A pinboard by
Anthony Lapsansky

I am a PhD student at the University of Montana currently studying the biomechanics of locomotion.


Read the latest in bird flight mechanics to learn how birds use their muscles to stay aloft.

How do birds use their muscles to fly?

In 10 seconds? Bird flight requires the coordination of a complex suite of muscles. The pectoralis and supracoracoideus work together to generate...well, work, but they're not alone!

Don't believe it? Birds rely on muscles in their chest, hand-wing, back, and even legs muscles to achieve liftoff and perform complex aerial movements. Furthermore, the roles these muscles play in flight change with velocity and the exact style of motion. Read more.

But don't birds flight in pretty much the same way as airplanes do? Not quite! Unlike and airplane, which generates lift with its wings and thrust with an engine, birds create lift and thrust with the same structure. This means that minor changes in posture, controlled by muscle contraction, can drastically affect the way a bird flies.

Scientists around the world are investigating how specifics of muscle architecture allow and constrain the range of possible motion we see in birds.


Contractile activity of the pectoralis in the zebra finch according to mode and velocity of flap-bounding flight.

Abstract: We studied flying zebra finch (Taeniopygia guttata, N = 12), to provide a new test of a long-standing ;fixed-gear' hypothesis that flap-bounding birds use only intermittent non-flapping phases, instead of variation in muscle activity, to vary mechanical power output in flight. Using sonomicrometry and electromyography, we measured in vivo fascicle length and neuromuscular recruitment in the pectoralis as the birds flew in different flight modes (level, ascending, descending; mean velocity 1.6+/-0.3 m s(-1)) and across velocities in a new, variable-speed wind tunnel (0-12 m s(-1)). Synchronized high-speed digital video (250 Hz) provided a record of wing kinematics. Flight mode had a significant effect upon pectoralis strain, strain rate, fractional shortening and the relative timing of muscle activity (onset, offset and duration). Among flight velocities, we observed significant variation in pectoralis strain, fractional lengthening and shortening, strain rate, relative electromyographic (EMG) amplitude, and EMG duration and offset. In particular, variation in strain rate and relative EMG amplitude indicates that the fixed-gear hypothesis should be rejected. Instead, it appears that zebra finch vary work and power output within wingbeats by modulating muscle contractile behavior and between wingbeats using intermittent bounds. Muscle activity patterns and wing kinematics were similar between free flight and wind tunnel flight at similar speeds. Comparing flights with and without surgically implanted transducers and electrodes, zebra finch exhibited a reduction in maximum velocity (from 14 to 12 m s(-1)) and a significant increase in wingbeat frequency and percent time flapping. This identifies a potential limitation of in vivo flight measurements, and similar studies of bird flight should, therefore, include measurements of the extent to which flight performance is compromised by experimental protocol.

Pub.: 27 Jul '05, Pinned: 23 Apr '17

Contractile properties of the pigeon supracoracoideus during different modes of flight.

Abstract: The supracoracoideus (SUPRA) is the primary upstroke muscle for avian flight and is the antagonist to the downstroke muscle, the pectoralis (PECT). We studied in vivo contractile properties and mechanical power output of both muscles during take-off, level and landing flight. We measured muscle length change and activation using sonomicrometry and electromyography, and muscle force development using strain recordings on the humerus. Our results support a hypothesis that the primary role of the SUPRA is to supinate the humerus. Antagonistic forces exerted by the SUPRA and PECT overlap during portions of the wingbeat cycle, thereby offering a potential mechanism for enhancing control of the wing. Among flight modes, muscle strain was approximately the same in the SUPRA (33-40%) and the PECT (35-42%), whereas peak muscle stress was higher in the SUPRA (85-126 N m(-2)) than in the PECT (50-58 N m(-2)). The SUPRA mainly shortened relative to resting length and the PECT mainly lengthened. We estimated that elastic energy storage in the tendon of the SUPRA contributed between 28 and 60% of the net work of the SUPRA and 6-10% of the total net mechanical work of both muscles. Mechanical power output in the SUPRA was congruent with the estimated inertial power required for upstroke, but power output from the PECT was only 42-46% of the estimated aerodynamic power requirements for flight. There was a significant effect of flight mode upon aspects of the contractile behavior of both muscles including strain, strain rate, peak stress, work and power.

Pub.: 01 Jan '08, Pinned: 23 Apr '17

Evolution of avian flight: muscles and constraints on performance.

Abstract: Competing hypotheses about evolutionary origins of flight are the 'fundamental wing-stroke' and 'directed aerial descent' hypotheses. Support for the fundamental wing-stroke hypothesis is that extant birds use flapping of their wings to climb even before they are able to fly; there are no reported examples of incrementally increasing use of wing movements in gliding transitioning to flapping. An open question is whether locomotor styles must evolve initially for efficiency or if they might instead arrive due to efficacy. The proximal muscles of the avian wing output work and power for flight, and new research is exploring functions of the distal muscles in relation to dynamic changes in wing shape. It will be useful to test the relative contributions of the muscles of the forearm compared with inertial and aerodynamic loading of the wing upon dynamic morphing. Body size has dramatic effects upon flight performance. New research has revealed that mass-specific muscle power declines with increasing body mass among species. This explains the constraints associated with being large. Hummingbirds are the only species that can sustain hovering. Their ability to generate force, work and power appears to be limited by time for activation and deactivation within their wingbeats of high frequency. Most small birds use flap-bounding flight, and this flight style may offer an energetic advantage over continuous flapping during fast flight or during flight into a headwind. The use of flap-bounding during slow flight remains enigmatic. Flap-bounding birds do not appear to be constrained to use their primary flight muscles in a fixed manner. To improve understanding of the functional significance of flap-bounding, the energetic costs and the relative use of alternative styles by a given species in nature merit study.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'.

Pub.: 17 Aug '16, Pinned: 23 Apr '17

Effects of flight speed upon muscle activity in hummingbirds.

Abstract: Hummingbirds have the smallest body size and highest wingbeat frequencies of all flying vertebrates, so they represent one endpoint for evaluating the effects of body size on sustained muscle function and flight performance. Other bird species vary neuromuscular recruitment and contractile behavior to accomplish flight over a wide range of speeds, typically exhibiting a U-shaped curve with maxima at the slowest and fastest flight speeds. To test whether the high wingbeat frequencies and aerodynamically active upstroke of hummingbirds lead to different patterns, we flew rufous hummingbirds (Selasphorus rufus, 3 g body mass, 42 Hz wingbeat frequency) in a variable-speed wind tunnel (0-10 m s(-1)). We measured neuromuscular activity in the pectoralis (PECT) and supracoracoideus (SUPRA) muscles using electromyography (EMG, N=4 birds), and we measured changes in PECT length using sonomicrometry (N=1). Differing markedly from the pattern in other birds, PECT deactivation occurred before the start of downstroke and the SUPRA was deactivated before the start of upstroke. The relative amplitude of EMG signal in the PECT and SUPRA varied according to a U-shaped curve with flight speed; additionally, the onset of SUPRA activity became relatively later in the wingbeat at intermediate flight speeds (4 and 6 m s(-1)). Variation in the relative amplitude of EMG was comparable with that observed in other birds but the timing of muscle activity was different. These data indicate the high wingbeat frequency of hummingbirds limits the time available for flight muscle relaxation before the next half stroke of a wingbeat. Unlike in a previous study that reported single-twitch EMG signals in the PECT of hovering hummingbirds, across all flight speeds we observed 2.9+/-0.8 spikes per contraction in the PECT and 3.8+/-0.8 spikes per contraction in the SUPRA. Muscle strain in the PECT was 10.8+/-0.5%, the lowest reported for a flying bird, and average strain rate was 7.4+/-0.2 muscle lengths s(-1). Among species of birds, PECT strain scales proportional to body mass to the 0.2 power (infinityM(b)(0.2)) using species data and infinityM(b)(0.3) using independent contrasts. This positive scaling is probably a physiological response to an adverse scaling of mass-specific power available for flight.

Pub.: 29 Jun '10, Pinned: 23 Apr '17

Muscle function in avian flight: achieving power and control.

Abstract: Flapping flight places strenuous requirements on the physiological performance of an animal. Bird flight muscles, particularly at smaller body sizes, generally contract at high frequencies and do substantial work in order to produce the aerodynamic power needed to support the animal's weight in the air and to overcome drag. This is in contrast to terrestrial locomotion, which offers mechanisms for minimizing energy losses associated with body movement combined with elastic energy savings to reduce the skeletal muscles' work requirements. Muscles also produce substantial power during swimming, but this is mainly to overcome body drag rather than to support the animal's weight. Here, I review the function and architecture of key flight muscles related to how these muscles contribute to producing the power required for flapping flight, how the muscles are recruited to control wing motion and how they are used in manoeuvring. An emergent property of the primary flight muscles, consistent with their need to produce considerable work by moving the wings through large excursions during each wing stroke, is that the pectoralis and supracoracoideus muscles shorten over a large fraction of their resting fibre length (33-42%). Both muscles are activated while being lengthened or undergoing nearly isometric force development, enhancing the work they perform during subsequent shortening. Two smaller muscles, the triceps and biceps, operate over a smaller range of contractile strains (12-23%), reflecting their role in controlling wing shape through elbow flexion and extension. Remarkably, pigeons adjust their wing stroke plane mainly via changes in whole-body pitch during take-off and landing, relative to level flight, allowing their wing muscles to operate with little change in activation timing, strain magnitude and pattern.

Pub.: 20 Apr '11, Pinned: 23 Apr '17

Muscle function during takeoff and landing flight in the pigeon (Columba livia).

Abstract: This study explored the muscle strain and activation patterns of several key flight muscles of the pigeon (Columba livia) during takeoff and landing flight. Using electromyography (EMG) to measure muscle activation, and sonomicrometry to quantify muscle strain, we evaluated the muscle function patterns of the pectoralis, biceps, humerotriceps and scapulotriceps as pigeons flew between two perches. These recordings were analyzed in the context of three-dimensional wing kinematics. To understand the different requirements of takeoff, midflight and landing, we compared the activity and strain of these muscles among the three flight modes. The pectoralis and biceps exhibited greater fascicle strain rates during takeoff than during midflight or landing. However, the triceps muscles did not exhibit notable differences in strain among flight modes. All observed strain, activation and kinematics were consistent with hypothesized muscle functions. The biceps contracted to stabilize and flex the elbow during the downstroke. The humerotriceps contracted to extend the elbow at the upstroke-downstroke transition, followed by scapulotriceps contraction to maintain elbow extension during the downstroke. The scapulotriceps also appeared to contribute to humeral elevation. Greater muscle activation intensity was observed during takeoff, compared with mid-flight and landing, in all muscles except the scapulotriceps. The timing patterns of muscle activation and length change differed among flight modes, yet demonstrated that pigeons do not change the basic mechanical actions of key flight muscles as they shift from flight activities that demand energy production, such as takeoff and midflight, to maneuvers that require absorption of energy, such as landing. Similarly, joint kinematics were consistent among flight modes. The stereotypy of these neuromuscular and joint kinematic patterns is consistent with previously observed stereotypy of wing kinematics relative to the pigeon's body (in the local body frame) across these flight behaviors. Taken together, these observations suggest that the control of takeoff and landing flight primarily involves modulation of overall body pitch to effect changes in stroke plane angle and resulting wing aerodynamics.

Pub.: 14 Sep '12, Pinned: 23 Apr '17

Kinematics and power requirements of ascending and descending flight in the pigeon (Columba livia).

Abstract: Ascending or descending locomotion involves a change in potential energy (PE) and a corresponding change in power requirement. We sought to test whether the mechanical power required for steady ascending or descending flight is a simple sum of the power required for level flight and the power necessary for potential energy change. Pigeons (Columba livia) were trained to fly at varying angles of ascent and descent (60 degrees , 30 degrees , 0 degrees , -30 degrees , -60 degrees ), and were recorded using high-speed video. Detailed three-dimensional kinematics were obtained from the recordings, allowing analysis of wing movement. Aerodynamic forces and power requirements were then estimated from kinematic data. As expected, ;PE flight power' increased significantly with angle of flight (0.234 W deg.(-1)), though there appeared to be a limit on the amount of PE that the birds could gain or dissipate per wingbeat. We found that the total power output for flight at various angles was not different from the sum of power required for level flight and the PE rate of change for a given angle, except for the steep -60 degrees descent. The total power for steep descent was higher than this sum because of a higher induced power due to the bird's deceleration and slower flight velocity. Aerodynamic force estimates during mid-downstroke did not differ significantly in magnitude or orientation among flight angles. Pigeons flew fastest during -30 degrees flights (4.9+/-0.1 m s(-1)) and slowest at 60 degrees (2.9+/-0.1 m s(-1)). Although wingbeat frequency ranged from 6.1 to 9.6 Hz across trials, the variation was not significant across flight angles. Stroke plane angle was more horizontal, and the wing more protracted, for both +60 degrees and -60 degrees flights, compared with other flight path angles.

Pub.: 18 Mar '08, Pinned: 23 Apr '17

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

Abstract: 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. Here, an account is made for such strategies by noting a well-described, general, physiological cost parameter of muscle: the cost of activation. Small birds, with brief downstrokes, experience disproportionately high costs due to muscle activation for power during contraction as opposed to work. Bounding flight may be an adaptation to modulate mean aerodynamic force production in response to 1) physiological pressure to extend the duration of downstroke to reduce power demands during contraction; 2) the prevention of a low-speed downstroke due to the geometric constraints of producing thrust; 3) an aerodynamic cost to flapping with very low lift coefficients. In contrast, flap-gliding birds, which tend to be larger, adopt a strategy that reduces the physiological cost of work due both to activation and contraction efficiency. Flap-gliding allows, despite constraints to modulation of aerodynamic force lever-arm, 1) adoption of moderately large wing-stroke amplitudes to achieve suitable muscle strains, thereby reducing the activation costs for work; 2) reasonably quick downstrokes, enabling muscle contraction at efficient velocities, while being 3) prevented from very slow weight-supporting upstrokes due to the cost of performing ‘negative’ muscle work.

Pub.: 11 Jul '16, Pinned: 23 Apr '17