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.

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.

Living Systems often seem to follow, in addition to external constraints and interactions, an intrinsic predictive model of the world — a defining trait of Anticipatory Systems. Here we study rhythmic behaviour in Caulerpa, a marine green alga, which appears to predict the day/night light cycle. Caulerpa consists of differentiated organs resembling leaves, stems and roots. While an individual can exceed a meter in size, it is a single multinucleated giant cell. Active transport has been hypothesized to play a key role in organismal development. It has been an open question in the literature whether rhythmic transport phenomena in this organism are of autonomous circadian nature. Using Raspberry-Pi cameras, we track over weeks the morphogenesis of tens of samples concurrently, while tracing at resolution of tens of seconds the variation of the green coverage. The latter reveals waves propagating over centimeters within few hours, and is attributed to chloroplast redistribution at whole-organism scale. Our observations of algal segments regenerating under 12-hour light/dark cycles indicate that the initiation of the waves precedes the external light change. Using time-frequency analysis, we find that the temporal spectrum of these green pulses contains a circadian period. The latter persists over days even under constant illumination, indicative of its autonomous nature. We further explore the system under non-circadian periods, to reveal how the spectral content changes in response. Time-keeping and synchronization are recurring themes in biological research at various levels of description — from subcellular components to ecological systems. We present a seemingly primitive living system that exhibits apparent anticipatory behaviour. This research offers quantitative constraints for theoretical frameworks of such systems.

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.

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.

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.

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.

Chromosomes are extremely long, active polymers that are spatially organized across multiple scales to promote cellular functions, such as gene transcription and genetic inheritance. During each cell cycle, chromosomes are dramatically compacted as cells divide and dynamically reorganized into less compact, spatiotemporally patterned structures after cell division. These activities are facilitated by DNA/chromatin-binding protein motors called SMC complexes. Each of these motors can perform a unique activity known as “loop extrusion,” in which the motor binds the DNA/chromatin polymer, reels in the polymer fiber, and extrudes it as a loop. Using simulations and theory, I show how loop-extruding motors can collectively compact and spatially organize chromosomes in different scenarios. First, I show that loop-extruding complexes can generate sufficient compaction for cell division, provided that loop-extrusion satisfies stringent physical requirements. Second, while loop-extrusion alone does not uniquely spatially pattern the genome, interactions between SMC complexes and protein “boundary elements” can generate patterns that emerge in the genome after cell division. Intriguingly, these “boundary elements” are not necessarily stationary, which can generate a variety of patterns in the neighborhood of transcriptionally active genes. These predictions, along with supporting experiments, show how SMC complexes and other molecular machinery, such as RNA polymerase, can spatially organize the genome. More generally, this work demonstrates both the versatility of the loop extrusion mechanism for chromosome functional organization and how seemingly subtle microscopic effects can emerge in the spatiotemporal structure of nonequilibrium polymers.

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.

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.

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.

While the motion and collective behavior of cells are well-studied on flat surfaces or in unconfined liquid media, in most natural settings, cells thrive in complex 3D environments. Bioprinting processes are capable of structuring cells in 3D and conventional bioprinting approaches address this challenge by embedding cells in bio-degradable polymer networks. However, heterogeneity in network structure and biodegradation often preclude quantitative studies of cell behavior in specified 3D architectures. Here, I will present a new approach to 3D bioprinting of cellular communities that utilizes jammed, granular polyelectrolyte microgels as a support medium. The self-healing nature of this medium allows the creation of highly precise cellular communities and tissue-like structures by direct injection of cells inside the 3D medium. Further, the transparent nature of this medium enables precise characterization of cellular behavior. I will describe two examples of my work using this platform to study the behavior of two different classes of cells in 3D. First, I will describe how we interrogate the growth, viability, and migration of mammalian cells—ranging from epithelial cells, cancer cells, and T cells—in the 3D pore space. Second, I will describe how we interrogate the migration of E. coli bacteria through the 3D pore space. Direct visualization enables us to reveal a new mode of motility exhibited by individual cells, in stark contrast to the paradigm of run-and-tumble motility, in which cells are intermittently and transiently trapped as they navigate the pore space; further, analysis of these dynamics enables prediction of single-cell transport over large length and time scales. Moreover, we show that concentrated populations of E. coli can collectively migrate through a porous medium—despite being strongly confined—by chemotactically “surfing” a self-generated nutrient gradient. Together, these studies highlight how the jammed microgel medium provides a powerful platform to design and interrogate complex cellular communities in 3D—with implications for tissue engineering, microtissue mechanics, studies of cellular interactions, and biophysical studies of active matter.

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.

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

In vivo, the human genome folds into a characteristic ensemble of 3D structures. The mechanism driving the folding process remains unknown. A theoretical model for chromatin (the minimal chromatin model) explains the folding of interphase chromosomes and generates chromosome conformations consistent with experimental data is presented. The energy landscape of the model was derived by using the maximum entropy principle and relies on two experimentally derived inputs: a classification of loci into chromatin types and a catalog of the positions of chromatin loops. This model was generalized by utilizing a neural network to infer these chromatin types using epigenetic marks present at a locus, as assayed by ChIP-Seq. The ensemble of structures resulting from these simulations completely agree with HI-C data and exhibits unknotted chromosomes, phase separation of chromatin types, and a tendency for open chromatin to lie at the periphery of chromosome territories. Although this theoretical methodology was trained in one cell line, the human GM12878 lymphoblastoid cells, it has successfully predicted the structural ensembles of multiple human cell lines. Finally, going beyond Hi-C, our predicted structures are also consistent with microscopy measurements. Analysis of both structures from simulation and microscopy reveals that short segments of chromatin make two-state transitions between closed conformations and open dumbbell conformations. For gene active segments, the vast majority of genes appear clustered in the linker region of the chromatin segment, allowing us to speculate possible mechanisms by which chromatin structure and dynamics may be involved in controlling gene expression. * Supported by the NSF

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.

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.

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.

Living cells contain millions of enzymes and proteins, which carry out multiple reactions simultaneously. To optimize these processes, cells compartmentalize reactions in membraneless liquid condensates. Certain features of cellular condensates can be explained by principles of liquid-liquid phase separation studied in material science. However, biological condensates exist in the inherently out of equilibrium environment of a living cell, being driven by force-generating microscopic processes. These cellular conditions are fundamentally different than the equilibrium conditions of liquid-liquid phase separation studied in materials science and physics. How condensates function in the active riotous environment of a cell is essential for understanding of cellular functions, as well as to the onset of neurodegenerative diseases. Currently, we lack model systems that enable rigorous studies of these processes. Living cells are too complex for quantitative analysis, while reconstituted equilibrium condensates fail to capture the non-equilibrium environment of biological cells. To bridge this gap, we reconstituted a DNA based membraneless condensates in an active environment that mimics the conditions of a living cell. We combine condensates with a reconstituted network of cytoskeletal filaments and molecular motors, and study how the mechanical interactions change the phase behavior and dynamics of membraneless structures. Studying these composite materials elucidates the fundamental physics rules that govern the behavior of liquid-liquid phase separation away from equilibrium while providing insight into the mechanism of condensate phase separation in cellular environments.

Cells and microorganisms produce and consume all sorts of chemicals, from nutrients to signalling molecules. The same happens at the nanoscale inside cells themselves, where enzymes catalyse the production and consumption of the chemicals needed for life. In this work, we have found a generic mechanism by which such chemically-active particles, be it cells or enzymes or engineered synthetic colloids, can "sense" each other and ultimately self- organize in a multitude of ways. A peculiarity of these chemical-mediated interactions is that they break action-reaction symmetry : for example, one particle may be repelled from a second particle, which is in turn attracted to the first one, so that it ends up "chasing" it. Such chasing interactions allow for the formation of large clusters of particles that "swim" autonomously. Regarding enzymes, we find that they can spontaneously aggregate into clusters with precisely the right composition, so that the product of one enzyme is passed on, without lack or excess, to the next enzyme in the metabolic cascade.

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

Granular matter can serve as a prototype for exploring the rich physics of many-body systems driven far from equilibrium. This talk will outline a new direction for granular physics with macroscopic particles, where acoustic levitation compensates the forces due to gravity and eliminates frictional interactions with supporting surfaces in order to focus on particle interactions. Levitating small particles by intense ultrasound fields in air makes it possible to manipulate and control their positions and assemble them into larger aggregates. The small air viscosity implies that the regime of underdamped dynamics can be explored, where inertial effects are important, in contrast to typical colloids in a liquid, where inertia can be neglected. Sound scattered off individual, levitated solid particles gives rise to controllable attractive forces with neighboring particles. I will discuss some of the key concepts underlying acoustic levitation, describe how detuning an acoustic cavity can introduce active fluctuations that control the assembly statistics of small levitated particles clusters, and give examples of how interactions between neighboring levitated objects can be controlled by their shape.

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.

A key challenge in modern soft matter is to identify the principles that govern the organisation and functionality in non-equilibrium systems. Current research efforts largely focus on non-equilibrium processes that occur either at the single-molecule scale (e.g. protein and DNA conformations under driving forces), or at the scale of whole tissues, organisms, and active colloidal and microscopic objects. However, the range of the scales in-between — from molecules to large-scaled molecular assemblies that consume energy and perform work — remains under-explored. This is, nevertheless, the scale that is crucial for the function of a living cell, where molecular self-assembly driven far from equilibrium produces mechanical work needed for cell reshaping, transport, motility, division, and healing. Today I will discuss physical modelling of active elastic filaments, called ESCRT-III filaments, that dynamically assemble and disassemble on cell membranes. This dynamic assembly changes the filaments’ shape and mechanical properties and leads to the remodelling and cutting of cells. I will present a range of experimental comparisons of our simulation results: from ESCRT-III-driven trafficking in eukaryotes to division of evolutionary simple archaeal cells.

At small length-scales, capillary effects are significant, and thus the mechanics of soft material interfaces may be dominated by solid surface stresses or liquid surface tensions. The balance between surface and bulk properties is described by an elasto-capillary length-scale, in which equilibrium interfacial energies are constant. However, at small length-scales in biological materials, including living cells and tissues, interfacial energies are not constant but are actively regulated and driven far from equilibrium. Thus, the balance between surface and bulk properties depends upon the distance from equilibrium. Here, we model the spreading (wetting) of living cell aggregates as ‘active droplets’, with a non-equilibrium surface tension that depends upon internal stress generated by the actomyosin cytoskeleton. Depending upon the extent of activity, droplet surface properties adapt to the mechanics of their surroundings. The impact of this adaptation challenges contemporary models of interfacial mechanics, including extensively used models of contact mechanics and wetting.

Bacterial motility is central to processes in agriculture, the environment, and medicine. While motility is typically studied in homogeneous environments, many bacterial habitats—e.g., soils, sediments, and biological gels/tissues—are heterogeneous porous media. Here, through studies of E. coli in transparent 3D porous media, we demonstrate that confinement in a heterogeneous medium fundamentally alters motility. In particular, we show how the paradigm of run-and-tumble motility is dramatically altered by pore-scale confinement, both for cells performing undirected motion and those performing chemotaxis, directed motion in response to a chemical stimulus. Our porous media also enable precisely structured multi-cellular communities to be 3D printed. Using this capability, we show how confinement-dependent chemotaxis enables populations to stabilize large-scale perturbations in their overall morphology. Together, our work thus reveals new principles to predict and control the behavior of bacteria, and active matter in general, in heterogeneous environments.

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.

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.

From slithering snakes, to entangling robots, self-deforming (shape changing) active systems display surprising dynamics. This is particularly true when such systems interact with environments or other agents to generate self-propulsion (movement). In this talk, I will discuss a few projects from my group illustrating unexpected effects in individual and collectives of self-deformers. For example, snakes and snake-like robots mechanically “diffract” from fixed environmental heterogeneities, collections of smart-active robots (smarticles) can locomote (and phototax) as a collective despite individual immobility, and geometrically actively entangling ensembles of blackworms and robots can self-propel as a unit to thermo or phototax without centralized control.

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.

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.

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.

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.