Rapid behavioral transitions produce chaotic mixing by a planktonic microswimmer

Despite their vast morphological diversity, many invertebrates have similar larval forms characterized by ciliary bands, innervated arrays of beating cilia that facilitate swimming and feeding. Hydrodynamics suggests that these bands should tightly constrain the behavioral strategies available to the larvae; however, their apparent ubiquity suggests that these bands also confer substantial adaptive advantages. Here, we use hydrodynamic techniques to investigate "blinking," an unusual behavioral phenomenon observed in many invertebrate larvae in which ciliary bands across the body rapidly change beating direction and produce transient rearrangement of the local flow field. Using a general theoretical model combined with quantitative experiments on starfish larvae, we find that the natural rhythm of larval blinking is hydrodynamically optimal for inducing strong mixing of the local fluid environment due to transient streamline crossing, thereby maximizing the larvae's overall feeding rate. Our results are consistent with previous hypotheses that filter feeding organisms may use chaotic mixing dynamics to overcome circulation constraints in viscous environments, and it suggests physical underpinnings for complex neurally-driven behaviors in early-divergent animals.Comment: 20 pages, 4 figure

The Hungry Fly: Hydrodynamics of feeding in the common house fly

A large number of insect species feed primarily on a fluid diet. To do so, they must overcome the numerous challenges that arise in the design of high-efficiency, miniature pumps. Although the morphology of insect feeding structures has been described for decades, their dynamics remain largely unknown even in the most well studied species (e.g. fruit fly). Here, in the fluid dynamics video, we demonstrate in-vivo imaging and microsurgery to elucidate the design principles of feeding structures of the common house fly. Using high-resolution X-ray absorption microscopy, we record in-vivo flow of sucrose solutions through the body over many hours during fly feeding. Borrowing from microsurgery techniques common in neurophysiology, we are able to perturb the pump to a stall position and thus evaluate function under load conditions. Furthermore, fluid viscosity-dependent feedback is observed for optimal pump performance. As the gut of the fly starts to fill up, feedback from the stretch receptors in the cuticle dictates the effective flow rate. Finally, via comparative analysis between the housefly, blow fly, fruit fly and bumble bees, we highlight the common design principles and the role of interfacial phenomena in feeding.Comment: Two videos are included with this submissio

Generation of droplet arrays with rational number spacing patterns driven by a periodic energy landscape

The generation of droplets at low Reynolds numbers is driven by non-linear dynamics that give rise to complex patterns concerning both the droplet-to-droplet spacing and the individual droplet sizes. Here we demonstrate an experimental system in which a time-varying energy landscape provides a periodic magnetic force that generates an array of droplets from an immiscible mixture of ferrofluid and silicone oil. The resulting droplet patterns are periodic, owing to the nature of the magnetic force, yet the droplet spacing and size can vary greatly by tuning a single bias pressure applied on the ferrofluid phase; for a given cycle period of the magnetic force, droplets can be generated either at integer multiples (1, 2, etc.), or at rational fractions (3/2, 5/3, 5/2, etc.) of this period with mono- or multidisperse droplet sizes. We develop a discrete-time dynamical systems model not only to reproduce the phenotypes of the observed patterns but also provide a framework for understanding systems driven by such periodic energy landscapes

Surface Tension dominates Insect Flight on Fluid Interfaces

Flight on the two-dimensional air-water interface, with body weight supported by surface tension, is a unique locomotion strategy well adapted for the environmental niche on the surface of water. Although previously described in phylogenetically basal aquatic insects like stone flies, the biomechanics of interfacial flight has never been analyzed. Here, we report interfacial flight as an adapted behaviour in water-lily beetles (Galerucella nymphaeae, Linnaeus 1758) which are also dexterous airborne fliers. We present the first quantitative biomechanical model of interfacial flight in insects, uncovering an intricate interplay of capillary, aerodynamic and neuromuscular forces. We show that water-lily beetles use their tarsal claws to attach themselves to the interface, via a fluid contact line pinned at the claw. We investigate the kinematics of interfacial flight trajectories using high-speed imaging and construct a mathematical model describing the flight dynamics. Our results show that nonlinear surface tension forces make interfacial flight energetically expensive compared to airborne flight at the relatively high speeds characteristic of water-lily beetles, and cause chaotic dynamics to arise naturally in these regimes. We identify the crucial roles of capillary-gravity wave drag and oscillatory surface tension forces which dominate interfacial flight, showing that the air-water interface presents a radically modified force landscape for flapping wing flight compared to air.Comment: 7 figures, 4 supplementary figures, 12 videos (link given in Supplementary Information