Light-regulated adsorption and desorption of Chlamydomonas cells at surfaces.

Microbial colonization of surfaces represents the first step towards biofilm formation, which is a recurring phenomenon in nature with beneficial and detrimental implications in technological and medical settings. Consequently, there is interest in elucidating the fundamental aspects of the initial stages of biofilm formation of microorganisms on solid surfaces. While most of the research is oriented to understand bacterial surface colonization, the fundamental principles of surface colonization of motile, photosynthetic microbes remain largely unexplored so far. Recent single-cell studies showed that the flagellar adhesion of Chlamydomonas reinhardtii is switched on in blue light and switched off under red light [Kreis et al., Nat. Phys., 2018, 14, 45-49]. Here, we study this light-switchable surface association on the population level and measure the kinetics of adsorption and desorption of suspensions of motile C. reinhardtii cells on glass surfaces using bright-field optical microscopy. We observe that both processes exhibit a response lag relative to the time at which the blue- and red-light conditions are set and model this feature using time-delayed Langmuir-type kinetics. We find that cell adsorption occurs significantly faster than desorption, which we attribute to the protein-mediated molecular adhesion mechanism of the cells. Adsorption experiments using phototactically blind C. reinhardtii mutants demonstrate that phototaxis does not affect the cell adsorption kinetics. Hence, this framework can be used as an assay for characterizing the dynamics of the surface colonization of microbial species exhibiting light-regulated surface adhesion under precisely controlled environmental conditions.

Capillary-Based Microfluidics-Coflow, Flow-Focusing, Electro-Coflow, Drops, Jets, and Instabilities.

Capillary-based microfluidics is a great technique to produce monodisperse and complex emulsions and particulate suspensions. In this review, the current understanding of drop and jet formation in capillary-based microfluidic devices for two primary flow configurations, coflow and flow-focusing is summarized. The experimental and theoretical description of fluid instabilities is discussed and conditions for controlled drop breakup in different modes of drop generation are provided. Current challenges in drop breakup with low interfacial tension systems and recent progress in overcoming drop size limitations using electro-coflow are addressed. In each scenario, the physical mechanisms for drop breakup are revisited, and simple scaling arguments proposed in the literature are introduced.

Shrinking instability of toroidal droplets.

Toroidal droplets are inherently unstable due to surface tension. They can break up, similar to cylindrical jets, but also exhibit a shrinking instability, which is inherent to the toroidal shape. We investigate the evolution of shrinking toroidal droplets using particle image velocimetry. We obtain the flow field inside the droplets and show that as the torus evolves, its cross-section significantly deviates from circular. We then use the experimentally obtained velocities at the torus interface to theoretically reconstruct the internal flow field. Our calculation correctly describes the experimental results and elucidates the role of those modes that, among the many possible ones, are required to capture all of the relevant experimental features.

Dynamic assembly of ultrasoft colloidal networks enables cell invasion within restrictive fibrillar polymers.

In regenerative medicine, natural protein-based polymers offer enhanced endogenous bioactivity and potential for seamless integration with tissue, yet form weak hydrogels that lack the physical robustness required for surgical manipulation, making them difficult to apply in practice. The use of higher concentrations of protein, exogenous cross-linkers, and blending synthetic polymers has all been applied to form more mechanically robust networks. Each relies on generating a smaller network mesh size, which increases the elastic modulus and robustness, but critically inhibits cell spreading and migration, hampering tissue regeneration. Here we report two unique observations; first, that colloidal suspensions, at sufficiently high volume fraction (ϕ), dynamically assemble into a fully percolated 3D network within high-concentration protein polymers. Second, cells appear capable of leveraging these unique domains for highly efficient cell migration throughout the composite construct. In contrast to porogens, the particles in our system remain embedded within the bulk polymer, creating a network of particle-filled tunnels. Whereas this would normally physically restrict cell motility, when the particulate network is created using ultralow cross-linked microgels, the colloidal suspension displays viscous behavior on the same timescale as cell spreading and migration and thus enables efficient cell infiltration of the construct through the colloidal-filled tunnels.

Toroidal Droplets: Growth Rates, Dispersion Relations, and Behavior in the Thick-Torus Limit.

Toroidal droplets in a viscous liquid are unstable and transform into single or multiple spherical droplets. For thin tori, this can happen via the Rayleigh-Plateau instability causing the breakup of cylindrical jets. In contrast, for thick tori, this can happpen via the shrinking of the "hole". In this work, we use the thin-torus limit to directly measure the growth rate associated with capillary disturbances. In the case of toroidal droplets inside a much more viscous liquid, we even obtain the full dispersion relation, which is in agreement with theoretical results for cylindrical jets. For thick tori, we employ particle image velocimetry to determine the flow field of a sinking toroidal drop inside another viscous liquid. We find that the presence of the "hole" greatly suppresses one of the circulation loops expected for sinking cylinders. Finally, using the flow field of a shrinking toroidal droplet and the time-reversal symmetry of the Stokes equations, we theoretically predict the expected shape deformation of an expanding torus and confirm the result experimentally using charged toroidal droplets.

Self-generated oxygen gradients control collective aggregation of photosynthetic microbes.

For billions of years, photosynthetic microbes have evolved under the variable exposure to sunlight in diverse ecosystems and microhabitats all over our planet. Their abilities to dynamically respond to alterations of the luminous intensity, including phototaxis, surface association and diurnal cell cycles, are pivotal for their survival. If these strategies fail in the absence of light, the microbes can still sustain essential metabolic functionalities and motility by switching their energy production from photosynthesis to oxygen respiration. For suspensions of motile C. reinhardtii cells above a critical density, we demonstrate that this switch reversibly controls collective microbial aggregation. Aerobic respiration dominates over photosynthesis in conditions of low light, which causes the microbial motility to sensitively depend on the local availability of oxygen. For dense microbial populations in self-generated oxygen gradients, microfluidic experiments and continuum theory based on a reaction-diffusion mechanism show that oxygen-regulated motility enables the collective emergence of highly localized regions of high and low cell densities.

Breakup dynamics of toroidal droplets in shear-thinning fluids.

We use toroidal droplets to study the breakup dynamics of a Newtonian liquid jet in the presence of rheologically nonlinear materials. We find that the droplets resist breakup for times that are longer than those in the presence of Newtonian liquids. More importantly, we show that our experiments can be explained by incorporating the nonlinearities into the linear treatment of the problem through the strain-rate-dependent viscosity. Finally, we show that the scaling factor required to relate the viscosity to the growth rate associated to unstable modes is given by the elastic modulus of the outside material, illustrating that both the viscoelastic and shear-thinning nature of the outside material play a crucial role in the dynamics of the problem.

Toroidal-droplet instabilities in the presence of charge.

Neutral toroidal droplets can break via the surface-tension-driven Rayleigh-Plateau instability. They can additionally exhibit a shrinking instability, which is also driven by surface tension, whereby the handle progressively disappears. We find that charging a toroidal droplet can qualitatively change the behavior and cause the droplet to expand. We successfully model the transition from shrinking to expanding, considering the pressure balance across the interface of the torus. However, despite the change in behavior, charged toroidal droplets end up breaking into spherical droplets. We quantify how the wavelength of the fastest unstable mode associated to this breakup depends on the applied voltage and compare the results with theoretical expectations for charged cylindrical jets.