Flexible filaments and fibres are essential components of important complex fluids that appear in many biological and industrial settings. Direct simulations of these systems that capture the motion and deformation of many immersed filaments in suspension remain a formidable computational challenge due to the complex, coupled fluid-structure interactions of all filaments, the numerical stiffness associated with filament bending, and the various constraints that must be maintained as the filaments deform. In this paper, we address these challenges by first describing filament kinematics using quaternions to resolve both bending and twisting, applying implicit time-integration to alleviate numerical stiffness, and using quasi-Newton methods to obtain solutions to the resulting system of nonlinear equations. In particular, we employ geometric time integration to ensure that the quaternions remain unit as the filaments move. We also show that our framework can be used with a variety of models and methods, including matrix-free fast methods, that resolve low Reynolds number hydrodynamic interactions. We provide a series of tests and example simulations to demonstrate the performance and possible applications of our method. Finally, we provide a link to a MATLAB/Octave implementation of our framework that can be used to learn more about our approach and as a tool for filament simulation.
Swimming cells and microorganisms are as diverse in their collective dynamics as they are in their individual shapes and propulsion mechanisms. Even for sperm cells, which have a stereotyped shape consisting of a cell body connected to a flexible flagellum, a wide range of collective dynamics is observed spanning from the formation of tightly packed groups to the display of larger-scale, turbulence-like motion. Using a detailed mathematical model that resolves flagellum dynamics, we perform simulations of sperm suspensions containing up to 1000 cells and explore the connection between individual and collective dynamics. We find that depending on the level of variation in individual dynamics from one swimmer to another, the sperm exhibit either a strong tendency to aggregate, or the suspension exhibits large-scale swirling. Hydrodynamic interactions govern the formation and evolution of both states. In addition, a quantitative analysis of the states reveals that the flows generated at the time scale of flagellum undulations contribute significantly to the overall energy in the surrounding fluid, highlighting the importance of resolving these flows.
While only a single sperm may fertilize the egg, getting to the egg can be facilitated, and possibly enhanced, by sperm group dynamics. Examples range from the trains formed by wood mouse sperm to the bundles exhibited by echidna sperm. In addition, observations of wave-like patterns exhibited by ram semen are used to score prospective sample fertility for artificial insemination in agriculture. In this review, we discuss these experimental observations of collective dynamics, as well as describe recent mechanistic models that link the motion of individual sperm cells and their flagella to observed collective dynamics. Establishing this link in models involves negotiating the disparate time- and length scales involved, typically separated by a factor of 1000, to capture the dynamics at the greatest length scales affected by mechanisms at the shortest time scales. Finally, we provide some outlook on the subject, in particular, the open questions regarding how collective dynamics impacts fertility. This article is part of the theme issue ‘Multi-scale analysis and modelling of collective migration in biological systems’.
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