Anomalous propulsion regime in axonemal-propelled cargoes

Bio-actuated micro-swimmers provide a platform to understand physical principles related to the motion of micro-organisms at low Reynolds numbers. Here, we used isolated and demembranated flagella from green algae Chlamydomonas reinhardtii as an ATP-fueled bio-actuator for propulsion of micron-sized beads. Chlamydomonas flagella have an asymmetric waveform, which can be accurately described as a superposition of a static component corresponding to an arc-shaped intrinsic curvature, a mode describing the global oscillations of the axonemal curvature, and a main base- to-tip traveling wave component. By decomposing experimental beat patterns in Fourier modes, and applying resistive force theory, we performed numerical simulations and obtained analytical approximations for the mean rotational and translational velocities of a flagellum-propelled bead. Our analysis reveals the existence of a counter- intuitive anomalous propulsion regime where the speed of the flagellum-driven cargo increases with increasing the cargo size. Further, it demonstrates that in addition to the intrinsic curvature and even harmonics, asymmetric bead-flagellum attachment also contributes in the rotational velocity of the micro-swimmer. This turning mechanism induced by sideways cargo attachment has potential applications in fabrication of bio- actuated medical micro-robots in the subject of targeted drug delivery and synthetic micro-swimmers.

Mechanisms and fluid dynamics of foraging in heterotrophic nanoflagellates

ABSTRACTHeterotrophic nanoflagellates are the main consumers of bacteria and picophytoplankton in the ocean. In their micro-scale world, viscosity impedes predator-prey contact, and the mechanisms that allow flagellates to daily clear a volume of water for prey corresponding to 106 times their own volume is unclear. It is also unclear what limits observed maximum ingestion rates of about 104 bacterial prey per day. We used high-speed video-microscopy to describe feeding flows, flagellum kinematics, and prey searching, capture, and handling in four species with different foraging strategies. In three species, prey-handling times limit ingestion rates and account well for their reported maximum values. Similarly, observed feeding flows match reported clearance rates. Simple point-force models allowed us to estimate the forces required to generate the feeding flows, between 4-13 pN, and consistent with the force produced by the hairy (hispid) flagellum, as estimated using resistive force theory. Hispid flagella can produce a force that is much higher than the force produced by a naked flagellum with similar kinematics, and the hairy flagellum is therefore key to foraging in most nanoflagellates. Our findings provide a mechanistic underpinning of observed functional responses of prey ingestion rates in nanoflagellates.

Surprising simplicity in the modeling of dynamic granular intrusion

Granular intrusions, such as dynamic impact or wheel locomotion, are complex multiphase phenomena where the grains exhibit solid-like and fluid-like characteristics together with an ejected gas-like phase. Despite decades of modeling efforts, a unified description of the physics in such intrusions is as yet unknown. Here, we show that a continuum model based on the simple notions of frictional flow and tension-free separation describes complex granular intrusions near free surfaces. This model captures dynamics in a variety of experiments including wheel locomotion, plate intrusions, and running legged robots. The model reveals that one static and two dynamic effects primarily give rise to intrusion forces in such scenarios. We merge these effects into a further reduced-order technique (dynamic resistive force theory) for rapid modeling of granular locomotion of arbitrarily shaped intruders. The continuum-motivated strategy we propose for identifying physical mechanisms and corresponding reduced-order relations has potential use for a variety of other materials.

Enhancing Swimming Performance by Optimizing Structure of Helical Swimmers

Untethered microrobots provide the prospect for performing minimally invasive surgery and targeted delivery of drugs in hard-to-reach areas of the human body. Recently, inspired by the way the prokaryotic flagella rotates to drive the body forward, numerous studies have been carried out to study the swimming properties of helical swimmers. In this study, the resistive force theory (RFT) was applied to analyze the influence of dimensional and kinematical parameters on the propulsion performance of conventional helical swimmers. The propulsion efficiency index was applied to quantitatively evaluate the swimming performance of helical swimmers. Quantitative analysis of the effect of different parameters on the propulsion performance was performed to optimize the design of structures. Then, RFT was modified to explore the tapered helical swimmers with the helix radius changing uniformly along the axis. Theoretical results show that the helical swimmer with a constant helix angle exhibits excellent propulsion performance. The evaluation index was found to increase with increased tapering, indicating that the tapered structures can produce more efficient motion. Additionally, the analysis method extended from RFT can be used to analyze the motion of special-shaped flagella in microorganisms.

Resistive force theory and wave dynamics in swimming flagellar apparatus isolated from C. reinhardtii

In basal apparatus isolated from C. reinhardtii, if the frequency mismatch is high, mechanical or hydrodynamic coupling cannot synchronize them.

Resistive force theory and wave dynamics in swimming isolated flagellar apparatus

Cilia-driven motility and fluid transport is ubiquitous in nature and essential for many biological processes, including swimming of eukaryotic unicellular organisms, mucus transport in airway apparatus or fluid flow in brain. The-biflagellated micro-swimmer Chlamydomonas reinhardtii is a model organism to study dynamics of flagellar synchronization. Hydrodynamic interactions, intracellular mechanical coupling or cell body rocking are believed to play crucial role in synchronization of flagellar beating in green algae. Here, we use freely swimming intact flagellar apparatus isolated from wall-less strain of Chlamydomonas to investigate wave dynamics. Our analysis in phase coordinates show that, when the frequency difference between the flagella is high, neither mechanical coupling via basal body nor hydrodynamics interactions are strong enough to synchronize two flagella, indicating that beating frequency is controlled internally by the cell. We also examined the validity of resistive force theory for a flagellar apparatus swimming freely in the vicinity of a substrate and found a quantitative agreement between experimental data and simulations with drag anisotropy of ratio 2. Finally, using a simplified wave form, we investigated the influence of phase and frequency differences, intrinsic curvature and wave amplitude on the swimming trajectory of flagellar apparatus. Our analysis shows that by controlling phase or frequency differences between two flagella, steering can occur.

Crash landing of Vibrio cholerae by MSHA pili-assisted braking and anchoring in a viscous environment

Mannose-sensitive hemagglutinin (MSHA) pili and flagellum are critical for the surface attachment of Vibrio cholerae. However, the cell landing mechanism remains largely unknown. Here, combining the cysteine-substitution-based labelling method with single-cell tracking techniques, we quantitatively characterized the landing of V. cholerae by directly observing both pili and flagellum of cells in viscous solutions. MSHA pili are evenly distributed along the cell length and can stick to surfaces at any point along the filament. With such properties, MSHA pili are observed to act as a brake and anchor during cell landing which include three phases: running, lingering, and attaching. Resistive-force-theory based models are proposed to describe near-surface motion. Importantly, the role of MSHA pili during cell landing is more apparent in viscous solutions. Our work provides a detailed picture of the landing dynamics of V. cholerae under viscous conditions, which can provide insights into ways to better control V. cholerae infections.