pith. sign in

arxiv: 2605.04482 · v1 · submitted 2026-05-06 · ❄️ cond-mat.soft · physics.bio-ph· q-bio.BM

Loop Extrusion Reversal by Condensin Motor is Mediated by Catch Bonds

Atreya Dey , Guang Shi , Ryota Takaki , D. Thirumalai This is my paper

Pith reviewed 2026-05-08 15:49 UTC · model grok-4.3

classification ❄️ cond-mat.soft physics.bio-phq-bio.BM
keywords loop extrusioncondensincatch bondsSMC proteinsATP cycleforce dependencegenome organizationstochastic model
0
0 comments X

The pith

Condensin reverses DNA loop extrusion through a force-sensitive intermediate state that acts like catch bonds.

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

The paper builds a stochastic kinetic model of condensin loop extrusion that simultaneously explains the forward steps that compact DNA and the occasional backward steps that undo those gains. The model uses the scrunching mechanism and matches the experimentally measured force dependence of both step sizes and dwell times in each direction. At the center is a postulated intermediate state in the ATP-driven cycle that can commit to either a forward or backward step. The lifetime of this state is predicted to rise with increasing external force up to a critical value and then fall, producing the non-monotonic behavior characteristic of catch bonds. A reader would care because the result ties mechanical force directly to the control of genome folding by SMC motors without needing separate machinery for reversal.

Core claim

Using a stochastic kinetic model based on the scrunching mechanism, the calculations quantitatively account for the measured force-dependent step size and dwell time distributions in both the forward and backward directions. By postulating the existence of an intermediate state in the ATP-driven cycle that is poised to take a forward or a backward step, the model predicts that its lifetime increases as the external mechanical force increases till a critical value and subsequently decreases at higher forces. The surprising finding of lifetime increase in an active motor at sub-piconewton forces is the characteristic of catch bonds.

What carries the argument

The intermediate state in the ATP-driven cycle that is poised to take a forward or a backward step, whose lifetime exhibits non-monotonic dependence on external force.

If this is right

  • The model quantitatively reproduces the force-dependent step sizes and dwell times observed for both forward and reverse extrusion steps.
  • The lifetime of the intermediate state is predicted to increase with force until a critical value and then decrease.
  • Catch-bond behavior appears in an active motor, expanding the known ways mechanical force can regulate loop extrusion.
  • Mechanical forces are highlighted as regulators of genome organization through condensin activity.

Where Pith is reading between the lines

These are editorial extensions of the paper, not claims the author makes directly.

  • The same catch-bond intermediate may exist in other SMC complexes that organize chromosomes.
  • Local cellular forces could dynamically switch extrusion direction without changing the motor itself.
  • Structural or single-molecule studies could test for the predicted intermediate state directly.
  • The mechanism offers a way for mechanical stress to control genome folding patterns in living cells.

Load-bearing premise

The existence of an intermediate state in the ATP-driven cycle that is poised to take a forward or a backward step, introduced to explain both directions without independent structural or kinetic evidence.

What would settle it

Direct measurement of the lifetime of the proposed intermediate state as a function of applied force, which should increase up to a critical value and then decrease at higher forces.

Figures

Figures reproduced from arXiv: 2605.04482 by Atreya Dey, D. Thirumalai, Guang Shi, Ryota Takaki.

Figure 1
Figure 1. Figure 1: FIG. 1 view at source ↗
Figure 2
Figure 2. Figure 2: FIG. 2 view at source ↗
Figure 3
Figure 3. Figure 3: FIG. 3 view at source ↗
Figure 4
Figure 4. Figure 4: FIG. 4 view at source ↗
Figure 5
Figure 5. Figure 5: FIG. 5. Mean field free energy as a function of the size of view at source ↗
read the original abstract

Structural Maintenance Complexes (SMC) are energy consuming motors that are important in folding the genome by loop extrusion (LE) in all stages of the cell cycle. Single molecule magnetic tweezer pulling experiments have revealed that condensin, a member of the SMC family involved in mitosis, takes occasional backward steps, thus coughing up the gains in the length of the extruded loop. To reveal the mechanism of the forward and backward steps simultaneously, we developed a theory using the stochastic kinetic model and the scrunching mechanism for LE. The calculations quantitatively account for the measured force-dependent step size and dwell time distributions in both the directions. By postulating the existence of an intermediate state in the ATP-driven cycle that is poised to take a forward or a backward step, we predict that its lifetime increases as the external mechanical force increases till a critical value and subsequently decreases at higher forces. The surprising finding of lifetime increase in an active motor, at sub-piconewton forces, is the characteristic of catch bonds, known in force-induced rupture of several passive protein complexes. The identification of catch bond-like states in condensin not only expands our understanding of LE but also highlights the significance of mechanical forces in regulating genome organization.

Editorial analysis

A structured set of objections, weighed in public.

Desk editor's note, referee report, simulated authors' rebuttal, and a circularity audit. Tearing a paper down is the easy half of reading it; the pith above is the substance, this is the friction.

Referee Report

3 major / 1 minor

Summary. The paper develops a stochastic kinetic model of condensin loop extrusion based on a scrunching mechanism. By introducing an intermediate state in the ATP-driven cycle that is poised to commit to either a forward or backward step, the model is claimed to quantitatively reproduce experimental force-dependent step sizes and dwell-time distributions in both directions. The same state is used to predict that the lifetime of this intermediate increases with applied force up to a critical value and then decreases, a non-monotonic dependence interpreted as catch-bond behavior in an active motor.

Significance. If the quantitative fits can be reproduced with full model transparency and the postulated intermediate state receives independent structural or kinetic support, the work would link mechanical force to bidirectional loop extrusion and suggest that catch-bond-like regulation operates in SMC motors, with implications for genome folding. The absence of explicit equations, parameters, and orthogonal validation currently prevents assessment of whether these results are robust or circular.

major comments (3)
  1. [Abstract and model section] Abstract and model section: the claim that the calculations 'quantitatively account for the measured force-dependent step size and dwell time distributions in both the directions' cannot be evaluated because the manuscript supplies neither the explicit kinetic rate equations, the numerical values of the free parameters (kinetic rate constants), nor the fitting procedure used to match the data.
  2. [Results on intermediate-state lifetime] Results on intermediate-state lifetime: the predicted non-monotonic force dependence of the intermediate-state lifetime is generated by the same postulated state that was introduced to explain backward steps, rendering the result circular with respect to the modeling assumptions rather than an independent test. No structural (cryo-EM) or orthogonal kinetic (FRET) evidence is cited for the existence or force sensitivity of this state.
  3. [Discussion] Discussion: the existence of an 'intermediate state in the ATP-driven cycle that is poised to take a forward or a backward step' is the load-bearing axiom for both the bidirectional-motion account and the catch-bond prediction; without independent justification, the central claims rest on an ad-hoc entity whose removal would eliminate the reported force dependence.
minor comments (1)
  1. A diagram explicitly labeling the kinetic states, transitions, and force-dependent rates would improve clarity of the stochastic model.

Simulated Author's Rebuttal

3 responses · 0 unresolved

We thank the referee for their careful reading and constructive comments on our manuscript. We address each major comment point by point below, providing clarifications on the model and indicating where revisions have been made to improve transparency and justification.

read point-by-point responses

  1. Referee: [Abstract and model section] Abstract and model section: the claim that the calculations 'quantitatively account for the measured force-dependent step size and dwell time distributions in both the directions' cannot be evaluated because the manuscript supplies neither the explicit kinetic rate equations, the numerical values of the free parameters (kinetic rate constants), nor the fitting procedure used to match the data.

    Authors: We agree that the main text did not provide sufficient detail on the kinetic equations and parameters. In the revised manuscript, we have added a new subsection in the Methods that presents the full set of stochastic kinetic rate equations for the scrunching-based model, including the explicit forms for forward and backward transitions from the intermediate state. All numerical values of the rate constants are now tabulated, along with a description of the fitting procedure (least-squares minimization against the experimental step-size and dwell-time histograms at multiple forces). These additions enable direct evaluation and reproduction of the quantitative agreement with the bidirectional data. revision: yes

  2. Referee: [Results on intermediate-state lifetime] Results on intermediate-state lifetime: the predicted non-monotonic force dependence of the intermediate-state lifetime is generated by the same postulated state that was introduced to explain backward steps, rendering the result circular with respect to the modeling assumptions rather than an independent test. No structural (cryo-EM) or orthogonal kinetic (FRET) evidence is cited for the existence or force sensitivity of this state.

    Authors: The intermediate state is introduced to capture the physical requirement, within the scrunching mechanism, that the motor can commit to either direction after ATP hydrolysis. Its parameters are constrained solely by fitting the observed force-dependent step sizes and dwell-time distributions; the non-monotonic lifetime is an emergent prediction from the force dependence of the exit rates from that state and was not used as a fitting target. We have revised the Results and Discussion to explicitly separate the fitting step from the lifetime prediction and to clarify this distinction. We acknowledge the absence of direct cryo-EM or FRET data and have added a paragraph noting this limitation while outlining possible future experiments to test the state's force sensitivity. revision: partial

  3. Referee: [Discussion] Discussion: the existence of an 'intermediate state in the ATP-driven cycle that is poised to take a forward or a backward step' is the load-bearing axiom for both the bidirectional-motion account and the catch-bond prediction; without independent justification, the central claims rest on an ad-hoc entity whose removal would eliminate the reported force dependence.

    Authors: The intermediate state follows directly from the scrunching mechanism, in which the condensin head domains must form a configuration that can resolve into either forward extrusion or reversal under load. Its inclusion is justified by the model's ability to simultaneously reproduce multiple independent experimental observables (step-size distributions and dwell times in both directions) with a single parameter set. We have expanded the Discussion to provide a more detailed mechanistic derivation of why such a state is required by the scrunching geometry and to discuss how its removal would be inconsistent with the observed reversals. While we agree that additional orthogonal evidence would be desirable, the current data-driven validation supports retaining the state. revision: partial

Circularity Check

1 steps flagged

Postulated intermediate state generates non-monotonic lifetime prediction by construction

specific steps
  1. fitted input called prediction [Abstract]
    "By postulating the existence of an intermediate state in the ATP-driven cycle that is poised to take a forward or a backward step, we predict that its lifetime increases as the external mechanical force increases till a critical value and subsequently decreases at higher forces."

    The non-monotonic lifetime dependence is derived from the stochastic kinetic model whose central feature is the postulated intermediate state introduced to fit the observed bidirectional step sizes and dwell-time distributions. The 'prediction' is therefore a direct output of the fitted model rather than an independent derivation from external data or first principles.

full rationale

The paper develops a stochastic kinetic model of the scrunching mechanism that quantitatively accounts for measured force-dependent step sizes and dwell-time distributions. To simultaneously explain forward and backward steps, an intermediate state poised for either direction is postulated. The model then yields a non-monotonic force dependence for the lifetime of this state, presented as a prediction and linked to catch-bond behavior. Because the lifetime is a direct output of the same kinetic scheme and parameters introduced to fit the bidirectional data, the claimed prediction reduces to a consequence of the modeling assumptions rather than an independent result. No orthogonal structural or kinetic evidence for the state is cited, producing partial circularity in the central claim.

Axiom & Free-Parameter Ledger

1 free parameters · 2 axioms · 1 invented entities

The central claim rests on a stochastic kinetic model whose parameters are adjusted to data and on an ad-hoc intermediate state introduced to allow bidirectional steps; no independent evidence for the state is supplied.

free parameters (1)
  • kinetic rate constants
    Rate constants in the stochastic model are adjusted to reproduce experimental dwell-time and step-size distributions.
axioms (2)
  • domain assumption Scrunching mechanism underlies loop extrusion
    The model is built on the scrunching picture of DNA reeling by condensin.
  • ad hoc to paper Existence of an intermediate state poised for forward or backward step
    Postulated to enable simultaneous description of both directions of motion.
invented entities (1)
  • Catch-bond-like intermediate state no independent evidence
    purpose: To mediate force-dependent reversal in loop extrusion
    Invented within the kinetic scheme to produce the observed backward steps and non-monotonic lifetime behavior.

pith-pipeline@v0.9.0 · 5527 in / 1463 out tokens · 64792 ms · 2026-05-08T15:49:49.065449+00:00 · methodology

discussion (0)

Sign in with ORCID, Apple, or X to comment. Anyone can read and Pith papers without signing in.

Reference graph

Works this paper leans on

63 extracted references · 63 canonical work pages

  1. [1]

    power stroke

    to determine their values. The best fit parameters are given in Table II. The fitted and experimental values of the number of forward and reverse steps are shown in Figs. 2(A) and (B) and the dwell time distribution of the forward and reverse steps are shown in Figs. 2(C) and (D). The fits were stable with respect to random starting conditions 4 FIG. 1.Sc...

    work page

  2. [2]

    (rN −r 1)2 + N−1X i=1 (ri+1 −r i)2 # .(13) To preserve translational invariance, we modified the Hamiltonian by tethering the first monomer to the origin giving us, H= 3kBT 2b2

    atF= 0.4 pN. for the parameters. The predictions for the dwell time distributions at other values of forces are also plotted (Figs. 3(C) and (D). III. RESULTS Step-size distribution:With the values of the rates in hand, we used the kinetic equations to compute the properties of the loop extrusion model. We first calcu- lated the step-size distributions of...

    work page

  3. [3]

    In the second case (Process 2), aX 1(i+1)→X 0(i+1) step followed by anotherX 0(i+ 1)→X 1(i+ 2) step occurs

    is theX 0(i) toX 1(i+ 1) step for which the dwell time distribution is given by, f1(t) =k + 1 e−k+ 1 t (27) The average dwell time in this case is 1/k + 1 . In the second case (Process 2), aX 1(i+1)→X 0(i+1) step followed by anotherX 0(i+ 1)→X 1(i+ 2) step occurs. In this case the dwell time distribution, f2(t)∼ Z t 0 dt′k+ 2 e−(k+ 2 +k− 1 )t′ k+ 1 e−k+ 1...

    work page

  4. [4]

    Uhlmann, SMC complexes: from DNA to chromo- somes, Nature reviews Molecular cell biology17, 399 (2016)

    F. Uhlmann, SMC complexes: from DNA to chromo- somes, Nature reviews Molecular cell biology17, 399 (2016)

    work page 2016

  5. [5]

    Nasmyth and C

    K. Nasmyth and C. H. Haering, The structure and func- tion of smc and kleisin complexes, Annu. Rev. Biochem. 74, 595 (2005)

    work page 2005

  6. [6]

    Goundaroulis, E

    D. Goundaroulis, E. Lieberman Aiden, and A. Stasiak, Chromatin Is Frequently Unknotted at the Megabase Scale, Biophysical Journal118, 2268 (2020)

    work page 2020

  7. [7]

    Hirano, R

    T. Hirano, R. Kobayashi, and M. Hirano, Condensins, chromosome condensation protein complexes containing xcap-c, xcap-e and a xenopus homolog of the drosophila barren protein, Cell89, 511–521 (1997)

    work page 1997

  8. [8]

    Michaelis, R

    C. Michaelis, R. Ciosk, and K. Nasmyth, Cohesins: Chro- mosomal proteins that prevent premature separation of sister chromatids, Cell91, 35–45 (1997)

    work page 1997

  9. [9]

    Pradhan, T

    B. Pradhan, T. Kanno, M. Umeda Igarashi, M. S. Loke, M. D. Baaske, J. S. K. Wong, K. Jeppsson, C. Bj¨ orkegren, and E. Kim, The smc5/6 complex is a dna loop-extruding motor, Nature616, 843 (2023)

    work page 2023

  10. [10]

    Takaki, A

    R. Takaki, A. Dey, G. Shi, and D. Thirumalai, Theory and simulations of condensin mediated loop extrusion in dna, Nature Communications12, 5865 (2021)

    work page 2021

  11. [11]

    J. F. Marko, P. De Los Rios, A. Barducci, and S. Gruber, DNA-segment-capture model for loop extrusion by struc- tural maintenance of chromosome (SMC) protein com- plexes, Nucleic acids research47, 6956 (2019)

    work page 2019

  12. [12]

    Hassler, I

    M. Hassler, I. A. Shaltiel, and C. H. Haering, Towards a unified model of smc complex function, Current Biology 28, R1266 (2018)

    work page 2018

  13. [14]

    Dekker, C

    C. Dekker, C. H. Haering, J.-M. Peters, and B. D. Row- land, How do molecular motors fold the genome?, Science 382, 646 (2023)

    work page 2023

  14. [15]

    Ganji, I

    M. Ganji, I. A. Shaltiel, S. Bisht, E. Kim, A. Kalichava, C. H. Haering, and C. Dekker, Real-time imaging of dna loop extrusion by condensin, Science360, 102 (2018)

    work page 2018

  15. [16]

    I. F. Davidson, B. Bauer, D. Goetz, W. Tang, G. Wutz, and J.-M. Peters, DNA loop extrusion by human cohesin, Science366, 1338 (2019)

    work page 2019

  16. [17]

    Y. Kim, Z. Shi, H. Zhang, I. J. Finkelstein, and H. Yu, Human cohesin compacts DNA by loop extrusion, Sci- ence366, 1345 (2019)

    work page 2019

  17. [19]

    Alipour and J

    E. Alipour and J. F. Marko, Self-organization of do- main structures by DNA-loop-extruding enzymes, Nu- cleic acids research40, 11202 (2012)

    work page 2012

  18. [20]

    Fudenberg, M

    G. Fudenberg, M. Imakaev, C. Lu, A. Goloborodko, N. Abdennur, and L. A. Mirny, Formation of chromo- somal domains by loop extrusion, Cell reports15, 2038 (2016)

    work page 2038

  19. [21]

    Revyakin, C

    A. Revyakin, C. Liu, R. H. Ebright, and T. R. Strick, Abortive initiation and productive initiation by rna polymerase involve DNA scrunching, Science314, 1139 (2006)

    work page 2006

  20. [22]

    A. N. Kapanidis, E. Margeat, S. O. Ho, E. Kortkhonjia, S. Weiss, and R. H. Ebright, Initial transcription by RNA polymerase proceeds through a DNA-scrunching mecha- nism, Science314, 1144 (2006)

    work page 2006

  21. [23]

    A. Dey, G. Shi, R. Takaki, and D. Thirumalai, Structural changes in chromosomes driven by multiple condensin motors during mitosis, Cell Reports42(2023)

    work page 2023

  22. [24]

    Golfier, T

    S. Golfier, T. Quail, H. Kimura, and J. Brugu´ es, Cohesin and condensin extrude dna loops in a cell cycle-dependent manner, Elife9, e53885 (2020)

    work page 2020

  23. [25]

    R. C. Tolman, The principle of microscopic reversibility, Proceedings of the National Academy of Sciences11, 436 (1925)

    work page 1925

  24. [26]

    M. L. Mugnai, C. Hyeon, M. Hinczewski, and D. Thiru- malai, Theoretical perspectives on biological machines, Reviews of Modern Physics92, 025001 (2020)

    work page 2020

  25. [27]

    A. B. Kolomeisky and M. E. Fisher, Molecular motors: a theorist’s perspective, Annu. Rev. Phys. Chem.58, 675 (2007)

    work page 2007

  26. [28]

    Ryu, S.-H

    J.-K. Ryu, S.-H. Rah, R. Janissen, J. W. Kerssemakers, A. Bonato, D. Michieletto, and C. Dekker, Condensin extrudes DNA loops in steps up to hundreds of base pairs that are generated by ATP binding events, Nucleic acids research50, 820 (2022)

    work page 2022

  27. [29]

    Yamamoto and H

    T. Yamamoto and H. Schiessel, Osmotic mechanism of the loop extrusion process, Physical Review E96, 10.1103/physreve.96.030402 (2017)

    work page doi:10.1103/physreve.96.030402 2017

  28. [30]

    Brackley, J

    C. Brackley, J. Johnson, D. Michieletto, A. Morozov, M. Nicodemi, P. Cook, and D. Marenduzzo, Nonequilib- rium chromosome looping via molecular slip links, Phys- ical Review Letters119, 10.1103/physrevlett.119.138101 (2017)

    work page doi:10.1103/physrevlett.119.138101 2017

  29. [31]

    C. A. Brackley, J. Johnson, D. Michieletto, A. N. Mo- rozov, M. Nicodemi, P. R. Cook, and D. Marenduzzo, Extrusion without a motor: a new take on the loop ex- trusion model of genome organization, Nucleus9, 95–103 (2018)

    work page 2018

  30. [32]

    Bonato and D

    A. Bonato and D. Michieletto, Three-dimensional loop extrusion, Biophysical Journal120, 5544–5552 (2021)

    work page 2021

  31. [33]

    T. L. Higashi, G. Pobegalov, M. Tang, M. I. Molodtsov, and F. Uhlmann, A brownian ratchet model for dna loop extrusion by the cohesin complex, eLife10, 10.7554/elife.67530 (2021)

    work page doi:10.7554/elife.67530 2021

  32. [34]

    Dembo, D

    M. Dembo, D. Torney, K. Saxman, and D. Hammer, The reaction-limited kinetics of membrane-to-surface ad- hesion and detachment, Proceedings of the Royal Society of London. Series B. Biological Sciences234, 55 (1988)

    work page 1988

  33. [35]

    Barsegov and D

    V. Barsegov and D. Thirumalai, Dynamics of unbinding of cell adhesion molecules: Transition from catch to slip bonds, Proc. Natl. Acad. Sci. USA102, 1835 (2005)

    work page 2005

  34. [36]

    Barsegov and D

    V. Barsegov and D. Thirumalai, Dynamic competition between catch and slip bonds in selectins bound to lig- ands, The Journal of Physical Chemistry B110, 26403 (2006)

    work page 2006

  35. [37]

    Thomas, V

    W. Thomas, V. Vogel, and E. Sokurenko, Biophysics of catch bonds, Annu. Rev. Biophys.37, 399 (2008)

    work page 2008

  36. [38]

    B. T. Marshall, M. Long, J. W. Piper, T. Yago, R. P. McEver, and C. Zhu, Direct observation of catch bonds involving cell-adhesion molecules, Nature423, 190 (2003)

    work page 2003

  37. [39]

    D. L. Huang, N. A. Bax, C. D. Buckley, W. I. Weis, and A. R. Dunn, Vinculin forms a directionally asymmetric 14 catch bond with F-actin, Science357, 703 (2017)

    work page 2017

  38. [40]

    V. C. Luca, B. C. Kim, C. Ge, S. Kakuda, D. Wu, M. Roein-Peikar, R. S. Haltiwanger, C. Zhu, T. Ha, and K. C. Garcia, Notch-jagged complex structure implicates a catch bond in tuning ligand sensitivity, Science355, 1320 (2017)

    work page 2017

  39. [41]

    Choi and C

    H.-K. Choi and C. Zhu, Catch bonds in immunology, An- nual Review of Immunology43(2025)

    work page 2025

  40. [42]

    S. M. Block, Kinesin motor mechanics: Binding, step- ping, tracking, gating and limping, Biophys. J.92, 2986 (2007)

    work page 2007

  41. [43]

    Svoboda and S

    K. Svoboda and S. M. Block, Force and velocity measured for single kinesin molecules, Cell77, 773 (1994)

    work page 1994

  42. [44]

    Svoboda and S

    K. Svoboda and S. M. Block, Biological applications of optical forces, Annual review of biophysics and biomolec- ular structure23, 247 (1994)

    work page 1994

  43. [45]

    E. L. Holzbaur and Y. E. Goldman, Coordination of molecular motors: from in vitro assays to intracellular dynamics, Current opinion in cell biology22, 4 (2010)

    work page 2010

  44. [46]

    Isojima, R

    H. Isojima, R. Iino, Y. Niitani, H. Noji, and M. Tomishige, Direct observation of intermediate states during the stepping motion of kinesin-1, Nature chemical biology12, 290 (2016)

    work page 2016

  45. [47]

    J.-K. Ryu, A. J. Katan, E. O. van der Sluis, T. Wisse, R. de Groot, C. H. Haering, and C. Dekker, The con- densin holocomplex cycles dynamically between open and collapsed states, Nature structural & molecular biology 27, 1134 (2020)

    work page 2020

  46. [48]

    Yildiz, M

    A. Yildiz, M. Tomishige, R. D. Vale, and P. R. Selvin, Kinesin walks hand-over-hand, Science303, 676 (2004)

    work page 2004

  47. [49]

    A. B. Kolomeisky and M. E. Fisher, A simple kinetic model describes the processivity of myosin-V, Biophysi- cal journal84, 1642 (2003)

    work page 2003

  48. [50]

    Kodera, D

    N. Kodera, D. Yamamoto, R. Ishikawa, and T. Ando, Video imaging of walking myosin v by high-speed atomic force microscopy, Nature468, 72 (2010)

    work page 2010

  49. [51]

    Hinczewski, R

    M. Hinczewski, R. Tehver, and D. Thirumalai, Design principles governing the motility of yosin V, Proceedings of the National Academy of Sciences110, E4059 (2013)

    work page 2013

  50. [52]

    G. I. Bell, Models for the specific adhesion of cells to cells, Science200, 618 (1978)

    work page 1978

  51. [53]

    T. R. Strick, T. Kawaguchi, and T. Hirano, Real-time de- tection of single-molecule DNA compaction by condensin I, Current biology14, 874 (2004)

    work page 2004

  52. [54]

    D. T. Gillespie, Exact stochastic simulation of coupled chemical reactions, The journal of physical chemistry81, 2340 (1977)

    work page 1977

  53. [55]

    H. T. Vu, S. Chakrabarti, M. Hinczewski, and D. Thiru- malai, Discrete step sizes of molecular motors lead to bimodal non-gaussian velocity distributions under force, Physical review letters117, 078101 (2016)

    work page 2016

  54. [56]

    Dembo, Lectures on mathematics in the life sciences, some mathematical problems in biology, American Math- ematical Society, Providence, RI51, 51 (1994)

    M. Dembo, Lectures on mathematics in the life sciences, some mathematical problems in biology, American Math- ematical Society, Providence, RI51, 51 (1994)

    work page 1994

  55. [57]

    Chakrabarti, M

    S. Chakrabarti, M. Hinczewski, and D. Thirumalai, Phe- nomenological and microscopic theories for catch bonds, Journal of structural biology197, 50 (2017)

    work page 2017

  56. [58]

    Harder, A.-K

    A. Harder, A.-K. M¨ oller, F. Milz, P. Neuhaus, V. Wal- horn, T. Dierks, and D. Anselmetti, Catch bond interac- tion between cell-surface sulfatase sulf1 and glycosamino- glycans, Biophysical journal108, 1709 (2015)

    work page 2015

  57. [60]

    Chakrabarti, M

    S. Chakrabarti, M. Hinczewski, and D. Thiru- malai, Plasticity of hydrogen bond networks regulates mechanochemistry of cell adhesion complexes, Proceed- ings of the National Academy of Sciences111, 9048 (2014)

    work page 2014

  58. [61]

    C. O. Barkan and R. F. Bruinsma, Topology of molecular deformations induces triphasic catch bonding in selectin– ligand bonds, Proceedings of the National Academy of Sciences121, e2315866121 (2024)

    work page 2024

  59. [62]

    C. M. Johnson, J. D. Fenn, A. Brown, and P. Jung, Dy- namic catch-bonding generates the large stall forces of cytoplasmic dynein, Physical biology17, 046004 (2020)

    work page 2020

  60. [63]

    Pobegalov, L.-Y

    G. Pobegalov, L.-Y. Chu, J.-M. Peters, and M. I. Molodtsov, Single cohesin molecules generate force by two distinct mechanisms, Nature Communications14, 3946 (2023)

    work page 2023

  61. [64]

    Ha and D

    B.-Y. Ha and D. Thirumalai, A mean-field model for semiflexible chains, The Journal of chemical physics103, 9408 (1995)

    work page 1995

  62. [65]

    A. L. Nord, E. Gachon, R. Perez-Carrasco, J. A. Nirody, A. Barducci, R. M. Berry, and F. Pedaci, Catch bond drives stator mechanosensitivity in the bacterial flagellar motor, Proceedings of the National Academy of Sciences 114, 12952 (2017)

    work page 2017

  63. [66]

    M. Doi, S. F. Edwards, and S. F. Edwards,The theory of polymer dynamics, Vol. 73 (oxford university press, 1988)

    work page 1988