Vectored jets power arms-first and tail-first turns differently in brief squid with assistance from fins and keeled arms.

Squids maneuver to capture prey, elude predators, navigate complex habitats and deny rivals access to mates. Despite the ecological importance of this essential locomotive function, limited quantitative data on turning performance and wake dynamics of squids are available. To better understand the contribution of the jet, fins and arms to turns, the role of orientation (i.e. arms first versus tail first) in maneuvering, and the relationship between jet flow and turning performance, kinematic and 3D velocimetry data were collected in tandem from brief squid, Lolliguncula brevis. The pulsed jet, which can be vectored to direct flows, was the primary driver of most turning behaviors, producing flows with the highest impulse magnitude and angular impulse about the main axis of the turn (yaw) and secondary axes (roll and pitch). The fins and keeled arms played subordinate but important roles in turning performance, contributing to angular impulse, stabilizing the maneuver along multiple axes and/or reducing rotational resistance. Orientation affected turning performance and dynamics, with tail-first turns being associated with greater impulse and angular impulse, longer jet structures, higher jet velocities and greater angular turning velocities than arms-first turns. Conversely, arms-first turns involved shorter, slower jets with less impulse, but these directed short pulses resulted in lower minimum length-specific turning radii. Although the length-to-diameter ratio (L/D) of ejected jet flow was a useful metric for characterizing vortical flow features, it, by itself, was not a reliable predictor of angular velocity or turning radii, which reflects the complexity of the squid multi-propulsor system.

Rheology of particulate suspensions with non-Newtonian fluids in capillaries.

Particulate suspensions occur in situations from blood flow to slurries in drilling applications. Existing investigations of these suspensions generally concentrate on the impact of particle volume fraction for suspensions in Newtonian fluids under free-flow conditions. Recently, particulate-polymer composites have been used in additive manufacturing (AM). Here, the polymer becomes a shear-thinning non-Newtonian fluid during extrusion, creating a particulate suspension. Motivated by the challenges in AM of particulate composites, this study investigates the rheology of suspensions of micrometre-sized particles in shear-thinning silicone while extruded through AM-scaled nozzles (millimetre-scale diameters). The suspensions were observed to follow a power-law behaviour and their rheology was investigated through the measured flow consistency ( K ) and behaviour ( n ) indices. The impact of the particle volume fraction ( ϕ ) and the ratio ( ω ) of the capillary inside diameter to the particle diameter on both indices were measured. n was found to be only impacted by the suspension fluid type and ϕ . K was found to be constant at large ω , but decreased and then increased to infinity with ω decreasing. Based on its behaviour, K was categorized into two conditions and analysed separately with semi-empirical models. The impact of particle size distribution was also investigated.

Squids use multiple escape jet patterns throughout ontogeny.

Throughout their lives, squids are both predators and prey for a multitude of animals, many of which are at the top of ocean food webs, making them an integral component of the trophic structure of marine ecosystems. The escape jet, which is produced by the rapid expulsion of water from the mantle cavity through a funnel, is central to a cephalopod's ability to avoid predation throughout its life. Although squid undergo morphological and behavioral changes and experience remarkably different Reynolds number regimes throughout their development, little is known about the dynamics and propulsive efficiency of escape jets throughout ontogeny. We examine the hydrodynamics and kinematics of escape jets in squid throughout ontogeny using 2D/3D velocimetry and high-speed videography. All life stages of squid produced two escape jet patterns: (1) 'escape jet I' characterized by short rapid pulses resulting in vortex ring formation and (2) 'escape jet II' characterized by long high-volume jets, often with a leading-edge vortex ring. Paralarvae exhibited higher propulsive efficiency than adult squid during escape jet ejection, and propulsive efficiency was higher for escape jet I than escape jet II in juveniles and adults. These results indicate that although squid undergo major ecological transitions and morphology changes from paralarvae to adults, all life stages demonstrate flexibility in escape jet responses and produce escape jets of surprisingly high propulsive efficiency.This article has an associated First Person interview with the first author of the paper.

Quantitative classification of vortical flows based on topological features using graph matching.

Vortical flow patterns generated by swimming animals or flow separation (e.g. behind bluff objects such as cylinders) provide important insight to global flow behaviour such as fluid dynamic drag or propulsive performance. The present work introduces a new method for quantitatively comparing and classifying flow fields using a novel graph-theoretic concept, called a weighted Gabriel graph, that employs critical points of the velocity vector field, which identify key flow features such as vortices, as graph vertices. The edges (connections between vertices) and edge weights of the weighted Gabriel graph encode local geometric structure. The resulting graph exhibits robustness to minor changes in the flow fields. Dissimilarity between flow fields is quantified by finding the best match (minimum difference) in weights of matched graph edges under relevant constraints on the properties of the edge vertices, and flows are classified using hierarchical clustering based on computed dissimilarity. Application of this approach to a set of artificially generated, periodic vortical flows demonstrates high classification accuracy, even for large perturbations, and insensitivity to scale variations and number of periods in the periodic flow pattern. The generality of the approach allows for comparison of flows generated by very different means (e.g. different animal species).

New approaches for assessing squid fin motions: coupling proper orthogonal decomposition with volumetric particle tracking velocimetry.

Squid, which swim using a coupled fin/jet system powered by muscular hydrostats, pose unique challenges for the study of locomotion. The high flexibility of the fins and complex flow fields generated by distinct propulsion systems require innovative techniques for locomotive assessment. For this study, we used proper orthogonal decomposition (POD) to decouple components of the fin motions and defocusing digital particle tracking velocimetry (DDPTV) to quantify the resultant 3D flow fields. Kinematic footage and DDPTV data were collected from brief squid, Lolliguncula brevis [3.1-6.5 cm dorsal mantle length (DML)], swimming freely in a water tunnel at speeds of 0.39-7.20 DML s-1 Both flap and wave components were present in all fin motions, but the relative importance of the wave components was higher for arms-first swimming than for tail-first swimming and for slower versus higher speed swimming. When prominent wave components were present, more complex interconnected vortex ring wakes were observed, while fin movements dominated by flapping resulted in more spatially separated vortex ring patterns. Although the jet often produced the majority of the thrust for steady rectilinear swimming, our results demonstrated that the fins can contribute more thrust than the jet at times, consistently produce comparable levels of lift to the jet during arms-first swimming, and can boost overall propulsive efficiency. By producing significant drag signatures, the fins can also aid in stabilization and maneuvering. Clearly, fins play multiple roles in squid locomotion, and when coupled with the jet, allow squid to perform a range of swimming behaviors integral to their ecological success.

Turning performance of brief squid Lolliguncula brevis during attacks on shrimp and fish.

Although squid are generally considered to be effective predators, little is currently known of how squid maneuver and position themselves during prey strikes. In this study, high-speed video and kinematic analyses were used to study attacks by the brief squid Lolliguncula brevis on both shrimp and fish. Squid attack success was high (>80%) and three behavioral phases were identified: (1) approach, (2) strike and (3) recoil. Lolliguncula brevis demonstrated greater maneuverability (i.e. a smaller length-specific turning radius) and employed more body adjustments (i.e. mantle angle posturing) during approaches toward shrimp versus fish. Squid exhibited higher linear approach/strike velocities and accelerations with faster-swimming fish prey compared with slower shrimp prey. Agility (i.e. turning rate) during prey encounters was comparable to performance extremes observed during non-predatory turns, and did not differ according to prey type or distance. Despite having the ability to modulate tentacle extension velocity, squid instead increased their own swimming velocity rather than increasing tentacle velocity when targeting faster fish prey during the strike phase, but this was not the case for shrimp prey. Irrespective of prey type, L. brevis consistently positioned themselves above the prey target prior to the tentacle strike, possibly to facilitate a more advantageous downward projection of the tentacles. During the recoil, L. brevis demonstrated length-specific turning radii similar to those recorded during the approach despite vigorous escape attempts by some prey. Clearly, turning performance is integral to prey attacks in squid, with differences in attack strategy varying depending on the prey target.

Multiple sensory modalities used by squid in successful predator evasion throughout ontogeny.

Squid rely on multiple sensory systems for predator detection. In this study we examine the role of two sensory systems, the lateral line analogue and vision, in successful predator evasion throughout ontogeny. Squid Doryteuthis pealeii and Lolliguncula brevis were recorded using high-speed videography in the presence of natural predators under light and dark conditions with their lateral line analogue intact or ablated via a pharmacological technique. Paralarval squid showed reduced escape responses when ablated; however, no differences were found between light and dark conditions in non-ablated paralarvae, as was previously shown in juveniles and adults, indicating that the lateral line analogue is integral for predator detection early in life. However, vision does play a role in survival because ablated squid in dark conditions had lower levels of survival than all other treatments. Throughout ontogeny, squid oriented themselves anteriorly towards the oncoming predator, maximizing sensory input to the lateral line analogue system and providing better positioning for tail-first escape jetting, the preferred escape mode. Ablated juveniles and adults had lower response times, escape velocities and peak acceleration than non-ablated individuals, indicating that the lateral line analogue enables squid to respond quicker and with more powerful jets to a predator and maximize escape success. Our findings reveal that the lateral line analogue plays a role in predator detection and successful escape response at the earliest life stages, and continues to contribute to successful evasion by aiding visual cues in juvenile and adult squid.

Turning performance in squid and cuttlefish: unique dual-mode, muscular hydrostatic systems.

Although steady swimming has received considerable attention in prior studies, unsteady swimming movements represent a larger portion of many aquatic animals' locomotive repertoire and have not been examined extensively. Squids and cuttlefishes are cephalopods with unique muscular hydrostat-driven, dual-mode propulsive systems involving paired fins and a pulsed jet. These animals exhibit a wide range of swimming behavior, but turning performance has not been examined quantitatively. Brief squid, Lolliguncula brevis, and dwarf cuttlefish, Sepia bandensis, were filmed during turns using high-speed cameras. Kinematic features were tracked, including the length-specific radius of the turn (R/L), a measure of maneuverability, and angular velocity (ω), a measure of agility. Both L. brevis and S. bandensis demonstrated high maneuverability, with (R/L)min values of 3.4×10(-3)±5.9×10(-4) and 1.2×10(-3)±4.7×10(-4) (mean±s.e.m.), respectively, which are the lowest measures of R/L reported for any aquatic taxa. Lolliguncula brevis exhibited higher agility than S. bandensis (ωa,max=725.8 versus 485.0 deg s(-1)), and both cephalopods have intermediate agility when compared with flexible-bodied and rigid-bodied nekton of similar size, reflecting their hybrid body architecture. In L. brevis, jet flows were the principal driver of angular velocity. Asymmetric fin motions played a reduced role, and arm wrapping increased turning performance to varying degrees depending on the species. This study indicates that coordination between the jet and fins is important for turning performance, with L. brevis achieving faster turns than S. bandensis and S. bandensis achieving tighter, more controlled turns than L. brevis.

Volumetric flow imaging reveals the importance of vortex ring formation in squid swimming tail-first and arms-first.

Squids use a pulsed jet and fin movements to swim both arms-first (forward) and tail-first (backward). Given the complexity of the squid multi-propulsor system, 3D velocimetry techniques are required for the comprehensive study of wake dynamics. Defocusing digital particle tracking velocimetry, a volumetric velocimetry technique, and high-speed videography were used to study arms-first and tail-first swimming of brief squid Lolliguncula brevis over a broad range of speeds [0-10 dorsal mantle lengths (DML) s(-1)] in a swim tunnel. Although there was considerable complexity in the wakes of these multi-propulsor swimmers, 3D vortex rings and their derivatives were prominent reoccurring features during both tail-first and arms-first swimming, with the greatest jet and fin flow complexity occurring at intermediate speeds (1.5-3.0 DML s(-1)). The jet generally produced the majority of thrust during rectilinear swimming, increasing in relative importance with speed, and the fins provided no thrust at speeds >4.5 DML s(-1). For both swimming orientations, the fins sometimes acted as stabilizers, producing negative thrust (drag), and consistently provided lift at low/intermediate speeds (<2.0 DML s(-1)) to counteract negative buoyancy. Propulsive efficiency (η) increased with speed irrespective of swimming orientation, and η for swimming sequences with clear isolated jet vortex rings was significantly greater (η=78.6±7.6%, mean±s.d.) than that for swimming sequences with clear elongated regions of concentrated jet vorticity (η=67.9±19.2%). This study reveals the complexity of 3D vortex wake flows produced by nekton with hydrodynamically distinct propulsors.

Effect of vehicle configuration on the performance of a submersible pulsed-jet vehicle at intermediate Reynolds number.

Recent results have demonstrated that pulsed-jet propulsion can achieve propulsive efficiency greater than that for steady jets when short, high frequency pulses are used, and the pulsed-jet advantage increases as Reynolds number decreases into the intermediate range (∼50). An important aspect of propulsive performance, however, is the vehicle configuration. The nozzle configuration influences the jet speed and, in the case of pulsed-jets, the formation of the vortex rings with each jet pulse, which have important effects on thrust. Likewise, the hull configuration influences the vehicle speed through its effect on drag. To investigate these effects, several flow inlet, nozzle, and hull tail configurations were tested on a submersible, self-propelled pulsed-jet vehicle ('Robosquid' for short) for jet pulse length-to-diameter ratios (L/D) in the range 0.5-6 and pulsing duty cycles (St(L)) of 0.2 and 0.5. For the configurations tested, the vehicle Reynolds number (Re(υ)) ranged from 25 to 110. In terms of propulsive efficiency, changing between forward and aft-facing inlets had little effect for the conditions considered, but changing from a smoothly tapered aft hull section to a blunt tail increased propulsive efficiency slightly due to reduced drag for the blunt tail at intermediate Re(υ). Sharp edged orifices also showed increased vehicle velocity and propulsive efficiency in comparison to smooth nozzles, which was associated with stronger vortex rings being produced by the flow contraction through the orifice. Larger diameter orifices showed additional gains in propulsive efficiency over smaller orifices if the rate of mass flow was matched with the smaller diameter cases, but using the same maximum jet velocity with the larger diameter decreased the propulsive efficiency relative to the smaller diameter cases.

The effect of Reynolds number on the propulsive efficiency of a biomorphic pulsed-jet underwater vehicle.

The effect of Reynolds number on the propulsive efficiency of pulsed-jet propulsion was studied experimentally on a self-propelled, pulsed-jet underwater vehicle, dubbed Robosquid due to the similarity of its propulsion system with squid. Robosquid was tested for jet slug length-to-diameter ratios (L/D) in the range 2-6 and dimensionless frequency (St(L)) in the range 0.2-0.6 in a glycerin-water mixture. Digital particle image velocimetry was used for measuring the impulse and energy of jet pulses from the velocity and vorticity fields of the jet flow to calculate the pulsed-jet propulsive efficiency, and compare it with an equivalent steady jet system. Robosquid's Reynolds number (Re) based on average vehicle velocity and vehicle diameter ranged between 37 and 60. The current results for propulsive efficiency were compared to the previously published results in water where Re ranged between 1300 and 2700. The results showed that the average propulsive efficiency decreased by 26% as the average Re decreased from 2000 to 50 while the ratio of pulsed-jet to steady jet efficiency (η(P)/η(P, ss)) increased up to 0.15 (26%) as the Re decreased over the same range and for similar pulsing conditions. The improved η(P)/η(P, ss) at lower Re suggests that pulsed-jet propulsion can be used as an efficient propulsion system for millimeter-scale propulsion applications. The Re = 37-60 conditions in the present investigation, showed a reduced dependence of η(P) and η(P)/η(P, ss)on L/D compared to higher Re results. This may be due to the lack of clearly observed vortex ring pinch-off as L/D increased for this Re regime.

Propulsive efficiency of a biomorphic pulsed-jet underwater vehicle.

The effect of the velocity program and duty cycle (St(L)) on the propulsive efficiency of pulsed-jet propulsion was studied experimentally on a self-propelled, pulsed-jet underwater vehicle, dubbed Robosquid due to the similarity of essential elements of its propulsion system with squid jet propulsion. Robosquid was tested for jet slug length-to-diameter ratios (L/D) in the range 2-6 and St(L) in the range 0.2-0.6 with jet velocity programs commanded to be triangular or trapezoidal. Digital particle image velocimetry was used for measuring the impulse and energy of jet pulses to calculate the pulsed-jet propulsive efficiency and compare it with an equivalent steady jet system. Robosquid's Reynolds number (Re) based on average vehicle velocity and vehicle diameter ranged between 1300 and 2700 for the conditions tested. The results indicated better propulsive efficiency of the trapezoidal velocity program (up to 20% higher) compared to the triangular velocity program. Also, an increase in the ratio of the pulsed-jet propulsive efficiency to the equivalent steady jet propulsive efficiency (eta(P)/eta(P, ss)) was observed as St(L) increased and L/D decreased. For cases of short L/D and high St(L), eta(P)/eta(P, ss) was found to be as high as 1.2, indicating better performance of pulsed jets. This result demonstrates a case where propulsion using essential elements of a biological locomotion system can outperform the traditional mechanical system equivalent in terms of efficiency. It was also found that changes in St(L) had a proportionately larger effect on propulsive efficiency compared to changes in L/D. A simple model is presented to explain the results in terms of the contribution of over-pressure at the nozzle exit plane associated with the formation of vortex rings with each jet pulse.

Hydrodynamic fin function of brief squid, Lolliguncula brevis.

Although the pulsed jet is often considered the foundation of a squid's locomotive system, the lateral fins also probably play an important role in swimming, potentially providing thrust, lift and dynamic stability as needed. Fin morphology and movement vary greatly among squid species, but the locomotive role of the fins is not well understood. To begin to elucidate the locomotive role of the fins in squids, fin hydrodynamics were studied in the brief squid Lolliguncula brevis, a species that exhibits a wide range of fin movements depending on swimming speed. Individual squid were trained to swim in both the arms-first and tail-first orientations against currents in a water tunnel seeded with light-reflective particles. Particle-laden water around the fins was illuminated with lasers and videotaped so that flow dynamics around the fins could be analyzed using digital particle image velocimetry (DPIV). Time-averaged forces generated by the fin were quantified from vorticity fields of the fin wake. During the low swimming speeds considered in this study [<2.5 dorsal mantle lengths (DML) per second], L. brevis exhibited four unique fin wake patterns, each with distinctive vortical structures: (1) fin mode I, in which one vortex is shed with each downstroke, generally occurring at low speeds; (2) fin mode II, an undulatory mode in which a continuous linked chain of vortices is produced; (3) fin mode III, in which one vortex is shed with each downstroke and upstroke, and; (4) fin mode IV, in which a discontinuous chain of linked double vortex structures is produced. All modes were detected during tail-first swimming but only fin modes II and III were observed during arms-first swimming. The fins produced horizontal and vertical forces of varying degrees depending on stroke phase, swimming speed, and swimming orientation. During tail-first swimming, the fins functioned primarily as stabilizers at low speeds before shifting to propulsors as speed increased, all while generating net lift. During arms-first swimming, the fins primarily provided lift with thrust production playing a reduced role. These results demonstrate the lateral fins are an integral component of the complex locomotive system of L. brevis, producing lift and thrust forces through different locomotive modes.

Sensitivity of Lagrangian coherent structure identification to flow field resolution and random errors.

The effect of spatial and temporal resolutions and random errors on identification of Lagrangian coherent structures (LCSs) from Eulerian velocity fields is evaluated using two canonical flows: a two-dimensional vortex pair and a vortex ring formed by transient ejection of a jet from a tube. The flow field for the vortex pair case was steady and obtained analytically while the transient vortex ring flow was simulated using computational fluid dynamics. To evaluate resolution and random error effects, the flow fields were degraded by locally smoothing the flow and sampling it on a sparser grid to reduce spatial resolution, adding Gaussian distributed random noise to provide random errors, and/or subsampling the time series of vector fields to reduce the temporal resolution (the latter applying only for the vortex ring case). The degradation methods were meant to emulate distortions and errors introduced in common flow measurement methods such as digital particle image velocimetry. Comparing the LCS corresponding to the vortex boundary (separatrix) obtained from the degraded velocity fields with the true separatrix (obtained analytically for the vortex pair case or from high resolution, noise-free velocity fields for the vortex ring case) showed that noise levels as low as 5%-10% of the vortex velocity can cause the separatrix to significantly deviate from its true location in a random fashion, but the "mean" location still remained close to the true location. Temporal and spatial resolution degradations were found to primarily affect transient portions of the flow with strong spatial gradients. Significant deviations in the location of the separatrix were observed even for spatial resolutions as high as 2% of the jet diameter for the vortex ring case.

Hydrodynamics of pulsed jetting in juvenile and adult brief squid Lolliguncula brevis: evidence of multiple jet 'modes' and their implications for propulsive efficiency.

The dynamics of pulsed jetting in squids throughout ontogeny is not well understood, especially with regard to the development of vortex rings, which are common features of mechanically generated jet pulses (also known as starting jets). Studies of mechanically generated starting jets have revealed a limiting principle for vortex ring formation characterized in terms of a ;formation number' (F), which delineates the transition between the formation of isolated vortex rings and vortex rings that have; pinched off' from the generating jet. Near F, there exists an optimum in pulse-averaged thrust with (potentially) low energetic cost, raising the question: do squids produce vortex rings and if so, do they fall near F, where propulsive benefits presumably occur? To better understand vortex ring dynamics and propulsive jet efficiency throughout ontogeny, brief squid Lolliguncula brevis ranging from 3.3 to 9.1 cm dorsal mantle length (DML) and swimming at speeds of 2.43-22.2 cms(-1) (0.54-3.50 DMLs(-1)) were studied using digital particle image velocimetry (DPIV). A range of jet structures were observed but most structures could be classified as variations of two principal jet modes: (1) jet mode I, where the ejected fluid rolled up into an isolated vortex ring; and (2) jet mode II, where the ejected fluid developed into a leading vortex ring that separated or ;pinched off' from a long trailing jet. The ratio of jet length [based on the vorticity extent (L(omega))] to jet diameter [based on peak vorticity locations (D(omega))] was <3.0 for jet mode I and >3.0 for jet mode II, placing the transition between modes in rough agreement with F determined in mechanical jet studies. Jet mode II produced greater time-averaged thrust and lift forces and was the jet mode most heavily used whereas jet mode I had higher propulsive efficiency, lower slip, shorter jet periods and a higher frequency of fin activity associated with it. No relationship between L(omega)/D(omega) and speed was detected and there was no apparent speed preference for the jet modes within the speed range considered in this study; however, propulsive efficiency did increase with speed partly because of a reduction in slip and jet angle with speed. Trends in higher slip, lower propulsive efficiency and higher relative lift production were observed for squid <5.0 cm DML compared with squid >/=5.0 cm DML. While these trends were observed when jet mode I and II were equally represented among the size classes, there was also greater relative dependence on jet mode I than jet mode II for squid <5.0 cm DML when all of the available jet sequences were examined. Collectively, these results indicate that approximately 5.0 cm DML is an important ontogenetic transition for the hydrodynamics of pulsed jetting in squids. The significance of our findings is that from early juvenile through to adult life stages, L. brevis is capable of producing a diversity of vortex ring-based jet structures, ranging from efficient short pulses to high-force longer duration pulses. Given that some of these structures had L(omega)/D(omega)s near F, and F represented the delineation between the two primary jet modes observed, fluid dynamics probably played an integral role in the evolution of squid locomotive systems. When this flexibility in jet dynamics is coupled with the highly versatile fins, which are capable of producing multiple hydrodynamic modes as well, it is clear that squid have a locomotive repertoire far more complex than originally thought.

Pulsed jet dynamics of squid hatchlings at intermediate Reynolds numbers.

Squid paralarvae (hatchlings) rely predominantly on a pulsed jet for locomotion, distinguishing them from the majority of aquatic locomotors at low/intermediate Reynolds numbers (Re), which employ oscillatory/undulatory modes of propulsion. Although squid paralarvae may delineate the lower size limit of biological jet propulsion, surprisingly little is known about the hydrodynamics and propulsive efficiency of paralarval jetting within the intermediate Re realm. To better understand paralarval jet dynamics, we used digital particle image velocimetry (DPIV) and high-speed video to measure bulk vortex properties (e.g. circulation, impulse, kinetic energy) and other jet features [e.g. average and peak jet velocity along the jet centerline (U(j) and U(jmax), respectively), jet angle, jet length based on the vorticity and velocity extents (L(omega) and L(V), respectively), jet diameter based on the distance between vorticity peaks (D(omega)), maximum funnel diameter (D(F)), average and maximum swimming speed (U and U(max), respectively)] in free-swimming Doryteuthis pealeii paralarvae (1.8 mm dorsal mantle length) (Re(squid)=25-90). Squid paralarvae spent the majority of their time station holding in the water column, relying predominantly on a frequent, high-volume, vertically directed jet. During station holding, paralarvae produced a range of jet structures from spherical vortex rings (L(omega)/D(omega)=2.1, L(V)/D(F)=13.6) to more elongated vortex ring structures with no distinguishable pinch-off (L(omega)/D(omega)=4.6, L(V)/D(F)=36.0). To swim faster, paralarvae increased pulse duration and L(omega)/D(omega), leading to higher impulse but kept jet velocity relatively constant. Paralarvae produced jets with low slip, i.e. ratio of jet velocity to swimming velocity (U(j)/U or U(jmax)/U(max)), and exhibited propulsive efficiency [eta(pd)=74.9+/-8.83% (+/-s.d.) for deconvolved data] comparable with oscillatory/undulatory swimmers. As slip decreased with speed, propulsive efficiency increased. The detection of high propulsive efficiency in paralarvae is significant because it contradicts many studies that predict low propulsive efficiency at intermediate Re for inertial forms of locomotion.

Swimming dynamics and propulsive efficiency of squids throughout ontogeny.

Squids encounter vastly different flow regimes throughout ontogeny as they undergo critical morphological changes to their two locomotive systems: the fins and jet. Squid hatchlings (paralarvae) operate at low and intermediate Reynolds numbers (Re) and typically have rounded bodies, small fins, and relatively large funnel apertures, whereas juveniles and adults operate at higher Re and generally have more streamlined bodies, larger fins, and relatively small funnel apertures. These morphological changes and varying flow conditions affect swimming performance in squids. To determine how swimming dynamics and propulsive efficiency change throughout ontogeny, digital particle image velocimetry (DPIV) and kinematic data were collected from an ontogenetic range of long-finned squid Doryteuthis pealeii and brief squid Lolliguncula brevis swimming in a holding chamber or water tunnel (Re = 20-20 000). Jet and fin wake bulk properties were quantified, and propulsive efficiency was computed based on measurements of impulse and excess kinetic energy in the wakes. Paralarvae relied predominantly on a vertically directed, high frequency, low velocity jet as they bobbed up and down in the water column. Although some spherical vortex rings were observed, most paralarval jets consisted of an elongated vortical region of variable length with no clear pinch-off of a vortex ring from the trailing tail component. Compared with paralarvae, juvenile and adult squid exhibited a more diverse range of swimming strategies, involving greater overall locomotive fin reliance and multiple fin and jet wake modes with better defined vortex rings. Despite greater locomotive flexibility, jet propulsive efficiency of juveniles/adults was significantly lower than that of paralarvae, even when juvenile/adults employed their highest efficiency jet mode involving the production of periodic isolated vortex rings with each jet pulse. When the fins were considered together with the jet for several juvenile/adult swimming sequences, overall propulsive efficiency increased, suggesting that fin contributions are important and should not be overlooked in analyses of the swimming performance of squids. The fins produced significant thrust and consistently had higher propulsive efficiency than did the jet. One particularly important area of future study is the determination of coordinated jet/fin wake modes that have the greatest impact on propulsive efficiency. Although such research would be technically challenging, requiring new, powerful, 3D approaches, it is necessary for a more comprehensive assessment of propulsive efficiency of the squid dual-mode locomotive system.