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Mechanical Regulation of Neurite Polarization and Growth: A Computational Study.

ABSTRACT: The densely packed microtubule (MT) array found in neuronal cell projections (neurites) serves two fundamental functions simultaneously: it provides a mechanically stable track for molecular motor-based transport and produces forces that drive neurite growth. The local pattern of MT polarity along the neurite shaft has been found to differ between axons and dendrites. In axons, the neurons' dominating long projections, roughly 90% of the MTs orient with their rapidly growing plus end away from the cell body, whereas in vertebrate dendrites, their orientations are locally mixed. Molecular motors are known to be responsible for cytoskeletal ordering and force generation, but their collective function in the dense MT cytoskeleton of neurites remains elusive. We here hypothesized that both the polarity pattern of MTs along the neurite shaft and the shaft's global extension are simultaneously driven by molecular motor forces and should thus be regulated by the mechanical load acting on the MT array as a whole. To investigate this, we simulated cylindrical bundles of MTs that are cross-linked and powered by molecular motors by iteratively solving a set of force-balance equations. The bundles were subjected to a fixed load arising from actively generated tension in the actomyosin cortex enveloping the MTs. The magnitude of the load and the level of motor-induced connectivity between the MTs have been varied systematically. With an increasing load and decreasing motor-induced connectivity between MTs, the bundles became wider in cross section and extended more slowly, and the local MT orientational order was reduced. These results reveal two, to our knowledge, novel mechanical factors that may underlie the distinctive development of the MT cytoskeleton in axons and dendrites: the cross-linking level of MTs by motors and the load acting on this cytoskeleton during growth.

SUBMITTER: Jakobs MAH

PROVIDER: S-EPMC7175593 | biostudies-literature |

REPOSITORIES: biostudies-literature

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Project description:Metamaterials, rationally designed multiscale composite systems, have attracted extensive interest because of their potential for a broad range of applications due to their unique properties such as negative Poisson's ratio, exceptional mechanical performance, tunable photonic and phononic properties, structural reconfiguration, etc. Though they are dominated by an auxetic structure, the constituents of metamaterials also play an indispensable role in determining their unprecedented properties. In this vein, 2D materials such as graphene, silicene, and phosphorene with superior structural tunability are ideal candidates for constituents of metamaterials. However, the nanostructure-property relationship and composition-property relationship of these 2D material-based metamaterials remain largely unexplored. Mechanical anisotropy inherited from the 2D material constituents, for example, may substantially impact the physical stability and robustness of the corresponding metamaterial systems. Herein, classical molecular dynamics simulations are performed using a generic coarse-grained model to explore the deformation mechanism of these 2D material-based metamaterials with sinusoidally curved ligaments and the effect of mechanical anisotropy on mechanical properties, especially the negative Poisson's ratio. The results indicate that deformation under axial tensile load can be divided into two stages: bending-dominated and stretching-dominated, in which the rotation of junctions in the former stage results in auxetic behavior of the proposed metamaterials. In addition, the auxetic behavior depends heavily on both the amplitude/wavelength ratio of the sinusoidal ligament and the stiffness ratio between axial and transverse directions. The magnitude of negative Poisson's ratio increases from 0 to 0.625, with an associated increase of the amplitude/wavelength ratio from 0 to 0.225, and fluctuates at around 0.625, in good agreement with the literature, with amplitude/wavelength ratios greater than 0.225. More interestingly, the magnitude of negative Poisson's ratio increases from 0.47 to 0.87 with the increase of the stiffness ratio from 0.125 to 8, in good agreement with additional all-atom molecular dynamics simulations for phosphorene and molybdenum disulfide. Overall, these research findings shed light on the deformation mechanism of auxetic metamaterials, providing useful guidelines for designing auxetic 2D lattice structures made of 2D materials that can display a tunable negative Poisson's ratio.

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Mechanical Regulation of Neurite Polarization and Growth: A Computational Study.

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