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The cytoskeleton–a composite network of biopolymers, molecular motors, and associated binding proteins–is a paradigmatic example of active matter. Particle transport through the cytoskeleton can range from anomalous and heterogeneous subdiffusion to superdiffusion and advection. Yet, recapitulating and understanding these properties–ubiquitous to the cytoskeleton and other out-of-equilibrium soft matter systems–remains challenging. Here, we combine light sheet microscopy with differential dynamic microscopy and single-particle tracking to elucidate anomalous and advective transport in actomyosin-microtubule composites. We show that particles exhibit multi-mode transport that transitions from pronounced subdiffusion to superdiffusion at tunable crossover timescales. Surprisingly, while higher actomyosin content increases the range of timescales over which transport is superdiffusive, it also markedly increases the degree of subdiffusion at short timescales and generally slows transport. Corresponding displacement distributions display unique combinations of non-Gaussianity, asymmetry, and non-zero modes, indicative of directed advection coupled with caged diffusion and hopping. At larger spatiotemporal scales, particles in active composites exhibit superdiffusive dynamics with scaling exponents that are robust to changing actomyosin fractions, in contrast to normal, yet faster, diffusion in networks without actomyosin. Our specific results shed important new light on the interplay between non-equilibrium processes, crowding and heterogeneity in active cytoskeletal systems. More generally, our approach is broadly applicable to active matter systems to elucidate transport and dynamics across scales.
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Investigating Superdiffusive Shock Acceleration at a Parallel Shock with a Fractional Parker Equation for Energetic-particle Interaction with Small-scale Magnetic Flux Ropes
Abstract It has been suggested before that small-scale magnetic flux rope (SMFR) structures in the solar wind can temporarily trap energetic charged particles. We present the derivation of a new fractional Parker equation for energetic-particle interaction with SMFRs from our pitch-angle-dependent fractional diffusion-advection equation that can account for such trapping effects. The latter was derived previously in le Roux & Zank from the first principles starting with the standard focused transport equation. The new equation features anomalous advection and diffusion terms. It suggests that energetic-particle parallel transport occurs with a decaying efficiency of advection effects as parallel superdiffusion becomes more dominant at late times. Parallel superdiffusion can be linked back to underlying anomalous pitch-angle transport, which might be subdiffusive during interaction with quasi-helical coherent SMFRs. We apply the new equation to time-dependent superdiffusive shock acceleration at a parallel shock. The results show that the superdiffusive-shock-acceleration timescale is fractional, the net fractional differential particle flux is conserved across the shock ignoring particle injection at the shock, and the accelerated particle spectrum at the shock converges to the familiar power-law spectrum predicted by standard steady-state diffusive-shock-acceleration theory at late times. Upstream, as parallel superdiffusion progressively dominates the advection of energetic particles, their spatial distributions decay on spatial scales that grow with time. Furthermore, superdiffusive parallel shock acceleration is found to be less efficient if parallel anomalous diffusion is more superdiffusive, while perpendicular particle escape from the shock, thought to be subdiffusive during SMFR interaction, is reduced when increasingly subdiffusive.
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- Award ID(s):
- 2148653
- PAR ID:
- 10366823
- Publisher / Repository:
- DOI PREFIX: 10.3847
- Date Published:
- Journal Name:
- The Astrophysical Journal
- Volume:
- 930
- Issue:
- 2
- ISSN:
- 0004-637X
- Format(s):
- Medium: X Size: Article No. 125
- Size(s):
- Article No. 125
- Sponsoring Org:
- National Science Foundation
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