Comparative power curves in bird flight.

The relationship between mechanical power output and forward velocity in bird flight is controversial, bearing on the comparative physiology and ecology of locomotion. Applied to flying birds, aerodynamic theory predicts that mechanical power should vary as a function of forward velocity in a U-shaped curve. The only empirical test of this theory, using the black-billed magpie (Pica pica), suggests that the mechanical power curve is relatively flat over intermediate velocities. Here, by integrating in vivo measurements of pectoralis force and length change with quasi-steady aerodynamic models developed using data on wing and body movement, we present mechanical power curves for cockatiels (Nymphicus hollandicus) and ringed turtle-doves (Streptopelia risoria). In contrast to the curve reported for magpies, the power curve for cockatiels is acutely concave, whereas that for doves is intermediate in shape and shows higher mass-specific power output at most speeds. We also find that wing-beat frequency and mechanical power output do not necessarily share minima in flying birds. Thus, aspects of morphology, wing kinematics and overall style of flight can greatly affect the magnitude and shape of a species' power curve.

Bird maneuvering flight: blurred bodies, clear heads.

While useful in describing the efficiency of maneuvering flight, steady-state (i.e., fixed wing) models of maneuvering performance cannot provide insight to the efficacy of maneuvering, particularly during low-speed flapping flight. Contrasted with airplane-analogous gliding/high speed maneuvering, the aerodynamic and biomechanical mechanisms employed by birds at low flight speeds are violent, with rapidly alternating forces routinely being developed. The saltatory nature of this type of flight results in extreme linear and angular displacements of the bird's body; however, birds isolate their heads from these accelerations with cervical reflexes. Experiments with pigeons suggest this ability to isolate the visual and vestibular systems is critical to controlled flapping flight: birds wearing collars that prohibited the neck from isolating the head from the angular accelerations of induced rolls frequently exhibited (50% of flights) a loss of vestibular and/or visual horizon and were unable to maintain controlled flight.

Pectoralis muscle performance during ascending and slow level flight in mallards (Anas platyrhynchos).

In vivo measurements of pectoralis muscle length change and force production were obtained using sonomicrometry and delto-pectoral bone strain recordings during ascending and slow level flight in mallards (Anas platyrhynchos). These measurements provide a description of the force/length properties of the pectoralis under dynamic conditions during two discrete flight behaviors and allow an examination of the effects of differences in body size and morphology on pectoralis performance by comparing the results with those of a recent similar study of slow level flight in pigeons (Columbia livia). In the present study, the mallard pectoralis showed a distinct pattern of active lengthening during the upstroke. This probably enhances the rate of force generation and the magnitude of the force generated and, thus, the amount of work and power produced during the downstroke. The power output of the pectoralis averaged 17.0 W kg(-)(1 )body mass (131 W kg(-)(1 )muscle mass) during slow level flight (3 m s(-)(1)) and 23.3 W kg(-)(1 )body mass (174 W kg(-)(1 )muscle mass) during ascending flight. This increase in power was achieved principally via an increase in muscle strain (29 % versus 36 %), rather than an increase in peak force (107 N versus 113 N) or cycle frequency (8.4 Hz versus 8.9 Hz). Body-mass-specific power output of mallards during slow level flight (17.0 W kg(-)(1)), measured in terms of pectoralis mechanical power, was similar to that measured recently in pigeons (16.1 W kg(-)(1)). Mallards compensate for their greater body mass and proportionately smaller wing area and pectoralis muscle volume by operating with a high myofibrillar stress to elevate mechanical power output.

Effects of body size on take-off flight performance in the Phasianidae (Aves).

To evaluate the mechanisms responsible for relationships between body mass and maximum take-off performance in birds, we studied four species in the Phasianidae: northern bobwhite (Colinus virginianus), chukar (Alectoris chukar), ring-necked pheasant (Phasianus colchicus) and wild turkey (Meleagris gallopavo). These species vary in body mass from 0.2 to 5.3 kg, and they use flight almost solely to escape predators. During take-off, all the species used a similar wingbeat style that appeared to be a vortex-ring gait with a tip reversal during the upstroke. The tip reversal is unusual for birds with rounded wings; it may offer an aerodynamic advantage during rapid acceleration. Flight anatomy generally scaled geometrically, except for average wing chord and wing area, which increased more than expected as body mass (m) increased. Pectoralis strain varied from 19.1 to 35.2 % and scaled in proportion to m(0.23). This positive scaling is not consistent with the widely held assumption that muscle strain is independent of body mass among geometrically similar species. The anatomy of the species precluded measurements of in vivo pectoralis force using the strain-gauge technique that has been employed successfully in other bird species, so we could not directly test in vivo pectoralis force-velocity relationships. However, whole-body kinematics revealed that take-off power (P(ta)), the excess power available for climbing and accelerating in flight, scaled in proportion to m(0.75) and that pectoralis mass-specific P(ta) decreased in proportion to m(-)(0.26) and was directly proportional to wingbeat frequency. These trends suggest that mass-specific pectoralis work did not vary with body mass and that pectoralis stress and strain were inversely proportional, as expected from classical force-velocity models for skeletal muscle. Our observations of P(ta) were consistent with evidence from other species engaged in escape flight and, therefore, appear to contradict evidence from studies of take-off or hovering with an added payload.

Flight style of the black-billed magpie: variation in wing kinematics, neuromuscular control, and muscle composition.

Black-billed magpies (Pica pica; Corvidae) exhibit an unusual flight style with pronounced, cyclic variation in wingbeat frequency and amplitude during level, cruising flight. In an effort to better understand the underlying internal mechanisms associated with this flight style, we studied muscle activity patterns, fiber composition of the pectoralis muscle, and wingbeat kinematics using both laboratory and field techniques. Over a wide range of speeds in a windtunnel (0-13.4 m s-1), wingbeat frequency, wingtip elevation, and relative intensity of electromyographic (EMG) signals s-1 from the flight muscles were least at intermediate speeds, and increased at both slower and faster speeds, in approximate agreement with theoretical models that predict a U-shaped curve of power output with flight speed. Considerable variation was evident in kinematic and electromyographic variables, but variation was continuous, and, thus, was not adequately described by the simple two-gait system which is currently accepted as describing gait selection during vertebrate flight. Indirect evidence suggests that magpies vary their flight style consistent with reducing average power costs in comparison to costs associated with continuous flapping at a fixed level of power per wingbeat. The range of variation for the kinematic variables was similar in the field and lab; however, in the field, proportionally fewer flights showed significant correlations between wingbeat frequency and the other variables. Average flight speed in the field was 8.0 m s-1. Average wingbeat frequency was less in the field than in the windtunnel, but mean values for wingtip elevation and wingspan at midupstroke were not significantly different. Histological study revealed that the pectoralis muscle of magpies contained only fast-twitch (acid-stable) muscle fibers, which were classified as red (R) and intermediate (I) based on oxidative and glycolytic capacities along with fiber diameter. This fiber composition may be related to variation in wingbeat kinematics, but such composition is found in the pectoralis of other bird species. This suggests that the muscle fibers commonly found in the pectoralis of small to medium sized birds are capable of a wider range of efficient contractile velocities than predicted by existing theory. Future studies should explore alternative explanations for variation in wingbeat kinematics, including the potential role of nonverbal communication among cospecifics.

A cineradiographic analysis of bird flight: the wishbone in starlings is a spring.

High-speed x-ray movies of European starlings flying in a wind tunnel provide detailed documentation of avian skeletal movements during flapping flight. The U-shaped furcula (or "wishbone," which represents the fused clavicles) bends laterally during downstroke and recoils during upstroke; these movements may facilitate inflation and deflation of the clavicular air sac. Sternal movements are also coupled with wingbeat, ascending and retracting on downstroke and descending and protracting on upstroke in an approximately elliptical pathway. The coupled actions of the sternum and furcula appear to be part of a respiratory cycling mechanism between the lungs and air sacs.

A functional analysis of the primary upstroke and downstroke muscles in the domestic pigeon (Columba livia) during flight.

In domestic pigeons (Columba livia), the electrical activity of the major depressor muscle of the wing, the pectoralis (pars thoracicus), beings in late upstroke well before the wing begins its downstroke excursion. The two architecturally distinct heads of the pectoralis, the sternobrachialis and the thoracobrachialis, are differentially recruited during take-off, level flight and landing. In addition to wing depression, the sternobrachialis protracts the humerus and the thoracobrachialis retracts the humerus. At the point of transition from wing upstroke to downstroke, the pectoralis EMG signal typically exhibits a reduction in amplitude. The supracoracoideus, in addition to an expected EMG associated with wing elevation, is co-activated with the pectoralis about 50% of the time.

Structure and neural control of the pectoralis in pigeons: implications for flight mechanics.

The pectoralis muscle in pigeons (Columba livia) is composed of two heads (sternobrachialis, thoracobrachialis) that are separately innervated and have different fiber orientations. High-speed film and electromyographic studies of free-flying pigeons reveal that the pectoralis is activated prior to wing depression (the power stroke) and that its two heads are differentially recruited during takeoff, level flight, and landing. The electrical activity patterns of both heads support an interpretation that intramuscular elasticity provides energy storage. The pectoralis is not only the prime wing depressor but is also capable of adjusting the excursion of the wing during different phases of flight.