Comparative jet wake structure and swimming performance of salps

Salps are barrel-shaped marine invertebrates that swim by jet propulsion. Morphological variations among species and life-cycle stages are accompanied by differences in swimming mode. The goal of this investigation was to compare propulsive jet wakes and swimming performance variables among morphologically distinct salp species (Pegea confoederata, Weelia (Salpa) cylindrica, Cyclosalpa sp.) and relate swimming patterns to ecological function. Using a combination of in situ dye visualization and particle image velocimetry (PIV) measurements, we describe properties of the jet wake and swimming performance variables including thrust, drag and propulsive efficiency. Locomotion by all species investigated was achieved via vortex ring propulsion. The slow-swimming P. confoederata produced the highest weight-specific thrust (T =53 N kg^(–1)) and swam with the highest wholecycle propulsive efficiency (η_wc= 55%). The fast-swimming W. cylindrica had the most streamlined body shape but produced an intermediate weight-specific thrust (T=30 N kg^(–1)) and swam with an intermediate whole-cycle propulsive efficiency (η_wc =52%). Weak swimming performance variables in the slow-swimming C. affinis, including the lowest weight-specific thrust (T=25 N kg^(–1)) and lowest whole-cycle propulsive efficiency (η_wc=47%), may be compensated by low energetic requirements. Swimming performance variables are considered in the context of ecological roles and evolutionary relationships

Fish swimming in schools save energy regardless of their spatial position

For animals, being a member of a group provides various advantages, such as reduced vulnerability to predators, increased foraging opportunities and reduced energetic costs of locomotion. In moving groups such as fish schools, there are benefits of group membership for trailing individuals, who can reduce the cost of movement by exploiting the flow patterns generated by the individuals swimming ahead of them. However, whether positions relative to the closest neighbours (e.g. ahead, sided by side or behind) modulate the individual energetic cost of swimming is still unknown. Here, we addressed these questions in grey mullet Liza aurata by measuring tail-beat frequency and amplitude of 15 focal fish, swimming in separate schools, while swimming in isolation and in various positions relative to their closest neighbours, at three speeds. Our results demonstrate that, in a fish school, individuals in any position have reduced costs of swimming, compared to when they swim at the same speed but alone. Although fish swimming behind their neighbours save the most energy, even fish swimming ahead of their nearest neighbour were able to gain a net energetic benefit over swimming in isolation, including those swimming at the front of a school. Interestingly, this energetic saving was greatest at the lowest swimming speed measured in our study. Because any member of a school gains an energetic benefit compared to swimming alone, we suggest that the benefits of membership in moving groups may be more strongly linked to reducing the costs of locomotion than previously appreciated

Morphology, Swimming Performance and Propulsive Mode of Six Co-occurring Hydromedusae

Jet propulsion, based on examples from the Hydrozoa, has served as a valuable model for swimming by medusae. However, cnidarian medusae span several taxonomic classes (collectively known as the Medusazoa) and represent a diverse array of morphologies and swimming styles. Does one mode of propulsion appropriately describe swimming by all medusae? This study examined a group of co-occurring hydromedusae collected from the waters of Friday Harbor, WA, USA, to investigate relationships between swimming performance and underlying mechanisms of thrust production. The six species examined encompassed a wide range of bell morphologies and swimming habits. Swimming performance (measured as swimming acceleration and velocity) varied widely among the species and was positively correlated with bell streamlining (measured as bell fineness ratio) and velar structure development (measured as velar aperture ratio). Calculated thrust production due to jet propulsion adequately explained acceleration patterns of prolate medusae (Aglantha digitale, Sarsia sp. and Proboscidactyla flavicirrata) possessing well-developed velums. However, acceleration patterns of oblate medusae (Aequorea victoria, Mitrocoma cellularia and Phialidium gregarium) that have less developed velums were poorly described by jet thrust production. An examination of the wakes behind swimming medusae indicated that, in contrast to the clearly defined jet structures produced by prolate species, oblate medusae did not produce defined jets but instead produced prominent vortices at the bell margins. These vortices are consistent with a predominantly drag-based, rowing mode of propulsion by the oblate species. These patterns of propulsive mechanics and swimming performance relate to the role played by swimming in the foraging ecology of each medusa. These patterns appear to extend beyond hydromedusae and thus have important implications for other members of the Medusazoa

Kinematics of the swimming of Spiroplasma

\emph{Spiroplasma} swimming is studied with a simple model based on resistive-force theory. Specifically, we consider a bacterium shaped in the form of a helix that propagates traveling-wave distortions which flip the handedness of the helical cell body. We treat cell length, pitch angle, kink velocity, and distance between kinks as parameters and calculate the swimming velocity that arises due to the distortions. We find that, for a fixed pitch angle, scaling collapses the swimming velocity (and the swimming efficiency) to a universal curve that depends only on the ratio of the distance between kinks to the cell length. Simultaneously optimizing the swimming efficiency with respect to inter-kink length and pitch angle, we find that the optimal pitch angle is 35.5$^\circ$ and the optimal inter-kink length ratio is 0.338, values in good agreement with experimental observations.Comment: 4 pages, 5 figure

Swimming in curved space or The Baron and the cat

We study the swimming of non-relativistic deformable bodies in (empty) static curved spaces. We focus on the case where the ambient geometry allows for rigid body motions. In this case the swimming equations turn out to be geometric. For a small swimmer, the swimming distance in one stroke is determined by the Riemann curvature times certain moments of the swimmer.Comment: 19 pages 6 figure

Unsteady feeding and optimal strokes of model ciliates

The flow field created by swimming microorganisms not only enables their locomotion but also leads to advective transport of nutrients. In this paper we address analytically and computationally the link between unsteady feeding and unsteady swimming on a model microorganism, the spherical squirmer, actuating the fluid in a time-periodic manner. We start by performing asymptotic calculations at low P\'eclet number (Pe) on the advection-diffusion problem for the nutrients. We show that the mean rate of feeding as well as its fluctuations in time depend only on the swimming modes of the squirmer up to order Pe^(3/2), even when no swimming occurs on average, while the influence of non-swimming modes comes in only at order Pe^2. We also show that generically we expect a phase delay between feeding and swimming of 1/8th of a period. Numerical computations for illustrative strokes at finite Pe confirm quantitatively our analytical results linking swimming and feeding. We finally derive, and use, an adjoint-based optimization algorithm to determine the optimal unsteady strokes maximizing feeding rate for a fixed energy budget. The overall optimal feeder is always the optimal steady swimmer. Within the set of time-periodic strokes, the optimal feeding strokes are found to be equivalent to those optimizing periodic swimming for all values of the P\'eclet number, and correspond to a regularization of the overall steady optimal.Comment: 26 pages, 11 figures, to appear in Journal of Fluid Mechanic

Dispersion of biased swimming microorganisms in a fluid flowing through a tube

Classical Taylor-Aris dispersion theory is extended to describe the transport of suspensions of self-propelled dipolar cells in a tubular flow. General expressions for the mean drift and effective diffusivity are determined exactly in terms of axial moments, and compared with an approximation a la Taylor. As in the Taylor-Aris case, the skewness of a finite distribution of biased swimming cells vanishes at long times. The general expressions can be applied to particular models of swimming microorganisms, and thus be used to predict swimming drift and diffusion in tubular bioreactors, and to elucidate competing unbounded swimming drift and diffusion descriptions. Here, specific examples are presented for gyrotactic swimming algae.Comment: 20 pages, 4 figures. Published version available at http://rspa.royalsocietypublishing.org/content/early/2010/02/09/rspa.2009.0606.short?rss=

Effects of seasonal change on activity rhythms and swimming behavior of age-0 bluefish (Pomatomus saltatrix) and a description of gliding behavior

Daily and seasonal activity rhythms, swimming speed, and modes of swimming were studied in a school of spring-spawned age-0 bluefish (Pomatomus saltatrix) for nine months in a 121-kL research aquarium. Temperature was lowered from 20° to 15°C, then returned to 20°C to match the seasonal cycle. The fish grew from a mean 198 mm to 320 mm (n= 67). Bluefish swam faster and in a more organized school during day (overall mean 47 cm/s) than at night (31 cm/s). Swimming speed declined in fall as temperature declined and accelerated in spring in response to change in photoperiod. Besides powered swimming, bluefish used a gliding-upswimming mode, which has not been previously described for this species. To glide, a bluefish rolled onto its side, ceased body and tail beating, and coasted diagonally downward. Bluefish glided in all months of the study, usually in the dark, and most intensely in winter. Energy savings while the fish is gliding and upswimming may be as much as 20% of the energy used in powered swimming. Additional savings accrue from increased lift due to the hydrofoil created by the horizontal body orientation and slightly concave shape. Energy-saving swimming would be advantageous during migration and overwintering