Microalgal motility through day/night cycles
We have characterised the motility of the swimming microalga Chlamydomonas reinhardtii as a function of day/night cycles, to which the microalgal growth is entrained. Intriguingly, we find that the microalgae swim almost twice as fast during the night than during the day. I will connect this result with the bioenergetics of flagellar propulsion, discussing consequences for the distributions of cells in lab-based and environmental water columns.
Coordinated motion of active filaments on spherical surfaces
Filaments (slender, microscopic elastic bodies) are prevalent in biological and industrial settings. In the biological case, the filaments are often active, in that they are driven internally by motor proteins, with the prime examples being cilia and flagella. For cilia in particular, which can appear in dense arrays, their resulting motions are coupled through the surrounding fluid, as well as through surfaces to which they are attached. In this talk, I present numerical simulations exploring the coordinated motion of active filaments and how it depends on the driving force, density of filaments, as well as the attached surface. In particular, we find that when the surface is spherical, its topology introduces local defects in coordinated motion which can then feedback and alter the global state. This is particularly true when the surface is not held fixed and is free to move in the surrounding fluid. These simulations take advantage of a computational framework we developed for fully 3D filament motion that combines unit quaternions, implicit geometric time integration, quasi-Newton methods, and fast, matrix-free methods for hydrodynamic interactions and it will also be presented.
Swimming and crawling of Euglena gracilis: a tale with many twists
Euglena gracilis is an interesting unicellular protist, also because it can adopt different motility strategies: swimming by flagellar propulsion, or crawling thanks to large amplitude shape changes of the whole body (a behavior known as “metaboly”, or “amoeboid motion”). Swimming trajectories are helical. The are powered by the beating of a single emerging flagellum, which spans non-planar waveforms in the shape of a twisted lasso. Finally the harmoniously coordinated shape changes that make metaboly possible, reminiscent of peristaltic waves, arise form the relative sliding of its pellicle strips, resulting in twisted helical bundles. We will report on the most recent findings on these interconnected topics, for which helical shapes provide a striking fil rouge.
Trapping active particles up to the limiting case: bacteria enclosed in a biofilm
Active matter systems are composed of constituents, each one in nonequilibrium, that consume energy in order to move [1]. A characteristic feature of active matter is collective motion leading to nonequilibrium phase transitions or large scale directed motion [2]. A number of recent works have featured active particles interacting with obstacles, either moving or fixed [3,4,5]. When an active particle encounters an asymmetric obstacle, different behaviours are detected depending on the nature of its active motion. On the one side, rectification effects arise in a suspension of run-and-tumble particles interacting with a wall of funnelled-shaped openings, caused by particles persistence length [6]. The same trapping mechanism could be responsible for the intake of microorganisms in the underground leaves [7] of Carnivorous plants [8]. On the other side, for aligning particles [9] interacting with a wall of funnelled-shaped openings, trapping happens on the (opposite) wider opening side of the funnels [10,11]. Interestingly, when funnels are located on a circular array, trapping is more localised and depends on the nature of the Vicsek model. Active particles can be synthetic (such as synthetic active colloids) or alive (such as living bacteria). A prototypical model to study living microswimmers is P. fluorescens, a rod shaped and biofilm forming bacterium. Biofilms are microbial communities self-assembled onto external interfaces. Biofilms can be described within the Soft Matter physics framework [12] as a viscoelastic material consisting of colloids (bacterial cells) embedded in a cross-linked polymer gel (polysaccharides cross-linked via proteins/multivalent cations), whose water content vary depending on the environmental conditions. Bacteria embedded in the polymeric matrix control biofilm structure and mechanical properties by regulating its matrix composition. We have recently monitored structural features of Pseudomonas fluorescens biofilms grown with and without hydrodynamic stress [13,14]. We have demonstrated that bacteria are capable of self-adapting to hostile hydrodynamic stress by tailoring the biofilm chemical composition, thus affecting both the mesoscale structure of the matrix and its viscoelastic properties that ultimately regulate the bacteria-polymer interactions. REFERENCES [1] C. Bechinger et al. Rev. Mod. Phys. 88, 045006 (2016); [2] T. Vicsek, A. Zafeiris Phys. Rep. 517, 71 (2012); [3] C. Bechinger, R. Di Leonardo, H. Lowen, C. Reichhardt, G. Volpe, and G. Volpe, Reviews of Modern Physics 88, 045006 (2016); [4] R Martinez, F Alarcon, DR Rodriguez, JL Aragones, C Valeriani The European Physical Journal E 41, 1 (2018); [5] DR Rodriguez, F Alarcon, R Martinez, J Ramírez, C Valeriani, Soft matter 16 (5), 1162 (2020); [6] C. O. Reichhardt and C. Reichhardt, Annual Review of Condensed Matter Physics 8, 51 (2017); [7] W Barthlott, S Porembski, E Fischer, B Gemmel Nature 392, 447 (1998); [8] C B. Giuliano, R Zhang, R.Martinez Fernandez, C.Valeriani and L.Wilson (in preparation, 2021); [9] R Martinez, F Alarcon, JL Aragones, C Valeriani Soft matter 16 (20), 4739 (2020); [10] P. Galajada, J. Keymer, P. Chaikin and R.Austin, Journal of bacteriology, 189, 8704 (2007); [11] M. Wan, C.O. Reichhardt, Z. Nussinov, and C. Reichhardt, Physical Review Letters 101, 018102 (2008); [12] J N. Wilking , T E. Angelini , A Seminara , M P. Brenner , and D A. Weitz MRS Bulletin 36, 385 (2011); [13]J Jara, F Alarcón, A K Monnappa, J Ignacio Santos, V Bianco, P Nie, M Pica Ciamarra, A Canales, L Dinis, I López-Montero, C Valeriani, B Orgaz, Frontiers in microbiology 11, 3460 (2021); [14] P Nie, F Alarcon, I López-Montero, B Orgaz, C Valeriani, M Pica Ciamarra
Microorganism locomotion in viscoelastic fluids
Many microorganisms and cells function in complex (non-Newtonian) fluids, which are mixtures of different materials and exhibit both viscous and elastic stresses. For example, mammalian sperm swim through cervical mucus on their journey through the female reproductive tract, and they must penetrate the viscoelastic gel outside the ovum to fertilize. In micro-scale swimming the dynamics emerge from the coupled interactions between the complex rheology of the surrounding media and the passive and active body dynamics of the swimmer. We use computational models of swimmers in viscoelastic fluids to investigate and provide mechanistic explanations for emergent swimming behaviors. I will discuss how flexible filaments (such as flagella) can store energy from a viscoelastic fluid to gain stroke boosts due to fluid elasticity. I will also describe 3D simulations of model organisms such as C. Reinhardtii and mammalian sperm, where we use experimentally measured stroke data to separate naturally coupled stroke and fluid effects. We explore why strokes that are adapted to Newtonian fluid environments might not do well in viscoelastic environments.
Sperm have got the bends
The journey of development begins with sperm swimming through the female reproductive tract en-route to the egg. In order to successfully complete this journey sperm must beat a single flagellum, propelling themselves through a wide range of fluids, from liquified semen to viscous cervical mucus. It is well-known that the beating tail is driven by an array of 9 microtubule doublets surrounding a central pair, with interconnecting dynein motors generating shear forces and driving elastic wave propagation. Despite this knowledge, the exact mechanism by which coordination of these motors drives oscillating waves along the flagellum remains unknown; hypothesised mechanisms include curvature control, sliding control, and geometric clutch. In this talk we will discuss the mechanisms of flagellar bending, and present a simple model of active curvature that is able to produce many of the various sperm waveforms that are seen experimentally, including those in low and high viscosity fluids and after a cell has ‘hyperactivated’ (a chemical process thought to be key for fertilization). We will show comparisons between these simulated waveforms and sperm that have been experimentally tracked, and discuss methods for fitting simulated mechanistic parameters to these real cells.
Free-falling dynamically scaled models: Foraminifera as a test case
The settling speeds of small biological particles influence the distribution of organisms such as plants, corals, and phytoplankton, but these speeds are difficult to quantify without magnification. In this talk, I highlight my novel method, using dynamic scaling principles and 3D printed models to solve this problem. Dynamic scaling involves creating models with differ in size to the original system and match the physical forces acting upon the model to the original system. I discuss the methodology behind the technique and show how it differs to previous works using dynamically scaled models. I show the flexibility of the technique and suggest how it can be applied to other free-falling particles (e.g. seeds and spores).
Hydrodynamic shape of microorganisms: Generalised Jeffery orbits
'Shape' of microorganisms are diverse. However, we sometimes approximate them as a sphere or a spheroid when we mathematically model the hydrodynamics of motile and non-motile cells. Such a geometrical simplification can be theoretically validated for motions in a linear background flow, since the dynamics, known as the Jeffery orbit, only contain a single geometric parameter, called the Bretherton constant. In this talk, we generalise the Jeffery equations for a chiral axisymmetric object using the low-Reynolds-number hydrokinetic symmetry and then demonstrate that the dynamics of a certain type of chiral object in a fluid flow are characterised by a new chiral parameter in addition to the Bretherton constant. We also discuss how the generalised Jeffery orbits are applied to biased locomotion of bacteria in a bulk shear flow and we will share the idea of hydrodynamic `shape' of microorganisms to simplify the description of their dynamics.
Sperm Navigation: from hydrodynamic interactions to parameter estimation
Microorganisms can swim in a variety of environments, interacting with chemicals and other proteins in the fluid. In this talk, we will highlight recent computational methods and results for swimming efficiency and hydrodynamic interactions of swimmers in different fluid environments. Sperm are modeled via a centerline representation where forces are solved for using elastic rod theory. The method of regularized Stokeslets is used to solve the fluid-structure interaction where emergent swimming speeds can be compared to asymptotic analysis. In the case of fluids with extra proteins or cells that may act as friction, swimming speeds may be enhanced, and attraction may not occur. We will also highlight how parameter estimation techniques can be utilized to infer fluid and/or swimmer properties.
Exploring the evolution of motile curved bacteria using a regularized Stokeslet Boundary Element Method and Pareto optimality theory
Bacteria exhibit a bewildering diversity of morphologies, but despite their impact on nearly all aspects of life, they are frequently classified into a few general categories, usually just “spheres” and “rods.” Curved-rod bacteria are one simple variation observed in many environments, particularly the ocean. However, why so many species have evolved this shape is unknown. We used a regularized Stokeslet Boundary Element Method to model the motility of flagellated, curved bacteria. We show that curvature can increase swimming efficiency, revealing a widely applicable selective advantage. Furthermore, we show that the distribution of cell lengths and curvatures observed across bacteria in nature is predicted by evolutionary trade-offs between three tasks influenced by shape: efficient swimming, the ability to detect chemical gradients, and reduced cost of cell construction. We therefore reveal shape as an important component of microbial fitness.
Imposed flow in active liquid crystals
Inspired by ongoing experiments on three dimensional active gels composed of sliding microtubule bundles, we study a few idealized problems in a minimal hydrodynamic model for active liquid crystals. Our aim is to use flow to determine the value of the coefficient of activity in a continuum theory. We consider the case of apolar active particles that form a disordered phase in the absence of flow, and study how activity affects the swimming speed of a prescribed swimmer, as well as the stability of a fluid interface. We also consider flows of active matter in channels or past immersed objects.
Stochastic control of passive colloidal objects by micro-swimmers
The way single colloidal objects behave in presence of active forces arising from within the bulk of the system is crucial to many situations, notably biological and ecological (e.g. intra-cellular transport, predation), and potential medical or environmental applications (e.g. targeted delivery of cargoes, depollution of waters and soils). In this talk I will present experimental findings that my collaborators and I have obtained over the past years on the dynamics of single Brownian colloids in suspensions of biological micro-swimmers, especially the green alga Chlamydomonas reinhardtii. I'll show notably that spatial heterogeneities and anisotropies in the active particles statistics can control the preferential localisation of their passive counterparts. The results will be rationalized using theoretical approaches from hydrodynamics and stochastic processes.
The impact of elongation on transport in shear flow
I shall present two recent piece of work investigating how shape effects the transport of active particles in shear. Firstly we will consider the sedimentation of particles in 2D laminar flow fields of increasing complexity; and how insights from this can help explain why turbulence can enhance the sedimentation of negatively buoyant diatoms [1]. Secondly, we will consider the 3D transport of elongated active particles under the action of an aligning force (e.g. gyrotactic swimmers) in some simple flow fields; and will see how shape can influence the vertical distribution, for example changing the structure of thin layers [2]. [1] Enhanced sedimentation of elongated plankton in simple flows (2018). IMA Journal of Applied Mathematics W Clifton, RN Bearon, & MA Bees. [2] Elongation enhances migration through hydrodynamic shear (in Prep), RN Bearon & WM Durham.
Transport and dispersion of active particles in complex porous media
Understanding the transport of microorganisms and self-propelled particles in porous media has important consequences in human health as well as for microbial ecology. In this work, we explore models for the dispersion of active particles in both periodic and random porous media. In a first problem, we analyze the long-time transport properties in a dilute system of active Brownian particles swimming in a periodic lattice in the presence of an external flow. Using generalized Taylor dispersion theory, we calculate the mean transport velocity and dispersion dyadic and explain their dependence on flow strength, swimming activity and geometry. In a second approach, we address the case of run-and-tumble particles swimming through unstructured porous media composed of randomly distributed circular pillars. There, we show that the long-time dispersion is described by a universal hindrance function that depends on the medium porosity and ratio of the swimmer run length to the pillar size. An asymptotic expression for the hindrance function is derived in dilute media, and its extension to semi-dilute and dense media is obtained using stochastic simulations. We conclude by discussing the role of hydrodynamic interactions and swimmer concentration effects.
Motility-dependent pathogenicity of a spirochetal bacterium
Motility is a crucial virulence factor for many species of bacteria, but it is not fully understood how bacterial motility is practically involved in pathogenicity. This time I will give a talk on the association of motility with pathogenicity in the zoonotic spirochete bacterium Leptospira. Recently, we measured swimming force of individual leptospires using optical tweezers and found that they can generate ~30 times of the swimming force of E. coli. We also observed that leptospires increase the reversal frequency of swimming at the gel-liquid interface, resembling host dermis exposed to contaminated water (Abe et al., 2020, Sci Rep). These could be involved in percutaneous infection of the spirochete. We have shown that Leptospira not only swims in liquid but also moves over solid surfaces (Tahara et al., 2018, Sci Adv). We quantified the surface motility called “crawling” on cultured kidney tissues from various mammals, showing that pathogenic leptospires crawl over the tissue surfaces more persistently that non-pathogenic ones (Xu et al., 2020, Front Microbiol). I will discuss the spirochete motility related to pathogenicity from the biophysical viewpoint.
Motility control in biological microswimmers
It is often assumed that biological swimmers conform faithfully to certain stereotypes assigned to them by physicists and mathematicians, when the reality is in fact much more complicated. In this talk we will use a combination of theory, experiments, and robotics, to understand the physical and evolutionary basis of motility control in a number of distinguished organisms. These organisms differ markedly in terms of their size, shape, and arrangement of locomotor appendages, but are united in their use of cilia - the ultimate shape-shifting organelle - to achieve self-propulsion and navigation.
Continuum modelling of active fluids beyond the generalised Taylor dispersion
The Smoluchowski equation has often been used as the starting point of many continuum models of active suspensions. However, its six-dimensional nature depending on time, space and orientation requires a huge computational cost, fundamentally limiting its use for large-scale problems, such as mixing and transport of active fluids in turbulent flows. Despite the singular nature in strain-dominant flows, the generalised Taylor dispersion (GTD) theory (Frankel & Brenner 1991, J. Fluid Mech. 230:147-181) has been understood to be one of the most promising ways to reduce the Smoluchowski equation into an advection-diffusion equation, the mean drift and diffusion tensor of which rely on ‘local’ flow information only. In this talk, we will introduce an exact transformation of the Smoluchowski equation into such an advection-diffusion equation requiring only local flow information. Based on this transformation, a new advection-diffusion equation will subsequently be proposed by taking an asymptotic analysis in the limit of small particle velocity. With several examples, it will be demonstrated that the new advection-diffusion model, non-singular in strain-dominant flows, outperforms the GTD theory.
Synthetic swimmers: microorganism swimming without microorganisms
The effect of non Newtonian liquid rheology on the swimming performance of microorganisms is still poorly understood, despite numerous recent studies. In our effort to clarify some aspects of this problem, we have developed a series of magnetic synthetic swimmers that self-propel immersed in a fluid by emulating the swimming strategy of flagellated microorganisms. With these devices, it is possible to control some aspects of the motion with the objective to isolate specific effects. In this talk, recent results on the effects of shear-thinning viscosity and viscoelasticity on the motion of helical swimmers will presented and discussed. Also, a number of other new uses of the synthetic swimmers will be presented including swimming across gradients, swimming in sand, interactions and rheometry.