Investigation of models to estimate flight performance of gliding birds from wakes.

Mathematical models based on inviscid flow theory are effective at predicting the aerodynamic forces on large-scale aircraft. Avian flight, however, is characterized by smaller sizes, slower speeds, and increased influence of viscous effects associated with lower Reynolds numbers. Therefore, inviscid mathematical models of avian flight should be used with caution.

The functions of leg muscles, structures and mechanisms in running.

The actions of the major human leg muscles are well established; however, the functions of these muscle actions during steady running remain unclear. Here, leg structures and mechanisms are considered in terms of their functions in meeting the task of a vehicle acting as an effective machine, supporting body weight during translation with low mechanical work demand and in supplying mechanical work economically. Legs are modelled as a sequence of linkages that predict muscle actions and reveal the varying muscle functions within the integrated leg.

Dynamics of hinged wings in strong upward gusts.

A bird's wings are articulated to its body via highly mobile shoulder joints. The joints confer an impressive range of motion, enabling the wings to make broad, sweeping movements that can modulate quite dramatically the production of aerodynamic load. This is enormously useful in challenging flight environments, especially the gusty, turbulent layers of the lower atmosphere.

The collisional geometry of economical walking predicts human leg and foot segment proportions.

Human walking appears complicated, with many muscles and joints performing rapidly varying roles over the stride. However, the function of walking is simple: to support body weight as it translates economically. Here, a scenario is proposed for the sequence of joint and muscle actions that achieves this function, with the timing of muscle loading and unloading driven by simple changes in geometry over stance.

Legs as linkages: an alternative paradigm for the role of tendons and isometric muscles in facilitating economical gait.

Considerable attention has been given to the spring-like behaviour of stretching and recoiling tendons, and how this can reduce the work demanded from muscle for a given loss-return cycling of mechanical energy during high-speed locomotion. However, even completely isometric muscle-tendon units have the potential to act as tension struts, forming links in linkages that avoid the demand for mechanical work-cycling in the first place. Here, forelimb and hindlimb structures and geometries of quadrupeds are considered in terms of linkages that avoid mechanical work at the level of both the whole limb and the individual muscles.

Virtual manipulation of tail postures of a gliding barn owl () demonstrates drag minimization when gliding.

Aerodynamic functions of the avian tail have been studied previously using observations of bird flight, physical models in wind tunnels, theoretical modelling and flow visualization. However, none of these approaches has provided rigorous, quantitative evidence concerning tail functions because (i) appropriate manipulation and controls cannot be achieved using live animals and (ii) the aerodynamic interplay between the wings and body challenges reductive theoretical or physical modelling approaches. Here, we have developed a comprehensive analytical drag model, calibrated by high-fidelity computational fluid dynamics (CFD), and used it to investigate the aerodynamic action of the tail by virtually manipulating its posture.

Raptor wing morphing with flight speed.

In gliding flight, birds morph their wings and tails to control their flight trajectory and speed. Using high-resolution videogrammetry, we reconstructed accurate and detailed three-dimensional geometries of gliding flights for three raptors (barn owl, ; tawny owl, , and goshawk, ). Wing shapes were highly repeatable and shoulder actuation was a key component of reconfiguring the overall planform and controlling angle of attack.

Limb work and joint work minimization reveal an energetic benefit to the elbows-back, knees-forward limb design in parasagittal quadrupeds.

Quadrupedal animal locomotion is energetically costly. We explore two forms of mechanical work that may be relevant in imposing these physiological demands. Limb work, due to the forces and velocities between the stance foot and the centre of mass, could theoretically be zero given vertical limb forces and horizontal centre of mass path.

Bird wings act as a suspension system that rejects gusts.

Musculoskeletal systems cope with many environmental perturbations without neurological control. These passive preflex responses aid animals to move swiftly through complex terrain. Whether preflexes play a substantial role in animal flight is uncertain.

The Possibility of Zero Limb-Work Gaits in Sprawled and Parasagittal Quadrupeds: Insights from Linkages of the Industrial Revolution.

Animal legs are diverse, complex, and perform many roles. One defining requirement of legs is to facilitate terrestrial travel with some degree of economy. This could, theoretically, be achieved without loss of mechanical energy if the body could take a continuous horizontal path supported by vertical forces only-effectively a wheel-like translation, and a condition closely approximated by walking tortoises.

Why are the fastest runners of intermediate size? Contrasting scaling of mechanical demands and muscle supply of work and power.

The fastest land animals are of intermediate size. Cheetah, antelope, greyhounds and racehorses have been measured running much faster than reported for elephants or elephant shrews. Can this be attributed to scaling of physical demands and explicit physiological constraints to supply? Here, we describe the scaling of mechanical work demand each stride, and the mechanical power demand each stance.

Artificial mass loading disrupts stable social order in pigeon dominance hierarchies.

Dominance hierarchies confer benefits to group members by decreasing the incidences of physical conflict, but may result in certain lower ranked individuals consistently missing out on access to resources. Here, we report a linear dominance hierarchy remaining stable over time in a closed population of birds. We show that this stability can be disrupted, however, by the artificial mass loading of birds that typically comprise the bottom 50% of the hierarchy.

High aerodynamic lift from the tail reduces drag in gliding raptors.

Many functions have been postulated for the aerodynamic role of the avian tail during steady-state flight. By analogy with conventional aircraft, the tail might provide passive pitch stability if it produced very low or negative lift. Alternatively, aeronautical principles might suggest strategies that allow the tail to reduce inviscid, induced drag: if the wings and tail act in different horizontal planes, they might benefit from biplane-like aerodynamics; if they act in the same plane, lift from the tail might compensate for lift lost over the fuselage (body), reducing induced drag with a more even downwash profile.

Minimalist analogue robot discovers animal-like walking gaits.

Robots based on simplified or abstracted biomechanical concepts can be a useful tool for investigating how and why animals move the way they do. In this paper we present an extremely simple quadruped robot, which is able to walk with no form of software or controller. Instead, individual leg movements are triggered directly by switches on each leg which detect leg loading and unloading.

An instrumented centrifuge for studying mouse locomotion and behaviour under hypergravity.

Gravity may influence multiple aspects of legged locomotion, from the periods of limbs moving as pendulums to the muscle forces required to support the body. We present a system for exposing mice to hypergravity using a centrifuge and studying their locomotion and activity during exposure. Centrifuge-induced hypergravity has the advantages that it both allows animals to move freely, and it affects both body and limbs.

An extension to the collisional model of the energetic cost of support qualitatively explains trotting and the trot-canter transition.

The majority of terrestrial mammals adopt distinct, discrete gaits across their speed range. Though there is evidence that walk, trot and gallop may be selected at speeds consistent with minimizing metabolic cost (Hoyt and Taylor, 1981, Nature, 291, 239-240), the mechanical causes underlying these costs and their changes with speed are not well understood. In particular, the paired, near-simultaneous contacts of the trot is puzzling as it appears to demand a high mechanical work that could easily be avoided with distributed contacts, as with a "running walk" gait or "tolt.

The scaling or ontogeny of human gait kinetics and walk-run transition: The implications of work vs. peak power minimization.

A simple model is developed to find vertical force profiles and stance durations that minimize either limb mechanical work or peak power demands during bipedal locomotion. The model predicts that work minimization is achieved with a symmetrical vertical force profile, consistent with previous models and observations of adult humans, and data for 487 participants (predominantly 11-18 years old) required to walk at a range of speeds at a Science Fair. Work minimization also predicts the discrete walk-run transition, familiar for adult humans.

The grazing gait, and implications of toppling table geometry for primate footfall sequences.

Many medium and large herbivores locomote forwards very slowly and intermittently when grazing. While the footfall order during grazing is the same as for walking, the relative fore-hind timing-phasing-is quite different. Extended periods of static stability are clearly required during grazing; however, stability requirements are insufficient to account for the timing.

Work minimization accounts for footfall phasing in slow quadrupedal gaits.

Quadrupeds, like most bipeds, tend to walk with an even left/right footfall timing. However, the phasing between hind and forelimbs shows considerable variation. Here, we account for this variation by modeling and explaining the influence of hind-fore limb phasing on mechanical work requirements.

The muscle-mechanical compromise framework: Implications for the scaling of gait and posture.

Many aspects of animal and human gait and posture cannot be predicted from purely mechanical work minimization or entirely based on optimizing muscle efficiency. Here, the Muscle-Mechanical Compromise Framework is introduced as a conceptual paradigm for considering the interactions and compromises between these two objectives. Current assumptions in implementing the Framework are presented.

Physiological, aerodynamic and geometric constraints of flapping account for bird gaits, and bounding and flap-gliding flight strategies.

Aerodynamically economical flight is steady and level. The high-amplitude flapping and bounding flight style of many small birds departs considerably from any aerodynamic or purely mechanical optimum. Further, many large birds adopt a flap-glide flight style in cruising flight which is not consistent with purely aerodynamic economy.

Locomotor preferences in terrestrial vertebrates: An online crowdsourcing approach to data collection.

Understanding how animals move within their environment is a burgeoning field of research. Despite this, relatively basic data, such as the locomotor speeds that animals choose to walk at in the wild, are sparse. If animals choose to walk with dynamic similarity, they will move at equal dimensionless speeds, represented by Froude number (Fr).

Social density processes regulate the functioning and performance of foraging human teams.

Social density processes impact the activity and order of collective behaviours in a variety of biological systems. Much effort has been devoted to understanding how density of people affects collective human motion in the context of pedestrian flows. However, there is a distinct lack of empirical data investigating the effects of social density on human behaviour in cooperative contexts.

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

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.

Identification of mouse gaits using a novel force-sensing exercise wheel.

The gaits that animals use can provide information on neurological and musculoskeletal disorders, as well as the biomechanics of locomotion. Mice are a common research model in many fields; however, there is no consensus in the literature on how (and if) mouse gaits vary with speed. One of the challenges in studying mouse gaits is that mice tend to run intermittently on treadmills or overground; this paper attempts to overcome this issue with a novel exercise wheel that measures vertical ground reaction forces.