J.L. van Leeuwen

Knee muscle moment arms from MRI and from tendon travel

We tested magnetic resonance imaging (MRI) as a means to collect geometric data for moment arm estimation. A knee specimen in five successive flexion postures was scanned by MRI, while simultaneously tendon positions of loaded muscles were measured (long head of biceps femoris, lateral and medial gastrocnemius, gracilis, rectus femoris, sartorius, semimembranosus, semitendinosus, and tensor fasciae latae). Discrete rotation centres were derived from MRI pictures. Moment arms were estimated as the distances from these centres to the tendons. The ratio of tendon travel over the increment of joint angulation was the alternative, more reliable estimate of the moment arm. An important principal shortcoming of MRI is the impossibility of accounting for force distribution in taut tissue. As a consequence, for some muscles, considerable inaccuracies in moment arm estimation are found in a relatively small range of joint angulation (up to about 30% for the rectus femoris and semimembranosus). For the tensor fasciae latae, the moment arm cannot be estimated by MRI, while the estimate by tendon travel is unreliable owing to the deformability and attachments of the fascia lata.

Estimation of instantaneous moment arms of lower-leg muscles

Muscle moment arms at the human knee and ankle were estimated from muscle length changes measured as a function of joint flexion angle in cadaver specimens. Nearly all lower-leg muscles were studied: extensor digitorum longus, extensor hallucis longus, flexor digitorum longus, flexor hallucis longus, gastrocnemius lateralis, gastrocnemius medialis, peroneus brevis, peroneus longus, peroneus tertius, plantaris, soleus, tibialis anterior, and tibialis posterior. Noise in measured muscle length was filtered by means of quintic splines. Moment arms of the mm. gastrocnemii appear to be much more dependent on joint flexion angles than was generally assumed by other investigators. Some consequences for earlier analyses are mentioned.

Evidence for an elastic projection mechanism in the chameleon tongue.

To capture prey, chameleons ballistically project their tongues as far as 1.5 body lengths with accelerations of up to 500 m s⊃–2. At the core of a chameleons tongue is a cylindrical tongue skeleton surrounded by the accelerator muscle. Previously, the cylindrical accelerator muscle was assumed to power tongue projection directly during the actual fast projection of the tongue. However, high–speed recordings of Chamaeleo melleri and C. pardalis reveal that peak powers of 3000 W kg⊃–1 are necessary to generate the observed accelerations, which exceed the accelerator muscles capacity by at least five– to 10–fold. Extrinsic structures might power projection via the tongue skeleton. High–speed fluoroscopy suggests that they contribute less than 10% of the required peak instantaneous power. Thus, the projection power must be generated predominantly within the tongue, and an energy–storage–and–release mechanism must be at work. The key structure in the projection mechanism is probably a cylindrical connective–tissue layer, which surrounds the entoglossal process and was previously suggested to act as lubricating tissue. This tissue layer comprises at least 10 sheaths that envelop the entoglossal process. The outer portion connects anteriorly to the accelerator muscle and the inner portion to the retractor structures. The sheaths contain helical arrays of collagen fibres. Prior to projection, the sheaths are longitudinally loaded by the combined radial contraction and hydrostatic lengthening of the accelerator muscle, at an estimated mean power of 144 W kg⊃–1 in C. melleri. Tongue projection is triggered as the accelerator muscle and the loaded portions of the sheaths start to slide over the tip of the entoglossal process. The springs relax radially while pushing off the rounded tip of the entoglossal process, making the elastic energy stored in the helical fibres available for a simultaneous forward acceleration of the tongue pad, accelerator muscle and retractor structures. The energy release continues as the multilayered spring slides over the tip of the smooth and lubricated entoglossal process. This sliding–spring theory predicts that the sheaths deliver most of the instantaneous power required for tongue projection. The release power of the sliding tubular springs exceeds the work rate of the accelerator muscle by at least a factor of 10 because the elastic–energy release occurs much faster than the loading process. Thus, we have identified a unique catapult mechanism that is very different from standard engineering designs. Our morphological and kinematic observations, as well as the available literature data, are consistent with the proposed mechanism of tongue projection, although experimental tests of the sheath strain and the lubrication of the entoglossal process are currently beyond our technical scope.

Leading-Edge Vortices Elevate Lift of Autorotating Plant Seeds

Helicopter Seed Lift The “helicopter” seeds of maple trees and other similar autorotating seeds detach from their parent tree under windy conditions and gyrate as they are dispersed by the wind. The reproductive success of the tree depends on the flight performance of its seeds. Autorotating seeds are known to generate high lift as they slowly descend through the air, but the means by which they do so is unclear. Lentink et al. (p. 1438, see the cover) have elucidated the aerodynamic mechanism for high lift in autorotating seeds using a robot model seed and a three-dimensional flow measurement technique. Maple seeds and a hornbeam seed create a prominent leading-edge vortex that is similar to the flow structures that are responsible for the high lift generated by the wings of hovering insects and bats. Thus, both animals and plants have converged on an identical aerodynamic solution for generating lift. Winged plant seeds use leading edge vortices to create lift, in the same way that flying animals do. As they descend, the autorotating seeds of maples and some other trees generate unexpectedly high lift, but how they attain this elevated performance is unknown. To elucidate the mechanisms responsible, we measured the three-dimensional flow around dynamically scaled models of maple and hornbeam seeds. Our results indicate that these seeds attain high lift by generating a stable leading-edge vortex (LEV) as they descend. The compact LEV, which we verified on real specimens, allows maple seeds to remain in the air more effectively than do a variety of nonautorotating seeds. LEVs also explain the high lift generated by hovering insects, bats, and possibly birds, suggesting that the use of LEVs represents a convergent aerodynamic solution in the evolution of flight performance in both animals and plants.

Optimum power output and structural design of sarcomeres

A model of a “general” sarcomere is presented for the calculation of power output as a function of (i) contraction range, (ii) contraction velocity, (iii) muscle fibre stimulation (active state) and (iv) structural parameters of the sarcomere (i.e. lengths of actin, myosin, and bare zone on myosin, and thickness of the Z-disc). The model is applicable to virtually all types of striated muscle fibres. By computer simulation, particular combinations of actin and myosin lengths were found that maximize the specific power output for particular functional demands, specified in terms of contraction range and contraction velocity. The accuracy of the prediction of the optimum sarcomere design by the model depends on the quality of its input, i.e. the available knowledge of the in vivo spectrum of contraction velocities and sarcomere excursions. Predictions of sarcomere design from model simulations were compared with ultrastructural data from the literature. With the present model, the complete variation in the ratio of myosin length over actin length (from about 1·05 down to 0·65, as observed in insect and vertebrate sarcomeres, see Fig. 9) can be explained as a series of adaptations for optimum power output from a small to a large contraction range, respectively.

Usability of normal force distribution measurements to evaluate asymmetrical loading of the back of the horse and different rider positions on a standing horse

Pressure measurement devices in equine sports have primarily focused on tack (saddle pads and saddle fitting methods). However, saddle pressure devices may also be useful in evaluating the interaction and distribution of normal forces between the horse and rider, including rider position and riding technique. This study examined the validity, reliability, repeatability and possibilities of using a saddle pressure device to evaluate rider position. All measurements were performed using a standing horse. Validity was tested by calculating the correlation coefficient between measured normal force and the weight of the rider. Repeatability was tested by calculating intra-class correlation coefficients. The use of normal force measurements to evaluate horse-rider interaction was tested by adding a known weight to saddle or rider and collecting measurements with the rider sitting in four different positions. The device was found to be valid and reliable for force measurements when the measurement device was not replaced. The system could be used to determine the expected differences with added weight and in different rider positions. The normal force distribution measurement device proved to be a valid and reliable tool for studying the interaction between a rider and a static horse provided it is positioned carefully and consistently relative to both the horse and the saddle.

Vortex-wake interactions of a flapping foil that models animal swimming and flight.

SUMMARY The fluid dynamics of many swimming and flying animals involves the generation and shedding of vortices into the wake. Here we studied the dynamics of similar vortices shed by a simple two-dimensional flapping foil in a soap-film tunnel. The flapping foil models an animal wing, fin or tail in forward locomotion. The vortical flow induced by the foil is correlated to (the resulting) thickness variations in the soap film. We visualized these thickness variations through light diffraction and recorded it with a digital high speed camera. This set-up enabled us to study the influence of foil kinematics on vortex-wake interactions. We varied the dimensionless wavelength of the foil (λ*=4–24) at a constant dimensionless flapping amplitude (A*=1.5) and geometric angle of attack amplitude (Aα,geo=15°). The corresponding Reynolds number was of the order of 1000. Such values are relevant for animal swimming and flight. We found that a significant leading edge vortex (LEV) was generated by the foil at low dimensionless wavelengths (λ*<10). The LEV separated from the foil for all dimensionless wavelengths. The relative time (compared with the flapping period) that the unstable LEV stayed above the flapping foil increased for decreasing dimensionless wavelengths. As the dimensionless wavelength decreased, the wake dynamics evolved from a wavy von Kármán-like vortex wake shed along the sinusoidal path of the foil into a wake densely packed with large interacting vortices. We found that strongly interacting vortices could change the wake topology abruptly. This occured when vortices were close enough to merge or tear each other apart. Our experiments show that relatively small changes in the kinematics of a flapping foil can alter the topology of the vortex wake drastically.

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 effect of rising and sitting trot on back movements and head‐neck position of the horse

REASON FOR PERFORMING STUDY During trot, the rider can either rise from the saddle during every stride or remain seated. Rising trot is used frequently because it is widely assumed that it decreases the loading of the equine back. This has, however, not been demonstrated in an objective study. OBJECTIVE To determine the effects of rising and sitting trot on the movements of the horse. HYPOTHESIS Sitting trot has more extending effect on the horses back than rising trot and also results in a higher head and neck position. METHODS Twelve horses and one rider were used. Kinematic data were captured at trot during over ground locomotion under 3 conditions: unloaded, rising trot and sitting trot. Back movements were calculated using a previously described method with a correction for trunk position. Head-neck position was xpressed as extension and flexion of C1, C3 and C6, and vertical displacement of C1 and the bit. RESULTS Sitting trot had an overall extending effect on the back of horses when compared to the unloaded situation. In rising trot: the maximal flexion of the back was similar to the unloaded situation, while the maximal extension was similar to sitting trot; lateral bending of the back was larger than during the unloaded situation and sitting trot; and the horses held their heads lower than in the other conditions. The angle of C6 was more flexed in rising than in sitting trot. CONCLUSIONS AND CLINICAL RELEVANCE The back movement during rising trot showed characteristics of both sitting trot and the unloaded condition. As the same maximal extension of the back is reached during rising and sitting trot, there is no reason to believe that rising trot was less challenging for the back.

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.