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We study the surface fluctuations of a tissue with a dynamics dictated by cell-rearrangement, cell-division, and cell-death processes. Surface fluctuations are calculated in the homeostatic state, where cell division and cell death equilibrate on average. The obtained fluctuation spectrum can be mapped onto several other spectra such as those characterizing incompressible fluids, compressible Maxwell elastomers, or permeable membranes in appropriate asymptotic regimes. Since cell division and cell death are out-of-equilibrium processes, detailed balance is broken, but a generalized fluctuation-response relation is satisfied in terms of appropriate observables. Our work is a first step toward the description of the out-of-equilibrium fluctuations of the surface of a thick epithelium and its dynamical response to external perturbations.
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Many microorganisms and artificial microswimmers use helical appendages in order to generate locomotion. Though often rotated so as to produce thrust, some species of bacteria such Spiroplasma, Rhodobacter sphaeroides and Spirochetes induce movement by deforming a helical-shaped body. Recently, artificial devices have been created which also generate motion by deforming their helical body in a non-reciprocal way (Mourran et al., Adv. Mater., 29, 1604825, 2017). Inspired by these systems, we investigate the transport of a deforming helix within a viscous fluid. Specifically, we consider a swimmer that maintains a helical centreline and a single handedness while changing its helix radius, pitch and wavelength uniformly across the body. We first discuss how a deforming helix can create a non-reciprocal translational and rotational swimming stroke and identify its principle direction of motion. We then determine the leading-order physics for helices with small helix radius before considering the general behaviour for different configuration parameters and how these swimmers can be optimised. Finally, we explore how the presence of walls, gravity, and defects in the centreline allow the helical device to break symmetries, increase its speed, and generate transport in directions not available to helices in bulk fluids.
Groups of beating flagella or cilia often synchronize so that neighboring filaments have identical frequencies and phases. A prime example is provided by the unicellular biflagellate Chlamydomonas reinhardtii, which typically displays synchronous in-phase beating in a low-Reynolds number version of breaststroke swimming. We report here the discovery that ptx1, a flagellar dominance mutant of C. reinhardtii, can exhibit synchronization in precise antiphase, as in the freestyle swimming stroke. Long-duration high-speed imaging shows that ptx1 flagella switch stochastically between in-phase and antiphase states, and that the latter has a distinct waveform and significantly higher frequency, both of which are strikingly similar to those found during phase slips that stochastically interrupt in-phase beating of the wildtype. Possible mechanisms underlying these observations are discussed.
Sheep are gregarious animals, and they often aggregate into dense, cohesive flocks, especially under stress. In this paper, we use image processing tools to analyze a publicly available aerial video showing a dense sheep flock moving under the stimulus of a shepherding dog. Inspired by the fluidity of the motion, we implement a hydrodynamics approach, extracting velocity fields, and measuring their propagation and correlations in space and time. We find that while the flock overall is stationary, significant dynamics happens at the edges, notably in the form of fluctuations propagating like waves, and large-scale correlations spanning the entire flock. These observations highlight the importance of incorporating interfacial dynamics, for instance in the form of line tension, when using a hydrodynamics framework to model the dynamics of dense, non-polarized swarms.
Compared to agile legged animals, wheeled and tracked vehicles often suffer large performance loss on granular surfaces like sand and gravel. Understanding the mechanics of legged locomotion on granular media can aid the development of legged robots with improved mobility on granular surfaces; however, no general force model yet exists for granular media to predict ground reaction forces during complex limb intrusions. Inspired by a recent study of sand-swimming, we develop a resistive force model in the vertical plane for legged locomotion on granular media. We divide an intruder of complex morphology and kinematics, e.g., a bio-inspired robot L-leg rotated through uniform granular media (loosely packed ~ 1 mm diameter poppy seeds), into small segments, and measure stresses as a function of depth, orientation, and direction of motion using a model leg segment. Summation of segmental forces over the intruder predicts the net forces on both an L-leg and a reversed L-leg rotated through granular media with better accuracy than using simple one-dimensional penetration and drag force models. A multi-body dynamic simulation using the resistive force model predicts the speeds of a small legged robot (15 cm, 150 g) moving on granular media using both L-legs and reversed L-legs.
Interfaces between stratified epithelia and their supporting stromas commonly exhibit irregular shapes. Undulations are particularly pronounced in dysplastic tissues and typically evolve into long, finger-like protrusions in carcinomas. In a previous work (Basan et al., Phys. Rev. Lett. 106, 158101 (2011)), we demonstrated that an instability arising from viscous shear stresses caused by the constant flow due to cell turnover in the epithelium could drive this phenomenon. While interfacial tension between the two tissues as well as mechanical resistance of the stroma tend to maintain a flat interface, an instability occurs for sufficiently large viscosity, cell-division rate and thickness of the dividing region in the epithelium. Here, extensions of this work are presented, where cell division in the epithelium is coupled to the local concentration of nutrients or growth factors diffusing from the stroma. This enhances the instability by a mechanism similar to that of the Mullins-Sekerka instability in single-diffusion processes of crystal growth. We furthermore present the instability for the generalized case of a viscoelastic stroma.
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