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Register a new userGlobal metachronal synchronization and active noise in cilia carpets
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Added by
Benjamin M. Friedrich
Publication date
2020
fields
Physics
and research's language is
English
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Carpets of actively bending cilia can exhibit self-organized metachronal coordination. Past research proposed synchronization by hydrodynamic coupling, but if such coupling is strong enough to overcome active phase noise had been addressed only for pairs of cilia. Using a multi-scale model calibrated by experimental cilia beat patterns, we find local multi-stability of wave modes. Yet, a single mode, corresponding to a dexioplectic wave, has predominant basin-of-attraction. Beyond a characteristic noise strength, we observe an abrupt loss of global synchronization even in finite systems.
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We calculate the hydrodynamic flow field generated far from a cilium which is attached to a surface and beats periodically. In the case of two beating cilia, hydrodynamic interactions can lead to synchronization of the cilia, which are nonlinear oscillators. We present a state diagram where synchronized states occur as a function of distance of cilia and the relative orientation of their beat. Synchronized states occur with different relative phases. In addition, asynchronous solutions exist. Our work could be relevant for the synchronized motion of cilia generating hydrodynamic flows on the surface of cells.
Cilia and flagella are hair-like extensions of eukaryotic cells which generate oscillatory beat patterns that can propel micro-organisms and create fluid flows near cellular surfaces. The evolutionary highly conserved core of cilia and flagella consists of a cylindrical arrangement of nine microtubule doublets, called the axoneme. The axoneme is an actively bending structure whose motility results from the action of dynein motor proteins cross-linking microtubule doublets and generating stresses that induce bending deformations. The periodic beat patterns are the result of a mechanical feedback that leads to self-organized bending waves along the axoneme. Using a theoretical framework to describe planar beating motion, we derive a nonlinear wave equation that describes the fundamental Fourier mode of the axonemal beat. We study the role of nonlinearities and investigate how the amplitude of oscillations increases in the vicinity of an oscillatory instability. We furthermore present numerical solutions of the nonlinear wave equation for different boundary conditions. We find that the nonlinear waves are well approximated by the linearly unstable modes for amplitudes of beat patterns similar to those observed experimentally.
We study synchronization of locally coupled noisy phase oscillators which move diffusively in a one-dimensional ring. Together with the disordered and the globally synchronized states, the system also exhibits several wave-like states which display local order. We use a statistical description valid for a large number of oscillators to show that for any finite system there is a critical spatial diffusion above which all wave-like solutions become unstable. Through Langevin simulations, we show that the transition to global synchronization is mediated by the relative size of attractor basins associated to wave-like states. Spatial diffusion disrupts these states and paves the way for the system to attain global synchronization.
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.
Sperm swimming at low Reynolds number have strong hydrodynamic interactions when their concentration is high in vivo or near substrates in vitro. The beating tails not only propel the sperm through a fluid, but also create flow fields through which sperm interact with each other. We study the hydrodynamic interaction and cooperation of sperm embedded in a two-dimensional fluid by using a particle-based mesoscopic simulation method, multi-particle collision dynamics (MPC). We analyze the sperm behavior by investigating the relationship between the beating-phase difference and the relative sperm position, as well as the energy consumption. Two effects of hydrodynamic interaction are found, synchronization and attraction. With these hydrodynamic effects, a multi-sperm system shows swarm behavior with a power-law dependence of the average cluster size on the width of the distribution of beating frequencies.
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