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Odd dynamics of living chiral crystals
The emergent dynamics exhibited by collections of living organisms often shows signatures of symmetries that are broken at the single-organism level. At the same time, organism development itself encompasses a well-coordinated sequence of symmetry breaking events that successively transform a single, nearly isotropic cell into an animal with well-defined body axis and various anatomical asymmetries. Combining these key aspects of collective phenomena and embryonic development, we describe here the spontaneous formation of hydrodynamically stabilized active crystals made of hundreds of starfish embryos that gather during early development near fluid surfaces. We describe a minimal hydrodynamic theory that is fully parameterized by experimental measurements of microscopic interactions among embryos. Using this theory, we can quantitatively describe the stability, formation and rotation of crystals and rationalize the emergence of mechanical properties that carry signatures of an odd elastic material. Our work thereby quantitatively connects developmental symmetry breaking events on the single-embryo level with remarkable macroscopic material properties of a novel living chiral crystal system.
Active mechanics of sea star oocytes
The cytoskeleton has the remarkable ability to self-organize into active materials which underlie diverse cellular processes ranging from motility to cell division. Actomyosin is a canonical example of an active material, which generates cellularscale contractility in part through the forces exerted by myosin motors on actin filaments. While the molecular players underlying actomyosin contractility have been well characterized, how cellular-scale deformation in disordered actomyosin networks emerges from filament-scale interactions is not well understood. In this talk, I’ll present work done in collaboration with Sebastian Fürthauer and Nikta Fakhri addressing this question in vivo using the meiotic surface contraction wave seen in oocytes of the bat star Patiria miniata as a model system. By perturbing actin polymerization, we find that the cellular deformation rate is a nonmonotonic function of cortical actin density peaked near the wild type density. To understand this, we develop an active fluid model coarse-grained from filament-scale interactions and find quantitative agreement with the measured data. The model makes further predictions, including the surprising prediction that deformation rate decreases with increasing motor concentration. We test these predictions through protein overexpression and find quantitative agreement. Taken together, this work is an important step for bridging the molecular and cellular length scales for cytoskeletal networks in vivo.
Membrane mechanics meet minimal manifolds
Changes in the geometry and topology of self-assembled membranes underlie diverse processes across cellular biology and engineering. Similar to lipid bilayers, monolayer colloidal membranes studied by the Sharma (IISc Bangalore) and Dogic (UCSB) Labs have in-plane fluid-like dynamics and out-of-plane bending elasticity, but their open edges and micron length scale provide a tractable system to study the equilibrium energetics and dynamic pathways of membrane assembly and reconfiguration. First, we discuss how doping colloidal membranes with short miscible rods transforms disk-shaped membranes into saddle-shaped minimal surfaces with complex edge structures. Theoretical modeling demonstrates that their formation is driven by increasing positive Gaussian modulus, which in turn is controlled by the fraction of short rods. Further coalescence of saddle-shaped surfaces leads to exotic topologically distinct structures, including shapes similar to catenoids, tri-noids, four-noids, and higher order structures. We then mathematically explore the mechanics of these catenoid-like structures subject to an external axial force and elucidate their intimate connection to two problems whose solutions date back to Euler: the shape of an area-minimizing soap film and the buckling of a slender rod under compression. A perturbation theory argument directly relates the tensions of membranes to the stability properties of minimal surfaces. We also investigate the effects of including a Gaussian curvature modulus, which, for small enough membranes, causes the axial force to diverge as the ring separation approaches its maximal value.
The Equation of State of a Tissue
An equation of state is something you hear about in introductory thermodynamics, for example, the Ideal gas equation. The ideal gas equation relates the pressure, volume, and the number of particles of the gas, to its temperature, uniquely defining its state. This description is possible in physics when the system under investigation is in equilibrium or near equilibrium. In biology, a tissue is modeled as a fluid composed of cells. These cells are constantly interacting with each other through mechanical and chemical signaling, driving them far from equilibrium. Can an equation of state exist for such a messy interacting system? In this talk, I show that the presence of strong cell-cell interaction in tissues gives rise to a novel non-equilibrium, size-dependent surface tension, something unheard of for classical fluids. This surface tension, in turn, modifies the packing of cells inside the tissue generating a size-dependent density and pressure. Finally, we show that a combination of these non-equilibrium pressure and densities can yield an equation of state for biological tissues arbitrarily far from equilibrium. In the end, I discuss how this new paradigm of size-dependent biological properties gives rise to novel modes of cellular motion in tissues
Exact coherent structures and transition to turbulence in a confined active nematic
Active matter describes a class of systems that are maintained far from equilibrium by driving forces acting on the constituent particles. Here I will focus on confined active nematics, which exhibit especially rich flow behavior, ranging from structured patterns in space and time to disordered turbulent flows. To understand this behavior, I will take a deterministic dynamical systems approach, beginning with the hydrodynamic equations for the active nematic. This approach reveals that the infinite-dimensional phase space of all possible flow configurations is populated by Exact Coherent Structures (ECS), which are exact solutions of the hydrodynamic equations with distinct and regular spatiotemporal structure; examples include unstable equilibria, periodic orbits, and traveling waves. The ECS are connected by dynamical pathways called invariant manifolds. The main hypothesis in this approach is that turbulence corresponds to a trajectory meandering in the phase space, transitioning between ECS by traveling on the invariant manifolds. Similar approaches have been successful in characterizing high Reynolds number turbulence of passive fluids. Here, I will present the first systematic study of active nematic ECS and their invariant manifolds and discuss their role in characterizing the phenomenon of active turbulence.
Towards model-based control of active matter: active nematics and oscillator networks
The richness of active matter's spatiotemporal patterns continues to capture our imagination. Shaping these emergent dynamics into pre-determined forms of our choosing is a grand challenge in the field. To complicate matters, multiple dynamical attractors can coexist in such systems, leading to initial condition-dependent dynamics. Consequently, non-trivial spatiotemporal inputs are generally needed to access these states. Optimal control theory provides a general framework for identifying such inputs and represents a promising computational tool for guiding experiments and interacting with various systems in soft active matter and biology. As an exemplar, I first consider an extensile active nematic fluid confined to a disk. In the absence of control, the system produces two topological defects that perpetually circulate. Optimal control identifies a time-varying active stress field that restructures the director field, flipping the system to its other attractor that rotates in the opposite direction. As a second, analogous case, I examine a small network of coupled Belousov-Zhabotinsky chemical oscillators that possesses two dominant attractors, two wave states of opposing chirality. Optimal control similarly achieves the task of attractor switching. I conclude with a few forward-looking remarks on how the same model-based control approach might come to bear on problems in biology.
Metachronal waves in swarms of nematode Turbatrix aceti
There is a recent surge of interest in the behavior of active particles that can at the same time align their direction of movement and synchronize their oscillations, known as swarmalators. While analytical and numerical models of such systems are now abundant, no real-life examples have been shown to date. I will present an experimental investigation of the collective motion of the nematode Turbatrix aceti, which self-propel by body undulation. I will show that under favorable conditions these nematodes can synchronize their body oscillations, forming striking traveling metachronal waves which, similar to the case of beating cilia, produce strong fluid flows. I will demonstrate that the location and strength of this collective state can be controlled through the shape of the confining structure; in our case the contact angle of a droplet. This opens a way for producing controlled work such as on-demand flows or displacement of objects. I will illustrate this by a practical example: showing that the force generated by the collectively moving nematodes is sufficient to change the mode of evaporation of fluid droplets, by counteracting the surface-tension force, which allow us to estimate its strength.
Growing in flows: from evolutionary dynamics to microbial jets
Biological systems can self-organize in complex structures, able to evolve and adapt to widely varying environmental conditions. Despite the importance of fluid flow for transporting and organizing populations, few laboratory systems exist to systematically investigate the impact of advection on their spatial evolutionary dynamics. In this talk, I will discuss how we can address this problem by studying the morphology and genetic spatial structure of microbial colonies growing on the surface of a viscous substrate. When grown on a liquid, I will show that S. cerevisiae (baker’s yeast) can behave like “active matter” and collectively generate a fluid flow many times larger than the unperturbed colony expansion speed, which in turn produces mechanical stresses and fragmentation of the initial colony. Combining laboratory experiments with numerical modeling, I will demonstrate that the coupling between metabolic activity and hydrodynamic flows can produce positive feedbacks and drive preferential growth phenomena leading to the formation of microbial jets. Our work provides rich opportunities to explore the interplay between hydrodynamics, growth and competition within a versatile system.
Theory of activity-powered interface
Interfaces and membranes are ubiquitous in cellular systems across various scales. From lipid membranes to the interfaces of biomolecular condensates inside the cell, these borders not only protect and segregate the inner components from the outside world, but also are actively participating in mechanical regulation and biochemical reaction of the cell. Being part of a living system, these interfaces (membranes) are usually active and away from equilibrium. Yet, it's still not clear how activity can tweak their equilibrium dynamics. Here, I will introduce a model system to tackle this problem. We put together a passive fluid and an active nematics, and study the behavior of this liquid-liquid interface. Whereas thermal fluctuation of such an interface is too weak to be observed, active stress can easily force the interface to fluctuate, overhang, and even break up. In the presence of a wall, the active phase exhibits superfluid-like behavior: it can climb up walls -- a phenomenon we call activity-induced wetting. I will show how to formulate theories to capture these phenomena, highlighting the nontrivial effects of active stress. Our work not only demonstrates that activity can introduce interesting features to an interface, but also sheds light on controlling interfacial properties using activity.
Flow singularities in soft materials: from thermal motion to active molecular stresses
The motion of passive or active agents in soft materials generates long ranged deformation fields with signatures informed by hydrodynamics and the properties of the soft matter host. These signatures are even more complex when the soft matter host itself is an active material. Measurement of these fields reveals mechanics of the soft materials and hydrodynamics central to understanding self-organization. In this talk, I first introduce a new method based on correlated displacement velocimetry, and use the method to measure flow fields around particles trapped at the interface between immiscible fluids. These flow fields, decomposed into interfacial hydrodynamic multipoles, including force monopole and dipole flows, provide key insights essential to understanding the interface’s mechanical response. I then extend this method to various actomyosin systems to measure local strain fields around myosin molecular motors. I show how active stresses propagate in 2d liquid crystalline structures and in disordered networks that are formed by the actin filaments. In particular, the response functions of contractile and stable gels are characterized. Through similar analysis, I also measure the retrograde flow fields of stress fibers in single cells to understand subcellular mechanochemical systems.
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.
Bacterial rheotaxis in bulk and at surfaces
Individual bacteria transported in viscous flows, show complex interactions with flows and bounding surfaces resulting from their complex shape as well as their activity. Understanding these transport dynamics is crucial, as they impact soil contamination, transport in biological conducts or catheters, and constitute thus a serious health threat. Here we investigate the trajectories of individual E-coli bacteria in confined geometries under flow, using microfluidic model systems in bulk flows as well as close to surfaces using a novel Langrangian 3D tracking method. Combining experimental observations and modelling we elucidate the origin of upstream swimming, lateral drift or persistent transport along corners. [1] Junot et al, EPL, 126 (2019) 44003 [2] Mathijssen et al. 10:3 (2019) Nature Comm. [3] Figueroa-Morales et al., Soft Matter, 2015,11, 6284-6293 [4] Darnige et al. Review of Scientific Instruments 88, 055106 (2017) [5] Jing et al, Science Advances, 2020; 6 : eabb2012 [6] Figueroa-Morales et al, Sci. Adv. 2020; 6 : eaay0155, 2020, 10.1126/sciadv.aay0155
The role of the fluid bilayer in kinesin-driven vesicle transport
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.
Flocking through complex environments
The spontaneous collective motion of self-propelled agents is ubiquitous in the natural world, and it often occurs in complex environments, be it bacteria and cells migrating through polymeric extracellular matrix or animal herds and human crowds navigating structured terrains. Much is known about flocking dynamics in pristine backgrounds, but how do spatio-temporal heterogeneities in the environment impact such collective self-organization? I will present two model systems, a colloidal active fluid negotiating disordered obstacles and a confined dense bacterial suspension in a viscoelastic medium, as controllable platforms to explore this question and highlight general mechanisms for active self-organization in complex environments. By combining theory and experiment, I will show how flocks on disordered substrates organize into a novel dynamic vortex glass phase, akin to vortex glasses in dirty superconductors, while the presence of viscoelasticity can calm the otherwise turbulent swarming of bacteria, allowing the emergence of a large scale coherent and even oscillatory vortex when confined on the millimetre scale.
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.
Flocks and crowds: a Gulliver travel
In the first part of my talk, combining experimental, numerical and theoretical results, I will explain how self-propelled colloidal particles self-organize in one of the most robust ordered state found in nature: flocks. I will explain how to describe macroscopic flocking motion as the spontaneous flows of an active fluid, and use this framework to elucidate the phase ordering dynamics of polar active matter. In the second part of my talk, I will show that the same tools and concepts can be effectively used to infer a hydrodynamic description of active fluids composed of particles 6 order of magnitude larger in size: pedestrian crowds.
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.
How can we learn from nature to build better polymer composites?
Nature is replete with extraordinary materials that can grow, move, respond, and adapt. In this talk I will describe our ongoing efforts to develop advanced polymeric materials, inspired by nature. First, I will describe my group’s efforts to develop ultrastiff, ultratough materials inspired by the byssal materials of marine mussels. These adhesive contacts allow mussels to secure themselves to rocks, wood, metals and other surfaces in the harsh conditions of the intertidal zone. By developing a foundational understanding of the structure-mechanics relationships and processing of the natural system, we can design high-performance materials that are extremely strong without compromising extensibility, as well as macroporous materials with tunable toughness and strength. In the second half of the talk, I will describe new efforts to exploit light as a means of remote control and power. By leveraging the phototransduction pathways of highly-absorbing, negatively photochromic molecules, we can drive the motion of amorphous polymeric materials as well as liquid flows. These innovations enable applications in packaging, connective tissue repair, soft robotics, and optofluidics.
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).
Tissue fluidization at the onset of zebrafish gastrulation
Embryo morphogenesis is impacted by dynamic changes in tissue material properties, which have been proposed to occur via processes akin phase transitions (PTs). Here, we show that rigidity percolation provides a simple and robust theoretical framework to predict material/structural PTs of embryonic tissues from local cell connectivity. By using percolation theory, combined with directly monitoring dynamic changes in tissue rheology and cell contact mechanics, we demonstrate that the zebrafish blastoderm undergoes a genuine rigidity PT, brought about by a small reduction in adhesion-dependent cell connectivity below a critical value. We quantitatively predict and experimentally verify hallmarks of PTs, including power-law exponents and associated discontinuities of macroscopic observables at criticality. Finally, we show that this uniform PT depends on blastoderm cells undergoing meta-synchronous divisions causing random and, consequently, uniform changes in cell connectivity. Collectively, our theoretical and experimental findings reveal the structural basis of material PTs in an organismal context.
Life in complex fluids
Inertial active soft matter
Active particles which are self-propelled by converting energy into mechanical motion represent an expanding research realm in physics and chemistry. For micron-sized particles moving in a liquid (``microswimmers''), most of the basic features have been described by using the model of overdamped active Brownian motion [1]. However, for macroscopic particles or microparticles moving in a gas, inertial effects become relevant such that the dynamics is underdamped. Therefore, recently, active particles with inertia have been described by extending the active Brownian motion model to active Langevin dynamics which include inertia [2]. In this talk, recent developments of active particles with inertia (``microflyers'', ``hoppers'' or ``runners'') are summarized including: inertial delay effects between particle velocity and self-propulsion direction [3], tuning of the long-time self-diffusion by the moment of inertia [3], the influence of inertia on motility-induced phase separation and the cluster growth exponent [4], and the formation of active micelles (“rotelles”) by using inertial active surfactants. References [1] C. Bechinger, R. di Leonardo, H. Löwen, C. Reichhardt, G. Volpe, G. Volpe, Reviews of Modern Physics 88, 045006 (2016). [2] H. Löwen, Journal of Chemical Physics 152, 040901 (2020). [3] C. Scholz, S. Jahanshahi, A. Ldov, H. Löwen, Nature Communications 9, 5156 (2018). [4] S. Mandal, B. Liebchen, H. Löwen, Physical Review Letters 123, 228001 (2019). [5] C. Scholz, A. Ldov, T. Pöschel, M. Engel, H. Löwen, Surfactants and rotelles in active chiral fluids, will be published
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.
Mixed active-passive suspensions: from particle entrainment to spontaneous demixing
Understanding the properties of active matter is a challenge which is currently driving a rapid growth in soft- and bio-physics. Some of the most important examples of active matter are at the microscale, and include active colloids and suspensions of microorganisms, both as a simple active fluid (single species) and as mixed suspensions of active and passive elements. In this last class of systems, recent experimental and theoretical work has started to provide a window into new phenomena including activity-induced depletion interactions, phase separation, and the possibility to extract net work from active suspensions. Here I will present our work on a paradigmatic example of mixed active-passive system, where the activity is provided by swimming microalgae. Macro- and micro-scopic experiments reveal that microorganism-colloid interactions are dominated by rare close encounters leading to large displacements through direct entrainment. Simulations and theoretical modelling show that the ensuing particle dynamics can be understood in terms of a simple jump-diffusion process, combining standard diffusion with Poisson-distributed jumps. Entrainment length can be understood within the framework of Taylor dispersion as a competition between advection by the no-slip surface of the cell body and microparticle diffusion. Building on these results, we then ask how external control of the dynamics of the active component (e.g. induced microswimmer anisotropy/inhomogeneity) can be used to alter the transport of passive cargo. As a first step in this direction, we study the behaviour of mixed active-passive systems in confinement. The resulting spatial inhomogeneity in swimmers’ distribution and orientation has a dramatic effect on the spatial distribution of passive particles, with the colloids accumulating either towards the boundaries or towards the bulk of the sample depending on the size of the container. We show that this can be used to induce the system to de-mix spontaneously.
Light-bacteria interactions
In 1676, using candle light and a small glass sphere as the lens, van Leeuwenhoek discovered the microscopic world of living microorganisms. Today, using lasers, spatial light modulators, digital cameras and computers, we study the statistical and fluid mechanics of microswimmers in ways that were unimaginable only 50 years ago. With light we can image swimming bacteria in 3D, apply controllable force fields or sculpt their 3D environment. In addition to shaping the physical world outside cells we can use light to control the internal state of genetically modified bacteria. I will review our recent work with light-bacteria interactions, going from some fundamental problems in the fluid and statistical mechanics of microswimmers to the use of bacteria as propellers for micro-machines or as a "living" paint controlled by light.
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.
Soft matter physics and the COVID-19 pandemic
Much of the science underpinning the global response to the COVID-19 pandemic lies in the soft matter domain. Coronaviruses are composite particles with a core of nucleic acids complexed to proteins surrounded by a protein-studded lipid bilayer shell. A dominant route for transmission is via air-borne aerosols and droplets. Viral interaction with polymeric body fluids, particularly mucus, and cell membranes controls their infectivity, while their interaction with skin and artificial surfaces underpins cleaning and disinfection and the efficacy of masks and other personal protective equipment. The global response to COVID-19 has highlighted gaps in the soft matter knowledge base. I will survey these gaps, especially as pertaining to the transmission of the disease, and suggest questions that can (and need to) be tackled, both in response to COVID-19 and to better prepare for future viral pandemics.
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.
“Rigidity and Fluidity in Biological Tissue”
The coordinated migration of groups of cells underlies many biological processes, including embryo development, wound healing and cancer metastasis. In many of these situations, tissues are able to tune themselves between liquid-like states, where cells flow collectively as in a liquid, and solid-like states that can support shear stresses. In this talk I will describe mesoscopic models of cell assemblies inspired by active matter physics to examine the roles of cell motility, cell crowding and the interplay of contractility and adhesion in controlling the rheological state of biological tissue.
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.
Lab-on-a-chip and diagnostic tools for COVID-19
The SARS-CoV-2 virus has rapidly evolved into a pandemic that is threatening public health, economics, and quality of life worldwide. The gold-standard for testing individuals for COVID-19 is using traditional RT-qPCR, which is expensive and can take up to several hours. Expanding surveillance across a global scale will call for new strategies and tests that are inexpensive, require minimal reagents, decrease assay time, and allow for simple point-of-care (POC) monitoring without need of trained personnel and with quick turnaround time. To expand the speed of COVID-19 surveillance, we are working on a point-of-care microfluidic chip to enable significantly faster and easier testing. This is based upon digital drop loop-mediated isothermal amplification that will allow for rapid testing of large populations at a reasonable cost. The device will employ a nucleic-acid based test called reverse transcriptase LAMP (RT- LAMP) that operates at a temperature of 60-65°C. RT-LAMP removes the bottleneck of thermal cycling and high temperatures required by traditional RT-qPCR thermocycling. The simplicity, speed, and sensitivity will enable early treatment and response to infection.
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.
Spontaneous and driven active matter flows
Understanding individual and macroscopic transport properties of motile micro-organisms in complex environments is a timely question, relevant to many ecological, medical and technological situations. At the fundamental level, this question is also receiving a lot of attention as fluids loaded with swimming micro-organisms has become a rich domain of applications and a conceptual playground for the statistical physics of “active matter”. The existence of microscopic sources of energy borne by the motile character of these micro-swimmers is driving self-organization processes at the origin of original emergent phases and unconventional macroscopic properties leading to revisit many standard concepts in the physics of suspensions. In this presentation, I will report on a recent exploration on the question of spontaneous formation of large scale collective motion in relation with the rheological response of active suspensions. I will also present new experiments showing how the motility of bacteria can be controlled such as to extract work macroscopically.
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.
Flow, fluctuate and freeze: Epithelial cell sheets as soft active matter
Epithelial cell sheets form a fundamental role in the developing embryo, and also in adult tissues including the gut and the cornea of the eye. Soft and active matter provides a theoretical and computational framework to understand the mechanics and dynamics of these tissues.I will start by introducing the simplest useful class of models, active brownian particles (ABPs), which incorporate uncoordinated active crawling over a substrate and mechanical interactions. Using this model, I will show how the extended ’swirly’ velocity fluctuations seen in sheets on a substrate can be understood using a simple model that couples linear elasticity with disordered activity. We are able to quantitatively match experiments using in-vitro corneal epithelial cells.Adding a different source of activity, cell division and apoptosis, to such a model leads to a novel 'self-melting' dense fluid state. Finally, I will discuss a direct application of this simple particle-based model to the steady-state spiral flow pattern on the mouse cornea.
Design Principles of Living Matter
In this talk, I will describe my lab’s recent efforts to understand the design principles of the active, soft materials that drive cell morphogenesis. In particular, we are interested in how collections of myosin II motors and actin polymers generate, relax, sense and adapt to mechanical force. I will discuss how motor-filament interactions lead to either distributed extensile or contractile stresses as the mechanics of the system changes from fluid to solid. Using optical control of motors, we are now exploring how spatially structured stress can be used to drive local flows and motion. If time, I will also describe how feedbacks between local geometry and activity can be harnessed to drive morphogenetic changes in model systems.
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.
Dynamics of microbiota communities during physical perturbation
The consortium of microbes living in and on our bodies is intimately connected with human biology and deeply influenced by physical forces. Despite incredible gains in describing this community, and emerging knowledge of the mechanisms linking it to human health, understanding the basic physical properties and responses of this ecosystem has been comparatively neglected. Most diseases have significant physical effects on the gut; diarrhea alters osmolality, fever and cancer increase temperature, and bowel diseases affect pH. Furthermore, the gut itself is comprised of localized niches that differ significantly in their physical environment, and are inhabited by different commensal microbes. Understanding the impact of common physical factors is necessary for engineering robust microbiota members and communities; however, our knowledge of how they affect the gut ecosystem is poor. We are investigating how changes in osmolality affect the host and the microbial community and lead to mechanical shifts in the cellular environment. Osmotic perturbation is extremely prevalent in humans, caused by the use of laxatives, lactose intolerance, or celiac disease. In our studies we monitored osmotic shock to the microbiota using a comprehensive and novel approach, which combined in vivo experiments to imaging, physical measurements, computational analysis and highly controlled microfluidic experiments. By bridging several disciplines, we developed a mechanistic understanding of the processes involved in osmotic diarrhea, linking single-cell biophysical changes to large-scale community dynamics. Our results indicate that physical perturbations can profoundly and permanently change the competitive and ecological landscape of the gut, and affect the cell wall of bacteria differentially, depending on their mechanical characteristics.
Pancreatic α and β cells are globally phase-locked
The Ca2+ modulated pulsatile secretions of glucagon and insulin by pancreatic α and β cells play a key role in glucose metabolism and homeostasis. However, how different types of cells in the islet couple and coordinate to give rise to various Ca2+ oscillation patterns and how these patterns are being tuned by paracrine regulation are still elusive. Here we developed a microfluidic device to facilitate long-term recording of islet Ca2+ activity at single cell level and found that islets show heterogeneous but intrinsic oscillation patterns. The α and β cells in an islet oscillate in antiphase and are globally phase locked to display a variety of oscillation modes. A mathematical model of islet oscillation maps out the dependence of the oscillation modes on the paracrine interactions between α and β cells. Our study reveals the origin of the islet oscillation patterns and highlights the role of paracrine regulation in tuning them.
fluid coverage
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