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Many-body interactions in systems of active matter can cause particles to move collectively and self-organize into dynamic structures with long-range order. In cells, the self-assembly of cytoskeletal filaments is critical for cellular motility, structure, intracellular transport, and division. Semiflexible cytoskeletal filaments driven by polymerization or motor-protein interactions on a two-dimensional substrate, such as the cell cortex, can induce filament bending and curvature leading to interesting collective behavior. For example, the bacterial cell-division filament FtsZ is known to have intrinsic curvature that causes it to self-organize into rings and vortices, and recent experiments reconstituting the collective motion of microtubules driven by motor proteins on a surface have observed chiral symmetry breaking of the collective behavior due to motor-induced curvature of the filaments. Previous work on the self-organization of driven filament systems have not studied the effects of curvature and filament structure on collective behavior. In this work, we present Brownian dynamics simulation results of driven semiflexible filaments with intrinsic curvature and investigate how the interplay between filament rigidity and radius of curvature can tune the self-organization behavior in homochiral systems and heterochiral mixtures. We find a curvature-induced reorganization from polar flocks to self-sorted chiral clusters, which is modified by filament flexibility. This transition changes filament transport from ballistic to diffusive at long timescales.
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In cells, cytoskeletal filament networks are responsible for cell movement, growth, and division. Filaments in the cytoskeleton are driven and organized by crosslinking molecular motors. In reconstituted cytoskeletal systems, motor activity is responsible for far-from-equilibrium phenomena such as active stress, self-organized flow, and spontaneous nematic defect generation. How microscopic interactions between motors and filaments lead to larger-scale dynamics remains incompletely understood. To build from motor-filament interactions to predict bulk behavior of cytoskeletal systems, more computationally efficient techniques for modeling motor-filament interactions are needed. Here, we derive a coarse-graining hierarchy of explicit and continuum models for crosslinking motors that bind to and walk on filament pairs. We compare the steady-state motor distribution and motor-induced filament motion for the different models and analyze their computational cost. All three models agree well in the limit of fast motor binding kinetics. Evolving a truncated moment expansion of motor density speeds the computation by [Formula: see text]-[Formula: see text] compared to the explicit or continuous-density simulations, suggesting an approach for more efficient simulation of large networks. These tools facilitate further study of motor-filament networks on micrometer to millimeter length scales.
Assuntos
Citoesqueleto/metabolismo , Modelos Biológicos , Proteínas Motores Moleculares/metabolismo , Cinética , Microtúbulos/metabolismoRESUMO
In active matter systems, self-propelled particles can self-organize to undergo collective motion, leading to persistent dynamical behavior out of equilibrium. In cells, cytoskeletal filaments and motor proteins form complex structures important for cell mechanics, motility, and division. Collective dynamics of cytoskeletal systems can be reconstituted using filament gliding experiments, in which cytoskeletal filaments are propelled by surface-bound motor proteins. These experiments have observed diverse dynamical states, including flocks, polar streams, swirling vortices, and single-filament spirals. Recent experiments with microtubules and kinesin motor proteins found that the collective behavior of gliding filaments can be tuned by altering the concentration of the crowding macromolecule methylcellulose in solution. Increasing the methylcellulose concentration reduced filament crossing, promoted alignment, and led to a transition from active, isotropically oriented filaments to locally aligned polar streams. This emergence of collective motion is typically explained as an increase in alignment interactions by Vicsek-type models of active polar particles. However, it is not yet understood how steric interactions and bending stiffness modify the collective behavior of active semiflexible filaments. Here we use simulations of driven filaments with tunable soft repulsion and rigidity in order to better understand how the interplay between filament flexibility and steric effects can lead to different active dynamic states. We find that increasing filament stiffness decreases the probability of filament alignment, yet increases collective motion and long-range order, in contrast to the assumptions of a Vicsek-type model. We identify swirling flocks, polar streams, buckling bands, and spirals, and describe the physics that govern transitions between these states. In addition to repulsion and driving, tuning filament stiffness can promote collective behavior, and controls the transition between active isotropic filaments, locally aligned flocks, and polar streams.
Assuntos
Citoesqueleto , Microtúbulos , Cinesinas , Movimento (Física) , MiosinasRESUMO
Newly processed global imaging and topographic mapping of Uranus's five major satellites reveal differences and similarities to mid-sized satellites at Saturn and Pluto. Three modes of internal heat redistribution are recognized. The broad similarity of Miranda's three oval resurfacing zones to those mapped on Enceladus and (subtly) on Dione are likely due to antipodal diapiric upwelling. Conversely, break-up and foundering of crustal blocks accompanied by extensive (cryo)volcanism is the dominant mode on both Charon and Ariel. Titania's fault network finds parallels on Rhea, Dione, Tethys and possibly Oberon. Differences in the geologic style of resurfacing in the satellite systems (e.g. plains on Charon, Dione, Tethys and perhaps Titania versus ridges on Miranda and Ariel) may be driven by differences in ice composition. Surface processes such as volatile transport may also be indicated by bright and dark materials on Oberon, Umbriel and Charon. The more complete and higher quality observations of the Saturnian and Plutonian mid-sized icy satellites by Cassini and New Horizons reveal a wealth of features and phenomena that cannot be perceived in the more limited Voyager coverage of the Uranian satellites, harbingers of many discoveries awaiting us on a return to Uranus. This article is part of a discussion meeting issue 'Future exploration of ice giant systems'.
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The Radar for Europa Assessment and Sounding: Ocean to Near-surface (REASON) is a dual-frequency ice-penetrating radar (9 and 60 MHz) onboard the Europa Clipper mission. REASON is designed to probe Europa from exosphere to subsurface ocean, contributing the third dimension to observations of this enigmatic world. The hypotheses REASON will test are that (1) the ice shell of Europa hosts liquid water, (2) the ice shell overlies an ocean and is subject to tidal flexing, and (3) the exosphere, near-surface, ice shell, and ocean participate in material exchange essential to the habitability of this moon. REASON will investigate processes governing this material exchange by characterizing the distribution of putative non-ice material (e.g., brines, salts) in the subsurface, searching for an ice-ocean interface, characterizing the ice shell's global structure, and constraining the amplitude of Europa's radial tidal deformations. REASON will accomplish these science objectives using a combination of radar measurement techniques including altimetry, reflectometry, sounding, interferometry, plasma characterization, and ranging. Building on a rich heritage from Earth, the moon, and Mars, REASON will be the first ice-penetrating radar to explore the outer solar system. Because these radars are untested for the icy worlds in the outer solar system, a novel approach to measurement quality assessment was developed to represent uncertainties in key properties of Europa that affect REASON performance and ensure robustness across a range of plausible parameters suggested for the icy moon. REASON will shed light on a never-before-seen dimension of Europa and - in concert with other instruments on Europa Clipper - help to investigate whether Europa is a habitable world.
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The New Horizons spacecraft returned images and compositional data showing that terrains on Pluto span a variety of ages, ranging from relatively ancient, heavily cratered areas to very young surfaces with few-to-no impact craters. One of the regions with very few impact craters is dominated by enormous rises with hummocky flanks. Similar features do not exist anywhere else in the imaged solar system. Here we analyze the geomorphology and composition of the features and conclude this region was resurfaced by cryovolcanic processes, of a type and scale so far unique to Pluto. Creation of this terrain requires multiple eruption sites and a large volume of material (>104 km3) to form what we propose are multiple, several-km-high domes, some of which merge to form more complex planforms. The existence of these massive features suggests Pluto's interior structure and evolution allows for either enhanced retention of heat or more heat overall than was anticipated before New Horizons, which permitted mobilization of water-ice-rich materials late in Pluto's history.
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The surface of Pluto is more geologically diverse and dynamic than had been expected, but the role of its tenuous atmosphere in shaping the landscape remains unclear. We describe observations from the New Horizons spacecraft of regularly spaced, linear ridges whose morphology, distribution, and orientation are consistent with being transverse dunes. These are located close to mountainous regions and are orthogonal to nearby wind streaks. We demonstrate that the wavelength of the dunes (~0.4 to 1 kilometer) is best explained by the deposition of sand-sized (~200 to ~300 micrometer) particles of methane ice in moderate winds (<10 meters per second). The undisturbed morphology of the dunes, and relationships with the underlying convective glacial ice, imply that the dunes have formed in the very recent geological past.
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New Horizons images of Pluto's companion Charon show a variety of terrains that display extensional tectonic features, with relief surprising for this relatively small world. These features suggest a global extensional areal strain of order 1% early in Charon's history. Such extension is consistent with the presence of an ancient global ocean, now frozen.
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NASA's New Horizons spacecraft has revealed the complex geology of Pluto and Charon. Pluto's encounter hemisphere shows ongoing surface geological activity centered on a vast basin containing a thick layer of volatile ices that appears to be involved in convection and advection, with a crater retention age no greater than ~10 million years. Surrounding terrains show active glacial flow, apparent transport and rotation of large buoyant water-ice crustal blocks, and pitting, the latter likely caused by sublimation erosion and/or collapse. More enigmatic features include tall mounds with central depressions that are conceivably cryovolcanic and ridges with complex bladed textures. Pluto also has ancient cratered terrains up to ~4 billion years old that are extensionally faulted and extensively mantled and perhaps eroded by glacial or other processes. Charon does not appear to be currently active, but experienced major extensional tectonism and resurfacing (probably cryovolcanic) nearly 4 billion years ago. Impact crater populations on Pluto and Charon are not consistent with the steepest impactor size-frequency distributions proposed for the Kuiper belt.
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The dusty jovian ring system must be replenished continuously from embedded source bodies. The New Horizons spacecraft has performed a comprehensive search for kilometer-sized moons within the system, which might have revealed the larger members of this population. No new moons were found, however, indicating a sharp cutoff in the population of jovian bodies smaller than 8-kilometer-radius Adrastea. However, the search revealed two families of clumps in the main ring: one close pair and one cluster of three to five. All orbit within a brighter ringlet just interior to Adrastea. Their properties are very different from those of the few other clumpy rings known; the origin and nonrandom distribution of these features remain unexplained, but resonant confinement by Metis may play a role.