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REVIEW 3 major objections 4 minor 71 references

Gliding microtubules exhibit tunable collective rotation driven by chiral active forces

T0 review · 3 major / 4 minor · reviewed 2026-08-06 · deepseek-v4-flash

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Pith's one-line read Chirality in the active forces motors exert on microtubules — not filament curvature — drives the coherent collective rotation seen in gliding assays, and filament stiffness sets its speed and handedness.

desk verdict Solid sufficiency result for chiral active forces in simulations; experimental confirmation of stiffness-tunable handedness rests on an admitted double-sign-flip that is not independently anchored. read the letter →

arxiv 2507.00245 v2 pith:OICBZAXU submitted 2025-06-30 cond-mat.soft physics.bio-ph

Madhuvanthi Guruprasad Athani , Nathan Prouse , Niranjan Sarpangala , Patrick Noerr , Guillaume Schiano-Lomoriello , Ankush Gargeshwari Kumar , Fereshteh L. Memarian , Jeremie Gaillard
show 6 more authors
Laurent Blanchoin Linda S. Hirst Kinjal Dasbiswas Ajay Gopinathan Ondřej Kučera Daniel A. Beller
This is my paper · ORCID
classification cond-mat.softphysics.bio-ph
keywords microtubuleglidingassaychiralactiveforcestimecholestericnematicorderfilamentbendingrigidityBrowniandynamicssimulationkinesinmotorchiralityhandednesspropagation
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

The paper proposes that the coherent rotation of the filament crowd in microtubule-kinesin gliding assays — a 'time cholesteric' state — comes from chirality in the active forces motors exert, not from curvature or shape chirality of the filaments. The authors model this as self-propulsion directed at a small fixed skew angle to the local filament tangent, and their Brownian dynamics simulations show that this single chiral ingredient is enough to produce rotating active nematic order (a long-range-aligned state whose average filament orientation steadily rotates), as long as colliding filaments can cross each other. The result is a tunable rotor: above a bending-rigidity threshold the director rotates with the same handedness as the molecular chirality, below it the rotation reverses, and at the threshold it stops. Experiments on glass and on lipid bilayers with two microtubule types (GMPCPP- and taxol-stabilized) support the collision-driven picture and a two-sign-flip account of why both rotate counterclockwise on lipids. If the mechanism is right, it supplies a minimal route for molecular handedness to reach the material scale and identifies filament stiffness as a control knob for chiral active matter.

What carries the argument

The carrying object is a two-dimensional Brownian dynamics model of self-propelled semiflexible filaments: each filament is a chain of 30 beads connected by Hookean springs, with a harmonic bending rigidity $\tilde{\kappa}$ that sets its stiffness, and every bead is driven by an active force of fixed magnitude oriented at a small skew angle $\alpha$ to the local tangent — the only chiral ingredient in the model. Filament–filament interactions are shifted Weeks–Chandler–Andersen repulsions that let colliding filaments interpenetrate at a modest energy cost, which prior work found necessary for long-range nematic order. The argument's second mechanism is collision-mediated rotation: two-filament scattering simulations show that the bisector of a colliding pair rotates with a handedness set by $\tilde{\kappa}$, because a filament hindered at its head tends to rotate opposite to the chiral bias while one hindered at its tail rotates with it; aligning collisions dominate for flexible filaments and crossovers for rigid ones. An iterative Markov-chain weighting of the pair-collision statistics predicts the bulk handedness reversal from two-filament data alone.

What would settle it

A decisive check is a gliding assay made only of 13-protofilament microtubules, which have no protofilament skew and therefore no chiral force angle from that source: if the aligned state still rotates coherently, chiral active forces of the modeled kind are not the operative mechanism — the authors note this experiment is impractical with stabilized filaments. A second check is a continuous stiffness scan on a single microtubule type, for instance by titrating a crowding agent or crosslinker: the director's rotation should slow, stop, and reverse as the stiffness threshold near the model's $\tilde{\kappa}\approx 200$ is crossed, and a handedness that never reverses with stiffness would falsify the tuning claim.

Watch

Extended reading notes

Core claim

On the paper's own terms, the central claim is that an off-tangent component of the motor force on each microtubule segment — a chiral active force at angle $\alpha$ to the local tangent — is sufficient to produce a bulk rotating nematic state in a gliding assay, with no need for intrinsic filament curvature. The supporting discovery is that collisions, rather than single-filament turning, transmit the chirality: the curvature measured on isolated filaments would predict rotation rates orders of magnitude below those observed, and lowering the filament density weakens nematic order while speeding up the director's rotation, as expected if collisions drive it. In the bead-spring simulation, a positive $\alpha$ gives counterclockwise director rotation at high bending rigidity ($\tilde{\kappa}=300$) but clockwise rotation for $\tilde{\kappa}\lesssim 200$, with the rotation vanishing at the threshold; pair-collision statistics reproduce the reversal by weighing aligning events (favored by flexible filaments, rotating against the molecular chirality) against crossovers (favored by rigid filaments, rotating with it). The authors also predict that chiral activity makes nematic order the stable long-range-ordered state for flexible filaments — reversing the achiral trend — and that some parameters admit bistable nematic and polar states rotating with opposite handedness.

Load-bearing premise

The experimental interpretation hangs on the assumption that the sign of the effective chiral motor-force angle is fixed by the handedness of the helical protofilament twist — positive for GMPCPP microtubules, which have an excess of 14 protofilaments, and negative for taxol-stabilized ones, which have an excess of 12 — together with the assumption that taxol-stabilized microtubules sit in the low-stiffness regime where collective rotation reverses; the authors concede that if leftward motor sidestepping makes the effective angle positive for all protofilament numbers, their specific account of the taxol data would be invalidated.

Editorial extensions

If this is right

  • Coherent collective rotation can arise in gliding assays without shape chirality, so observing rotation does not by itself license claims about intrinsic filament curvature.
  • Filament bending rigidity is a single-knob control for macroscopic chirality: crossing the stiffness threshold near $\tilde{\kappa}\approx 200$ reverses the rotation's handedness, and at the threshold the rotation stops.
  • Collision frequency is the transmission channel: lower nematic order means more collisions and faster director rotation, so rotation rate and order parameter should be anti-correlated across conditions.
  • Chiral activity stabilizes nematic order over polar order in flexible-filament regimes, and produces initial-condition-dependent bistability in which nematic and polar states rotate with opposite handedness.
  • The same stiffness-controlled reversal is predicted for any chiral active system of penetrable semiflexible filaments driven at a fixed skew angle, a class the authors suggest extends beyond microtubule-kinesin assays.

Reading between the lines

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

  • Editorial inference — the head- versus tail-pinning picture implies a clean single-filament test the authors do not run: a microtubule blocked at its leading end by an obstacle should rotate opposite to one blocked at its tail, checkable in dilute assays with engineered barriers.
  • Editorial inference — if motor-driven chiral forces do contribute to cortical microtubule reorganization in plant cells as the authors speculate, their stiffness-tuned handedness predicts that interventions changing lattice rigidity (crosslinkers, acetylation, severing) should alter the sense or rate of array rotation.
  • Editorial inference — because the handedness reversal is tied to flexibility rather than to the sign of the chiral bias, synthetic chiral active rods pushed at a skew angle should show the same switch, making stiffness a general design parameter for rotor direction in active materials.
  • Editorial inference — the model's claim that collective rotation stabilizes nematic order implies that, across experimental conditions, the most persistently ordered nematic states should be those with the fastest director rotation; this correlation is measurable from existing assay movies.
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Editorial analysis

A structured set of objections, weighed in public.

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

Referee Report

3 major / 4 minor

Summary. The manuscript combines gliding assays of GMPCPP- and taxol-stabilized microtubules on glass and lipid substrates with Brownian dynamics simulations of semiflexible filaments propelled by a chiral active force tilted at a fixed angle α to the local tangent. The authors report that the emergent long-range nematic state rotates coherently (a 'time cholesteric'), that the rotation rate anti-correlates with nematic order in experiments, that chiral active forces without shape chirality are sufficient to produce such rotating states in simulations, and that the collective angular velocity reverses sign as bending stiffness κ̃ is varied below a threshold κ̃≈200. They present pair-collision simulations and an iterative Markov-chain extrapolation to support a collision-driven mechanism, and they interpret the CCW rotation of both GMPCPP- and taxol-stabilized microtubules on lipid substrates as resulting from a double sign flip in α and κ̃ for taxol.

Significance. The central simulation result—that off-tangent chiral active forces suffice to produce coherently rotating nematic order, with a non-trivial stiffness-controlled handedness reversal—is a credible and valuable contribution to active matter. The pair-collision analysis with an explicit extrapolation to bulk rotation is a strong feature, as is the experimental demonstration that filament-shape curvature cannot account for the observed rotation. The stiffness-tuning prediction appears to have been made before the taxol experiments, which mitigates circularity concerns. However, the experimental evidence for stiffness-tunable handedness is only a consistency argument, not a confirmation, and the taxol interpretation carries an admitted double-sign-flip assignment.

major comments (3)
  1. [Experimental evidence for collision-driven, rigidity-modulated collective rotation (Fig. 4D,E; Discussion)] The experimental support for the stiffness-tunable handedness prediction rests on a double-sign-flip assignment that is not independently anchored. The paper assumes taxol-stabilized microtubules have α<0 (12-protofilament excess) and assigns them to the low-κ̃ regime because the literature value κ̃≈250 is 'notably close' to the threshold κ̃≈200; neither the protofilament distribution nor the effective bending rigidity of the taxol microtubules in this assay is measured. Both observed states rotate CCW (Fig. 4D), so the taxol datum is equally consistent with the alternative, explicitly conceded hypothesis that kinesin sidestepping makes α>0 for all protofilament numbers and that taxol microtubules belong to the high-κ̃ regime; under that hypothesis there is no handedness reversal to explain. Since the paper states 'We cannot rule out this hypothesis. If true, this would invalidate our proposal...', the experiments should be described as a consistency check rather than as evidence for the stiffness-tuned handedness, unless direct measurements of the protofilament distribution and effective stiffness are added.
  2. [Collective rotation emerges from interactions, not filament curvature (Fig. 1E)] The reported anti-correlation between nematic order parameter and rotation rate is confounded by the simultaneous density change. The authors decreased the microtubule surface density from 0.20±0.02 to 0.13±0.01 filaments/µm² to obtain the lower-order state; lower density generally decreases, not increases, collision frequency, so the statement that 'collisions are expected to be more frequent when nematic order is lower' is not established by this manipulation. The observed increase in rotation rate could reflect a density-dependent change in the active nematic state rather than a collision-driven torque. A test at fixed density, or a direct measurement of collision events, is needed before this experimental result can be used to support the collision-driven mechanism.
  3. [Modeling supports chiral self-propulsion as driver of rotation, via collisions (Fig. 2D vs. Fig. 1F)] The model's parameter mapping to the experiments is strained by a single-filament discrepancy. The simulations show that high-κ̃ filaments, which are stated to correspond to the experimental microtubules, exhibit no significant average rotation, and the paper uses this to argue that the collective rotation arises from filament interactions. However, the single-filament gliding experiments on GMPCPP microtubules measure a statistically significant CCW turning bias (p=5.75×10^-11, 0.029 deg/s). If the experimental filaments rotate, they are not in the simulated high-κ̃ single-filament regime, so the claim that individual rotation does not contribute to the experimental collective rotation is not supported. Please report the simulated single-filament rotation rate at κ̃=300 in the same units as the experiments, or otherwise reconcile the measured individual turning with the model.
minor comments (4)
  1. [Fig. 4D caption vs. main text] The taxol lipid-bilayer experiments are described inconsistently: the main text reports n=2 experiments with density≈0.45 filaments/µm², while the Fig. 4D caption reports three experiments and mentions a second density of ≈0.95 filaments/µm²; please reconcile these numbers and describe the statistical treatment of multiple fields of view.
  2. [References [29]] Reference [29] is cited for the claim that nematic ordering correlates with surface density and crowding-agent concentration, but the listed reference (Saito et al., RSC Adv. 2017) is about glyme-based electrolytes and appears unrelated; this citation should be corrected.
  3. [Supporting Information, Section 2] The Supplementary Information contains a duplicated phrase: 'When when chirality is introduced' in the discussion of stationary matrices; please fix this typo.
  4. [Discussion] The sentence 'We find no overlap of the high-κ̃ regime in which LRO states emerge and the low-κ̃ regime in which individual filaments rotate with a consistent handedness' is potentially confusing because 'overlap' is used to describe disjoint intervals in κ̃; consider rewording to 'there is no κ̃ value belonging to both regimes.'

Circularity Check

1 steps flagged · score 4.0 of 10

Core simulation result is self-contained; circularity is confined to the post-hoc assignment of taxol microtubules to the low-κ regime, which turns the experimental 'confirmation' into a consistency fit.

  1. fitted input called prediction [Section 'Experimental evidence for collision-driven, rigidity-modulated collective rotation'; Fig. 4D,E]
    "This is consistent with our model if we identify taxol-stabilized microtubules as belonging to the low-bending-rigidity regime of Fig. 3C. There, with α=0.1 rad, we observed CW collective rotation for κ̃≲200; this result directly implies that with α=−0.1 rad, CCW rotation is predicted for the same κ̃ range. ... We cannot rule out this hypothesis. If true, this would invalidate our proposal that taxol-stabilized microtubules rotate opposite to the handedness of their chiral self-propulsion."

    The taxol experiment is presented as evidence for stiffness-tunable handedness, but the assignment of taxol to the low-κ regime is made after observing that taxol rotates CCW. For α<0, the model can produce either CW rotation (high-κ) or CCW rotation (low-κ), so any observed taxol handedness can be accommodated by choosing the κ regime. The authors concede this non-uniqueness by noting that if kinesin sidestepping makes α effective positive for all protofilament numbers, taxol could simply be re-classified as high-κ. Thus the 'prediction' of CCW taxol rotation is not forced by independently measured model inputs; the κ-regime assignment is selected to reproduce the observation, making the experimental validation a post-hoc consistency fit rather than an independent confirmation.

full rationale

The central Brownian-dynamics result is self-contained: a fixed off-tangent active force α produces coherently rotating nematic order even without filament curvature, and the handedness reversal near κ̃≈200 is an emergent simulation finding corroborated by two-filament scattering statistics and the phase diagram. The single-filament curvature measurements independently disfavor shape-chirality as the driver. The main circularity burden is confined to the experimental validation of the stiffness-tuned handedness claim. Taxol-stabilized microtubules rotate CCW like GMPCPP microtubules, so the authors interpret them as (α<0, low-κ) after the fact, combining two sign flips to match the data. They explicitly admit the alternative hypothesis that effective α>0 for all N would invalidate their taxol proposal, in which case taxol could be re-classified as high-κ. This means the taxol experiment does not independently confirm the stiffness-tuning prediction; it is a consistency argument with adjustable regime assignment. However, this does not reduce the core simulation derivation to its inputs, so the overall circularity is partial rather than total.

Assumptions & free parameters 5 free parameters · 4 assumptions · 0 invented entities

The model's central claim rests on the choice of an off-tangent active force as the only chiral ingredient, on parameters inherited from the authors' prior achiral filament study (ε, r_shift, bead number), and on interpretative assignments of experimental microtubule types to model regimes (κ̃, α sign). No new physical entities are introduced.

free parameters (5)
  • chiral skew angle α = 0.1 rad
    Model parameter setting the off-tangent angle of active force; motivated by protofilament supertwist but not measured in this work; all main simulations use α=0.1 rad.
  • bending rigidity κ̃ = 30 to 300 (dimensionless)
    Control parameter; κ̃=300 chosen as typical for microtubules from Ref [33]; assignment of GMPCPP to high-κ̃ and taxol to low-κ̃ regimes is an interpretative choice used to match experimental handedness.
  • area fraction ϕ = 0.05 to 0.9
    Control parameter varied in the phase diagram; ϕ=0.3 used for the main simulations.
  • WCA softness parameters ε and r_shift = ε=1e-8, r_shift=0.1
    Taken from the authors' prior Ref [33]; tuned there to produce LRO active nematic states; not re-fit here but central to allowing crossover collisions.
  • filament discretization (30 beads, r0=0.5σ) = ℓ=31r0
    Numerical discretization choice inherited from Ref [33].
assumptions (4)
  • domain assumption Kinesin-1 exerts forces along protofilament direction, yielding an active self-propulsion at a fixed skew angle α to the local tangent.
    Invoked in 'Modeling supports chiral self-propulsion...' and Fig. 2B; based on Refs [20,21,23,24].
  • ad hoc to paper The only chiral element in the model is the off-tangent active force; filament shapes are achiral.
    This is the hypothesis under test; it is a modeling choice, not an established fact.
  • domain assumption Filaments may interpenetrate at modest energy cost (WCA with r_shift=0.1, ε=1e-8); no aligning or attractive inter-filament interactions.
    Needed for LRO nematic states; justified by delamination/overlap observations and by LRO without depletants on lipid substrates; parameters from Ref [33].
  • ad hoc to paper All filaments in a given simulation share the same α and κ̃.
    Simplification; real MTs have polydisperse protofilament numbers and heterogeneous stiffness (acknowledged in the text).

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Cite this review

Pith. "Pith review of Gliding microtubules exhibit tunable collective rotation driven by chiral active forces." pith.science (2026). https://pith.science/paper/OICBZAXU

@misc{pith2026250700245,
  author       = {Pith},
  title        = {Pith review of: Gliding microtubules exhibit tunable collective rotation driven by chiral active forces},
  year         = {2026},
  howpublished = {\url{https://pith.science/paper/OICBZAXU}},
  note         = {Machine review of arXiv:2507.00245}
}
read the original abstract

How chirality propagates across scales remains an open question in many biological and synthetic systems. An especially clear manifestation of this propagation is found in in vitro gliding assays of cytoskeletal filaments on surfaces, driven by molecular motors. These assays have become model systems of active matter dynamics, as they spontaneously organize into diverse dynamical states, including collective motions with chiral rotation. However, the microscopic mechanisms underlying these chiral collective dynamics have remained unclear. Here, we investigate rotating active nematic order in microtubule gliding assay experiments under two stabilization conditions, each on two types of substrates. We propose that chirality in active forces exerted by motors on microtubules represents a viable mechanism for this large-scale chirality. Using Brownian dynamics simulations of self-propelled, semiflexible filaments with chiral activity, we demonstrate that coherently rotating active nematic order emerges by this mechanism even in the absence of curvature, i.e. shape chirality, of the constituent filaments. Moreover, we predict that the angular speed and handedness of the collective rotation can be tuned by modulating filament stiffness. Our findings identify a new set of sufficient microscopic ingredients for predictable propagation of chiral handedness from the molecular to the material scale in living and active matter.

Figures

Figures reproduced from arXiv: 2507.00245 by the authors.

Figure 1
Figure 1. Microtubule gliding assay self-organizes into rotating nematic state. [PITH_FULL_IMAGE:figures/full_fig_p014_1.png] view at source ↗
Figure 2
Figure 2. A. Illustration of microtubule lattice structures with N = 12, 13 and 14 protofilaments. All examples shown have a 3-start helix. The skew angle α of the protofilaments from the tube axis is noted, with different signs for N = 12, 14 reflecting differing handedness of supertwist. B. Schematic illustration of the bead-spring chain model of a microtubule, with Hookean linear springs (blue coils) of rest length r0 and … view at source ↗
Figure 3
Figure 3. A. Simulation snapshots showing that the direction of the nematic director’s collective rotation depends on ˜κ. The two rows are time series from two simulations, with time t˜ measured in units of T˜ and with α = 0.1 rad CCW, area fraction ϕ = 0.3. B. Time evolution of global nematic and polar order parameters for ˜κ = 100, overlaying data from a 3-run ensemble with randomized isotropic initial conditions. Insets sh… view at source ↗
Figures from the paper (2 more)
Figure 4
Figure 4. Figure 4: Microtubule motility gliding assay system on lipid bilayer self-organizes into rotating nematic [PITH_FULL_IMAGE:figures/full_fig_p017_4.png]
Figure 5
Figure 5. Figure 5: A. Snapshots of time evolution in a typical simulation from a polar initialization toward a nematic steady state for α = 0.1rad CCW, ˜κ = 30, ϕ = 0.3, Lx = 150. B,C. Snapshots of the two different types of bistable steady-state scenarios. B. For higher ˜κ values, a sys…

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Pith tools

Reviewed August 6, 2026 · model on record in the stance chip above.