Pith. sign in

REVIEW 3 major objections 5 minor 84 references

Active Matter Invasion

T0 review · 3 major / 5 minor · reviewed 2026-08-14 · deepseek-v4-flash

Pith's one-line read A growing active fluid invading a narrow capillary switches among three distinct invasion patterns—flat coherent front, S-shaped wavy front, and detaching clusters—depending only on a single dimensionless activity number.

desk verdict A competent and largely convincing numerical study mapping three invasion regimes for growing active nematics in a capillary; the main weakness is an unsupported claim about robustness to the growth implementation. read the letter →

arxiv 1908.00768 v1 pith:UQRJGN5O submitted 2019-08-02 cond-mat.soft

classification cond-mat.soft
keywords activenematicsmattercollectiveinvasioncapillaryconfinementgrowthdynamicsspontaneousflowtopologicaldefectsphase-fieldmodel
verification ladder T0 review T1 audit T2 compute T3 formal

The pith

A machine-rendered reading of the paper's core claim, the machinery that carries it, and where it could break.

The reading

This paper uses computer simulations of a generic continuum model of active matter—fluid whose elongated constituents generate internal stresses—to ask how a growing colony invades a narrow fluid-filled capillary. It claims that the invasion organizes into three distinct regimes controlled by a single dimensionless number A, the ratio of capillary width to the activity-determined length scale. At low activity the front stays flat and coherent; at intermediate activity spontaneous flows bend it into an S-shape that periodically flips sides; at high activity active blobs detach and penetrate deeper into the channel. The paper matters because it suggests that biological invaders such as bacterial biofilms or cell sheets can change their mode of spread simply by changing how much mechanical stress their constituents generate, without changing geometry. It also gives concrete thresholds and mechanisms that experiments could test.

What carries the argument

The central object is a two-phase active nematohydrodynamics model: a scalar phase field φ marks active versus passive fluid, a nematic tensor Q_αβ tracks orientational order, and an active stress −ζφQ_αβ injects energy at the scale set by the activity ζ. The organizing dimensionless group is the activity number A = d/√(K_Q/ζ), the capillary width over the active length scale, which collapses data from different channel widths onto one phase map. The argument works by identifying two observables—interface deformation (h_max − h_min)/d and number of detached clusters N_c—and showing that both jump at A ≈ 16 and A ≈ 20, with the mechanisms being spontaneous flow onset and activity-versus-surface-tension pinch-off respectively.

What would settle it

Run the same two-phase active-nematic equations with a different growth source term, such as spatially uniform proliferation instead of random local events, and check whether the two crossovers in (h_max − h_min)/d and cluster count N_c stay at A ≈ 16 and A ≈ 20; a shift or disappearance would show the regime classification depends on the growth implementation. Alternatively, measure interface flatness and detached cluster count in an expanding bacterial or epithelial monolayer in a channel while varying available chemical energy, and see whether the same two abrupt changes appear.

Watch

Extended reading notes

Core claim

The paper claims that a growing active nematic invading a fluid-filled capillary from a reservoir passes through three qualitatively distinct invasion regimes as the dimensionless activity number A = d/√(K_Q/ζ) increases: a flat-interface, flow-free regime (A ≲ 16) where invasion is purely growth and diffusion controlled; an S-shaped-interface regime (roughly 16 ≲ A ≲ 20) driven by spontaneous flow generation that advects material, with periodic flipping of the front between capillary walls as vortices form; and a regime (A ≳ 20) where active clusters pinch off from the main body and penetrate 0.5 to 1 capillary widths deeper, even though total active material in the capillary is similar to regime II. The first crossover is traced to the well-known hydrodynamic instability to spontaneous flow in confined active nematics; the second is attributed to active stresses overcoming surface tension at the interface, aided by bulk dynamics and +1/2 topological defects.

Load-bearing premise

The random local growth events that feed the active phase are assumed not to change the invasion regime; the paper compares growth versus no growth and reservoir versus no reservoir, but never varies how growth is implemented, so the regime boundaries could in principle be artifacts of that particular source term.

Editorial extensions

If this is right

  • The activity number A, rather than activity, channel width, or elasticity separately, sets the invasion mode: the same crossovers appear for capillaries of different widths when plotted against A.
  • Before the first crossover, invasion is slow and nearly independent of activity because transport is purely diffusive and growth-driven, with no spontaneous flows in the capillary.
  • After the first crossover, activity-induced flows advect active material into the capillary and the invasion index rises approximately linearly with A.
  • After the second crossover, detached clusters add little to the total amount of active material in the channel but extend maximum reach by up to one capillary width.
  • Within regime II, the front's S-shape switches from one wall to the other when the most-forward vortex reverses its rotation, giving periodic front oscillations.

Reading between the lines

Editorial extensions of the paper, not claims the author makes directly.

  • If the three-regime classification holds generally, invasive efficiency in confined geometries could be regulated by biochemical energy supply alone: a colony could switch from coherent to cluster-shedding invasion by tuning its activity, suggesting a physical control point for slowing or aiding spread.
  • The ESI result that removing the reservoir shifts the second crossover to higher A implies that upstream geometry and growth pressure participate in setting cluster detachment thresholds; an experimental study varying reservoir size while holding capillary activity fixed would test this long-range influence.
  • A natural extension is to give the activity coefficient a curvature dependence at the interface, mimicking leader cells; one prediction of the model framework is that such a term would move the first crossover or alter the S-shape switching frequency, which could be checked in particle-based simulations.
  • Cluster detachment as a way to reach deeper suggests a generic physical rationale for the advantage of disseminating small groups during collective invasion, independent of specific biochemical signalling.
Share X Bluesky LinkedIn Reddit HN

Signed reviews

No signed human review yet.

Editorial analysis

A structured set of objections, weighed in public.

Desk editor's note, referee report, and a circularity audit.

Referee Report

3 major / 5 minor

Summary. The paper uses two-dimensional hybrid lattice Boltzmann simulations of a two-phase active nematohydrodynamic model to study the invasion of a growing active nematic from a reservoir into a narrower capillary filled with isotropic fluid. Growth is implemented as stochastic local increases of the phase field in the reservoir with logistic saturation. Varying the dimensionless activity number A = d/sqrt(KQ/zeta) reveals three qualitatively distinct invasion regimes: a flat-interface coherent regime (I), an S-shaped interface with spontaneous flows and vortex-induced switching (II), and a regime in which active clusters detach from the main body (III). The authors characterize the two crossovers using the interface height difference (hmax - hmin)/d and the number of detached clusters Nc, connect the first crossover to the spontaneous flow transition in confined active nematics, and propose that the second crossover arises from active stresses overcoming surface tension. They further show that cluster detachment lets the active material protrude about 0.5 to 1 capillary-width deeper into the channel.

Significance. If the classification is robust, the paper provides a useful organizing framework for how growth, activity, and confinement combine to produce distinct modes of collective invasion, with potential relevance to biofilm expansion and collective cell migration. The strengths of the manuscript are the fully specified governing equations and parameters, well-defined observables with error bars from repeated initial conditions, and explicit falsifiable predictions (the two crossover thresholds in A and the cluster-penetration depth). The ESI adds useful controls, including no-growth and no-reservoir comparisons and a check of two channel widths. The authors are also honest about the uncertainty in the second-crossover mechanism. The central risk is the untested sensitivity of the regime structure to the details of the stochastic growth protocol, which is the ingredient that makes the regimes exist.

major comments (3)
  1. [Section 2.2 and ESI A] The claim that 'the details of this implementation or even the geometry of a reservoir are not important for the qualitative dynamics in the capillary (see ESI section A)' is not supported by the evidence presented. ESI A tests only two binary variations: removing growth entirely and removing the reservoir. It does not vary the stochastic growth parameters r, tau_g, r_g, alpha, and phi_c, nor does it test growth placed inside the capillary. Since growth is the component that makes the three invasion regimes appear, a hidden sensitivity of the crossover positions or of cluster detachment to these parameters would directly undermine the central classification. I therefore request either additional simulations varying the growth protocol (at least for a representative set of parameters and for growth distributed in the capillary) or a reformulation of the claim to state that the classification is demonstrated for the specific growth implementation used.
  2. [Section 3.2.2 and Figure 4C] The mechanistic explanation for the crossover from regime II to III is that active stresses overcome the stabilizing effect of surface tension, leading to pinch-off of clusters. However, the text itself admits that 'it is not possible to decide to what degree this effect is purely interfacial and whether dynamics in the bulk are important.' Because the paper explicitly claims to characterize 'the mechanical mechanisms underlying the crossovers,' a quantitative test is needed. For example, varying the surface-tension-related coefficients D or K_phi at fixed A and measuring Nc and the threshold would distinguish interfacial from bulk contributions. Without such a test, the proposed mechanism remains a plausible hypothesis rather than a demonstrated result; the existence of the regimes is still supported by the observables, but the explanatory part of the central claim is not yet established.
  3. [Section 3.1 and Figure 2] The text describes the two transitions as 'well-defined crossovers,' but the data points are spaced by increments of order unity in A (e.g., A = 15.5, 16.7, 17.9, 19.2), so the sharpness of the transitions is not resolved. A denser sampling of A near the claimed thresholds, together with a quantitative criterion for locating the crossover (for instance, the value of A where (hmax - hmin)/d first exceeds a threshold or where Nc becomes non-zero), would strengthen the claim that the regimes are separated by well-defined crossovers rather than by gradual changes over a finite range.
minor comments (5)
  1. [Section 2.2] The stochastic growth algorithm is described verbally; please specify the time cadence of growth attempts (e.g., whether each lattice site in the reservoir is considered once per lattice-Boltzmann time step or once per tau_g) so that the implementation is unambiguous and reproducible.
  2. [ESI A and Supplementary Figure 1] The statement that 'growth is an essential factor to create the phenomena reported in the main text' appears to be in tension with the note in the caption of Supplementary Figure 1 that 'clusters are also present for a system lacking growth, but it takes longer for them to appear.' Please clarify in what sense growth is essential (e.g., for the S-shaped interface and the early appearance of clusters) and reconcile these two statements.
  3. [Figure 3A] The axis label in Figure 3A uses units of d over the active time scale tau_zeta, but the main text says 'rms-velocity v_rms in units of d' without specifying the time normalization until the caption; please make the axis label and text consistent.
  4. [ESI B and Supplementary Figure 2] For the channel-width comparison, please state explicitly whether the same activity coefficients zeta were used for both d values (with A varying through d) or whether zeta was adjusted; this affects how the collapse onto A in Supplementary Figure 2 should be interpreted.
  5. [References] References [10] and [19] appear to be the same reference (Conrad and Poling-Skutvik, Annu. Rev. Chem. Biomol. Eng. 9:175–200, 2018) cited twice with slightly different formatting; please merge them into a single entry.

Circularity Check

0 steps flagged · score 0.0 of 10

No significant circularity: the three-regime classification is read directly from simulation observables, the spontaneous-flow benchmark is an external literature result, and no fitted parameter is renamed as a prediction.

full rationale

The paper's central claim is a classification of invasion into three regimes as the activity number A is varied. The crossover observables (hmax-hmin)/d and Nc are direct measurements from the simulations, and the crossover positions are read off the data; they are not fitted parameters and are not constructed from the classification itself. The dimensionless activity number A = d/sqrt(KQ/zeta) is a standard combination of model parameters defined independently of the output regimes, so no self-definitional loop is present. The I-to-II crossover is explained by reference to the well-established spontaneous flow transition in confined active nematics [49,50]; although some cited authors overlap with the present paper, that transition is an external, previously established result and the paper explicitly notes that growth and reservoir geometry shift its location, showing it is not being asserted by construction. The II-to-III crossover is described mechanistically as a competition between active stresses and surface tension, with the paper honestly stating that 'it is not possible to decide to what degree this effect is purely interfacial and whether dynamics in the bulk are important'; this is a hedged interpretation, not a circular reduction. One robustness gap should be flagged: Section 2.2 claims that 'the details of this implementation or even the geometry of a reservoir are not important for the qualitative dynamics in the capillary (see ESI section A)', but ESI section A only tests growth versus no-growth and reservoir versus no-reservoir; it does not vary the stochastic growth parameters r, tau_g, r_g, alpha, phi_c, or the location of growth. This is a missing-support issue for the insensitivity claim, not a circularity issue, because the regime structure is not definitionally tied to those implementation details and the claim remains empirically testable. No equation in the paper reduces to its own input, no fitted quantity is renamed as a prediction, and the central qualitative result is a direct simulation outcome rather than an output forced by a self-citation chain.

Assumptions & free parameters 4 free parameters · 4 assumptions · 0 invented entities

The central result depends on a specific continuum model and a hand-chosen parameter set inherited from prior active-nematics simulations; no parameter was fitted to the invasion observables, but the location of the crossovers is conditional on the chosen interfacial and growth parameters. No new physical entities are introduced.

free parameters (4)
  • = 0.08
    Phase-field elastic constant contributing to surface tension; the II-to-III crossover is said to depend on surface tension, so the threshold A~20 is conditional on this choice and was not varied systematically.
  • D = 0.08
    Cahn-Hilliard double-well coefficient setting interface energy; the paper states variation of D did not shift the first crossover but shows no data; D affects interfacial width and could affect cluster detachment.
  • CLQ = 0.15
    Coupling constant between nematic order and phase field; fixes interfacial properties and is not varied.
  • growth parameters (r, τg, rg, α, φc) = r=0.001, τg=10000, rg=5, α=0.01, φc=1.2
    Growth source parameters chosen by hand; the paper asserts insensitivity to their details but provides no variation study.
assumptions (4)
  • domain assumption Continuum active nematohydrodynamics (Eqs. 1-16) captures the essential physics of growing biological active matter in confinement.
    The model neglects single-cell resolution, 3D effects, and viscoelastic extracellular media, as acknowledged in the Conclusions; if these missing ingredients shift the regime boundaries, the classification would not transfer to real tissues.
  • ad hoc to paper Growth can be represented by random local mass sources with logistic saturation (Section 2.2).
    This is a specific modeling choice; the ESI tests growth vs no-growth but not alternative growth mechanisms, yet the text claims implementation details are unimportant.
  • domain assumption The activity number A = d / sqrt(KQ/ζ) is the relevant dimensionless control parameter.
    Supported by ESI Fig. 2 for two channel widths (d=40 and d=60), but only ζ and d were varied; other dimensionless groups such as surface tension and viscosity are held fixed, so A alone may not fully determine the regimes.
  • domain assumption No-slip boundaries and von Neumann conditions for φ and Q (no anchoring) in the capillary.
    The authors state that variation of boundary conditions could lead to additional phases; the reported results are conditional on this choice.

how reviews work

0 comments
Cite this review

Pith. "Pith review of Active Matter Invasion." pith.science (2026). https://pith.science/paper/UQRJGN5O

@misc{pith2026190800768,
  author       = {Pith},
  title        = {Pith review of: Active Matter Invasion},
  year         = {2026},
  howpublished = {\url{https://pith.science/paper/UQRJGN5O}},
  note         = {Machine review of arXiv:1908.00768}
}
read the original abstract

Biological active materials such as bacterial biofilms and eukaryotic cells thrive in confined micro-spaces. Here, we show through numerical simulations that confinement can serve as a mechanical guidance to achieve distinct modes of collective invasion when combined with growth dynamics and the intrinsic activity of biological materials. We assess the dynamics of the growing interface and classify these collective modes of invasion based on the activity of the constituent particles of the growing matter. While at small and moderate activities the active material grows as a coherent unit, we find that blobs of active material collectively detach from the cohort above a well-defined activity threshold. We further characterise the mechanical mechanisms underlying the crossovers between different modes of invasion and quantify their impact on the overall invasion speed.

Figures

Figures reproduced from arXiv: 1908.00768 by the authors.

Figure 1
Figure 1. Simulation setup (A) Schematic drawing of simulation setup. The active phase (yellow) invades from the broader reser￾voir (region below black dashed line) where it can grow into the narrower capillary (width d) that is initially filled with isotropic liquid (green). The broad gray arrow illustrates the invasion direction. Different heights are marked for later discussion of observables: hmax, the highest point of th… view at source ↗
Figure 2
Figure 2. Phenomenological regimes for varying activity number. (A) Snapshots from simulations showing typical configurations of the different regimes. From left to right: Regime I with flat interface, regime II with a deformed interface, and regime III where clusters (red circle) start to detach from the main active phase. Corresponding ESI Movies are flows0020.mp4, flows0035.mp4, and flows0060.mp4, respectively, see ESI sec… view at source ↗
Figure 3
Figure 3. Comparison of flows across regimes. (A) Root mean-squared velocity in units of d over the active time scale τζ = η/ζ [56] plotted against A . Crossover from regime I to II coincides with the appearance of finite flows. Strength of flows increases inside regimes II and III with a small plateau at the crossover from regime II to regime III. Lower case letters indi￾cate the values at which examples in (B-D) were taken.… view at source ↗
Figures from the paper (2 more)
Figure 4
Figure 4. Figure 4: Interface dynamics in regimes II and III. (A) Character￾istic periodic motion in regime II (A = 17.9). The +1/2-defect like bend-deformation on the right wall moves backwards rela￾tive to the interface until it is pinched off and dissolved in the nematic bulk. Simultan…
Figure 5
Figure 5. Figure 5: Propensity of the system to invade the capillary. (A) Invasion index ΦT := R φ (r, T) d 2 r/d 2 defined as the amount of active material inside the capillary after 9.5 106 simulation steps plotted versus activity number A. In regime I, the invasion shows little depende…

Discussion (0). Continue with ORCID to comment.

Reference graph

Works this paper leans on

84 extracted references · 73 canonical work pages

  1. [1]

    A. G. Clark and D. M. Vignjevic. Modes of cancer cell invasion and the role of the microenvironment. Curr. Opin. Cell Biol., 36:13–22, 2015

  2. [2]

    Cancer cells in the tumor core exhibit spatially coordinated migration patterns

    Ralitza Staneva, Fatima El Marjou, Jorge Barbazan, De- nis Krndija, Sophie Richon, Andrew Clark, and Dani- jela Matic Vignjevic. Cancer cells in the tumor core exhibit spatially coordinated migration patterns. J. Cell Sci., 132(6):jcs.220277, 2019. ISSN 0021-9533. doi: 10.1242/jcs.220277. URL http://jcs.biologists. org/content/early/2019/02/11/jcs.220277

  3. [3]

    Collective motion of bacteria in two dimensions

    Yilin Wu. Collective motion of bacteria in two dimensions. Quant. Biol., 3:199–205, 2015

  4. [4]

    Re- cent advances in studying single bacteria and biofilm me- chanics

    Catherine Even, Christian Marlière, Jean-Marc Ghigo, Jean-Marc Allain, Alba Marcellan, and Eric Raspaud. Re- cent advances in studying single bacteria and biofilm me- chanics. Adv. Colloid Interface Sci., 247:573–588, 2017

  5. [5]

    Emergence of three-dimensional order and structure in growing biofilms

    Raimo Hartmann, Praveen K Singh, Philip Pearce, Rachel Mok, Boya Song, Francisco Díaz-Pascual, Jörn Dunkel, and Knut Drescher. Emergence of three-dimensional order and structure in growing biofilms. Nat. Phys., 15:251–256, 2018

  6. [6]

    Biomass accumulation and control strategies in gas biofiltration

    Chunping Yang, Hong Chen, Guangming Zeng, Guan- long Yu, and Shenglian Luo. Biomass accumulation and control strategies in gas biofiltration. Biotech- nol. Adv. , 28(4):531 – 540, 2010. ISSN 0734-

  7. [7]

    Gritsenko, Olga Ilina, and Peter Friedl

    Pavlo G. Gritsenko, Olga Ilina, and Peter Friedl. In- terstitial guidance of cancer invasion. The Journal of Pathology, 226(2):185–199, 2011. doi: 10.1002 /path

  8. [8]

    In- travital third harmonic generation microscopy of collective melanoma cell invasion

    Bettina Weigelin, Gert-Jan Bakker, and Peter Friedl. In- travital third harmonic generation microscopy of collective melanoma cell invasion. IntraVital, 1(1):32–43, 2012. doi: 10.4161/intv.21223. URL https://doi.org/10.4161/ intv.21223

Show all 84 references
  1. [9]

    William Costerton, and Paul Stoodley

    Luanne Hall-Stoodley, J. William Costerton, and Paul Stoodley. Bacterial biofilms: from the natural environ- ment to infectious diseases. Nat. Rev. Microbiol., 2:95– 108, Feb 2004. URL http://dx.doi.org/10.1038/ nrmicro821. Review Article

  2. [10]

    Confined flow: Consequences and implications for bacteria and biofilms

    Jacinta C Conrad and Ryan Poling-Skutvik. Confined flow: Consequences and implications for bacteria and biofilms. Annu. Rev. Chem. Biomol. Eng., 9:175–200, 2018

  3. [11]

    Kabla, Chwee Teck Lim, and Benoît Ladoux

    Sri Ram Krishna Vedula, Man Chun Leong, Tan Lei Lai, Pascal Hersen, Alexandre J. Kabla, Chwee Teck Lim, and Benoît Ladoux. Emerging modes of collective cell migration induced by geometrical constraints. Proc. Natl. Acad. Sci. U. S. A. , 109(32):12974–12979, 2012. ISSN 0027-842...

  4. [12]

    Anna Kristina Marel, Matthias Zorn, Christoph Klingner, Roland Wedlich-Söldner, Erwin Frey, and Joachim O. Rädler. Flow and di ffusion in channel-guided cell mi- gration. Biophys. J., 107(5):1054–1064, 2014. ISSN 15420086. doi: 10.1016 /j.bpj.2014.07.017

  5. [13]

    Alignment of cell division axes in directed epithelial cell migration

    Anna-Kristina Marel, Nils Podewitz, Matthias Zorn, Joachim Oskar Rädler, and Jens Elgeti. Alignment of cell division axes in directed epithelial cell migration. New J. Phys., 16(11):115005, 2014. URL http://stacks. iop.org/1367-2630/16/i=11/a=115005

  6. [14]

    Dean, Reza Riahi, and Pak Kin Wong

    Yongliang Yang, Nima Jamilpour, Baoyin Yao, Zachary S. Dean, Reza Riahi, and Pak Kin Wong. Probing leader cells in endothelial collective migration by plasma lithography geometric confinement. Sci. Rep., 6:srep22707, Mar 2016. URL http://dx.doi.org/10.1038/srep22707. Arti- cle

  7. [15]

    Fardin, and Benoît Ladoux

    Estelle Gauquelin, Sham Tlili, Cyprien Gay, Grégoire Peyret, René-Marc Mège, Marc A. Fardin, and Benoît Ladoux. Influence of proliferation on the motions of ep- ithelial monolayers invading adherent strips. Soft Matter, 15:2798–2810, 2019. doi: 10.1039 /C9SM00105K. URL http://d...

  8. [16]

    Topological defects in con- fined populations of spindle-shaped cells

    Guillaume Duclos, Christoph Erlenkämper, Jean-François Joanny, and Pascal Silberzan. Topological defects in con- fined populations of spindle-shaped cells. Nat. Phys., 13: 58–62, Sep 2017. URL http://dx.doi.org/10.1038/ nphys3876

  9. [17]

    Duclos, C

    G. Duclos, C. Blanch-Mercader, V . Yashunsky, G. Sal- breux, J.-F. Joanny, J. Prost, and P. Silberzan. Spon- taneous shear flow in confined cellular nematics. Nat. Phys., 14(7):728–732, 2018. ISSN 1745-2481. doi: 10.1038/s41567-018-0099-7. URL https://doi.org/ 10.1038/s41567-018-0099-7

  10. [18]

    Directed collec- tive motion of bacteria under channel confinement

    H Wioland, E Lushi, and R E Goldstein. Directed collec- tive motion of bacteria under channel confinement. New J. Phys., 18(7):075002, 2016. URL http://stacks.iop. org/1367-2630/18/i=7/a=075002

  11. [19]

    Conrad and Ryan Poling-Skutvik

    Jacinta C. Conrad and Ryan Poling-Skutvik. Confined flow: Consequences and implications for bacteria and biofilms. Annu. Rev. Chem. Biomol. Eng., 9(1):175–200,

  12. [20]

    Spatial confinement of active microtubule networks induces large-scale rotational cytoplasmic flow

    Kazuya Suzuki, Makito Miyazaki, Jun Takagi, Takeshi Itabashi, and Shin’ichi Ishiwata. Spatial confinement of active microtubule networks induces large-scale rotational cytoplasmic flow. Proc. Natl. Acad. Sci. U. S. A., 114(11): 2922–2927, 2017. ISSN 0027-8424. doi: 10.1073 /pnas...

  13. [21]

    Kun-Ta Wu, Jean Bernard Hishamunda, Daniel T. N. Chen, Stephen J. DeCamp, Ya-Wen Chang, Alberto Fernández-Nieves, Seth Fraden, and Zvonimir Dogic. Transition from turbulent to coherent flows in confined three-dimensional active fluids. Science, 355(6331): eaal1979, 2017. ISSN 003...

  14. [22]

    Woodhouse, Jörn Dunkel, John O

    Hugo Wioland, Francis G. Woodhouse, Jörn Dunkel, John O. Kessler, and Raymond E. Goldstein. Confine- ment stabilizes a bacterial suspension into a spiral vor- tex. Phys. Rev. Lett. , 110:268102, Jun 2013. doi: 10.1103/PhysRevLett.110.268102. URL https://link. aps.org/doi/10.110...

  15. [23]

    Reconfigurable flows and defect landscape of confined active nematics

    Jérôme Hardoüin, Rian Hughes, Amin Doostmohammadi, Justine Laurent, Teresa Lopez-Leon, Julia M Yeomans, Jordi Ignés-Mullol, and Francesc Sagués. Reconfigurable flows and defect landscape of confined active nematics. page arXiv:1903.01787, 2019

  16. [24]

    Norton, Michael P

    Achini Opathalage, Michael M. Norton, Michael P. N. Juniper, Blake Langeslay, S. Ali Aghvami, Seth Fraden, and Zvonimir Dogic. Self-organized dynamics and the transition to turbulence of confined active nematics. Proc. Natl. Acad. Sci. U. S. A., 116(11):4788–4797, 2019. ISSN 00...

  17. [25]

    E ffective diffusivity of microswimmers in a crowded environment

    Marvin Brun-Cosme-Bruny, Eric Bertin, Benoît Coasne, Philippe Peyla, and Salima Rafaï. E ffective diffusivity of microswimmers in a crowded environment. J. Chem. Phys., 150(10):104901, 2019. doi: 10.1063 /1.5081507. URL https://doi.org/10.1063/1.5081507

  18. [26]

    Transport and dispersion of active parti- cles in periodic porous media

    Roberto Alonso-Matilla, Brato Chakrabarti, and David Saintillan. Transport and dispersion of active parti- cles in periodic porous media. Phys. Rev. Fluids , 4: 043101, Apr 2019. doi: 10.1103 /PhysRevFluids.4. 043101. URL https://link.aps.org/doi/10.1103/ PhysRevFluids.4.043101

  19. [27]

    Wensink, Jörn Dunkel, Sebastian Heiden- reich, Knut Drescher, Raymond E

    Henricus H. Wensink, Jörn Dunkel, Sebastian Heiden- reich, 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(36): 14308–14313, 2012. ISSN 0027-8424. doi: 10.1073 /pnas. 12020321...

  20. [28]

    Thampi, Ramin Golestanian, and Julia M

    Sumesh P. Thampi, Ramin Golestanian, and Julia M. Yeomans. V orticity, defects and correlations in ac- tive turbulence. Philos. Trans. R. Soc., A , 372 (2029):20130366, 2014. doi: 10.1098 /rsta.2013.0366. URL https://royalsocietypublishing.org/doi/ abs/10.1098/rsta.2013.0366

  21. [29]

    Geometry and topology of turbulence in active nematics

    Luca Giomi. Geometry and topology of turbulence in active nematics. Phys. Rev. X, 5:031003, Jul 2015. doi: 10.1103/PhysRevX.5.031003. URL https://link.aps. org/doi/10.1103/PhysRevX.5.031003

  22. [30]

    Martin James, Wouter J. T. Bos, and Michael Wilczek. Turbulence and turbulent pattern formation in a min- imal model for active fluids. Phys. Rev. Fluids , 3: 061101, Jun 2018. doi: 10.1103 /PhysRevFluids.3. 061101. URL https://link.aps.org/doi/10.1103/ PhysRevFluids.3.061101

  23. [31]

    New class of turbulence in active fluids

    Vasil Bratanov, Frank Jenko, and Erwin Frey. New class of turbulence in active fluids. Proc. Natl. Acad. Sci. U. S. A., 112(49):15048–15053, 2015. ISSN 0027-8424. doi: 10. 1073/pnas.1509304112. URL https://www.pnas.org/ content/112/49/15048

  24. [32]

    Elgeti, M

    J. Elgeti, M. E. Cates, and D. Marenduzzo. Defect hy- drodynamics in 2d polar active fluids. Soft Matter , 7: 3177–3185, 2011. doi: 10.1039 /C0SM01097A. URL http://dx.doi.org/10.1039/C0SM01097A

  25. [33]

    Bowick, Xu Ma, and M

    Luca Giomi, Mark J. Bowick, Xu Ma, and M. Cristina Marchetti. Defect annihilation and proliferation in active nematics. Phys. Rev. Lett., 110:228101, May 2013. doi: 10.1103/PhysRevLett.110.228101. URL https://link. aps.org/doi/10.1103/PhysRevLett.110.228101

  26. [34]

    Defect dynam- ics in active nematics

    Luca Giomi, Mark J Bowick, Prashant Mishra, Rastko Sknepnek, and M Cristina Marchetti. Defect dynam- ics in active nematics. Philos. Trans. R. Soc., A , 372 (2029):20130365, 2014. doi: 10.1098 /rsta.2013.0365. URL https://royalsocietypublishing.org/doi/ abs/10.1098/rsta.2013.0365

  27. [35]

    Yeomans, and Benoit Ladoux

    Thuan Beng Saw, Amin Doostmohammadi, Vincent Nier, Leyla Kocgozlu, Sumesh Thampi, Yusuke Toyama, Philippe Marcq, Chwee Teck Lim, Julia M. Yeomans, and Benoit Ladoux. Topological defects in epithelia govern cell death and extrusion. Nature, 544:212–216, Apr 2017. URL http://dx....

  28. [36]

    Modeling collective cell migration in geometric confine- ment

    Victoria Tarle, Estelle Gauquelin, S R K Vedula, Joseph D’Alessandro, C T Lim, Benoit Ladoux, and Nir S Gov. Modeling collective cell migration in geometric confine- ment. Phys. Biol. , 14(3):035001, 2017. URL http: //stacks.iop.org/1478-3975/14/i=3/a=035001

  29. [37]

    Blow, Sumesh P

    Matthew L. Blow, Sumesh P. Thampi, and Julia M. Yeo- mans. Biphasic, lyotropic, active nematics.Phys. Rev. Lett., 113:248303, Dec 2014. doi: 10.1103 /PhysRevLett.113. 248303. URL http://link.aps.org/doi/10.1103/ PhysRevLett.113.248303

  30. [38]

    Marenduzzo, E

    D. Marenduzzo, E. Orlandini, M. E. Cates, and J. M. Yeo- mans. Steady-state hydrodynamic instabilities of active liquid crystals: Hybrid lattice boltzmann simulations. Phys. Rev. E, 76:031921, Sep 2007. doi: 10.1103 /PhysRevE.76. 031921. URL https://link.aps.org/doi/10.1103/ P...

  31. [39]

    Aditi Simha and Sriram Ramaswamy

    R. Aditi Simha and Sriram Ramaswamy. Hydrody- namic fluctuations and instabilities in ordered suspen- sions of self-propelled particles. Phys. Rev. Lett. , 89:058101, Jul 2002. doi: 10.1103 /PhysRevLett.89. 058101. URL https://link.aps.org/doi/10.1103/ PhysRevLett.89.058101

  32. [40]

    Thampi, Ramin Golestanian, and Julia M

    Sumesh P. Thampi, Ramin Golestanian, and Julia M. Yeo- mans. Velocity correlations in an active nematic.Phys. Rev. Lett., 111:118101, Sep 2013. doi: 10.1103 /PhysRevLett. 111.118101. URL https://link.aps.org/doi/10. 1103/PhysRevLett.111.118101

  33. [41]

    DeCamp, Gabriel S

    Stephen J. DeCamp, Gabriel S. Redner, Aparna Baskaran, Michael F. Hagan, and Zvonimir Dogic. Orientational order of motile defects in active nematics. Nat. Mater., 14: 1110–1115, Aug 2015. URL http://dx.doi.org/10. 1038/nmat4387

  34. [42]

    Liverpool

    Dario Cortese, Jens Eggers, and Tanniemola B. Liverpool. Pair creation, motion, and annihilation of topological de- fects in two-dimensional nematic liquid crystals. Phys. Rev. E, 97:022704, Feb 2018. doi: 10.1103 /PhysRevE.97. Preprint – Active Matter Inv asion 10 022704. URL...

  35. [43]

    Yeomans, and Francesc Sagués

    Amin Doostmohammadi, Jordi Ignés-Mullol, Julia M. Yeomans, and Francesc Sagués. Active nematics. Nat. Commun., 9(1):3246, 2018. ISSN 2041-1723. doi: 10.1038/s41467-018-05666-8. URL https://doi.org/ 10.1038/s41467-018-05666-8

  36. [44]

    Tim Sanchez, Daniel T. N. Chen, Stephen J. DeCamp, Michael Heymann, and Zvonimir Dogic. Spontaneous motion in hierarchically assembled active matter. Nature, 491:431–434, Nov 2012. URL http://dx.doi.org/10. 1038/nature11591

  37. [45]

    Dynamics of anisotropic tissue growth

    Thomas Bittig, Ortrud Wartlick, Anna Kicheva, Mar- cos González-Gaitán, and Frank Jülicher. Dynamics of anisotropic tissue growth. New J. Phys., 10(6):063001,

  38. [46]

    Duclos, S

    G. Duclos, S. Garcia, H. G. Yevick, and P. Silberzan. Perfect nematic order in confined monolayers of spindle- shaped cells. Soft Matter , 10:2346–2353, 2014. doi: 10.1039/C3SM52323C. URL http://dx.doi.org/10. 1039/C3SM52323C

  39. [47]

    Thampi, Thuan B

    Amin Doostmohammadi, Sumesh P. Thampi, Thuan B. Saw, Chwee T. Lim, Benoit Ladoux, and Julia M. Yeomans. Celebrating soft matter’s 10th anniversary: Cell division: a source of active stress in cellular monolayers. Soft Matter, 11:7328–7336, 2015. doi: 10.1039 /C5SM01382H. URL h...

  40. [48]

    Tsimring

    Dmitri V olfson, Scott Cookson, Je ff Hasty, and Lev S. Tsimring. Biomechanical ordering of dense cell popula- tions. Proc. Natl. Acad. Sci. U. S. A. , 105(40):15346– 15351, 2008. ISSN 0027-8424. doi: 10.1073 /pnas. 0706805105. URL http://www.pnas.org/content/ 105/40/15346

  41. [49]

    V oituriez, J

    R. V oituriez, J. F. Joanny, and J. Prost. Spontaneous flow transition in active polar gels. EPL, 70(3):404,

  42. [50]

    S. A. Edwards and J. M. Yeomans. Spontaneous flow states in active nematics: A unified picture. EPL, 85(1):18008,

  43. [51]

    Shendruk, Amin Doostmohammadi, Kristian Thi- jssen, and Julia M

    Tyler N. Shendruk, Amin Doostmohammadi, Kristian Thi- jssen, and Julia M. Yeomans. Dancing disclinations in con- fined active nematics. Soft Matter, 13:3853–3862, 2017. doi: 10.1039/C6SM02310J. URL http://dx.doi.org/ 10.1039/C6SM02310J

  44. [52]

    Shendruk, Kristian Thijssen, and Julia M

    Amin Doostmohammadi, Tyler N. Shendruk, Kristian Thijssen, and Julia M. Yeomans. Onset of meso- scale turbulence in active nematics. Nat. Commun. , 8:15326, 2017. URL http://dx.doi.org/10.1038/ ncomms15326. Article

  45. [53]

    Norton, Arvind Baskaran, Achini Opatha- lage, Blake Langeslay, Seth Fraden, Aparna Baskaran, and Michael F

    Michael M. Norton, Arvind Baskaran, Achini Opatha- lage, Blake Langeslay, Seth Fraden, Aparna Baskaran, and Michael F. Hagan. Insensitivity of active nematic liquid crystal dynamics to topological constraints. Phys. Rev. E, 97:012702, Jan 2018. doi: 10.1103 /PhysRevE.97. 01270...

  46. [54]

    de Gennes and J

    Pierre G. de Gennes and J. Prost. The Physics of Liquid Crystals. Oxford University Press, Oxford, 1995

  47. [55]

    Chaikin and Taylor C

    Paul M. Chaikin and Taylor C. Lubensky. Principles of Condensed Matter Physics. Cambridge University Press, Cambridge, 2000

  48. [56]

    Orlandini, M

    E. Orlandini, M. R. Swift, and J. M. Yeomans. A lattice boltzmann model of binary-fluid mixtures. EPL, 32(6):463,

  49. [57]

    R. G. Larson. The structure and rheology of complex fluids. Oxford University Press, New York (N.Y .), 1999

  50. [58]

    The mechanics and statistics of active matter

    Sriram Ramaswamy. The mechanics and statistics of active matter. Annu. Rev. Condens. Matter Phys. , 1:323–345, 2010

  51. [59]

    Rossen, Jens M

    Ninna S. Rossen, Jens M. Tarp, Joachim Mathiesen, Mo- gens H. Jensen, and Lene B. Oddershede. Long-range ordered vorticity patterns in living tissue induced by cell division. Nat. Commun., 5:5720, Dec 2014. URL http://dx.doi.org/10.1038/ncomms6720. Article

  52. [60]

    Stabilization of active matter by flow-vortex lattices and defect ordering

    Amin Doostmohammadi, Michael F Adamer, Sumesh P Thampi, and Julia M Yeomans. Stabilization of active matter by flow-vortex lattices and defect ordering. Nat. Commun., 7:10557, 2016

  53. [61]

    Hemingway, Prashant Mishra, M

    Ewan J. Hemingway, Prashant Mishra, M. Cristina Marchetti, and Suzanne M. Fielding. Correlation lengths in hydrodynamic models of active nematics. Soft Matter, 12:7943–7952, 2016. doi: 10.1039 /C6SM00812G. URL http://dx.doi.org/10.1039/C6SM00812G

  54. [62]

    Taming active turbulence with patterned soft interfaces

    Pau Guillamat, Jordi Ignés-Mullol, and Francesc Sagués. Taming active turbulence with patterned soft interfaces. Nat. Commun., 8:564, 2017

  55. [63]

    Phase separation and emergent structures in an active nematic fluid

    Elias Putzig and Aparna Baskaran. Phase separation and emergent structures in an active nematic fluid. Phys. Rev. E, 90:042304, Oct 2014. doi: 10.1103 /PhysRevE.90. 042304. URL https://link.aps.org/doi/10.1103/ PhysRevE.90.042304

  56. [64]

    Geometric control of active collective motion

    Maxime Theillard, Roberto Alonso-Matilla, and David Saintillan. Geometric control of active collective motion. Soft Matter, 13:363–375, 2017

  57. [65]

    Thampi, Amin Doostmohammadi, Ramin Golestanian, and Julia M

    Sumesh P. Thampi, Amin Doostmohammadi, Ramin Golestanian, and Julia M. Yeomans. Intrinsic free energy in active nematics. EPL, 112(2):28004, 2015. URL http: //stacks.iop.org/0295-5075/112/i=2/a=28004

  58. [66]

    He Li, Xia-qing Shi, Mingji Huang, Xiao Chen, Minfeng Xiao, Chenli Liu, Hugues Chaté, and H. P. Zhang. Data- driven quantitative modeling of bacterial active nematics. Proc. Natl. Acad. Sci. U. S. A. , 116(3):777–785, 2019. ISSN 0027-8424. doi: 10.1073 /pnas.1812570116. URL ht...

  59. [67]

    Nematic liquid crystals formed by living amoeboid cells

    H Gruler, U Dewald, and M Eberhardt. Nematic liquid crystals formed by living amoeboid cells. Euro. Phys. J. B, 11:187–192, 1999

  60. [68]

    Turbulent dynamics of epithelial cell cultures

    C Blanch-Mercader, V Yashunsky, S Garcia, G Duclos, L Giomi, and P Silberzan. Turbulent dynamics of epithelial cell cultures. Phys. Rev. Lett., 120:208101, 2018. Preprint – Active Matter Inv asion 11

  61. [69]

    Topological defects control collective dynam- ics in neural progenitor cell cultures

    Kyogo Kawaguchi, Ryoichiro Kageyama, and Masaki Sano. Topological defects control collective dynam- ics in neural progenitor cell cultures. Nature, 545:327– 331, Apr 2017. URL http://dx.doi.org/10.1038/ nature22321

  62. [70]

    Biological tissues as active nematic liq- uid crystals

    Thuan Beng Saw, Wang Xi, Benoit Ladoux, and Chwee Teck Lim. Biological tissues as active nematic liq- uid crystals. Adv. Mater., 30(47):1802579, 2018. doi: 10. 1002/adma.201802579. URL https://onlinelibrary. wiley.com/doi/abs/10.1002/adma.201802579

  63. [71]

    Computational model for cell morphodynamics

    Danying Shao, Wouter-Jan Rappel, and Herbert Levine. Computational model for cell morphodynamics. Phys. Rev. Lett., 105:108104, Sep 2010. doi: 10.1103 /PhysRevLett. 105.108104. URL https://link.aps.org/doi/10. 1103/PhysRevLett.105.108104

  64. [72]

    Igor S. Aranson. Physical Models of Cell Motility. Springer International Publishing, Switzerland, 2016

  65. [73]

    Yeomans, and Amin Doost- mohammadi

    Romain Mueller, Julia M. Yeomans, and Amin Doost- mohammadi. Emergence of active nematic behavior in monolayers of isotropic cells. Phys. Rev. Lett. , 122: 048004, Feb 2019. doi: 10.1103 /PhysRevLett.122. 048004. URL https://link.aps.org/doi/10.1103/ PhysRevLett.122.048004

  66. [74]

    Bridging the gap between single cell migra- tion and collective dynamics

    Florian Thueroff, Andriy Goychuk, Matthias Reiter, and Erwin Frey. Bridging the gap between single cell migra- tion and collective dynamics. page bioRxiv/548677, 2019. doi: 10.1101 /548677. URL https://www.biorxiv. org/content/early/2019/02/13/548677

  67. [75]

    Collective migration of an epithelial monolayer in response to a model wound

    Mathieu Poujade, Erwan Grasland-Mongrain, A Hertzog, J Jouanneau, Philippe Chavrier, Benoît Ladoux, Axel Buguin, and Pascal Silberzan. Collective migration of an epithelial monolayer in response to a model wound. Proc. Natl. Acad. Sci. U. S. A., 104:15988–15993, 2007

  68. [76]

    Physical forces during collective cell migration

    Xavier Trepat, Michael R Wasserman, Thomas E Angelini, Emil Millet, David A Weitz, James P Butler, and Jeffrey J Fredberg. Physical forces during collective cell migration. Nat. Phys., 5(6):426, 2009

  69. [77]

    Physical model of the dynamic instability in an expanding cell culture

    Shirley Mark, Roie Shlomovitz, Nir S Gov, Mathieu Pou- jade, Erwan Grasland-Mongrain, and Pascal Silberzan. Physical model of the dynamic instability in an expanding cell culture. Biophys. J., 98:361–370, 2010. Preprint – Active Matter Inv asion 12 SupplementaryMaterials A R o...

  70. [1995]

    URL http://stacks.iop.org/0295-5075/32/ i=6/a=001

  71. [2005]

    URL http://stacks.iop.org/0295-5075/70/ i=3/a=404

  72. [2008]

    URL http://stacks.iop.org/1367-2630/10/ i=6/a=063001

  73. [2009]

    URL http://stacks.iop.org/0295-5075/85/ i=1/a=18008

  74. [2018]

    URL https://doi.org/10.1146/ annurev-chembioeng-060817-084006

    doi: 10.1146 /annurev-chembioeng-060817-084006. URL https://doi.org/10.1146/ annurev-chembioeng-060817-084006 . PMID: 29561646

  75. [3031]

    URL https://onlinelibrary.wiley.com/ doi/abs/10.1002/path.3031

  76. [9750]

    doi: https: //doi.org/10.1016/j.biotechadv.2010. 04.002. URL http://www.sciencedirect.com/ science/article/pii/S0734975010000418

Pith tools

Reviewed August 14, 2026 · model on record in the stance chip above.