Tatjana Y. Hubel

Upwash exploitation and downwash avoidance by flap phasing in ibis formation flight.

Many species travel in highly organized groups. The most quoted function of these configurations is to reduce energy expenditure and enhance locomotor performance of individuals in the assemblage. The distinctive V formation of bird flocks has long intrigued researchers and continues to attract both scientific and popular attention. The well-held belief is that such aggregations give an energetic benefit for those birds that are flying behind and to one side of another bird through using the regions of upwash generated by the wings of the preceding bird, although a definitive account of the aerodynamic implications of these formations has remained elusive. Here we show that individuals of northern bald ibises (Geronticus eremita) flying in a V flock position themselves in aerodynamically optimum positions, in that they agree with theoretical aerodynamic predictions. Furthermore, we demonstrate that birds show wingtip path coherence when flying in V positions, flapping spatially in phase and thus enabling upwash capture to be maximized throughout the entire flap cycle. In contrast, when birds fly immediately behind another bird—in a streamwise position—there is no wingtip path coherence; the wing-beats are in spatial anti-phase. This could potentially reduce the adverse effects of downwash for the following bird. These aerodynamic accomplishments were previously not thought possible for birds because of the complex flight dynamics and sensory feedback that would be required to perform such a feat. We conclude that the intricate mechanisms involved in V formation flight indicate awareness of the spatial wake structures of nearby flock-mates, and remarkable ability either to sense or predict it. We suggest that birds in V formation have phasing strategies to cope with the dynamic wakes produced by flapping wings.

Wake structure and wing kinematics: the flight of the lesser dog-faced fruit bat, Cynopterus brachyotis

SUMMARY We investigated the detailed kinematics and wake structure of lesser dog-faced fruit bats (Cynopterus brachyotis) flying in a wind tunnel. High speed recordings of the kinematics were conducted to obtain three-dimensional reconstructions of wing movements. Simultaneously, the flow structure in the spanwise plane perpendicular to the flow stream was visualized using time-resolved particle image velocimetry. The flight of four individuals was investigated to reveal patterns in kinematics and wake structure typical for lower and higher speeds. The wake structure identified as typical for both speed categories was a closed-loop ring vortex consisting of the tip vortex and the limited appearance of a counter-rotating vortex near the body, as well as a small distally located vortex system at the end of the upstroke that generated negative lift. We also investigated the degree of consistency within trials and looked at individual variation in flight parameters, and found distinct differences between individuals as well as within individuals.

Changes in kinematics and aerodynamics over a range of speeds in Tadarida brasiliensis, the Brazilian free-tailed bat

To date, wake measurements using particle image velocimetry (PIV) of bats in flight have studied only three bat species, all fruit and nectar feeders. In this study, we present the first wake structure analysis for an insectivorous bat. Tadarida brasiliensis, the Brazilian free-tailed bat, is an aerial hunter that annually migrates long distances and also differs strikingly from the previously investigated species morphologically. We compare the aerodynamics of T. brasiliensis with those of other, frugivorous bats and with common swifts, Apus apus, a bird with wing morphology, kinematics and flight ecology similar to that of these bats. The comparison reveals that, for the range of speeds evaluated, the cyclical pattern of aerodynamic forces associated with a wingbeat shows more similarities between T. brasiliensis and A. apus than between T. brasiliensis and other frugivorous bats.

The importance of leading edge vortices under simplified flapping flight conditions at the size scale of birds

SUMMARY Over the last decade, interest in animal flight has grown, in part due to the possible use of flapping propulsion for micro air vehicles. The importance of unsteady lift-enhancing mechanisms in insect flight has been recognized, but unsteady effects were generally thought to be absent for the flapping flight of larger animals. Only recently has the existence of LEVs (leading edge vortices) in small vertebrates such as swifts, small bats and hummingbirds been confirmed. To study the relevance of unsteady effects at the scale of large birds [reduced frequency k between 0.05 and 0.3, k=(πfc)/U∞; f is wingbeat frequency, U∞ is free-stream velocity, and c is the average wing chord], and the consequences of the lack of kinematic and morphological refinements, we have designed a simplified goose-sized flapping model for wind tunnel testing. The 2-D flow patterns along the wing span were quantitatively visualized using particle image velocimetry (PIV), and a three-component balance was used to measure the forces generated by the wings. The flow visualization on the wing showed the appearance of LEVs, which is typically associated with a delayed stall effect, and the transition into flow separation. Also, the influence of the delayed stall and flow separation was clearly visible in measurements of instantaneous net force over the wingbeat cycle. Here, we show that, even at reduced frequencies as low as those of large bird flight, unsteady effects are present and non-negligible and have to be addressed by kinematic and morphological adaptations.

Bats go head-under-heels: the biomechanics of landing on a ceiling

SUMMARY Bats typically roost head-under-heels but they cannot hover in this position, thus, landing on a ceiling presents a biomechanical challenge. To land, a bat must perform an acrobatic flip that brings the claws of the toes in contact with the ceiling and do so gently enough as to avoid injury to its slender hindlimbs. In the present study, we sought to determine how bats land, to seek a link between landing kinematics and ceiling impact forces, and to determine whether landing strategies vary among bat species. To do this, we measured the kinematics and kinetics of landing behaviour in three species of bats as they landed on a force-measuring platform (Cynopterus brachyotis, N=3; Carollia perspicillata, N=5; Glossophaga soricina, N=5). Kinematics were similar for all bats within a species but differed among species. C. brachyotis performed four-point landings, during which body pitch increased until the ventral surface of the body faced the ceiling and the thumbs and hindlimbs simultaneously grasped the surface. Bats of the other two species performed two-point landings, whereby only the hindlimbs made contact with the ceiling. During these two-point landings, the hindlimbs were drawn up the side of the body to come in contact with the ceiling, causing simultaneous changes in body pitch, roll and yaw over the course of the landing sequence. Right-handed and left-handed forms of the two-point landing were observed, with individuals often switching back and forth between them among landing events. The four-point landing of C. brachyotis resulted in larger peak forces (3.7±2.4 body weights; median ± interquartile range) than the two-point landings of C. perspicillata (0.8±0.6 body weights) or G. soricina (0.8±0.2 body weights). Our results demonstrate that the kinematics and kinetics of landing vary among bat species and that there is a correlation between the way a bat moves its body when it lands and the magnitude of peak impact force it experiences during that landing. We postulate that these interspecific differences in impact force could result because of stronger selective pressure for gentle landing in cave-roosting (C. perspicillata, G. soricina) versus foliage-roosting (C. brachyotis) species.

The human foot and heel-sole - toe walking strategy: a mechanism enabling an inverted pendular gait with low isometric muscle force?

Mechanically, the most economical gait for slow bipedal locomotion requires walking as an ‘inverted pendulum’, with: I, an impulsive, energy-dissipating leg compression at the beginning of stance; II, a stiff-limbed vault; and III, an impulsive, powering push-off at the end of stance. The characteristic ‘M’-shaped vertical ground reaction forces of walking in humans reflect this impulse–vault–impulse strategy. Humans achieve this gait by dissipating energy during the heel-to-sole transition in early stance, approximately stiff-limbed, flat-footed vaulting over midstance and ankle plantarflexion (powering the toes down) in late stance. Here, we show that the ‘M’-shaped walking ground reaction force profile does not require the plantigrade human foot or heel–sole–toe stance; it is maintained in tip–toe and high-heel walking as well as in ostriches. However, the unusual, stiff, human foot structure—with ground-contacting heel behind ankle and toes in front—enables both mechanically economical inverted pendular walking and physiologically economical muscle loading, by producing extreme changes in mechanical advantage between muscles and ground reaction forces. With a human foot, and heel–sole–toe strategy during stance, the shin muscles that dissipate energy, or calf muscles that power the push-off, need not be loaded at all—largely avoiding the ‘cost of muscle force’—during the passive vaulting phase.

Energy cost and return for hunting in African wild dogs and cheetahs

African wild dogs (Lycaon pictus) are reported to hunt with energetically costly long chase distances. We used high-resolution GPS and inertial technology to record 1,119 high-speed chases of all members of a pack of six adult African wild dogs in northern Botswana. Dogs performed multiple short, high-speed, mostly unsuccessful chases to capture prey, while cheetahs (Acinonyx jubatus) undertook even shorter, higher-speed hunts. We used an energy balance model to show that the energy return from group hunting and feeding substantially outweighs the cost of multiple short chases, which indicates that African wild dogs are more energetically robust than previously believed. Comparison with cheetah illustrates the trade-off between sheer athleticism and high individual kill rate characteristic of cheetahs, and the energetic robustness of frequent opportunistic group hunting and feeding by African wild dogs.

Additive opportunistic capture explains group hunting benefits in African wild dogs.

African wild dogs (Lycaon pictus) are described as highly collaborative endurance pursuit hunters based on observations derived primarily from the grass plains of East Africa. However, the remaining population of this endangered species mainly occupies mixed woodland savannah where hunting strategies appear to differ from those previously described. We used high-resolution GPS and inertial technology to record fine-scale movement of all members of a single pack of six adult African wild dogs in northern Botswana. The dogs used multiple short-distance hunting attempts with a low individual kill rate (15.5%), but high group feeding rate due to the sharing of prey. Use of high-level cooperative chase strategies (coordination and collaboration) was not recorded. In the mixed woodland habitats typical of their current range, simultaneous, opportunistic, short-distance chasing by dogs pursuing multiple prey (rather than long collaborative pursuits of single prey by multiple individuals) could be the key to their relative success in these habitats.

Children and adults minimise activated muscle volume by selecting gait parameters that balance gross mechanical power and work demands

ABSTRACT Terrestrial locomotion on legs is energetically expensive. Compared with cycling, or with locomotion in swimming or flying animals, walking and running are highly uneconomical. Legged gaits that minimise mechanical work have previously been identified and broadly match walking and running at appropriate speeds. Furthermore, the ‘cost of muscle force’ approaches are effective in relating locomotion kinetics to metabolic cost. However, few accounts have been made for why animals deviate from either work-minimising or muscle-force-minimising strategies. Also, there is no current mechanistic account for the scaling of locomotion kinetics with animal size and speed. Here, we report measurements of ground reaction forces in walking children and adult humans, and their stance durations during running. We find that many aspects of gait kinetics and kinematics scale with speed and size in a manner that is consistent with minimising muscle activation required for the more demanding between mechanical work and power: spreading the duration of muscle action reduces activation requirements for power, at the cost of greater work demands. Mechanical work is relatively more demanding for larger bipeds – adult humans – accounting for their symmetrical M-shaped vertical force traces in walking, and relatively brief stance durations in running compared with smaller bipeds – children. The gaits of small children, and the greater deviation of their mechanics from work-minimising strategies, may be understood as appropriate for their scale, not merely as immature, incompletely developed and energetically sub-optimal versions of adult gaits. Highlighted Article: The gross mechanics of walking and running children and adults support a new model for the costs dominating level terrestrial locomotion – muscle activation for mechanical work or power.

Energetically optimal running requires torques about the centre of mass

Bipedal animals experience ground reaction forces (GRFs) that pass close to the centre of mass (CoM) throughout stance, first decelerating the body, then re-accelerating it during the second half of stance. This results in fluctuations in kinetic energy, requiring mechanical work from the muscles. However, here we show analytically that, in extreme cases (with a very large body pitch moment of inertia), continuous alignment of the GRF through the CoM requires greater mechanical work than a maintained vertical force; we show numerically that GRFs passing between CoM and vertical throughout stance are energetically favourable under realistic conditions; and demonstrate that the magnitude, if not the precise form, of actual CoM-torque profiles in running is broadly consistent with simple mechanical work minimization for humans with appropriate pitch moment of inertia. While the potential energetic savings of CoM-torque support strategies are small (a few per cent) over the range of human running, their importance increases dramatically at high speeds and stance angles. Fast, compliant runners or hoppers would benefit considerably from GRFs more vertical than the zero-CoM-torque strategy, especially with bodies of high pitch moment of inertia—suggesting a novel advantage to kangaroos of their peculiar long-head/long-tail structure.