Realizing the physics of motile cilia synchronization with driven colloids

Cilia and flagella in biological systems often show large scale cooperative behaviors such as the synchronization of their beats in "metachronal waves". These are beautiful examples of emergent dynamics in biology, and are essential for life, allowing diverse processes from the motility of eukaryotic microorganisms, to nutrient transport and clearance of pathogens from mammalian airways. How these collective states arise is not fully understood, but it is clear that individual cilia interact mechanically,and that a strong and long ranged component of the coupling is mediated by the viscous fluid. We review here the work by ourselves and others aimed at understanding the behavior of hydrodynamically coupled systems, and particularly a set of results that have been obtained both experimentally and theoretically by studying actively driven colloidal systems. In these controlled scenarios, it is possible to selectively test aspects of the living motile cilia, such as the geometrical arrangement, the effects of the driving profile and the distance to no-slip boundaries. We outline and give examples of how it is possible to link model systems to observations on living systems, which can be made on microorganisms, on cell cultures or on tissue sections. This area of research has clear clinical application in the long term, as severe pathologies are associated with compromised cilia function in humans.Comment: 31 pages, to appear in Annual Review of Condensed Matter Physic

Programmable interactions with biomimetic DNA linkers at fluid membranes and interfaces

At the heart of the structured architecture and complex dynamics of biological systems are specific and timely interactions operated by biomolecules. In many instances, biomolecular agents are spatially confined to flexible lipid membranes where, among other functions, they control cell adhesion, motility and tissue formation. Besides being central to several biological processes, \emph{multivalent interactions} mediated by reactive linkers confined to deformable substrates underpin the design of synthetic-biological platforms and advanced biomimetic materials. Here we review recent advances on the experimental study and theoretical modelling of a heterogeneous class of biomimetic systems in which synthetic linkers mediate multivalent interactions between fluid and deformable colloidal units, including lipid vesicles and emulsion droplets. Linkers are often prepared from synthetic DNA nanostructures, enabling full programmability of the thermodynamic and kinetic properties of their mutual interactions. The coupling of the statistical effects of multivalent interactions with substrate fluidity and deformability gives rise to a rich emerging phenomenology that, in the context of self-assembled soft materials, has been shown to produce exotic phase behaviour, stimuli-responsiveness, and kinetic programmability of the self-assembly process. Applications to (synthetic) biology will also be reviewed.Comment: 63 pages, revie

Studies of a weak polyampholyte at the air-buffer interface: The effect of varying pH and ionic strength

We have carried out experiments to probe the static and dynamic interfacial properties of $\beta$--casein monolayers spread at the air-buffer interface, and analysed these results in the context of models of weak polyampholytes. Measurements have been made systematically over a wide range of ionic strength and pH. In the semi-dilute regime of surface concentration a scaling exponent, which can be linked to the degree of chain swelling, is found. This shows that at pH close to the isoelectric point, the protein is compact. At pH away from the isoelectric pH the protein is extended. The transition between compact and extended states is continuous. As a function of increasing ionic strength, we observe swelling of the protein at the isoelectric pH but contraction of the protein at pH values away from it. These behaviours are typical of a those predicted theoretically for a weak polyampholyte. Dilational moduli measurements, made as a function of surface concentration exhibit maxima that are linked to the collapse of hydrophilic regions of the protein into the subphase. Based on this data we present a configuration map of the protein configuration in the monolayer. These findings are supported by strain (surface pressure) relaxation measurements and surface quasi-elastic light scattering (SQELS) measurements which suggest the existence of loops and tails in the subphase at higher surface concentrations.Comment: Submitted to J. Chem. Phy

Correlation between crystalline order and vitrification in colloidal monolayers

We investigate experimentally the relationship between local structure and dynamical arrest in a quasi-2d colloidal model system which approximates hard discs. We introduce polydispersity to the system to suppress crystallisation. Upon compression, the increase in structural relaxation time is accompanied by the emergence of local hexagonal symmetry. Examining the dynamical heterogeneity of the system, we identify three types of motion : "zero-dimensional" corresponding to beta-relaxation, "one-dimensional" or stringlike motion and "two-dimensional" motion. The dynamic heterogeneity is correlated with the local order, that is to say locally hexagonal regions are more likely to be dynamically slow. However we find that lengthscales corresponding to dynamic heterogeneity and local structure do not appear to scale together approaching the glass transition.Comment: 13 papes, to appear in J. Phys.: Condens. Matte

Changes in geometrical aspects of a simple model of cilia synchronization control the dynamical state, a possible mechanism for switching of swimming gaits in microswimmers.

Active oscillators, with purely hydrodynamic coupling, are useful simple models to understand various aspects of motile cilia synchronization. Motile cilia are used by microorganisms to swim and to control the flow fields in their surroundings; the patterns observed in cilia carpets can be remarkably complex, and can be changed over time by the organism. It is often not known to what extent the coupling between cilia is due to just hydrodynamic forces, and neither is it known if it is biological or physical triggers that can change the dynamical collective state. Here we treat this question from a very simplified point of view. We describe three possible mechanisms that enable a switch in the dynamical state, in a simple scenario of a chain of oscillators. We find that shape-change provides the most consistent strategy to control collective dynamics, but also imposing small changes in frequency produces some unique stable states. Demonstrating these effects in the abstract minimal model proves that these could be possible explanations for gait switching seen in ciliated micro organisms like Paramecium and others. Microorganisms with many cilia could in principle be taking advantage of hydrodynamic coupling, to switch their swimming gait through either a shape change that manifests in decreased coupling between groups of cilia, or alterations to the beat style of a small subset of the cilia

Changes in geometrical aspects of a simple model of cilia synchronization control the dynamical state, a possible mechanism for switching of swimming gaits in microswimmers.

Active oscillators, with purely hydrodynamic coupling, are useful simple models to understand various aspects of motile cilia synchronization. Motile cilia are used by microorganisms to swim and to control the flow fields in their surroundings; the patterns observed in cilia carpets can be remarkably complex, and can be changed over time by the organism. It is often not known to what extent the coupling between cilia is due to just hydrodynamic forces, and neither is it known if it is biological or physical triggers that can change the dynamical collective state. Here we treat this question from a very simplified point of view. We describe three possible mechanisms that enable a switch in the dynamical state, in a simple scenario of a chain of oscillators. We find that shape-change provides the most consistent strategy to control collective dynamics, but also imposing small changes in frequency produces some unique stable states. Demonstrating these effects in the abstract minimal model proves that these could be possible explanations for gait switching seen in ciliated micro organisms like Paramecium and others. Microorganisms with many cilia could in principle be taking advantage of hydrodynamic coupling, to switch their swimming gait through either a shape change that manifests in decreased coupling between groups of cilia, or alterations to the beat style of a small subset of the cilia