Fast-swimming hydromedusae exploit velar kinematics to form an optimal vortex wake

Fast-swimming hydromedusan jellyfish possess a characteristic funnel-shaped velum at the exit of their oral cavity that interacts with the pulsed jets of water ejected during swimming motions. It has been previously assumed that the velum primarily serves to augment swimming thrust by constricting the ejected flow in order to produce higher jet velocities. This paper presents high-speed video and dye-flow visualizations of free-swimming Nemopsis bachei hydromedusae, which instead indicate that the time-dependent velar kinematics observed during the swimming cycle primarily serve to optimize vortices formed by the ejected water rather than to affect the speed of the ejected flow. Optimal vortex formation is favorable in fast-swimming jellyfish because, unlike the jet funnelling mechanism, it allows for the minimization of energy costs while maximizing thrust forces. However, the vortex `formation number' corresponding to optimality in N. bachei is substantially greater than the value of 4 found in previous engineering studies of pulsed jets from rigid tubes. The increased optimal vortex formation number is attributable to the transient velar kinematics exhibited by the animals. A recently developed model for instantaneous forces generated during swimming motions is implemented to demonstrate that transient velar kinematics are required in order to achieve the measured swimming trajectories. The presence of velar structures in fast-swimming jellyfish and the occurrence of similar jet-regulating mechanisms in other jet-propelled swimmers (e.g. the funnel of squid) appear to be a primary factor contributing to success of fast-swimming jetters, despite their primitive body plans

Morphological diversity of medusan lineages constrained by animal–fluid interactions

Cnidarian medusae, commonly known as jellyfish, represent the earliest known animal taxa to achieve locomotion using muscle power. Propulsion by medusae requires the force of bell contraction to generate forward thrust. However, thrust production is limited in medusae by the primitive structure of their epitheliomuscular cells. This paper demonstrates that constraints in available locomotor muscular force result in a trade-off between high-thrust swimming via jet propulsion and high-efficiency swimming via a combined jet-paddling propulsion. This trade-off is reflected in the morphological diversity of medusae, which exhibit a range of fineness ratios (i.e. the ratio between bell height and diameter) and small body size in the high-thrust regime, and low fineness ratios and large body size in the high-efficiency regime. A quantitative model of the animal–fluid interactions that dictate this trade-off is developed and validated by comparison with morphological data collected from 660 extant medusan species ranging in size from 300 µm to over 2 m. These results demonstrate a biomechanical basis linking fluid dynamics and the evolution of medusan bell morphology. We believe these to be the organising principles for muscle-driven motility in Cnidaria

A Wake-Based Correlate of Swimming Performance and Foraging Behavior in Seven Co-Occurring Jellyfish Species

It is generally accepted that animal–fluid interactions have shaped the evolution of animals that swim and fly. However, the functional ecological advantages associated with those adaptations are currently difficult to predict on the basis of measurements of the animal–fluid interactions. We report the identification of a robust, fluid dynamic correlate of distinct ecological functions in seven jellyfish species that represent a broad range of morphologies and foraging modes. Since the comparative study is based on properties of the vortex wake – specifically, a fluid dynamical concept called optimal vortex formation – and not on details of animal morphology or phylogeny, we propose that higher organisms can also be understood in terms of these fluid dynamic organizing principles. This enables a quantitative, physically based understanding of how alterations in the fluid dynamics of aquatic and aerial animals throughout their evolution can result in distinct ecological functions

Flow patterns generated by oblate medusan jellyfish: field measurements and laboratory analyses

Flow patterns generated by medusan swimmers such as jellyfish are known to differ according the morphology of the various animal species. Oblate medusae have been previously observed to generate vortex ring structures during the propulsive cycle. Owing to the inherent physical coupling between locomotor and feeding structures in these animals, the dynamics of vortex ring formation must be robustly tuned to facilitate effective functioning of both systems. To understand how this is achieved, we employed dye visualization techniques on scyphomedusae (Aurelia aurita) observed swimming in their natural marine habitat. The flow created during each propulsive cycle consists of a toroidal starting vortex formed during the power swimming stroke, followed by a stopping vortex of opposite rotational sense generated during the recovery stroke. These two vortices merge in a laterally oriented vortex superstructure that induces flow both toward the subumbrellar feeding surfaces and downstream. The lateral vortex motif discovered here appears to be critical to the dual function of the medusa bell as a flow source for feeding and propulsion. Furthermore, vortices in the animal wake have a greater volume and closer spacing than predicted by prevailing models of medusan swimming. These effects are shown to be advantageous for feeding and swimming performance, and are an important consequence of vortex interactions that have been previously neglected

3D Particle Tracking Velocimetry Method: Advances and Error Analysis

A full three-dimensional particle tracking system was developed and tested. By using three separate CCDs placed at the vertices of an equilateral triangle, the threedimensional location of particles can be determined. Particle locations measured at two different times can then be used to create a three-component, three-dimensional velocity field. Key developments are: the ability to accurately process overlapping particle images, offset CCDs to significantly improve effective resolution, allowance for dim particle images, and a hybrid particle tracking technique ideal for three-dimensional flows when only two sets of images exist. An in-depth theoretical error analysis was performed which gives the important sources of error and their effect on the overall system. This error analysis was verified through a series of experiments, which utilized a test target with 100 small dots per square inch. For displacements of 2.54mm the mean errors were less than 2% and the 90% confidence limits were less than 5.2 μm in the plane perpendicular to the camera axis, and 66 μm in the direction of the camera axis. The system was used for flow measurements around a delta wing at an angle of attack. These measurements show the successful implementation of the system for three-dimensional flow velocimetry

Electric Circuit Simulation of Floquet Topological Insulators

We present a method for simulating any non-interacting and time-periodic tight-binding Hamiltonian in Fourier space using electric circuits made of inductors and capacitors. We first map the time-periodic Hamiltonian to a Floquet Hamiltonian, which converts the time dimension into a Floquet dimension. In electric circuits, this Floquet dimension is simulated as an extra spatial dimension without any time dependency in the electrical elements. The number of replicas needed in the Floquet Hamiltonian depends on the frequency and strength of the drive. We also demonstrate that we can detect the topological edge states (including the anomalous edge states in the dynamical gap) in an electric circuit by measuring the two-point impedance between the nodes. Our method paves a simple and promising way to explore and control Floquet topological phases in electric circuits.Comment: 6 pages, 5 figure

Phenotypic Plasticity in Juvenile Jellyfish Medusae Facilitates Effective Animal–Fluid Interaction

Locomotion and feeding in marine animals are intimately linked to the flow dynamics created by specialized body parts. This interaction is of particular importance during ontogeny, when changes in behaviour and scale challenge the organism with shifts in fluid regimes and altered functionality. Previous studies have indicated that Scyphozoan jellyfish ontogeny accommodates the changes in fluid dynamics associated with increasing body dimensions and velocities during development. However, in addition to scale and behaviour that—to a certain degree—underlie the control of the animal, flow dynamics are also dependent on external factors such as temperature. Here, we show phenotypic plasticity in juvenile Aurelia aurita medusae, where morphogenesis is adapted to altered fluid regimes imposed by changes in ambient temperature. In particular, differential proportional growth was found to compensate for temperature-dependent changes in viscous effects, enabling the animal to use adhering water boundary layers as ‘paddles’—and thus economize tissue—at low temperatures, while switching to tissue-dominated propulsion at higher temperatures where the boundary layer thickness is insufficient to serve for paddling. This effect was predicted by a model of animal–fluid interaction and confirmed empirically by flow-field visualization and assays of propulsion efficiency

Floquet states and optical conductivity of an irradiated two dimensional topological insulator

We study the topology of the Floquet states and time-averaged optical conductivity of the lattice model of a thin topological insulator subject to a circularly polarized light using the extended Kubo formalism. Two driving regimes, the off-resonant and on-resonant, and two models for the occupation of the Floquet states, the ideal and mean-energy occupation, are considered. In the ideal occupation, the real part of DC optical Hall conductivity is shown to be quantized while it is not quantized for the mean energy distribution. The optical transitions in the Floquet band structure depend strongly on the occupation and also the optical weight which consequently affect all components of optical conductivity. At high frequency regime, we present an analytical calculation of the effective Hamiltonian and also its phase diagram which depends on the tunneling energy between two surfaces. The topology of the system shows rich phases when it is irradiated by a weak on-resonant drive giving rise to emergence of anomalous edge states.Comment: 11 pages, 8 figure

Functional Morphology and Fluid Interactions During Early Development of the Scyphomedusa Aurelia aurita

Scyphomedusae undergo a predictable ontogenetic transition from a conserved, universal larval form to a diverse array of adult morphologies. This transition entails a change in bell morphology from a highly discontinuous ephyral form, with deep clefts separating eight discrete lappets, to a continuous solid umbrella-like adult form. We used a combination of kinematic, modeling, and flow visualization techniques to examine the function of the medusan bell throughout the developmental changes of the scyphomedusa Aurelia aurita. We found that flow around swimming ephyrae and their lappets was relatively viscous (1 < Re < 10) and, as a result, ephyral lappets were surrounded by thick, overlapping boundary layers that occluded flow through the gaps between lappets. As medusae grew, their fluid environment became increasingly influenced by inertial forces (10 < Re < 10,000) and, simultaneously, clefts between the lappets were replaced by organic tissue. Hence, although the bell undergoes a structural transition from discontinuous (lappets with gaps) to continuous (solid bell) surfaces during development, all developmental stages maintain functionally continuous paddling surfaces. This developmental pattern enables ephyrae to efficiently allocate tissue to bell diameter increase via lappet growth, while minimizing tissue allocation to inter-lappet spaces that maintain paddle function due to boundary layer overlap