Accipiters (Accipiter spp.) and falcons (Falco spp.) both use their feet to seize prey, but falcons kill primarily with their beaks, whereas accipiters kill with their feet. This study examines the mechanistic basis to differences in their modes of dispatching prey, by focusing on the myology and biomechanics of the jaws, digits, and distal hindlimb. Bite, grip, and distal hindlimb flexion forces were estimated from measurements of physiological cross-sectional area (PCSA) and indices of mechanical advantage (MA) for the major jaw adductors, and digit and tarsometatarsal flexors. Estimated bite force, total jaw adductor PCSA, and jaw MA (averaged over adductors) tended to be relatively and absolutely greater in falcons, reflecting their emphasis on biting for dispatching their prey. Differences between genera in estimated grip force, total digit flexor PCSA, and digit MA (averaged over inter-phalangeal joints and digits) were not as clear-cut; each of these parameters scaled positively allometric in accipiters, which may reflect the scaling of both prey size, and the proportion of mammalian prey consumed by this lineage with increasing body size. Estimated tarsometatarsal force was greater in falcons than in accipiters, due to their greater MA, which may reflect selection for incurring greater forces during prey strikes. Conversely, the comparatively lower tarsometatarsal MA in accipiters reflects their capacity for greater foot speed potentially necessary for grasping elusive prey. Thus, this study elucidates how differences in jaw and hindlimb musculoskeletal morphology of accipiters and falcons are reflected in differences in their killing modes, and through differences in their force-generating capacities.

Musculoskeletal underpinnings to differences in killing behavior between North American accipiters (Falconiformes: Accipitridae) and falcons (Falconidae).

Studies of medium- and large-bodied avian species have suggested that variation in flight muscle composition is related to differences in flight behavior. For example, slow-twitch or tonic fibers are generally found only in the flight muscles of non-volant or soaring/gliding birds. However, we know comparatively little about fiber composition of the muscles of the smallest birds. Here we describe the fiber composition of muscles from the wings, shoulders, and legs of two small avian species, which also display very high wingbeat frequencies: Anna's hummingbirds (Calypte anna) and zebra finches (Taeniopygia guttata). All flight muscles examined in both species contained exclusively fast oxidative glycolytic (FOG) fibers. These unique results suggest that fast oxidative fibers are both necessary and sufficient for the full range of flight behaviors in these small-bodied birds. Like all other studied birds, the zebra finch gastrocnemius, a tarsometatarsal extensor, contained a mixture of FOG (27.1%), slow oxidative (SO, 12.7%), and fast glycolytic (FG, 60.2%) fibers. By contrast, the hummingbird gastrocnemius lacked FG fibers (85.5% FOG, 14.5% SO), which may reflect the reduced role of the hindlimb during take-off. We further hypothesize that thermogenic requirements constrain fiber type heterogeneity in these small endothermic vertebrates.

Fiber type homogeneity of the flight musculature in small birds.

Twenty-three species within the avian family Alcidae are capable of wing-propelled flight in the air and underwater. Alcids have been viewed as Northern Hemisphere parallels to penguins, and have often been studied to see if their underwater flight comes at a cost, compromising their aerial flying ability. We examined the anatomy and histochemistry of select wing muscles (Mm. pectoralis, supracoracoideus, latissimus dorsi caudalis, coracobrachialis caudalis, triceps scapularis, and scapulohumeralis caudalis) from Atlantic puffins (Fratercula arctica) to assess if the muscle fiber types reveal the existence of a compromise associated with "dual-medium" flight. Pectoralis was found to be proportional in size with that of nondiving species, although the supracoracoideus was proportionally larger in puffins. Muscle fiber types were largely aerobic in both muscles, with two distinct fast-twitch types demonstrable: a smaller, aerobic, moderately glycolytic population (FOg), and a larger, moderately aerobic, glycolytic population (FoG). The presence of these two fiber types in the primary flight muscles of puffins suggests that aerial and underwater flight necessitate a largely aerobic fiber complement. We suggest that alcids do not represent an adaptive compromise, but a stable adaptation for wing-propelled locomotion both in the air and underwater.

/pdf/anatomy-and-histochemistry-of-flight-muscles-in-a-wing-4nhjn2b40m.pdf

Anatomy and histochemistry of flight muscles in a wing-propelled diving bird, the Atlantic puffin, Fratercula arctica.

As a postural behavior, gliding and soaring flight in birds requires less energy than flapping flight. Slow tonic and slow twitch muscle fibers are specialized for sustained contraction with high fatigue resistance and are typically found in muscles associated with posture. Albatrosses are the elite of avian gliders; as such, we wanted to learn how their musculoskeletal system enables them to maintain spread-wing posture for prolonged gliding bouts. We used dissection and immunohistochemistry to evaluate muscle function for gliding flight in Laysan and Black-footed albatrosses. Albatrosses possess a locking mechanism at the shoulder composed of a tendinous sheet that extends from origin to insertion throughout the length of the deep layer of the pectoralis muscle. This fascial "strut" passively maintains horizontal wing orientation during gliding and soaring flight. A number of muscles, which likely facilitate gliding posture, are composed exclusively of slow fibers. These include Mm. coracobrachialis cranialis, extensor metacarpi radialis dorsalis, and deep pectoralis. In addition, a number of other muscles, including triceps scapularis, triceps humeralis, supracoracoideus, and extensor metacarpi radialis ventralis, were found to have populations of slow fibers. We believe that this extensive suite of uniformly slow muscles is associated with sustained gliding and is unique to birds that glide and soar for extended periods. These findings suggest that albatrosses utilize a combination of slow muscle fibers and a rigid limiting tendon for maintaining a prolonged, gliding posture.

/pdf/anatomy-and-histochemistry-of-spread-wing-posture-in-birds-3-32jn8bo7cl.pdf

Anatomy and histochemistry of spread-wing posture in birds. 3. Immunohistochemistry of flight muscles and the "shoulder lock" in albatrosses.

Two styles of bird locomotion, hovering and intermittent flight, have great potential to inform future development of autonomous flying vehicles Hummingbirds are the smallest flying vertebrates, and they are the only birds that can sustain hovering Their ability to hover is due to their small size, high wingbeat frequency, relatively large margin of mass-specific power available for flight and a suite of anatomical features that include proportionally massive major flight muscles (pectoralis and supracoracoideus) and wing anatomy that enables them to leave their wings extended yet turned over (supinated) during upstroke so that they can generate lift to support their weight Hummingbirds generate three times more lift during downstroke compared with upstroke, with the disparity due to wing twist during upstroke Much like insects, hummingbirds exploit unsteady mechanisms during hovering including delayed stall during wing translation that is manifest as a leading-edge vortex (LEV) on the wing and rotational circulation at the end of each half stroke Intermittent flight is common in small- and medium-sized birds and consists of pauses during which the wings are flexed (bound) or extended (glide) Flap-bounding appears to be an energy-saving style when flying relatively fast, with the production of lift by the body and tail critical to this saving Flap-gliding is thought to be less costly than continuous flapping during flight at most speeds Some species are known to shift from flap-gliding at slow speeds to flap-bounding at fast speeds, but there is an upper size limit for the ability to bound (~03 kg) and small birds with rounded wings do not use intermittent glides

/pdf/hovering-and-intermittent-flight-in-birds-5btruk2dx5.pdf

Hovering and intermittent flight in birds.

Gliding flight is a postural activity which requires the wings to be held in a horizontal position to support the weight of the body. Postural behaviors typically utilize isometric contractions in which no change in length takes place. Due to longer actin-myosin interactions, slow contracting muscle fibers represent an economical means for this type of contraction. In specialized soaring birds, such as vultures and pelicans, a deep layer of the pectoralis muscle, composed entirely of slow fibers, is believed to perform this function. Muscles involved in gliding posture were examined in California gulls (Larus californicus) and tested for the presence of slow fibers using myosin ATPase histochemistry and antibodies. Surprisingly small numbers of slow fibers were found in the M. extensor metacarpi radialis, M. coracobrachialis cranialis, and M. coracobrachialis caudalis, which function in wrist extension, wing protraction, and body support, respectively. The low number of slow fibers in these muscles and the absence of slow fibers in muscles associated with wing extension and primary body support suggest that gulls do not require slow fibers for their postural behaviors. Gulls also lack the deep belly to the pectoralis found in other gliding birds. Since bird muscle is highly oxidative, we hypothesize that fast muscle fibers may function to maintain wing position during gliding flight in California gulls.

/pdf/anatomy-and-histochemistry-of-spread-wing-posture-in-birds-2-1byqa4kyus.pdf

Anatomy and histochemistry of spread‐wing posture in birds. 2. Gliding flight in the California Gull, Larus californicus: A paradox of fast fibers and posture

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Anatomy and histochemistry of spread‐wing posture in birds. 2. Gliding flight in the California Gull, Larus californicus: A paradox of fast fibers and posture