SUMMARY Fish larvae, like most adult fish, undulate their bodies to propel themselves. A detailed kinematic study of the larval body wave is a prerequisite to formulate a set of functional requirements that the locomotor system must fulfil to generate the observed swimming kinematics. Lateral displacement and curvature profiles were obtained for zebrafish (Danio rerio) larvae at 2–21 days post-fertilisation for three swimming behaviours (cyclic swimming, slow starts and fast startle responses) using high-speed video. During cyclic swimming, fish larvae maintain tail beat frequencies of up to 100 Hz. The corresponding longitudinal strains, estimated from the peak curvatures of the midline, reach up to 0.19 in superficial tissue. The strain rate can reach 120 s–1. The wave of curvature travels along the body at a near-constant rate. Posterior to the stiff head, body-lengthspecific curvature is high and rises gently along the entire trunk to a maximum value of 6. Burst-and-coast swimming generates similar peak curvatures to cyclic swimming, but curvature rises more steeply from head to tail. Fish larvae exhibit phase shifts of 57–63°between the wave of lateral displacement and the wave of curvature, resulting in a 1:1.2 ratio of body wave length to curvature wave length. During C-starts, muscle strain can reach 0.19 and superficial longitudinal strain rates approach 30 s–1. Fish larvae do not initiate their escape response with a standing wave of curvature, although their C-starts approach a standing wave as the larvae grow older. The performance demands derived from swimming kinematics suggest that larval axial muscles have very short contraction cycles (10 ms), experience considerable strains (up to 0.2)and strain rates (up to 30 s–1 in white muscle fibres) yet are able to power swimming for several seconds.

Gliding birds continually change the shape and size of their wings, presumably to exploit the profound effect of wing morphology on aerodynamic performance. That birds should adjust wing sweep to suit glide speed has been predicted qualitatively by analytical glide models, which extrapolated the wing's performance envelope from aerodynamic theory. Here we describe the aerodynamic and structural performance of actual swift wings, as measured in a wind tunnel, and on this basis build a semi-empirical glide model. By measuring inside and outside swifts' behavioural envelope, we show that choosing the most suitable sweep can halve sink speed or triple turning rate. Extended wings are superior for slow glides and turns; swept wings are superior for fast glides and turns. This superiority is due to better aerodynamic performance-with the exception of fast turns. Swept wings are less effective at generating lift while turning at high speeds, but can bear the extreme loads. Finally, our glide model predicts that cost-effective gliding occurs at speeds of 8-10 m s(-1), whereas agility-related figures of merit peak at 15-25 m s(-1). In fact, swifts spend the night ('roost') in flight at 8-10 m s(-1) (ref. 11), thus our model can explain this choice for a resting behaviour. Morphing not only adjusts birds' wing performance to the task at hand, but could also control the flight of future aircraft.

One of the mysteries of the animal kingdom is the longdistance migration (5000-6000·km) of the European eel Anguilla anguilla L. from the coasts of Europe to its spawning grounds in the Sargasso Sea. The only evidence for the location of the spawning site of the European eel in the Sargasso Sea is the discovery by Johannes Schmidt at the beginning of the previous century of the smallest eel larvae (leptocephali) near the Sargasso Sea. For years it has been questioned whether the fasting eels have sufficient energy reserves to cover this enormous distance. We have tested Schmidt's theory by placing eels in swim tunnels in the laboratory and allowing them to make a simulated migration of 5500·km. We find that eels swim 4-6 times more efficiently than non-eel-like fish. Our findings are an important advance in this field because they remove a central objection to Schmidt's theory by showing that their energy reserves are, in principle, sufficient for the migration. Conclusive proof of the Sargasso Sea theory is likely to come from satellite tracking technology.

SummaryFish larvae, like many adult fish, swim by undulating their body. However, their body size and swimming speeds put them in the intermediate flow regime, where viscous and inertial forces both play an important role in the interaction between fish and water. To study the influence of the relatively high viscous forces compared with adult fish, we mapped the flow around swimming zebrafish (Danio rerio) larvae using two-dimensional digital particle image velocimetry (2D-DPIV) in the horizontal and transverse plane of the fish. Fish larvae initiate a swimming bout by bending their body into a C shape. During this initial tail-beat cycle, larvae shed two vortex pairs in the horizontal plane of their wake, one during the preparatory and one during the subsequent propulsive stroke. When they swim ʻcyclicallyʼ (mean swimming speed does not change significantly between tail beats), fish larvae generate a wide drag wake along their head and anterior body. The flow along the posterior body is dominated by the undulating body movements that cause jet flows into the concave bends of the body wave. Patches of elevated vorticity form around the jets, and travel posteriorly along with the body wave, until they are ultimately shed at the tail near the moment of stroke reversal. Behind the larva, two vortex pairs are formed per tail-beat cycle (the tail beating once left-to-right and then rightto-left) in the horizontal plane of the larval wake. By combining transverse and horizontal cross sections of the wake, we inferred that the wake behind a cyclically swimming zebrafish larva contains two diverging rows of vortex rings to the left and right of the mean path of motion, resembling the wake of steadily swimming adult eels. When the fish larva slows down at the end of a swimming bout, it gradually reduces its tail-beat frequency and amplitude, while the separated boundary layer and drag wake of the anterior body extend posteriorly to envelope the entire larva. This drag wake is considerably wider than the larval body. The effects of the intermediate flow regime manifest as a thick boundary layer and in the quick dying-off of the larval wake within less than half a second.

SUMMARYTo understand the mechanics of fish swimming, we need to know the forces exerted by the fluid and how these forces affect the motion of the fish. To this end, we developed a 3-D computational approach that integrates hydrodynamics and body dynamics. This study quantifies the flow around a swimming zebrafish (Danio rerio) larva. We used morphological and kinematics data from actual fish larvae aged 3 and 5days post fertilization as input for a computational model that predicted free-swimming dynamics from prescribed changes in body shape. We simulated cyclic swimming and a spontaneous C-start. A rigorous comparison with 2-D particle image velocimetry and kinematics data revealed that the computational model accurately predicted the motion of the fishʼs centre of mass as well as the spatial and temporal characteristics of the flow. The distribution of pressure and shear forces along the body showed that thrust is mainly produced in the posterior half of the body. We also explored the effect of the body wave amplitude on swimming performance by considering wave amplitudes that were up to 40% larger or smaller than the experimentally observed value. Increasing the body wave amplitude increased forward swimming speed from 7 to 21 body lengths per second, which is consistent with experimental observations. The model also predicted a non-linear increase in propulsive efficiency from 0.22 to 0.32 while the required mechanical power quadrupled. The efficiency increase was only minor for wave amplitudes above the experimental reference value, whereas the cost of transport rose significantly.Supplementary material available online at http://jeb.biologists.org/cgi/content/full/215/22/4015/DC1

SUMMARYThe zebrafish Danio rerio is a widely used model organism in studies of genetics, developmental biology, and recently, biomechanics. In order to quantify changes in swimming during all stages of development, we have developed a visual tracking system that estimates the posture of fish. Our current approach assumes planar motion of the fish, given image sequences taken from a top view. An accurate geometric fish model is automatically designed and fit to the images at each time frame. Our approach works across a range of fish shapes and sizes and is therefore well suited for studying the ontogeny of fish swimming, while also being robust to common environmental occlusions. Our current analysis focuses on measuring the influence of vertebra development on the swimming capabilities of zebrafish. We examine wild-type zebrafish and mutants with stiff vertebrae (stocksteif ) and quantify their body kinematics as a function of their development from larvae to adult (mutants made available by the Hubrecht laboratory, The Netherlands). By tracking the fish, we are able to measure the curvature and net acceleration along the body that result from the fishʼs body wave. Here, we demonstrate the capabilities of the tracking system for the escape response of wild-type zebrafish and stocksteif mutant zebrafish. The response was filmed with a digital high-speed camera at 1500·frames·s -1 . Our approach enables biomechanists and ethologists to process much larger datasets than possible at present. Our automated tracking scheme can therefore accelerate insight in the swimming behavior of many species of (developing) fish. Supplementary material available online at

Bird songs frequently contain trilling sounds that demand extremely fast vocalization control. Here we show that doves control their syrinx, a vocal organ that is unique to birds, by using superfast muscles. These muscles, which are similar to those that operate highly specialist acoustic organs such as the rattle of the rattlesnake, are among the fastest vertebrate muscles known and could be much more widespread than previously thought if they are the principal muscle type used to control bird songs.

Small undulatory swimmers such as larval zebrafish experience both inertial and viscous forces, the relative importance of which is indicated by the Reynolds number (Re). Re is proportional to swimming speed (v swim ) and body length; faster swimming reduces the relative effect of viscous forces. Compared with adults, larval fish experience relatively high (mainly viscous) drag during cyclic swimming. To enhance thrust to an equally high level, they must employ a high product of tail-beat frequency and ( peakto-peak) amplitude fA tail , resulting in a relatively high fA tail / v swim ratio (Strouhal number, St), and implying relatively high lateral momentum shedding and low propulsive efficiency. Using kinematic and inverse-dynamics analyses, we studied cyclic swimming of larval zebrafish aged 2-5 days post-fertilization (dpf ). Larvae at 4-5 dpf reach higher f (95 Hz) and A tail (2.4 mm) than at 2 dpf (80 Hz, 1.8 mm), increasing swimming speed and Re, indicating increasing muscle powers. As Re increases (60 ! 1400), St (2.5 ! 0.72) decreases nonlinearly towards values of large swimmers (0.2-0.6), indicating increased propulsive efficiency with v swim and age. Swimming at high St is associated with high-amplitude body torques and rotations. Low propulsive efficiencies and large yawing amplitudes are unavoidable physical constraints for small undulatory swimmers.