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Register a new userActin filaments pushing against a barrier: Comparison between two force generation mechanisms
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To theoretically understand force generation properties of actin filaments, many models consider growing filaments pushing against a movable obstacle or barrier. In order to grow, the filaments need space and hence it is necessary to move the barrier. Two different mechanisms for this growth are widely considered in literature. In one class of models (type $A$), the filaments can directly push the barrier and move it, thereby performing some work in the process. In another type of models (type $B$), the filaments wait till thermal fluctuations of the barrier position create enough space between the filament tip and the barrier, and then they grow by inserting one monomer in that gap. The difference between these two types of growth seems microscopic and rather a matter of modelling details. However, we find that this difference has important effect on many qualitative features of the models. In particular, how the relative time-scale between the barrier dynamics and filament dynamics influences the force generation properties, are significantly different for type $A$ and $B$ models. We illustrate these differences for three types of barrier: a rigid wall-like barrier, an elastic barrier and a barrier with Kardar-Parisi-Zhang dynamics. Our numerical simulations match well with our analytical calculations. Our study highlights the importance of taking the details of filament-barrier interaction into account while modelling force generation properties of actin filaments.
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We study the force generation by a set of parallel actin filaments growing against an elastic membrane. The elastic membrane tries to stay flat and any deformation from this flat state, either caused by thermal fluctuations or due to protrusive polymerization force exerted by the filaments, costs energy. We study two lattice models to describe the membrane dynamics. In one case, the energy cost is assumed to be proportional to the absolute magnitude of the height gradient (gradient model) and in the other case it is proportional to the square of the height gradient (Gaussian model). For the gradient model we find that the membrane velocity is a non-monotonic function of the elastic constant $mu$, and reaches a peak at $mu=mu^ast$. For $mu < mu^ast$ the system fails to reach a steady state and the membrane energy keeps increasing with time. For the Gaussian model, the system always reaches a steady state and the membrane velocity decreases monotonically with the elastic constant $ u$ for all nonzero values of $ u$. Multiple filaments give rise to protrusions at different regions of the membrane and the elasticity of the membrane induces an effective attraction between the two protrusions in the Gaussian model which causes the protrusions to merge and a single wide protrusion is present in the system. In both the models, the relative time-scale between the membrane and filament dynamics plays an important role in deciding whether the shape of elasticity-velocity curve is concave or convex. Our numerical simulations agree reasonably well with our analytical calculations.
We investigate the mechanical interplay between the spatial organization of the actin cytoskeleton and the shape of animal cells adhering on micropillar arrays. Using a combination of analytical work, computer simulations and in vitro experiments, we demonstrate that the orientation of the stress fibers strongly influences the geometry of the cell edge. In the presence of a uniformly aligned cytoskeleton, the cell edge can be well approximated by elliptical arcs, whose eccentricity reflects the degree of anisotropy of the cells internal stresses. Upon modeling the actin cytoskeleton as a nematic liquid crystal, we further show that the geometry of the cell edge feeds back on the organization of the stress fibers by altering the length scale at which these are confined. This feedback mechanism is controlled by a dimensionless number, the anchoring number, representing the relative weight of surface-anchoring and bulk-aligning torques. Our model allows to predict both cellular shape and the internal structure of the actin cytoskeleton and is in good quantitative agreement with experiments on fibroblastoid (GD$beta$1,GD$beta$3) and epithelioid (GE$beta$1, GE$beta$3) cells.
By exerting mechanical force it is possible to unfold/refold RNA molecules one at a time. In a small range of forces, an RNA molecule can hop between the folded and the unfolded state with force-dependent kinetic rates. Here, we introduce a mesoscopic model to analyze the hopping kinetics of RNA hairpins in an optical tweezers setup. The model includes different elements of the experimental setup (beads, handles and RNA sequence) and limitations of the instrument (time lag of the force-feedback mechanism and finite bandwidth of data acquisition). We investigated the influence of the instrument on the measured hopping rates. Results from the model are in good agreement with the experiments reported in the companion article (1). The comparison between theory and experiments allowed us to infer the values of the intrinsic molecular rates of the RNA hairpin alone and to search for the optimal experimental conditions to do the measurements. We conclude that long handles and soft laser traps represent the best conditions to extract rate estimates that are closest to the intrinsic molecular rates. The methodology and rationale presented here can be applied to other experimental setups and other molecules.
Respiration in bacteria involves a sequence of energetically-coupled electron and proton transfers creating an electrochemical gradient of protons (a proton-motive force) across the inner bacterial membrane. With a simple kinetic model we analyze a redox loop mechanism of proton-motive force generation mediated by a molecular shuttle diffusing inside the membrane. This model, which includes six electron-binding and two proton-binding sites, reflects the main features of nitrate respiration in E. coli bacteria. We describe the time evolution of the proton translocation process. We find that the electron-proton electrostatic coupling on the shuttle plays a significant role in the process of energy conversion between electron and proton components. We determine the conditions where the redox loop mechanism is able to translocate protons against the transmembrane voltage gradient above 200 mV with a thermodynamic efficiency of about 37%, in the physiologically important range of temperatures from 250 to 350 K.
We consider a one-dimensional elastic membrane, which is pushed by growing filaments. The filaments tend to grow by creating local protrusions in the membrane and this process has surface energy and bending energy costs. Although it is expected that with increasing surface tension and bending rigidity, it should become more difficult to create a protrusion, we find that for a fixed bending rigidity, as the surface tension increases, protrusions are more easily formed. This effect also gives rise to nontrivial dependence of membrane velocity on the surface tension, characterized by a dip and a peak. We explain this unusual phenomenon by studying in detail the interplay of the surface and the bending energy and show that this interplay is responsible for a qualitative shape change of the membrane, which gives rise to the above effect.
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