1597 (2019).
- Sumesh P. Thampi, Channel confined active nematics, Curr. Opin. Colloid Interface Sci. 61, 101613 (2022).
- Abhik Samui, Julia M. Yeomans, and Sumesh P. Thampi, Flow transitions and length scales of a channel-confined active nematic, Soft Matter 17, 10640 (2021).
- Ryan R. Keogh, Santhan Chandragiri, Benjamin Loewe, Tapio Ala-Nissila, Sumesh P. Thampi, and Tyler N. Shendruk, Helical flow states in active nematics, Phys. Rev. E 106, L012602 (2022).
- Louise C. Head, Giuseppe Negro, Livio N. Carenza, Nathan Johnson, Ryan R. Keogh, Giuseppe Gonnella, Alexander Morozov, Enzo Orlandini, Tyler N. Shendruk, Adriano Tiribocchi et al., Majorana quasiparticles and topological phases in 3D active nematics, Proc. Natl. Acad. Sci. U.S.A. 121, e2405304121 (2024).
- Ricard Alert, Jean-François Joanny, and Jaume Casademunt, Universal scaling of active nematic turbulence, Nat. Phys. 16, 682 (2020).
- Amin Doostmohammadi, Tyler N. Shendruk, Kristian Thijssen, and Julia M. Yeomans, Onset of meso-scale turbulence in active nematics, Nat. Commun. 8, 15326 (2017).
- Tyler N. Shendruk, Amin Doostmohammadi, Kristian Thijssen, and Julia M. Yeomans, Dancing disclinations in confined active nematics, Soft Matter 13, 3853 (2017).
- Tyler N. Shendruk, Kristian Thijssen, Julia M. Yeomans, and Amin Doostmohammadi, Twist-induced crossover from two-dimensional to three-dimensional turbulence in active nematics, Phys. Rev. E 98, 010601(R) (2018).
- Yuli Liu, Dongdong Li, Wei Feng, Hao Luo, Yanan Liu, and Guangyin Jing, Enhanced diffusion of stretching DNA chains in active baths, New J. Phys. 27, 033023 (2025).
- Hong Shen, Yueyue Hu, and W. M. Saltzman, DNA diffusion in mucus: Effect of size, topology of DNAs, and transfection reagents, Biophys. J. 91, 639 (2006).
- Rae M. Robertson, Stephan Laib, and Douglas E. Smith, Diffusion of isolated DNA molecules: Dependence on length and topology, Proc. Natl. Acad. Sci. U.S.A. 103, 7310 (2006).
- Chih-Chen Hsieh, Anthony Balducci, and Patrick S. Doyle, An experimental study of DNA rotational relaxation time in nanoslits, Macromolecules 40, 5196 (2007).
- Mehdi Shafiei Aporvari, Steven Dang, Juexin Marfai, Kara Coursey, Ryan McGorty, and Rae M. Robertson-Anderson, Crowding and confinement act in concert to slow DNA diffusion within cell-sized droplets, Iscience 25, 105122 (2022).
- Warren M. Mardoum, Stephanie M. Gorczyca, Kathryn E. Regan, Tsai-Chin Wu, and Rae M. Robertson-Anderson, Crowding induces entropically-driven changes to DNA dynamics that depend on crowder structure and ionic conditions, Front. Phys. 6, 53 (2018).
- Devynn M. Wulstein, Kathryn E. Regan, Jonathan Garamella, Ryan J. McGorty, and Rae M. Robertson-Anderson, Topology-dependent anomalous dynamics of ring and linear DNA are sensitive to cytoskeleton crosslinking, Sci. Adv. 5, eaay5912 (2019).
- Mehdi Shafiei Aporvari, Mustafa Utkur, Emine Ulku Saritas, Giovanni Volpe, and Joakim Stenhammar, Anisotropic dynamics of a self-assembled colloidal chain in an active bath, Soft Matter 16, 5609 (2020).
- Kelsey M. Hallinen, Steven P. Bodine, Howard A. Stone, Tom W. Muir, Ned S. Wingreen, and Zemer Gitai, Bacterial species with different nanocolony morphologies have distinct flow-dependent colonization behaviors, Proc. Natl. Acad. Sci. U.S.A. 122, e2419899122 (2025).
- Dario Vincenzi, Effect of internal friction on the coil–stretch transition in turbulent flows, Soft Matter 17, 2421 (2021).
- Dario Vincenzi, Takeshi Watanabe, Samriddhi Sankar Ray, and Jason R. Picardo, Polymer scission in turbulent flows, J. Fluid Mech. 912, A18 (2021).
- Francesco Serafini, Francesco Battista, Paolo Gualtieri, and Carlo M. Casciola, Polymers in turbulence: Any better than dumbbells?, J. Fluid Mech. 987, R1 (2024).
- Fenghui Lin, Jiaxing Song, Nansheng Liu, Zhenhua Wan, Xi-Yun Lu, and Bamin Khomami, Maximum drag enhancement asymptote in turbulent Taylor–Couette flow of dilute polymeric solutions, J. Non-Newtonian Fluid Mech. 323, 105172 (2024).
- Wen Yan, Saad Ansari, Adam Lamson, Matthew A. Glaser, Robert Blackwell, Meredith D. Betterton, and Michael Shelley, Toward the cellular-scale simulation of motor-driven cytoskeletal assemblies, eLife 11, e74160 (2022).
- Tim Sanchez, Daniel T. N. Chen, Stephen J. DeCamp, Michael Heymann, and Zvonimir Dogic, Spontaneous motion in hierarchically assembled active matter, Nature (London) 491, 431 (2012).
- Julio M. Belmonte, Maria Leptin, and François Nédélec, A theory that predicts behaviors of disordered cytoskeletal networks, Mol. Syst. Biol. 13, 941 (2017).
- Johanna Roostalu, Jamie Rickman, Claire Thomas, François Nédélec, and Thomas Surrey, Determinants of polar versus nematic organization in networks of dynamic microtubules and mitotic motors, Cell 175, 796 (2018).
- Charles R. Martin and Zuzanna S. Siwy, Learning nature’s way: Biosensing with synthetic nanopores, Science 317, 331 (2007).
- Basit Yameen, Mubarak Ali, Reinhard Neumann, Wolfgang Ensinger, Wolfgang Knoll, and Omar Azzaroni, Synthetic proton-gated ion channels via single solid-state nanochannels modified with responsive polymer brushes, Nano Lett. 9, 2788 (2009).
- Tingting Mei, Hongjie Zhang, and Kai Xiao, Bioinspired artificial ion pumps, ACS Nano 16, 13323 (2022).
- Jian Wang, Yahong Zhou, and Lei Jiang, Bio-inspired track-etched polymeric nanochannels: Steady-state biosensors for detection of analytes, ACS Nano 15, 18974 (2021).
- Jacques Prost, Frank Jülicher, and Jean-François Joanny, Active gel physics, Nat. Phys. 11, 111 (2015).
- Hervé Turlier, Basile Audoly, Jacques Prost, and Jean-François Joanny, Furrow constriction in animal cell cytokinesis, Biophys. J. 106, 114 (2014).
- Hudson Borja da Rocha, Jeremy Bleyer, and Hervé Turlier, A viscous active shell theory of the cell cortex, J. Mech. Phys. Solids 164, 104876 (2022).
- Tatyana M. Svitkina, Actin cell cortex: Structure and molecular organization, Trends Cell Biol. 30, 556 (2020).