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Register a new userBistability in the synchronization of actuated microfilaments
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Cilia and flagella are essential building blocks for biological fluid transport and locomotion at the micron scale. They often beat in synchrony and may transition between different synchronization modes in the same cell type. Here, we investigate the behavior of elastic microfilaments, protruding from a surface and driven at their base by a configuration-dependent torque. We consider full hydrodynamic interactions among and within filaments and no slip at the surface. Isolated filaments exhibit periodic deformations, with increasing waviness and frequency as the magnitude of the driving torque increases. Two nearby but independently-driven filaments synchronize their beating in-phase or anti-phase. This synchrony arises autonomously via the interplay between hydrodynamic coupling and filament elasticity. Importantly, in-phase and anti-phase synchronization modes are bistable and co-exist for a range of driving torques and separation distances. These findings are consistent with experimental observations of in-phase and anti-phase synchronization in the biflagellate textit{Chlamydomonas reinhardtii} and could have important implications on understanding the biophysical mechanisms underlying transitions between multiple synchronization modes.
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Cilia and flagella are highly conserved slender organelles that exhibit a variety of rhythmic beating patterns from non-planar cone-like motions to planar wave-like deformations. Although their internal structure, composed of a microtubule-based axoneme driven by dynein motors, is known, the mechanism responsible for these beating patterns remains elusive. Existing theories suggest that the dynein activity is dynamically regulated, via a geometric feedback from the ciliums mechanical deformation to the dynein force. An alternative, open-loop mechanism based on a flutter instability was recently proven to lead to planar oscillations of elastic filaments under follower forces. Here, we show that an elastic filament in viscous fluid, clamped at one end and acted on by an external distribution of compressive axial forces, exhibits a Hopf bifurcation that leads to non-planar spinning of the buckled filament at a locked curvature. We also show the existence of a second bifurcation, at larger force values, that induces a transition from non-planar spinning to planar wave-like oscillations. We elucidate the nature of these instabilities using a combination of nonlinear numerical analysis, linear stability theory, and low-order bead-spring models. Our results show that away from the transition thresholds, these beating patterns are robust to perturbations in the distribution of axial forces and in the filament configuration. These findings support the theory that an open-loop, instability-driven mechanism could explain both the sustained oscillations and the wide variety of periodic beating patterns observed in cilia and flagella.
Buoyancy-driven exchange flows are common to a variety of natural and engineering systems ranging from persistently active volcanoes to counterflows in oceanic straits. Experiments of exchange flows in closed vertical tubes have been used as surrogates to elucidate the basic features of such flows. The resulting data have historically been analyzed and interpreted through core-annular flow solutions, the most common flow configuration at finite viscosity contrasts. These models have been successful in fitting experimental data, but less effective at explaining the variability observed in natural systems. In this paper, we formulate a core-annular solution to the classical problem of buoyancy-driven exchange flows in vertical tubes. The model posits the existence of two mathematically valid solutions, i.e. thin- and thick-core solutions. The theoretical existence of two solutions, however, does not necessarily imply that the system is bistable in the sense that flow switching may occur. Using direct numerical simulations, we test the hypothesis that core-annular flow in vertical tubes is bistable, which implies that the realized flow field is not uniquely defined by the material parameters of the flow. Our numerical experiments, which fully predict experimental data without fitting parameters, demonstrate that buoyancy-driven exchange flows are indeed inherently bistable systems. This finding is consistent with previous experimental data, but in contrast to the underlying hypothesis of previous analytical models that the solution is unique and can be identified by maximizing the flux or extremizing the dissipation in the system. These results have important implications for data interpretation by analytical models, and may also have relevant ramifications for understanding volcanic degassing.
Periodically forced, oscillatory fluid flows have been the focus of intense research for decades due to their richness as a nonlinear dynamical system and their relevance to applications in transportation, aeronautics, and energy conversion. Recently, it has been observed that turbulent bluff-body wakes exhibit a subharmonic resonant response when excited with specific spatial symmetries at twice the natural vortex shedding frequency, which is hypothesized to be caused by triadic interactions. The focus of this paper is to provide new physical insight into the dynamics of turbulent oscillator flows, based on improved mechanistic models informed by a comprehensive experimental study of the turbulent wake behind a D-shaped body under periodic forcing. We confirm for the first time the role of resonant triadic interactions in the forced flow by studying the dominant components in the power spectra across multiple excitation frequencies and amplitudes. We then develop an extended Stuart-Landau model for the forced global wake mode, incorporating parametric and non-harmonic forcing. This model captures the system dynamics and reveals the boundaries of multiple synchronization regions. Further, it is possible to identify model coefficients from sparse measurement data, making it applicable to a wide range of turbulent oscillator flows. We believe these generalized synchronization models will be valuable for prediction, control, and understanding of the underlying physics in this ubiquitous class of flows.
The human cortex is never at rest but in a state of sparse and noisy neural activity that can be detected at broadly diverse resolution scales. It has been conjectured that such a state is best described as a critical dynamical process -- whose nature is still not fully understood -- where scale-free avalanches of activity emerge at the edge of a phase transition. In particular, some works suggest that this is most likely a synchronization transition, separating synchronous from asynchronous phases. Here, by investigating a simplified model of coupled excitable oscillators describing the cortex dynamics at a mesoscopic level, we investigate the possible nature of such a synchronization phase transition. Within our modeling approach, we conclude that -- in order to reproduce all key empirical observations, such as scale-free avalanches and bistability, on which fundamental functional advantages rely -- the transition to collective oscillatory behavior needs to be of an unconventional hybrid type, with mixed features of type-I and type-II excitability, opening the possibility for a particularly rich dynamical repertoire.
The ultimate goal of a sound theory of turbulence in fluids is to close in a rational way the Reynolds equations, namely to express the tensor of turbulent stress as a function of the time average of the velocity field. Based on the idea that dissipation in fully developed turbulence is by singular events resulting from an evolution described by the Euler equations, it has been recently observed that the closure problem is strongly restricted, and that it implies that the turbulent stress is a non local function in space of the average velocity field, a kind of extension of classical Boussinesq theory of turbulent viscosity. This leads to rather complex nonlinear integral equation(s) for the time averaged velocity field. This one satisfies some symmetries of the Euler equations. Such symmetries were used by Prandtl and Landau to make various predictions about the shape of the turbulent domain in simple geometries. We explore specifically the case of mixing layer for which the average velocity field only depends on the angle in the wedge behind the splitter plate. This solution yields a pressure difference between the two sides of the splitter which contributes to the lift felt by the plate. Moreover, because of the structure of the equations for the turbulent stress, one can satisfy the Cauchy-Schwarz inequalities, also called the realizability conditions, for this turbulent stress. Such realizability conditions cannot be satisfied with a simple turbulent viscosity.
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