Kakani Katija

A viscosity-enhanced mechanism for biogenic ocean mixing

Recent observations of biologically generated turbulence in the ocean have led to conflicting conclusions regarding the significance of the contribution of animal swimming to ocean mixing. Measurements indicate elevated turbulent dissipation—comparable with levels caused by winds and tides—in the vicinity of large populations of planktonic animals swimming together. However, it has also been noted that elevated turbulent dissipation is by itself insufficient proof of substantial biogenic mixing, because much of the turbulent kinetic energy of small animals is injected below the Ozmidov buoyancy length scale, where it is primarily dissipated as heat by the fluid viscosity before it can affect ocean mixing. Ongoing debate regarding biogenic mixing has focused on comparisons between animal wake turbulence and ocean turbulence. Here, we show that a second, previously neglected mechanism of fluid mixing—first described over 50 years ago by Charles Darwin— is the dominant mechanism of mixing by swimming animals. The efficiency of mixing by Darwin’s mechanism is dependent on animal shape rather than fluid length scale and, unlike turbulent wake mixing, is enhanced by fluid viscosity. Therefore, it provides a means of biogenic mixing that can be equally effective in small zooplankton and large mammals. A theoretical model for the relative contributions of Darwinian mixing and turbulent wake mixing is created and validated by in situ field measurements of swimming jellyfish using a newly developed scuba-based laser velocimetry device. Extrapolation of these results to other animals is straightforward given knowledge of the animal shape and orientation during vertical migration. On the basis of calculations of a broad range of aquatic animal species, we conclude that biogenic mixing via Darwin’s mechanism can be a significant contributor to ocean mixing and nutrient transport.

Transport and stirring induced by vortex formation

The purpose of this study is to analyse the transport and stirring of fluid that occurs owing to the formation and growth of a laminar vortex ring. Experimental data was collected upstream and downstream of the exit plane of a piston-cylinder apparatus by particle-image velocimetry. This data was used to compute Lagrangian coherent structures to demonstrate how fluid is advected during the transient process of vortex ring formation. Similar computations were performed from computational fluid dynamics (CFD) data, which showed qualitative agreement with the experimental results, although the CFD data provides better resolution in the boundary layer of the cylinder. A parametric study is performed to demonstrate how varying the piston-stroke length-to-diameter ratio affects fluid entrainment during formation. Additionally, we study how regions of fluid are stirred together during vortex formation to help establish a quantitative understanding of the role of vortical flows in mixing. We show that identification of the flow geometry during vortex formation can aid in the determination of efficient stirring. We compare this framework with a traditional stirring metric and show that the framework presented in this paper is better suited for understanding stirring/mixing in transient flow problems. A movie is available with the online version of the paper.

A wake-based correlate of swimming performance and foraging behavior in seven co-occurring jellyfish species.

SUMMARY 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.

Biogenic inputs to ocean mixing

Summary Recent studies have evoked heated debate about whether biologically generated (or biogenic) fluid disturbances affect mixing in the ocean. Estimates of biogenic inputs have shown that their contribution to ocean mixing is of the same order as winds and tides. Although these estimates are intriguing, further study using theoretical, numerical and experimental techniques is required to obtain conclusive evidence of biogenic mixing in the ocean. Biogenic ocean mixing is a complex problem that requires detailed understanding of: (1) marine organism behavior and characteristics (i.e. swimming dynamics, abundance and migratory behavior), (2) mechanisms utilized by swimming animals that have the ability to mix stratified fluids (i.e. turbulence and fluid drift) and (3) knowledge of the physical environment to isolate contributions of marine organisms from other sources of mixing. In addition to summarizing prior work addressing the points above, observations on the effect of animal swimming mode and body morphology on biogenic fluid transport will also be presented. It is argued that to inform the debate on whether biogenic mixing can contribute to ocean mixing, our studies should focus on diel vertical migrators that traverse stratified waters of the upper pycnocline. Based on our understanding of mixing mechanisms, body morphologies, swimming modes and body orientation, combined with our knowledge of vertically migrating populations of animals, it is likely that copepods, krill and some species of gelatinous zooplankton and fish have the potential to be strong sources of biogenic mixing.

Propulsion in Cubomedusae: Mechanisms and Utility

Evolutionary constraints which limit the forces produced during bell contractions of medusae affect the overall medusan morphospace such that jet propulsion is limited to only small medusae. Cubomedusae, which often possess large prolate bells and are thought to swim via jet propulsion, appear to violate the theoretical constraints which determine the medusan morphospace. To examine propulsion by cubomedusae, we quantified size related changes in wake dynamics, bell shape, swimming and turning kinematics of two species of cubomedusae, Chironex fleckeri and Chiropsella bronzie. During growth, these cubomedusae transitioned from using jet propulsion at smaller sizes to a rowing-jetting hybrid mode of propulsion at larger sizes. Simple modifications in the flexibility and kinematics of their velarium appeared to be sufficient to alter their propulsive mode. Turning occurs during both bell contraction and expansion and is achieved by generating asymmetric vortex structures during both stages of the swimming cycle. Swimming characteristics were considered in conjunction with the unique foraging strategy used by cubomedusae.

Circulation Generation and Vortex Ring Formation by Conic Nozzles

Vortex rings are one of the fundamental flow structures in nature. In this paper, the generation of circulation and vortex rings by a vortex generator with a static converging conic nozzle exit is studied numerically. Conic nozzles can manipulate circulation and other flow invariants by accelerating the flow, increasing the Reynolds number, and by establishing a two-dimensional flow at the exit. The increase in the circulation efflux is accompanied by an increase in the vortex circulation. A novel normalization method is suggested to differentiate between two contributions to the circulation generation: a one-dimensional slug-type flow contribution and an inherently two-dimensional flow contribution. The one-dimensional contribution to the circulation increases with the square of the centerline exit velocity, while the two-dimensional contribution increases linearly with the decrease in the exit diameter. The two-dimensional flow contribution to the circulation production is not limited to the impulsive initiation of the flow only (as in straight tube vortex generators), but it persists during the entire ejection. The two-dimensional contribution can reach as much as 44% of the total circulation (in the case of an orifice). The present study offers evidences on the importance of the vortex generator geometry, and in particular, the exit configuration on the emerging flow, circulation generation, and vortex ring formation. It is shown that both total and vortex ring circulations can be controlled to some extent by the shape of the exit nozzle.

Ontogenetic propulsive transitions by Sarsia tubulosa medusae

ABSTRACT While swimming in their natural environment, marine organisms must successfully forage, escape from predation, and search for mates to reproduce. In the process, planktonic organisms interact with their fluid environment, generating fluid signatures around their body and in their downstream wake through ontogeny. In the early stages of their life cycle, marine organisms operate in environments where viscous effects dominate and govern physical processes. Ontogenetic propulsive transitions in swimming organisms often involve dramatic changes in morphology and swimming behavior. However, for organisms that do not undergo significant changes in morphology, swimming behavior or propulsive mode, how is their swimming performance affected? We investigated the ontogenetic propulsive transitions of the hydromedusa Sarsia tubulosa, which utilizes jet propulsion and possesses a similar bell morphology throughout its life cycle. We used digital particle image velocimetry and high-speed imaging to measure the body kinematics, velocity fields and wake structures induced by swimming S. tubulosa with bell exit diameters from 1 to 10 mm. Our experimental observations revealed three distinct classes of hydrodynamic wakes: elongated vortex rings for 1030 (larger than 2 mm bell exit diameter) and elliptical vortex rings (or leading vortex rings) followed by trailing jets for most instances where Re>100 (larger than 4 or 5 mm bell exit diameter). The relative travel distance and propulsive efficiency remained unchanged throughout ontogeny, and the swimming proficiency and hydrodynamic cost of transport decreased non-linearly. Summary: For organisms that maintain morphology and swimming behaviour over ontogeny, we show that distinct propulsive wake structures alter swimming performance and energetics.

Fluid interactions that enable stealth predation by the upstream-foraging hydromedusa Craspedacusta sowerbyi.

Unlike most medusae that forage with tentacles trailing behind their bells, several species forage upstream of their bells using aborally located tentacles. It has been hypothesized that these medusae forage as stealth predators by placing their tentacles in more quiescent regions of flow around their bells. Consequently, they are able to capture highly mobile, sensitive prey. We used digital particle image velocimetry (DPIV) to quantitatively characterize the flow field around Craspedacusta sowerbyi, a freshwater upstream-foraging hydromedusa, to evaluate the mechanics of its stealth predation. We found that fluid velocities were minimal in front and along the sides of the bell where the tentacles are located. As a result, the deformation rates in the regions where the tentacles are located were low, below the threshold rates required to elicit an escape response in several species of copepods. Estimates of their encounter volume rates were examined on the basis of flow past the tentacles, and trade-offs associated with tentacle characteristics were evaluated.

Quantitatively Measuring In situ Flows using a Self-Contained Underwater Velocimetry Apparatus (SCUVA)

The ability to directly measure velocity fields in a fluid environment is necessary to provide empirical data for studies in fields as diverse as oceanography, ecology, biology, and fluid mechanics. Field measurements introduce practical challenges such as environmental conditions, animal availability, and the need for field-compatible measurement techniques. To avoid these challenges, scientists typically use controlled laboratory environments to study animal-fluid interactions. However, it is reasonable to question whether one can extrapolate natural behavior (i.e., that which occurs in the field) from laboratory measurements. Therefore, in situ quantitative flow measurements are needed to accurately describe animal swimming in their natural environment. We designed a self-contained, portable device that operates independent of any connection to the surface, and can provide quantitative measurements of the flow field surrounding an animal. This apparatus, a self-contained underwater velocimetry apparatus (SCUVA), can be operated by a single scuba diver in depths up to 40 m. Due to the added complexity inherent of field conditions, additional considerations and preparation are required when compared to laboratory measurements. These considerations include, but are not limited to, operator motion, predicting position of swimming targets, available natural suspended particulate, and orientation of SCUVA relative to the flow of interest. The following protocol is intended to address these common field challenges and to maximize measurement success.