U.K. Müller

Eel migration to the Sargasso: remarkably high swimming efficiency and low energy costs

SUMMARY One of the mysteries of the animal kingdom is the long-distance 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 Schmidts 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 Schmidts 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.

Swimming of larval zebrafish: ontogeny of body waves and implications for locomotory development

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.

How swifts control their glide performance with morphing wings

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.

Automated visual tracking for studying the ontogeny of zebrafish swimming

SUMMARY The 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 fishs 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.

Riding the waves: the role of the body wave in undulatory fish swimming

Abstract A continuously swimming mullet modulates its thrust production by changing slip-the ratio between its swimming speed U and the speed V with which the body wave travels down its body. This variation in thrust is reflected in the wake of the fish. We obtained 2-dimensional impressions of the wake behind a mullet swimming at a slip of 0.7 equivalent to active swimming, at a slip of 0.9 close to free-wheeling, and at a slip of 1.1 when the fish is braking. Independent of the slip, vortices are shed at the tail when the tail tip reaches its maximum lateral excursion. The manner in which the wake changes as it decays depends on the degree of slip: At a slip well below unity, the wake decays without any qualitative changes in shape, the medio-frontal cross section of the mature wake consists of a double row of alternating vortices separated by an undulating jet, and the angle between the jet flow and the mean path of motion is close to 45°; at a slip above unity, the vortices stretch out laterally and the mature wake resembles a single row of oval vortices with two vortex cores, and the jet between the vortices is almost perpendicular to the mean path of motion; the wake at slip of 0.9 exhibits a pattern intermediate between the wakes at slips 0.7 and 0.9 with slightly elongate vortices and a jet angle of 61°.

Bird song: Superfast muscles control dove's trill

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.

Body dynamics and hydrodynamics of swimming fish larvae: a computational study

SUMMARY To 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 fishs 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.

The scaling and structure of aquatic animal wakes.

Abstract Animal generated water movements are visualized and quantified using two-dimensional particle image velocimetry (PIV). The resulting vector flow fields allow for the study of the distribution of velocity, vorticity and vortices. Structural and temporal aspects of animal-induced flows covering a range of Reynolds (Re) numbers between less than 1 to more than 104 are presented. Maps of flow induced by continuous foraging and intermittent escape responses of tethered nauplius and copepodid stages of the marine copepod Temora longicornis offer insight in viscosity-dominated flow regimes. Fast escape responses of the equally sized largest nauplius stage and the smallest copepodid stage are compared. The nauplius moves by generating a viscous flow pattern with high velocities and vorticity; the copepodid moves by using inertial effects to produce a vortex ring with a rearward jet through the center. Larvae and small adult fish (zebra danio) use a burst-and-coast-swimming mode at Re numbers up to 6,000, shedding a vortex ring with the associated jet at the tail during the burst phase. Flow patterns during the coasting phase differ between the small larvae and larger adults due to the changes in importance of viscosity. A 12 cm long mullet swimming in a continuous mode generates a chain of vortex rings with a backward undulating jet through the centers of the rings at Re numbers of 4 × 104 in inertia-dominated regimes. Our empirical results provide realistic insight in the scale effects determining the morphology of the interactions between animals and water.

Superfast muscles control dove's trill

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.

Syringeal muscles fit the trill in ring doves (Streptopelia risoria L.)

SUMMARY In contrast to human phonation, the virtuoso vocalizations of most birds are modulated at the level of the sound generator, the syrinx. We address the hypothesis that syringeal muscles are physiologically capable of controlling the sound-generating syringeal membranes in the ring dove (Streptopelia risoria) syrinx. We establish the role of the tracheolateralis muscle and propose a new function for the sternotrachealis muscle. The tracheolateralis and sternotrachealis muscles have an antagonistic mechanical effect on the syringeal aperture. Here, we show that both syringeal muscles can dynamically control the full syringeal aperture. The tracheolateralis muscle is thought to directly alter position and tension of the vibrating syringeal membranes that determine the gating and the frequency of sound elements. Our measurements of the muscles contractile properties, combined with existing electromyographic and endoscopic evidence, establish its modulating role during the doves trill. The muscle delivers the highest power output at cycle frequencies that closely match the repetition rates of the fastest sound elements in the coo. We show that the two syringeal muscles share nearly identical contraction characteristics, and that sternotrachealis activity does not clearly modulate during the rapid trill. We propose that the sternotrachealis muscle acts as a damper that stabilizes longitudinal movements of the sound-generating system induced by tracheolateralis muscle contraction. The extreme performance of both syringeal muscles implies that they play an important role in fine-tuning membrane position and tension, which determines the quality of the sound for a conspecific mate.