Large-scale vortex lattice emerging from collectively moving microtubules.

Spontaneous collective motion, as in some flocks of bird and schools of fish, is an example of an emergent phenomenon. Such phenomena are at present of great interest and physicists have put forward a number of theoretical results that so far lack experimental verification. In animal behaviour studies, large-scale data collection is now technologically possible, but data are still scarce and arise from observations rather than controlled experiments. Multicellular biological systems, such as bacterial colonies or tissues, allow more control, but may have many hidden variables and interactions, hindering proper tests of theoretical ideas. However, in systems on the subcellular scale such tests may be possible, particularly in in vitro experiments with only few purified components. Motility assays, in which protein filaments are driven by molecular motors grafted to a substrate in the presence of ATP, can show collective motion for high densities of motors and attached filaments. This was demonstrated recently for the actomyosin system, but a complete understanding of the mechanisms at work is still lacking. Here we report experiments in which microtubules are propelled by surface-bound dyneins. In this system it is possible to study the local interaction: we find that colliding microtubules align with each other with high probability. At high densities, this alignment results in self-organization of the microtubules, which are on average 15 µm long, into vortices with diameters of around 400 µm. Inside the vortices, the microtubules circulate both clockwise and anticlockwise. On longer timescales, the vortices form a lattice structure. The emergence of these structures, as verified by a mathematical model, is the result of the smooth, reptation-like motion of single microtubules in combination with local interactions (the nematic alignment due to collisions)--there is no need for long-range interactions. Apart from its potential relevance to cortical arrays in plant cells and other biological situations, our study provides evidence for the existence of previously unsuspected universality classes of collective motion phenomena.

Photoinduced Fusion of Lipid Bilayer Membranes.

We have developed a novel system for photocontrol of the fusion of lipid vesicles through the use of a photosensitive surfactant containing an azobenzene moiety (AzoTAB). Real-time microscopic observations clarified a change in both the surface area and internal volume of vesicles during fusion. We also determined the optimal cholesterol concentrations and temperature for inducing fusion. The mechanism of fusion can be attributed to a change in membrane tension, which is caused by the solubilization of lipids through the isomerization of AzoTAB. We used a micropipet technique to estimate membrane tension and discuss the mechanism of fusion in terms of membrane elastic energy. The obtained results regarding this novel photoinduced fusion could lead to a better understanding of the mechanism of membrane fusion in living cells and may also see wider applications, such as in drug delivery and biomimetic material design.

Lateral Diffusion of a Submicrometer Particle on a Lipid Bilayer Membrane.

In past decades, nanoparticles and nanomaterials have been actively used for applications such as visualizing nano/submicrometer cell structure, killing cancer cells, and using drug delivery systems. It is important to understand the physicochemical mechanisms that govern the motion of nanoparticles on a plasma membrane surface. However, the motion of small particles of <1000 nm on lipid membranes is poorly understood. In this study, we investigated the diffusion of particles with a diameter of 200-800 nm on a lipid membrane using cell-sized liposomes. Particle-associated liposomes were obtained by applying centrifugal force to a mixture of liposomes and particle solutions. We measured the thermal motion of the particles by phase-contrast microscopy. We found that (i) the particle-size dependence of the diffusion of particles adhering to membranes was better described by the DADL model rather than the Einstein-Stokes model, (ii) the diffusion coefficient of a particle strongly depends on the adsorption state of the particle, such as fully or partially wrapped by the membrane, and (iii) anomalous diffusion was induced by the localization of particles on the neck of budded vesicles.

Mathematical model for self-propelled droplets driven by interfacial tension.

We propose a model for the spontaneous motion of a droplet induced by inhomogeneity in interfacial tension. The model is derived from a variation of the Lagrangian of the system and we use a time-discretized Morse flow scheme to perform its numerical simulations. Our model can naturally simulate the dynamics of a single droplet, as well as that of multiple droplets, where the volume of each droplet is conserved. We reproduced the ballistic motion and fission of a droplet, and the collision of two droplets was also examined numerically.

Collective motion of self-propelled particles with memory.

We show that memory, in the form of underdamped angular dynamics, is a crucial ingredient for the collective properties of self-propelled particles. Using Vicsek-style models with an Ornstein-Uhlenbeck process acting on angular velocity, we uncover a rich variety of collective phases not observed in usual overdamped systems, including vortex lattices and active foams. In a model with strictly nematic interactions the smectic arrangement of Vicsek waves giving rise to global polar order is observed. We also provide a calculation of the effective interaction between vortices in the case where a telegraphic noise process is at play, explaining thus the emergence and structure of the vortex lattices observed here and in motility assay experiments.

C. elegans collectively forms dynamical networks.

Understanding physical rules underlying collective motions requires perturbation of controllable parameters in self-propelled particles. However, controlling parameters in animals is generally not easy, which makes collective behaviours of animals elusive. Here, we report an experimental system in which a conventional model animal, Caenorhabditis elegans, collectively forms dynamical networks of bundle-shaped aggregates. We investigate the dependence of our experimental system on various extrinsic parameters (material of substrate, ambient humidity and density of worms). Taking advantage of well-established C. elegans genetics, we also control intrinsic parameters (genetically determined motility) by mutations and by forced neural activation via optogenetics. Furthermore, we develop a minimal agent-based model that reproduces the dynamical network formation and its dependence on the parameters, suggesting that the key factors are alignment of worms after collision and smooth turning. Our findings imply that the concepts of active matter physics may help us to understand biological functions of animal groups.

Collective motion of rod-shaped self-propelled particles through collision.

Self-propelled rods, which propel by themselves in the direction from the tail to the head and align nematically through collision, have been well-investigated theoretically. Various phenomena including true long-range ordered phase with the Giant number fluctuations, and the collective motion composed of many vorices were predicted using the minimal mathematical models of self-propelled rods. Using filamentous bacteria and running microtubules, we found that the predicted phenomena by the minimal models occur in the real world. This strongly indicates that there exists the unified description of self-propelled rods independent of the details of the systems. The theoretically predicted phenomena and the experimental results concerning the phenomena are reviewed.

Controlled Construction of Stable Network Structure Composed of Honeycomb-Shaped Microhydrogels.

Recently, the construction of models for multicellular systems such as tissues has been attracting great interest. These model systems are expected to reproduce a cell communication network and provide insight into complicated functions in living systems./Such network structures have mainly been modelled using a droplet and a vesicle. However, in the droplet and vesicle network, there are difficulties attributed to structural instabilities due to external stimuli and perturbations. Thus, the fabrication of a network composed of a stable component such as hydrogel is desired. In this article, the construction of a stable network composed of honeycomb-shaped microhydrogels is described. We produced the microhydrogel network using a centrifugal microfluidic technique and a photosensitive polymer. In the network, densely packed honeycomb-shaped microhydrogels were observed. Additionally, we successfully controlled the degree of packing of microhydrogels in the network by changing the centrifugal force. We believe that our stable network will contribute to the study of cell communication in multicellular systems.

Lateral Tension-Induced Penetration of Particles into a Liposome.

It is important that we understand the mechanism of the penetration of particles into a living cell to achieve advances in bionanotechnology, such as for treatment, visualization within a cell, and genetic modification. Although there have been many studies on the application of functional particles to cells, the basic mechanism of penetration across a biological membrane is still poorly understood. Here we used a model membrane system to demonstrate that lateral membrane tension drives particle penetration across a lipid bilayer. After the application of osmotic pressure, fully wrapped particles on a liposome surface were found to enter the liposome. We discuss the mechanism of the tension-induced penetration in terms of narrow constriction of the membrane at the neck part. The present findings are expected to provide insight into the application of particles to biological systems.

Long-range nematic order and anomalous fluctuations in suspensions of swimming filamentous bacteria.

We study the collective dynamics of elongated swimmers in a very thin fluid layer by devising long filamentous nontumbling bacteria. The strong confinement induces weak nematic alignment upon collision, which, for large enough density of cells, gives rise to global nematic order. This homogeneous but fluctuating phase, observed on the largest experimentally accessible scale of millimeters, exhibits the properties predicted by standard models for flocking, such as the Vicsek-style model of polar particles with nematic alignment: true long-range nematic order and nontrivial giant number fluctuations.

Complex-shaped three-dimensional multi-compartmental microparticles generated by diffusional and Marangoni microflows in centrifugally discharged droplets.

We report a versatile method for the generation of complex-shaped three-dimensional multi-compartmental (3D-MC) microparticles. Complex-shaped microparticles have recently received much attention for potential application in self-assemblies, micromachines, and biomedical and environmental engineering. Here, we have developed a method based on 3D nonequilibrium-induced microflows (Marangoni and diffusional flows) of microdroplets that are discharged from the tip of a thin capillary in a simple centrifugal microfluidic device. The microparticle shapes can be tuned by the partial dissolution of specific compartments and by the deformation of the precursor microdroplets by manipulating the 3D microflows. We believe that this method will have wide applications in nano- and microscience and technologies.

Influence of Asymmetry and Driving Forces on the Propulsion of Bubble-Propelled Catalytic Micromotors.

Bubble-propelled catalytic micromotors have recently been attracting much attention. A bubble-propulsion mechanism has the advantage of producing a stronger force and higher speed than other mechanisms for catalytic micromotors, but the nature of the fluctuated bubble generation process affects the motions of the micromotors, making it difficult to control their motions. Thus, understanding of the influence of fluctuating bubble propulsion on the motions of catalytic micromotors is important in exploiting the advantages of bubble-propelled micromotors. Here, we report experimental demonstrations of the bubble-propelled motions of propeller-shaped micromotors and numerical analyses of the influence of fluctuating bubble propulsion on the motions of propeller-shaped micromotors. We found that motions such as trochoid-like motion and circular motion emerged depending on the magnitude or symmetricity of fluctuations in the bubble-propulsion process. We hope that those results will help in the construction and application of sophisticated bubble-propelled micromotors in the future.

Rotational motion of a droplet induced by interfacial tension.

Spontaneous rotation of a droplet induced by the Marangoni flow is analyzed in a two-dimensional system. The droplet with the small particle which supplies a surfactant at the interface is considered. We calculated flow field around the droplet using the Stokes equation and found that advective nonlinearity breaks symmetry for rotation. Theoretical calculation indicates that the droplet spontaneously rotates when the radius of the droplet is an appropriate size. The theoretical results were validated through comparison with the experiments.

Drift instability in the motion of a fluid droplet with a chemically reactive surface driven by Marangoni flow.

We theoretically derive the amplitude equations for a self-propelled droplet driven by Marangoni flow. As advective flow driven by surface tension gradient is enhanced, the stationary state becomes unstable and the droplet starts to move. The velocity of the droplet is determined from a cubic nonlinear term in the amplitude equations. The obtained critical point and the characteristic velocity are well supported by numerical simulations.

Spontaneous motion of a droplet coupled with a chemical wave.

We propose a framework for the spontaneous motion of a droplet coupled with internal dynamic patterns generated in a reaction-diffusion system. The spatiotemporal order of the chemical reaction gives rise to inhomogeneous surface tension and results in self-propulsion driven by the surrounding flow due to the Marangoni effect. Numerical calculations of internal patterns together with theoretical results of the flow fields at low Reynolds number reproduce well the experimental results obtained using a droplet of the Belousov-Zhabotinsky reaction medium.

Noise-induced synchronization of a large population of globally coupled nonidentical oscillators.

We study a large population of globally coupled phase oscillators subject to common white Gaussian noise and find analytically that the critical coupling strength between oscillators for synchronization transition decreases with an increase in the intensity of common noise. Thus, common noise promotes the onset of synchronization. Our prediction is confirmed by numerical simulations of the phase oscillators as well as of limit-cycle oscillators.