Fibre types in primary ‘flight’ muscles of the African Penguin (Spheniscus demersus)

The African penguin (Spheniscus demersus) is an endangered seabird that resides on the temperate southern coast of Africa. Like all penguins it is flightless, instead using its specialized wings for underwater locomotion termed ‘aquatic flight’. While musculature and locomotion of the large Antarctic penguins have been well studied, smaller penguins show different biochemical and behavioural adaptations to their habitats. We used histochemical and immunohistochemical methods to characterize fibre type composition of the African penguin primary flight muscles, the pectoralis and supracoracoideus. We hypothesized the pectoralis would contain predominantly fast oxidative–glycolytic (FOG) fibres, with mainly aerobic subtypes. As the supracoracoideus and pectoralis both power thrust, we further hypothesized these muscles would have a similar fibre type complement. Our results supported these hypotheses, also showing an unexpected slow fibre population in the deep parts of pectoralis and supracoracoideus. The latissimus dorsi was also examined as it may contribute to thrust generation during aquatic flight, and in other avian species typically contains definitive fibre types. Unique among birds studied to date, the African penguin anterior latissimus dorsi was found to consist mainly of fast fibres. This study shows the African penguin has specialized flight musculature distinct from other birds, including large Antarctic penguins.     

Adaptations for rapid and forceful contraction in wing muscles of the male golden-collared manakin: sex and species comparisons

The courtship display of the male golden-collared manakin (Manacus vitellinus) of Panamanian rainforests is noteworthy for several types of whip-crack-like sounds created by a rapid overhead flip of the wings. We have hypothesized that this courtship behavior. which is not performed by females, is associated with steroid-sensitive and sexually dimorphic neuromuscular systems. Presumably, muscles creating the motion of the wingsnap in males are specialized for greater force generation and speed of contraction. We tested this hypothesis by examining mass, fiber diameter, metabolic enzyme activity, and myosin isoform expression in several muscles of male and female manakins and in both sexes of a non-wingsnapping bird, the zebra finch (Taenopygia guttata). We have identified three wing muscles, the scapulohumeralis caudalis, the supracoracoideus, and the pectoralis major, that differ in one or more of these characteristics across sex and species, suggesting they are specialized for faster contraction and greater force production in male manakins. These muscles normally function to raise and lower the wings during flight. As this movement is the principal motion of the wingsnap, these adaptations presumably underlie the performance of the wingsnap display.     

New data on the humerotriceps of penguins and its implications in the evolution of the fossa tricipitalis

A paddle-shaped wing, the general morphology of the humerus, and the muscles involved in wing movement are among the most characteristic adaptations to diving in penguins. Particularly, the humeral fossa tricipitalis and the musculus humerotriceps are clear examples of muscular rearrangement accompanying skeletal changes. In extant Spheniscidae, we were able to identify two heads of this muscle attaching within a different compartment of the bipartite fossa. Since the partition of the fossa appeared as a novelty during the Miocene, we propose that this might have had implications for underwater flight contributing to wing-propelled diving efficiency.     

Endocranial Anatomy of the Charadriiformes: Sensory System Variation and the Evolution of Wing-Propelled Diving

Just as skeletal characteristics provide clues regarding behavior of extinct vertebrates, phylogenetically-informed evaluation of endocranial morphology facilitates comparisons among extinct taxa and extant taxa with known behavioral characteristics. Previous research has established that endocranial morphology varies across Aves; however, variation of those systems among closely related species remains largely unexplored. The Charadriiformes (shorebirds and allies) are an ecologically diverse clade with a comparatively rich fossil record, and therefore, are well suited for investigating interspecies variation, and potential links between endocranial morphology, phylogeny, ecology and other life history attributes. Endocranial endocasts were rendered from high resolution X-ray computed tomography data for 17 charadriiforms (15 extant and two flightless extinct species). Evaluation of endocranial character state changes on a phylogeny for Charadriiformes resulted in identification of characters that vary in taxa with distinct feeding and locomotor ecologies. In comparison with all other charadriiforms, stem and crown clade wing-propelled diving Pan-Alcidae displayed compressed semicircular canals, and indistinct occipital sinuses and cerebellar fissures. Flightless wing-propelled divers have relatively smaller brains for their body mass and smaller optic lobes than volant pan-alcids. Observed differences between volant and flightless wing-propelled sister taxa are striking given that flightless pan-alcids continue to rely on the flight stroke for underwater propulsion. Additionally, the brain of the Black Skimmer Rynchops niger, a taxon with a unique feeding ecology that involves continuous forward aerial motion and touch-based prey detection used both at day and night, is discovered to be unlike that of any other sampled charadriiform in having an extremely large wulst as well as a small optic lobe and distinct occipital sinus. Notably, the differences between the Black Skimmer and other charadriiforms are more pronounced than between wing-propelled divers and other charadriiforms. Finally, aspects of endosseous labyrinth morphology are remarkably similar between divers and non-divers, and may deserve further evaluation.     

Twofold seasonal variation in the supposedly constant, species-specific, ratio of upstroke to downstroke flight muscles in red knots Calidris canutus

We show that in a long-distance migrant shorebird species with outspoken seasonal changes in body mass and composition, the red knot Calidris canutus , the ratio between the masses of the small flight muscle ( musculus supracoracoideus , powering twists and active upstrokes of the wings) and the larger flight muscle ( musculus pectoralis , for the downstrokes) is far from constant. During an annual cycle the supracoracoideus / pectoralis ratio varied more than twofold between values of 0.058 (±0.005 SE) in early winter period and of 0.124 (±0.05 SE) on the High Arctic tundra breeding grounds. The ratios thus spanned a range from those typical of soaring raptors and seabirds to those of fast and agile fliers and birds with rapid take-offs. The overall average ratio was 0.102 (±0.001 SE, for non-starved knots, and 0.103±0.001 including starved knots) and did not differ between males and females. As predicted from the known functions of supracoracoideus and pectoralis , the ratio was a negative function of body mass. However, after arrival on the breeding grounds (0.124) and during winter starvation (0.135) particularly high ratios were reached: these may be times when wing-manoeuvrability (in flight display and during the evasive 'rodent run' away from predators at the nest) and an ability for rapid take-off and active up-strokes (from –near– the nest, and in times of depletion of flight muscle mass during winter starvation) may be at premium. The particularly low ratio of 0.06 in early winter is puzzling. Many aspects of avian phenotypes have recently been shown to be intraindividually variable. To a twofold seasonal variation in flight muscle mass ( Dietz et al. 2007 ), we can now add the twofold variation in the ratio between the muscles for the upstroke and the downstroke.     

REPTILIAN PHYSIOLOGY AND THE FLIGHT CAPACITY OF ARCHAEOPTERYX

Current scenarios frequently interpret the Late Jurassic bird Archaeopteryx as having had an avian-type physiology and as having been capable of flapping flight, but only from “the trees downward.” It putatively lacked capacity for takeoff and powered flight from the ground upward. Data from extant reptiles indicate that if Archaeopteryx were physiologically reptilian, it would have been capable of ground upward takeoff from a standstill, as well as “trees downward” powered flight. This conclusion is based largely on a previously unrecognized attribute of locomotory (skeletal) muscle in a variety of extant reptiles: During “burst-level” activity, major locomotory muscles of a number of active terrestrial taxa generate at least twice the power (watts kg^?1 muscle tissue) as those of birds and mammals. Reptilian physiological status also helps resolve the apparently uneven development of various flight support structures in Archaeopteryx (e.g., well-developed flight features but relatively unspecialized pectoral girdle, supracoracoideus muscles, etc.). Endothermy and capacity for longer-distance powered flight probably evolved only in Early Cretaceous birds, which were the first birds to have a keeled sternum, strap-like coracoid, and hypocleidium-bearing furcula.     

Capillarity and fiber types in locomotory muscles of wild common coots,Fulica atra

Six locomotory muscles of wild common coots, Fulica atra, were analyzed histochemically. Capillarity and fiber-type distributions were correlated to the functional implications and physiological needs of each muscle. Leg muscles exhibit three unevenly distributed fiber types, a pattern that reflects the great variety of terrestrial and aquatic locomotory performances that coots are able to develop. Aerobic zones are presumably recruited during steady swimming and diving, while regions with anaerobic characteristics may be used for bursts of activity such as sprint swimming or during take off, when coots run along the water's surface. Fiber types and capillarization in wing muscles have a marked oxidative trend. High wing beat frequencies, short and broad wings, and the long distance migrations that these birds perform indicate that the presence of high numbers of oxidative fibers and the well developed capillary supply are needed for enhanced oxygen uptake. The pectoralis muscle, except in its deep part, has exclusively fast oxidative fibers with a very high staining intensity for succinate dehydrogenase assay as compared to the same fiber type of other muscles. Its predominant role in flapping flight justifies these characteristics that are typical of fibers with high aerobic metabolism. The deep part of the pectoralis muscle presents a low proportion of an unusual slow anaerobic fiber type. These fibers could play a role during feeding dives when the bird presses the air out of the feathers by tightening the wings against the body. A linear relationship between capillary and fiber densities in all coot muscles studied reflects an adjustment between fiber diameter and vascularization in order to obtain the oxygen for mitochondrial supply. This strategy seems a suitable way to cope with the rigid aerobic constraints that flying and diving impose upon the coot's physiology. J. Morphol. 237:147–164, 1998. © 1998 Wiley-Liss, Inc.     

Muscle function in avian flight: achieving power and control

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.     

Histochemical and biochemical properties of flight muscle fibers in the little brown bat,Myotis lucifugus

Summary The purpose of this investigation was (1) to determine the fiber composition of pectoralis muscle of the little brown bat,Myotis lucifugus; (2) to compare the fiber composition of this muscle with two of the animal's accessory flight muscles; and (3) to study the effect of hibernation on pectoralis muscle fiber composition. Bat skeletal muscle fibers were also compared with those of white laboratory rats (Rattus norvegicus). Bat pectoralis muscles possessed exceptionally high oxidative capacities as indicated by their succinate dehydrogenase activities, but relatively low glycolytic potentials (phosphofructokinase activities). Muscle histochemistry demonstrated that fiber composition of bat pectorlis muscle was homogeneous; all fibers possessed high aerobic and low glycolytic potentials, and high myofibrillar ATPase activities indicating fast contractile properties. In contrast, accessory flight muscles possessed three distinguishable fiber types. During hibernation there was a significant decline in oxidative potential, no change in glycolytic potential, and no alteration in basic fiber composition of bat pectoralis muscle. The findings of this study suggest that pectoralis muscles ofM. lucifugus may approach the ultimate adaptation of a mammalian locomotory muscle for aerobic generation of muscular power.Abbreviations FG fast-twich glycolytic - FOG fast-twitch-oxydative-glycolytic - -GPDH -glycerophosphate dehydrogenase - LDH lactate dehydrogenase - NADH-D reduced nicotinamide adenine dinucleotide diaphorase - PFK phosphofructokinase - SDH succinate dehydrogenase - SO slowtwich-oxidative     

Structural basis for oxygen delivery: Muscle capillaries and manifolds in tuna red muscle

We summarize our morphometric data on fiber vascularization and aerobic capacity in red muscle of tuna (Katsuwonus pelamis), compared to intensely aerobic flight muscles of hummingbird (Selasphorus rufus, BW 3–4 g) and bat (Eptesius fuscus, BW 15–16 g, Pipistrellus hesperus, BW 3–5 g). Three characteristic features of high flux paths for oxygen: (a) small fiber size, (b) dense capillary network and (c) high mitochondrial volume density were found in tuna, but they were not as pronounced as in hummingbird and bat flight muscles. A particular arrangement of capillary manifolds, also seen in flight muscle of birds but not in bats, was found in tuna, forming dense envelopes of capillary branches around portions of muscle fibers. However, all indexes of fiber capillarization were relatively low in tuna red muscle for its mitochondrial volume, compared with other intensely aerobic muscles. Capillary length per unit volume of mitochondria, and capillary surface per mitochondrial inner (and outer) membrane surface area, were about one half of those in hummingbird or bat flight muscles. Consistent differences exist in the size of the capillary network for the size of the mitochondrial compartment in highly aerobic red muscle of tuna compared with bird and mammal.     

A histochemical comparison of fibre types in the M. pectoralis and M. supracoracoideus of the great skua Catharacta skua and the herring gull Larus argentatus with reference to kleptoparasitic capabilities

Skuas, which are closely related to gulls, frequently use a specialized feeding method (kleptoparasitism) by which they rob other seabirds of their food. This paper tests the idea that skuas have evolved as specialist kleptoparasites.
The fibre type composition of the M. pectoralis, M. supracoracoideus and M. iliofibularis of a great skua Catharacta skua (Brünn.) and a herring gull Larus argentatus (Pontopp.) was determined by three enzyme histochemical methods commonly used for mammalian fibre classification; the reactions for alphaGPDH, NADH-TR and mATPase activity.
In both species slow fibres were present only in the M. iliofibularis, and fast twitch glycolytic fibres were not present in any of the muscles. The M. pectoralis and M. supracoracoideus of both species consisted entirely of the fast twitch oxidative-glycolytic fibres.
The overall metabolic enzyme activities of the muscles were assessed in terms of the proportions of fibres with high, intermediate and low metabolic enzyme activity. The overall levels of oxidative and glycolytic enzyme activity were significantly higher in the M. pectoralis than in the M. supracoracoideus and significantly higher in both of these than in the M. iliofibularis. This was true of both species.
The oxidative and glycolytic activities of all three muscles of the great skua were significantly higher than those of the homologous muscles of the herring gull. This was particularly true of the M. pectoralis and M. supracoracoideus. It is suggested that this difference between great skuas and herring gulls enables the former to be more effective aerial kleptoparasites than the latter.     

Myoglobin levels and mATPase activity in pectoral muscles of spruce and ruffed grouse (Aves: Tetraoninae)

Myoglobin concentration and myosin ATPase activity were measured in the pectoral muscle of wild spruce grouse (Dendragapus canadensis) and ruffed grouse (Bonasa umbellus), together with the weight of the Mm. pectoralis, supracoracoideus and heart. mATPase activities were similar in both species, but spruce grouse contained 15 times more myoglobin in the pectoralis muscle and the heart was three times heavier than that of the ruffed grouse. The relative mass of the flight muscles and wing loading were similar between species. Characteristics of the pectoral muscle of both grouse species reflect adaptations to predation and advertising displays. The glycolytic nature of the ruffed grouse pectoral muscle and small heart size is an adaptation to a sedentary existence within a small home range. The more oxidative pectoral muscle of spruce grouse together with its larger heart are adaptations to seasonal dispersals requiring more sustained flight.     

Effects of the sex-linked dwarf gene (dw) on the expression of the muscular dystrophy gene (am) in chicken.

The sex-linked dwarf gene (dw) was introduced into companion muscular dystrophic (am) and nondystrophic (Am+) New Hampshire chicken lines to investigate influences of the dwarf gene on breast muscle weights, muscle fiber area, and the histological expression of muscular dystrophy. Dystrophic and nondystrophic chickens within dwarf or nondwarf genotypes were similar in body and carcass weights. Pectoralis and supracoracoideus muscle weights (as a percentage of adjusted carcass weight) were similar in nondystrophic dwarf and nondwarf males and females. In addition, pectoralis weight was similar in dystrophic dwarf males and dystrophic nondwarf males and females. However, pectoralis weight was significantly smaller in dystrophic dwarf females than in dystrophic nondwarf females, whereas supracoracoideus weight was significantly larger in dystrophic dwarf males than in dystrophic nondwarf males. Supracoracoideus weight was similar in dystrophic dwarf males and females and dystrophic nondwarf females. Pectoralis muscle fiber area was influenced by sex and by dwarf and dystrophy genotype. Muscle fiber area was larger in females than in males, smaller in dwarfs than in nondwarfs, and smaller in dystrophic than in nondystrophic muscles. Muscle fiber degeneration and adipose infiltration was more extensive in dystrophic than in nondystrophic females and males, and it was more advanced in dwarfs than in nondwarfs. Excessive acetylcholinesterase staining patterns were characteristic of dystrophic muscle in both dwarf and nondwarf genotypes. Nondystrophic and dystrophic dwarf male and female chickens are comparable substitutes for nondwarfs as biomedical models with respect to pectoralis histology, acetylcholinesterase staining pattern, and pectoralis muscle hypertrophy.     

Uniform myosin isoforms in the flight muscles for little brown bats, Myotis lucifugus

The present study used muscle histochemistry and polyacrylamide gel electrophoresis of native myosin and myosin heavy chains to establish a correlation, if any, between chiropteran histochemical fiber types and myosin isoform composition. Histochemical analysis of the primary flight muscle, the pectoralis profundus, documented the presence of a single histochemical fiber type, here termed Type II. Electrophoresis of native myosin isolated from pectoralis muscle yielded a single isoform that comigrated with the FM-3 isoform of rat diaphragm. Heavy chain analysis of the Myotis pectoralis demonstrated a single heavy chain with comparable electrophoretic mobility to rat IIa myosin heavy chain. These data demonstrate unique histochemical and biochemical homogeneity in the myosin composition of the pectoralis muscle of Myotis lucifugus. Thus this muscle is extremely specialized for flight at histochemical, morphologic, and molecular levels. These data contrast with the mixed myosin and histochemical fiber types found in other mammals, as well as in other muscles of Myotis lucifugus.     

Histochemistry of flight muscles in the Jamaican fruit bat, Artibeus jamaicensis: implications for motor control

Two fast-twitch fiber types are histochemically identified in the primary flight muscles of Artibeus jamaicensis. These are classified as type IIa and IIb according to an acid-preincubation staining protocol for myosin ATPase. All fibers in the bat flight muscles exhibit relatively intense staining properties for NADH-TR, suggesting a high oxidative capacity. The glycolytic potential of all fibers is rather low, as assessed by stains for alpha-GPD. This two-type histochemical profile appears to parallel biphasic electromyographic patterns observed in these muscles and leads us to propose that flight muscle histochemistry and activation are mediated by a "two-gear" neuromuscular control system. In contrast, earlier studies on Tadarida brasiliensis demonstrate the existence of a "one-gear" neuromuscular control system, exemplified by the presence of one fiber type. These observations are discussed with respect to the natural history and flight styles of several species.     

Morphologic-functional study of the locomotor system of penguins as a general model of movement in under-water flight. I

Regarding several theories of the evolution of the Sphenisciformes the specific morpho-physiological alterations for the changeover from aerial to underwater life are discussed. The peculiarities in the Penguin's "construction" become comprehensible as strong adjustments to the subaquatic locomotion. Surely they took their origin from the equipment of flying birds. The present data of the kinematics of the underwater locomotion show, that propulsion is produced in the same principal way by the flapping wings as in aerial flight. Therefore the short term "underwater flight" for the Penguin's style of locomotion is justified. Known data of swimming performance suggest that its essential adaptation is not that to top achievements but more to an economical use of energy budget. The favourable hydrodynamic characteristics of the Penguin body may be well interpreted from this point of view. The peculiarity of underwater flight is the absence of the necessity to produce a weight-compensating force. In order to create thrust forces in an appropriate magnitude during up- and downstroke of the beating cycle the upstroke must be powered. The anatomical architecture and the mode of operation of the parts of the muscle system must be adjusted to this demand. Based on these statements, the anatomy of active and passive apparatus of movement was studied by dissection of 26 individuals of Pygoscelis papua, P. antarctica, P. adeliae, Eudyptes chrysolophus, and Aptenodytes forsteri. Besides the functional explanation of the Articulatio sternocoracoidea (diverging considerably from the usual type in birds), a new interpretation is given for the structures of the Articulatio humeri. In this context, the role of the Ligamentum acrocoracohumerale as an important element for coordination of the motion processes in the shoulder joint is elucidated. The essential curvature of the Caput humeri is found to be satisfactorily approximated by a logarithmic spiral. The understanding of the mechanics of bones and tendons leads to a reinterpretation of the role of several groups of muscles which is described in detail. Besides of the preponderant thrust producing (flapping) muscles working mainly in the isotonic manner, muscles can be distinguished which are managing the transfer of the produced forces to the body operating thereby in the isometric way. Another group of muscles has to control the position of the humerus adjusting in this way the hydrodynamic angle of attack corresponding to the respective flow conditions.(ABSTRACT TRUNCATED AT 400 WORDS)     

Ratios,adaptations, and the differential metabolic capability of avian flight muscles

The eared grebe Podiceps nigricollis shows seasonal variation in the relative size of the major flight muscles that lift and lower the wing: respectively, supracoracoideus (s) and pectoralis (p). S/p ratios are low (≈0.07–0.12) when grebes are in flying condition, higher (≈0.11–0.15) when staging and flightless, and extreme (to 0.29) when starving. Shifts were driven by changes in the protein content in the pectoralis; intramuscular fat had little effect. S/p ratios also vary seasonally in the red knot Calidris canutus and are higher in birds newly arrived in breeding areas than at other times. If that increase was an adaptive response to promote wing‐lifting in association with various breeding behaviors as suggested, one would expect it to result from an absolute increase in the post‐arrival size of the supracoracoideus, which was not observed. Instead, we propose that it is unrelated to enhancing the upstroke but results from a decrease in the size of the pectoralis, which is a consequence of the greater rate at which this muscle is catabolized in times of exertion and stress, as at the end of a long migration or during starvation. Fuller data on the size, morphology and physiology of individual muscles at various stages of the annual cycle and migration will help to clarify how ratio changes are achieved, and evaluate potential adaptive significance.     

Functional morphometric analysis of the furcula in mesozoic birds

The furcula displays enormous morphological and structural diversity. Acting as an important origin for flight muscles involved in the downstroke, the form of this element has been shown to vary with flight mode. This study seeks to clarify the strength of this form-function relationship through the use of eigenshape morphometric analysis coupled with recently developed phylogenetic comparative methods (PCMs), including phylogenetic Flexible Discriminant Analysis (pFDA). Additionally, the morphospace derived from the furculae of extant birds is used to shed light on possible flight adaptations of Mesozoic fossil taxa. While broad conclusions of earlier work are supported (U-shaped furculae are associated with soaring, strong anteroposterior curvature with wing-propelled diving), correlations between form and function do not appear to be so clear-cut, likely due to the significantly larger dataset and wider spectrum of flight modes sampled here. Interclavicular angle is an even more powerful discriminator of flight mode than curvature, and is positively correlated with body size. With the exception of the close relatives of modern birds, the ornithuromorphs, Mesozoic taxa tend to occupy unique regions of morphospace, and thus may have either evolved unfamiliar flight styles or have arrived at similar styles through divergent musculoskeletal configurations.     

Presence in chicken tibialis anterior and extensor digitorum longus muscle spindles of reactive and unreactive intrafusal fibers after incubation with monoclonal antibodies against myosin heavy chains

Cross and longitudinal sections from the encapsulated portions of chicken tibialis anterior and extensor digitorum longus muscle spindles were examined to determine whether their intrafusal fibers were a structurally homogeneous or heterogeneous population. The techniques used were the histochemical actomyosin (mATPase) reaction, and fluorescence immunohistochemistry employing two monoclonal antibodies, CA-83 and CCM-52, that are specific for myosin heavy chains. After incubation with antibody CCM-52, intrafusal fibers fluoresced either strongly or weakly to moderately. Antibody CA-83 was even more selective. In addition to identifying the strongly reactive category, it clearly separated the remaining fibers into unreactive and moderately reactive groups. As a whole, after incubation for mATPase, pH 9.6 preincubation, unreactive fibers stained darker than strongly reactive fibers. Moreover, the cross-sectional area of the unreactive fibers was significantly larger than that of the strongly reactive fibers. In the average-size muscle spindle with six intrafusal fibers, there were four unreactive fibers and two strongly reactive fibers. In about one-third of the receptors examined, one moderately reactive fiber was present. Taken together, the data indicate that intrafusal fibers of chicken tibialis anterior and extensor digitorum longus muscles are not structurally homogeneous. The observed variations can be better explained in terms of different fiber types than of continuous gradients within one type of fiber.