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A semi-flexible model prediction for the polymerization force exerted by a living F-actin filament on a fixed wall

par Pierleoni, Carlo ;Ciccotti, Giovanni;Ryckaert, Jean-Paul
Référence The Journal of Chemical Physics, 143, 14, 145101
Publication Publié, 2015-10

Article révisé par les pairs

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Résumé :

We consider a single living semi-flexible filament with persistence length ℓp in chemical equilibrium with a solution of free monomers at fixed monomer chemical potential μ1 and fixed temperature T. While one end of the filament is chemically active with single monomer (de)polymerization steps, the other end is grafted normally to a rigid wall to mimic a rigid network from which the filament under consideration emerges. A second rigid wall, parallel to the grafting wall, is fixed at distance L < < ℓp from the filament seed. In supercritical conditions where monomer density ρ1 is higher than the critical density ρ1c, the filament tends to polymerize and impinges onto the second surface which, in suitable conditions (non-escaping filament regime), stops the filament growth. We first establish the grand-potential Ω(μ1, T, L) of this system treated as an ideal reactive mixture, and derive some general properties, in particular the filament size distribution and the force exerted by the living filament on the obstacle wall. We apply this formalism to the semi-flexible, living, discrete Wormlike chain model with step size d and persistence length ℓp, hitting a hard wall. Explicit properties require the computation of the mean force f i (L) exerted by the wall at L and associated potential f i (L) = - d W i (L) / d L on a filament of fixed size i. By original Monte-Carlo calculations for few filament lengths in a wide range of compression, we justify the use of the weak bending universal expressions of Gholami et al. [Phys. Rev. E 74, 041803 (2006)] over the whole non-escaping filament regime. For a filament of size i with contour length Lc = (i - 1) d, this universal form is rapidly growing from zero (non-compression state) to the buckling value f b (L c, ℓ p) = π 2 k B T ℓ p 4 L c 2 over a compression range much narrower than the size d of a monomer. Employing this universal form for living filaments, we find that the average force exerted by a living filament on a wall at distance L is in practice L independent and very close to the value of the stalling force F s H = (k B T / d) ln (ρ 1) predicted by Hill, this expression being strictly valid in the rigid filament limit. The average filament force results from the product of the cumulative size fraction x = x (L, ℓ p, ρ 1), where the filament is in contact with the wall, times the buckling force on a filament of size Lc ≈ L, namely, F s H = x f b (L; ℓ p). The observed L independence of F s H implies that x ∝ L-2 for given (ℓ p, ρ 1) and x ∝ ln ρ 1 for given (ℓp, L). At fixed (L, ρ 1), one also has x ∝ ℓ p - 1 which indicates that the rigid filament limit ℓp → ∞ is a singular limit in which an infinite force has zero weight. Finally, we derive the physically relevant threshold for filament escaping in the case of actin filaments.