Various phases of active matter emerging from bacteria and their implications
Pith reviewed 2026-05-10 12:22 UTC · model grok-4.3
The pith
Bacterial populations model active gas, liquid, glass and liquid crystal phases distinct from equilibrium matter.
A machine-rendered reading of the paper's core claim, the machinery that carries it, and where it could break.
Core claim
Bacterial populations allow for the exploration and characterization of various phases of active matter and bring rich implications for both physics and biology. Specifically, active gas, active liquid, active glass and active liquid crystal states observed in bacterial populations differ from their thermal counterparts.
What carries the argument
Bacterial populations as a model system realizing active analogs of gas, liquid, glass, and liquid crystal states through collective self-propulsion.
If this is right
- Bacterial systems enable direct observation of how activity alters phase boundaries compared with thermal equilibrium.
- Active liquid crystal states in bacteria may underlie oriented collective motion seen in certain biological processes.
- Active glass states could relate to jammed or arrested dynamics in dense bacterial communities.
- Implications extend to designing experiments that treat bacteria as tunable active materials.
Where Pith is reading between the lines
- The approach could be tested in other self-propelled systems like synthetic microswimmers to check if the same phase distinctions hold.
- If the mappings are robust, they might guide predictions for phase behavior in larger multicellular aggregates.
- Controlled removal of signaling pathways could provide a direct test of the active-matter classification.
Load-bearing premise
Observed bacterial collective behaviors can be classified as active matter phases without major confounding from biological signaling or genetic regulation.
What would settle it
An experiment isolating bacterial motion while suppressing chemotaxis and gene regulation, then showing the resulting states do not match active gas-liquid-glass-liquid-crystal classifications.
Figures
read the original abstract
In this perspective article, we discuss bacterial populations as a model system of active matter. It allows for the exploration and characterization of various phases of active matter and brings rich implications for both physics and biology. Specifically, we focus on active gas, active liquid, active glass and active liquid crystal states observed in bacterial populations and describe how these differ from their thermal counterparts. A few future directions are also discussed that will deepen the physical interest in active matter as a new type of material, with its implications for several life phenomena observed in bacterial populations and other biological systems.
Editorial analysis
A structured set of objections, weighed in public.
Referee Report
Summary. This perspective article argues that bacterial populations serve as an effective model system for exploring and characterizing various phases of active matter, including active gas, active liquid, active glass, and active liquid crystal states. It contrasts these with their thermal equilibrium counterparts, notes differences arising from self-propulsion and activity, and outlines implications for physics and biology along with suggested future directions.
Significance. If the phase classifications and distinctions hold under closer scrutiny, the perspective usefully synthesizes how bacterial collectives can illuminate active matter as a new class of materials while linking physical principles to biological collective behaviors, potentially guiding experiments that test activity-driven transitions in living systems.
major comments (2)
- [sections on active liquid and active glass states] The central mapping of observed bacterial states to active gas/liquid/glass/liquid-crystal phases (as stated in the abstract and the sections describing each phase) rests on the assumption that self-propulsion dominates over biological regulation; however, the text does not quantify or bound the relative contributions of quorum sensing, chemotaxis, or genetic motility control in the cited experiments, leaving the classification vulnerable to confounding.
- [implications and future directions] In the discussion of differences from thermal counterparts (abstract and implications section), the perspective notes qualitative distinctions but provides no concrete metrics or comparisons (e.g., effective temperatures, persistence lengths, or order parameters) that would allow readers to assess how robustly the bacterial observations align with minimal active-matter models versus biologically driven mechanisms.
minor comments (2)
- [abstract] The abstract and introduction could more explicitly state the scope as a perspective synthesizing prior work rather than presenting new derivations or data.
- [phase description sections] Figure captions (if present) or phase descriptions would benefit from clearer notation distinguishing bacterial-specific parameters from standard active-matter variables.
Simulated Author's Rebuttal
We thank the referee for their positive assessment of our perspective article and for the constructive comments, which help clarify the scope and limitations of our discussion. We address each major comment point by point below.
read point-by-point responses
-
Referee: [sections on active liquid and active glass states] The central mapping of observed bacterial states to active gas/liquid/glass/liquid-crystal phases (as stated in the abstract and the sections describing each phase) rests on the assumption that self-propulsion dominates over biological regulation; however, the text does not quantify or bound the relative contributions of quorum sensing, chemotaxis, or genetic motility control in the cited experiments, leaving the classification vulnerable to confounding.
Authors: We agree that the perspective would benefit from a more explicit acknowledgment of the potential confounding roles of biological regulation. As a synthesis of existing literature rather than a new experimental study, our classification draws on the standard active-matter interpretation of the cited bacterial experiments, where self-propulsion is taken as the dominant driver at the relevant scales. Nevertheless, we will add a concise paragraph in the introduction (or integrated into the phase sections) that reviews literature estimates on the relative magnitudes of quorum sensing, chemotaxis, and motility control versus physical activity in the density and length-scale regimes discussed. This will help readers assess the robustness of the mapping without changing the overall perspective character of the manuscript. revision: yes
-
Referee: [implications and future directions] In the discussion of differences from thermal counterparts (abstract and implications section), the perspective notes qualitative distinctions but provides no concrete metrics or comparisons (e.g., effective temperatures, persistence lengths, or order parameters) that would allow readers to assess how robustly the bacterial observations align with minimal active-matter models versus biologically driven mechanisms.
Authors: The referee correctly identifies that our treatment of distinctions from equilibrium systems remains largely qualitative. While this choice was deliberate to maintain accessibility in a perspective format, we accept that including illustrative quantitative anchors would strengthen the argument. In the revised implications section we will insert specific examples drawn from the literature, such as reported effective temperatures in bacterial baths, persistence lengths in bacterial active nematics, and orientational order parameters in bacterial liquid crystals, together with brief comparisons to minimal-model predictions. These additions will be kept concise and will not alter the forward-looking nature of the discussion. revision: yes
Circularity Check
No circularity: perspective article reviews observations without derivations or self-referential reductions
full rationale
This is a perspective article that summarizes and discusses existing experimental observations of bacterial collective behaviors mapped to active-matter phases (gas, liquid, glass, liquid crystal). No new equations, derivations, fitted parameters, or predictions are presented that could reduce to inputs by construction. Central claims rest on cited literature describing observed states and their differences from thermal systems, without load-bearing self-citations that would make the mapping circular. The text is self-contained as a review of external benchmarks and does not invoke uniqueness theorems, ansatzes, or renamings that collapse to prior author work.
Axiom & Free-Parameter Ledger
axioms (1)
- domain assumption Bacterial self-propulsion can be modeled within the active matter framework
Reference graph
Works this paper leans on
-
[1]
[Fig. 2(e)]. The realization and exploration of such phenomena that are not possible for thermal systems un- derscores the significant potential of active matter as an exotic material. TRANSITIONS TO GLASSY ST A TES As stated earlier, self-propulsion may not be the only active aspect of constituent particles of active matter. Growth and division, and resu...
-
[2]
[Fig. 4(a,b)]. This attraction and repulsion of cells for +1/2 and−1/2 defects is consistent with an earlier obser- vation for neural progenitor cells [55] and was successfully explained theoretically on the basis of two-dimensional active nematics [48, 55]. However, while bacteria grow two-dimensionally at the early stage of colony develop- ment, they st...
-
[3]
M. te Vrugt, B. Liebchen, and M. E. Cates, What exactly is ’active matter’?, arXiv:2507.21621
-
[4]
Y. Zhang and H. Hess, Enhanced diffusion of catalytically active enzymes, ACS Cent. Sci.5, 939 (2019)
work page 2019
-
[5]
O. Hallatschek, S. S. Datta, K. Drescher, J. Dunkel, J. El- geti, B. Waclaw, and N. S. Wingreen, Proliferating active matter, Nat. Rev. Phys.5, 407 (2023)
work page 2023
-
[6]
I. S. Aranson, Bacterial active matter, Rep. Prog. Phys. 85, 076601 (2022)
work page 2022
-
[7]
M. Miyata, R. C. Robinson, T. Q. P. Uyeda, Y. Fuku- mori, S. ichi Fukushima, S. Haruta, M. Homma, K. In- 6 aba, M. Ito, C. Kaito, K. Kato, T. Kenri, Y. Ki- nosita, S. Kojima, T. Minamino, H. Mori, S. Nakamura, D. Nakane, K. Nakayama, M. Nishiyama, S. Shibata, K. Shimabukuro, M. Tamakoshi, A. Taoka, Y. Tashiro, I. Tulum, H. Wada, and K. ichi Wakabayashi, T...
work page 2020
-
[8]
N. Wadhwa and H. C. Berg, Bacterial motility: machin- ery and mechanisms, Nat. Rev. Microbiol.20, 161 (2021)
work page 2021
-
[9]
Lauga,The Fluid Dynamics of Cell Motility(Cam- bridge Univ
E. Lauga,The Fluid Dynamics of Cell Motility(Cam- bridge Univ. Press, Cambridge, 2020)
work page 2020
-
[10]
Y. M. Bar-On, R. Phillips, and R. Milo, The biomass distribution on earth, Proc. Natl. Acad. Sci. USA115, 6506 (2018)
work page 2018
-
[11]
H.-C. Flemming, J. Wingender, U. Szewzyk, P. Stein- berg, S. A. Rice, and S. Kjelleberg, Biofilms: an emer- gent form of bacterial life, Nat. Rev. Microbiol.14, 563 (2016)
work page 2016
-
[12]
H. Lama, M. J. Yamamoto, Y. Furuta, T. Shimaya, and K. A. Takeuchi, Emergence of bacterial glass, PNAS Nexus3, pgae238 (2024)
work page 2024
-
[13]
T. Shimaya and K. A. Takeuchi, Tilt-induced polar order and topological defects in growing bacterial populations, PNAS Nexus1, pgac269 (2022)
work page 2022
-
[14]
S. E. Hulme, W. R. DiLuzio, S. S. Shevkoplyas, L. Turner, M. Mayer, H. C. Berg, and G. M. Whitesides, Using ratchets and sorters to fractionate motile cells of escherichia coli by length, Lab Chip8, 1888 (2008)
work page 2008
-
[15]
P. Galajda, J. Keymer, P. Chaikin, and R. Austin, A wall of funnels concentrates swimming bacteria, J. Bacteriol. 189, 8704 (2007)
work page 2007
-
[16]
D. Nishiguchi, S. Shiratani, K. A. Takeuchi, and I. S. Aranson, Vortex reversal is a precursor of confined bacterial turbulence, Proc. Natl. Acad. Sci. USA122, e2414446122 (2025)
work page 2025
-
[17]
R. Di Leonardo, L. Angelani, D. Dell’Arciprete, G. Ruocco, V. Iebba, S. Schippa, M. P. Conte, F. Mecarini, F. De Angelis, and E. Di Fabrizio, Bacterial ratchet motors, Proc. Natl. Acad. Sci. USA107, 9541 (2010)
work page 2010
-
[18]
A. Sokolov, M. M. Apodaca, B. A. Grzybowski, and I. S. Aranson, Swimming bacteria power microscopic gears, Proc. Natl. Acad. Sci. USA107, 969 (2010)
work page 2010
- [19]
-
[20]
A. P. Berke, L. Turner, H. C. Berg, and E. Lauga, Hy- drodynamic attraction of swimming microorganisms by surfaces, Phys. Rev. Lett.101, 038102 (2008)
work page 2008
- [21]
-
[22]
S. Bianchi, F. Saglimbeni, and R. Di Leonardo, Holo- graphic imaging reveals the mechanism of wall entrap- ment in swimming bacteria, Phys. Rev. X7, 011010 (2017)
work page 2017
-
[23]
Y. Takaha and D. Nishiguchi, Quasi-two-dimensional bacterial swimming around pillars: Enhanced trapping efficiency and curvature dependence, Phys. Rev. E107, 014602 (2023)
work page 2023
-
[24]
W. R. DiLuzio, L. Turner, M. Mayer, P. Garstecki, D. B. Weibel, H. C. Berg, and G. M. Whitesides, Es- cherichia coli swim on the right-hand side, Nature435, 1271 (2005)
work page 2005
-
[25]
R. Di Leonardo, D. Dell’Arciprete, L. Angelani, and V. Iebba, Swimming with an image, Phys. Rev. Lett. 106, 038101 (2011)
work page 2011
-
[26]
C. O. Reichhardt and C. Reichhardt, Ratchet effects in active matter systems, Annu. Rev. Condens. Matter Phys.8, 51 (2017)
work page 2017
- [27]
-
[28]
D. Nishiguchi, Deciphering long-range order in active matter: Insights from swimming bacteria in quasi-2d and electrokinetic janus particles, J. Phys. Soc. Jpn.92, 121007 (2023)
work page 2023
-
[29]
B. Perez-Estay, V. Martinez, C. Douarche, J. Schwarz- Linek, J. Arlt, P.-H. Delville, G. McConnell, W. C. K. Poon, A. Lindner, and E. Clement, Bacteria collective motion is scale-free, arXiv:2509.15918
-
[30]
H. Wioland, F. G. Woodhouse, J. Dunkel, J. O. Kessler, and R. E. Goldstein, Confinement stabilizes a bacterial suspension into a spiral vortex, Phys. Rev. Lett.110, 268102 (2013)
work page 2013
- [31]
-
[32]
D. Nishiguchi, I. S. Aranson, A. Snezhko, and A. Sokolov, Engineering bacterial vortex lattice via direct laser lithography, Nat. Commun.9, 4486 (2018)
work page 2018
-
[33]
J. P. Eckmann, Roads to turbulence in dissipative dy- namical systems, Rev. Mod. Phys.53, 643 (1981)
work page 1981
-
[34]
T. Shimaya, R. Okura, Y. Wakamoto, and K. A. Takeuchi, Scale invariance of cell size fluctuations in starving bacteria, Commun. Phys.4, 238 (2021)
work page 2021
- [35]
- [36]
-
[37]
M. Letz, R. Schilling, and A. Latz, Ideal glass transitions for hard ellipsoids, Phys. Rev. E62, 5173 (2000)
work page 2000
- [38]
- [39]
-
[40]
C. K. Mishra, A. Rangarajan, and R. Ganapathy, Two- step glass transition induced by attractive interactions in quasi-two-dimensional suspensions of ellipsoidal parti- cles, Phys. Rev. Lett.110, 188301 (2013)
work page 2013
- [41]
-
[42]
L. M. C. Janssen, Active glasses, J. Phys.: Condens. Mat- ter31, 503002 (2019)
work page 2019
-
[43]
K. Nishizawa, K. Fujiwara, M. Ikenaga, N. Nakajo, M. Yanagisawa, and D. Mizuno, Universal glass-forming behavior of in vitro and living cytoplasm, Sci. Rep.7, 15143 (2017)
work page 2017
-
[44]
T. C. Boothby, H. Tapia, A. H. Brozena, S. Piszkiewicz, A. E. Smith, I. Giovannini, L. Rebecchi, G. J. Pielak, D. Koshland, and B. Goldstein, Tardigrades use intrinsi- cally disordered proteins to survive desiccation, Mol. Cell 65, 975 (2017). 7
work page 2017
-
[45]
A. Doostmohammadi, J. Ign´ es-Mullol, J. M. Yeomans, and F. Sagu´ es, Active nematics, Nat. Commun.9, 3246 (2018)
work page 2018
-
[46]
A. Doostmohammadi and B. Ladoux, Physics of liquid crystals in cell biology, Trends Cell Biol.32, 140 (2022)
work page 2022
-
[47]
Y. Maroudas-Sacks, L. Garion, L. Shani-Zerbib, A. Livshits, E. Braun, and K. Keren, Topological defects in the nematic order of actin fibres as organization cen- tres of hydra morphogenesis, Nat. Phys.17, 251 (2021)
work page 2021
-
[48]
Y. Ravichandran, M. Vogg, K. Kruse, D. J. G. Pearce, and A. Roux, Topology changes of hydra define actin orientation defects as organizers of morphogenesis, Sci. Adv.11, eadr9855 (2025)
work page 2025
-
[49]
A. Doostmohammadi, S. P. Thampi, and J. M. Yeomans, Defect-mediated morphologies in growing cell colonies, Phys. Rev. Lett.117, 048102 (2016)
work page 2016
-
[50]
K. Copenhagen, R. Alert, N. S. Wingreen, and J. W. Shaevitz, Topological defects promote layer formation in myxococcus xanthus colonies, Nat. Phys.17, 211 (2021)
work page 2021
-
[51]
K. Drescher, J. Dunkel, C. D. Nadell, S. van Teeffelen, I. Grnja, N. S. Wingreen, H. A. Stone, and B. L. Bassler, Architectural transitions in vibrio cholerae biofilms at single-cell resolution, Proc. Natl. Acad. Sci. USA113, E2066 (2016)
work page 2016
-
[52]
R. Hartmann, P. K. Singh, P. Pearce, R. Mok, B. Song, F. D´ ıaz-Pascual, J. Dunkel, and K. Drescher, Emer- gence of three-dimensional order and structure in growing biofilms, Nat. Phys.15, 251 (2018)
work page 2018
- [53]
- [54]
- [55]
-
[56]
F. Yokoyama and K. A. Takeuchi, Biofilm initiation via extracellular matrix production driven by cell orientation patterning in growing escherichia coli populations (2026), DOI:10.64898/2026.03.26.714369, bioRxiv
-
[57]
K. Kawaguchi, R. Kageyama, and M. Sano, Topological defects control collective dynamics in neural progenitor cell cultures, Nature545, 327 (2017)
work page 2017
-
[58]
P. W. Anderson, More is different, Science177, 393 (1972)
work page 1972
discussion (0)
Sign in with ORCID, Apple, or X to comment. Anyone can read and Pith papers without signing in.