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1.
During jumping or falling in humans and various other mammals, limb muscles are activated before landing, and the intensity and timing of this pre-landing activity are scaled to the expected impact. In this study, we test whether similarly tuned anticipatory muscle activity is present in hopping cane toads. Toads use their forelimbs for landing, and we analysed pre-landing electromyographic (EMG) timing and intensity in relation to hop distance for the m. coracoradialis and m. anconeus, which act antagonistically at the elbow, and are presumably important in stabilizing the forelimb during landing. In most cases, a significant, positive relationship between hop distance and pre-landing EMG intensity was found. Moreover, pre-landing activation timing of m. anconeus was tightly linked to when the forelimbs touched down at landing. Thus, like mammals, toads appear to gauge the timing and magnitude of their impending impact and activate elbow muscles accordingly. To our knowledge these data represent the first demonstration of tuned pre-landing muscle recruitment in anurans and raise questions about how important the visual, vestibular and/or proprioceptive systems are in mediating this response.  相似文献   

2.
Moles have modified thoracic limbs with hypertrophied pectoral girdle muscles that allow them to apply remarkably high lateral out‐forces during the power stroke when burrowing. To further understand the high force capabilities of mole forelimbs, architectural properties of the thoracic limb muscles were quantified in the Eastern mole (Scalopus aquaticus). Architectural properties measured included muscle mass, moment arm, belly length, fascicle length, and pennation angle, and these were used to provide estimates of maximum isometric force, joint torque, and power. Measurements of muscle moment arms and limb lever lengths were additionally used to analyze the out‐force contributions of the major pectoral girdle muscles. Most muscles have relatively long fascicles and little‐to‐no pennation. The humeral abductor/rotators as a functional group are massive and are capable of relatively high force, power, and joint torque. Of this group, the bipennate m. teres major is the most massive and has the capacity to produce the highest force and joint torque to abduct and axially rotate the humerus. In general, the distal limb muscles are relatively small, but have the capacity for high force and mechanical work by fascicle shortening. The muscle architectural properties of the elbow extensors (e.g., m. triceps brachii) and carpal flexors (e.g., m. palmaris longus) are consistent with the function of these muscles to augment lateral out‐force application. The humeral abductor/rotators m. latissimus dorsi, m. teres major, m. pectoralis, and m. subscapularis are calculated to contribute 13.9 N to out‐force during the power stroke, and this force is applied in a ‘frontal’ plane causing abduction of the humerus about the sternoclavicular joint. Moles have several specializations of their digging apparatus that greatly enhance the application of out‐force, and these morphological features suggest convergence on limb form and burrowing function between New and Old World moles. J. Morphol. 274:1277–1287, 2013. © 2013 Wiley Periodicals, Inc.  相似文献   

3.
Homo floresiensis and the evolution of the hominin shoulder   总被引:1,自引:1,他引:0  
The holotype of Homo floresiensis, diminutive hominins with tiny brains living until 12,000 years ago on the island of Flores, is a partial skeleton (LB1) that includes a partial clavicle (LB1/5) and a nearly complete right humerus (LB1/50). Although the humerus appears fairly modern in most regards, it is remarkable in displaying only 110 degrees of humeral torsion, well below modern human average values. Assuming a modern human shoulder configuration, such a low degree of humeral torsion would result in a lateral set to the elbow. Such an elbow joint would function more nearly in a frontal than in a sagittal plane, and this is certainly not what anyone would have predicted for a tool-making Pleistocene hominin. We argue that Homo floresiensis probably did not have a modern human shoulder configuration: the clavicle was relatively short, and we suggest that the scapula was more protracted, resulting in a glenoid fossa that faced anteriorly rather than laterally. A posteriorly directed humeral head was therefore appropriate for maintaining a normally functioning elbow joint. Similar morphology in the Homo erectus Nariokotome boy (KNM-WT 15000) suggests that this shoulder configuration may represent a transitional stage in pectoral girdle evolution in the human lineage.  相似文献   

4.
The earliest eutherian mammals were small-bodied locomotor generalists with a forelimb morphology that strongly resembles that of extant rats. Understanding the kinematics of the humerus, radius, and ulna of extant rats can inform and constrain hypotheses concerning typical posture and mobility in early eutherian forelimbs. The locomotion of Rattus norvegicus has been extensively studied, but the three-dimensional kinematics of the bones themselves remains under-explored. Here, for the first time, we use markerless XROMM (Scientific Rotoscoping) to explore the three-dimensional long bone movements in Rattus norvegicus during a normal, symmetrical gait (walking). Our data show a basic kinematic profile that agrees with previous studies on rats and other small therians: rats maintain a crouched forelimb posture throughout the step cycle, and the ulna is confined to flexion/extension in a parasagittal plane. However, our three-dimensional data illuminate long-axis rotation (LAR) movements for both the humerus and the radius for the first time. Medial LAR of the humerus throughout stance maintains an adducted elbow with a caudally-facing olecranon process, which in turn maintains a cranially-directed manus orientation (pronation). The radius also shows significant LAR correlated with manus pronation and supination. Moreover, we report that elbow flexion and manus orientation are correlated in R. norvegicus: as the elbow angle becomes more acute, manus supination increases. Our data also suggest that manus pronation and orientation in R. norvegicus rely on a divided system of labor between the ulna and radius. Given that the radius follows the flexion and extension trajectory of the ulna, it must rotate at the elbow (on the capitulum) so that during the stance phase its distal end lies medial to ulna, ensuring that the manus remains pronated while the forelimb is supporting the body. We suggest that forelimb posture and kinematics in Juramaia, Eomaia, and other basal eutherians were grossly similar to those of rats, and that humerus and radius LAR may have always played a significant role in forelimb and manus posture in small eutherian mammals.  相似文献   

5.
The Indian Purple frog, Nasikabatrachus sahyadrensis, occupies a basal phylogenetic position among neobatrachian anurans and has a very unusual life history. Tadpoles have a large ventral oral sucker, which they use to cling to rocks in torrents, whereas metamorphs possess adaptations for life underground. The developmental changes that underlie these shifts in habits and habitats, and especially the internal remodeling of the cranial and postcranial skeleton, are unknown. Using a nearly complete metamorphic series from free-living larva to metamorph, we describe the postembryonic skeletal ontogeny of this ancient and unique monotypic lineage. The torrent-dwelling larva possesses a dorsoventrally flattened body and a head with tiny dorsal eyes, robust lower and upper jaw cartilages, well-developed trabecular horns, and a definable gap between the trabecular horns and the tip of the snout. Unlike tadpoles of many other frogs, those of Nasikabatrachus retain larval mouthparts into late metamorphic stages. This unusual feature enables the larvae to maintain their clinging habit until near the end of metamorphosis. The subsequent ontogenetic shift from clinging to digging is correlated with rapid morphological changes and behavioral modifications. Metamorphs are equipped with a shortened tibiafibula and ossified prehallical elements, which likely facilitate initial digging using the hind limbs. Subsequently, the frogs may shift to headfirst burrowing by using the wedge-shaped skull, anteriorly positioned pectoral girdle, well-developed humeral crests and spatula-shaped forelimbs. The transition from an aquatic life in torrents to a terrestrial life underground entails dramatic changes in skeletal morphology and function that represent an extreme in metamorphic remodeling. Our analysis enhances the scope for detailed comparative studies across anurans, a group renowned for the diversity of its life history strategies.  相似文献   

6.

Background

Theria (marsupials and placental mammals) are characterized by a highly mobile pectoral girdle in which the scapula has been shown to be an important propulsive element during locomotion. Shoulder function and kinematics are highly conservative during locomotion within quadrupedal therian mammals. In order to gain insight into the functional morphology and evolution of the pectoral girdle of the two-toed sloth we here analyze the anatomy and the three-dimensional (3D) pattern of shoulder kinematics during quadrupedal suspensory ('upside-down') locomotion.

Methods

We use scientific rotoscoping, a new, non-invasive, markerless approach for x-ray reconstruction of moving morphology (XROMM), to quantify in vivo the 3D movements of all constituent skeletal elements of the shoulder girdle. Additionally we use histologic staining to analyze the configuration of the sterno-clavicular articulation (SCA).

Results

Despite the inverse orientation of the body towards gravity, sloths display a 3D kinematic pattern and an orientation of the scapula relative to the thorax similar to pronograde claviculate mammalian species that differs from that of aclaviculate as well as brachiating mammals. Reduction of the relative length of the scapula alters its displacing effect on limb excursions. The configuration of the SCA maximizes mobility at this joint and demonstrates a tensile loading regime between thorax and limbs.

Conclusions

The morphological characteristics of the scapula and the SCA allow maximal mobility of the forelimb to facilitate effective locomotion within a discontinuous habitat. These evolutionary changes associated with the adoption of the suspensory posture emphasized humeral influence on forelimb motion, but allowed the retention of the plesiomorphic 3D kinematic pattern.  相似文献   

7.
This article describes the growth of the anuran pectoral girdle of Rana pipiens and compares skeletal development of the shoulder to that of long bones. The pectoral girdle chondrifies as two halves, each adjacent to a developing humerus. In each, the scapula and coracoid form as single foci of condensed chondrocytes that fuse, creating a cartilaginous glenoid bridge articulating with the humerus. Based on histological sections, both the dermal clavicle and cleithrum begin to ossify at approximately the same time as the periosteum forms around the endochondral bones. The dermal and endochondral bones of the girdle form immobile joints with neighboring girdle elements; however, the cellular organization and growth pattern of the scapula and coracoid closely resemble those of a long bone. Similar to a long bone epiphysis, distal margins of both endochondral elements have zones of hyaline, stratified, and hypertrophic cartilages. As a result, fused elements of the girdle can grow without altering the glenoid articulation with the humerus. Comparisons of anuran long bone and pectoral girdle growth suggest that different bones can have similar histology and development regardless of adult morphology.  相似文献   

8.
Although landing behaviour in anurans differs significantly among species, a take-off behaviour seems to be largely conserved in the evolution of frogs and toads. The ancestral mode of landing is hypothesized to involve the body crash-landing on the substrate, after which the anuran cycles the limbs forward and then backward to their resting position. The part of the body that first contacts the substrate may vary among taxa. The limbs and pectoral girdle muscles as well as those of the caudopelvic region, involved with landing and take-off behaviours, are investigated. The existence of a relationship exists between myology and different take-off and landing behaviours is explored. The results suggest that most of the muscles involved in both take-off and landing are conserved morphologically, with only few differing, depending on the locomotor behaviour. Two muscles tend to be longer; the m. coracobrachialis, which is involved with landing, and the m. coccygeosacralis, related to the take-off.  相似文献   

9.
The pectoral girdle and forelimb of the Late Triassic drepanosauromorph reptile Megalancosaurus are redescribed and their function reinterpreted. The whole skeleton of this diapsid is highly specialised for arboreal life, and also the peculiarities of the shoulder girdle and forelimb were interpreted as adaptations for a limb-based locomotion using gap-bridging to move from one support to another, as in chameleons. Re-examination of the pectoral girdle and forelimb revealed the presence of clavicles fused into a furcula-like structure, a saddle-shaped glenoid and a tight connection between the radius and ulna that strengthened the forearm but hindered pronation and supination movements at that joint. The new information plus a reconstruction of the pectoral and forelimb musculature suggests that the forelimb was also specialised for grasping and raking in addition to climbing and thus prey capture may have been an important function for the forelimb. The new functional interpretation fits well with the overall body architecture of Megalancosaurus’ skeleton, suggesting that this reptile was an ambush predator that may have assumed a stable tripodal position, secured by the hooked tail and hind limbs, freeing its forelimbs to catch prey by sudden extension of the arm and firm grasping with the pincer-like digits.  相似文献   

10.
It has been hypothesized that fluctuating asymmetry (FA) may provide an indication of the functional importance of structures within an organism, with structures that more strongly impact fitness being more symmetric. Based on this idea, we predicted that for tetrapods in which the forelimbs and hindlimbs play an unequal role in locomotion, the less functionally important limb set should display higher levels of FA. We conducted a multispecies test of this hypothesis in anurans (frogs and toads), whose saltatory locomotor mode is powered by the hindlimbs. We also tested whether FA in the forelimbs, which play a more important role during landing, differed between families that differ in the degree of forelimb use in locomotion (Bufonidae vs. Ranidae). We calculated FA from the lengths of humeri and femora measured from disarticulated skeletal specimens of four anuran taxa (Bufonidae: Anaxyrus americanus, Rhinella marina; Ranidae: Lithobates catesbeianus, Lithobates clamitans). Our findings were consistent with the hypothesis that natural selection for increased locomotor performance may influence patterns of FA seen in vertebrate limbs, with all species displaying lower mean FA in the hindlimbs. More subtle functional roles between the forelimbs of bufonids and ranids, however, did not elicit different levels of FA.  相似文献   

11.
Henosferida from the Middle-Upper Jurassic of Western Gondwana is the most probable sister group for monotremes. They share the derived pretribosphenic structure of lower molars combined with the presumably absent protocone on the upper molars and the plesiomorphic retention of postdentary bones and pseudangular process of the lower jaw. In addition, the two groups share the dental formula with three molars and the position of the Meckel’s groove, which passes ventral to the mandibular foramen. In the course of subsequent evolution, monotremes acquired the mammalian middle ear with three auditory ossicles independently of therian mammals and multituberculates. Jurassic Laurasian Shuotheriidae are probably a sister group of the Gondwanian clade Henosferida + Monotremata. The Jurassic shuotheriid Pseudotribos shows a great plesiomorphic similarity to monotremes in the structure of the pectoral girdle, with a large interclavicle immovably connected to the clavicle. In the lineages leading to therian mammals and multituberculates, the pectoral girdle changed probably independently and in parallel in connection with the establishment of the parasagittal posture of the forelimbs (reduction of the interclavicle, mobile articulation of the interclavicle with clavicle, reduction of the procoracoid, and development of a supraspinous fossa of the scapula) and formation of the mammalian middle ear with three auditory ossicles.  相似文献   

12.
The latest works on iniopterygians question their monophyly when considering only the neurocranium of the two families (Sibyrhynchidae and Iniopterygidae), which have different conditions of preservation. Some of the synapomorphies of the Iniopterygia concern the pectoral girdle and fins. However, the anatomy of these different elements is still poorly known in this taxon. Here we describe in details three dimensionally preserved cartilages of the pectoral girdle and fins of the sibyrhynchid Iniopera sp. These structures have been extracted virtually from phosphatised nodules thanks to conventional and synchrotron microtomography, using absorption and phase contrast based techniques in the later case. The pectoral girdle of Iniopera sp. consists of three elements, which are, from dorsal to ventral, a paired suprascapular cartilage, a pair of robust scapulocoracoids and an unpaired intercoracoid cartilage. The scapular part of the scapulocoracoids is extremely reduced and the suprascapular cartilages link the scapulcoracoids to the rear of the neurocranium. These characters may be iniopterygian synapomorphies. Iniopterygians, stem and crown-holocephalans share a basipterygium that articulates with the pectoral girdle and bears an enlarged first pectoral fin radial. Posteriorly, the basipterygium articulates with either a well-defined metapterygium (in crown-holocephalans) or a metapterygial axis (in stem-holocephalans).  相似文献   

13.
14.
Two types of pectoral girdles occur among frogs. Arciferal girdles have overlapping epicoracoid cartilages; in firmisternal girdles the epicoracoid cartilages are fused along the ventral midline. Cineradiographic experiments of jumping frogs show that the epicoracoid cartilages of arciferal girdles move relative to each other at the time of landing. Recordings of landings on a force platform reveal that the pectoral girdle of frogs is loaded compressively through the glenoid. This loading regime coupled with differential mobility between firmisternal and arciferal girdles results in differences in stress distribution in the two girdles during landing. The patterns of stress distribution suggest that variation seen among frogs in other aspects of pectoral morphology in addition to the condition of the epicoracoid cartilages may be best understood when analysed from a biomechanical perspective.  相似文献   

15.
Zhu M  Yu X  Choo B  Qu Q  Jia L  Zhao W  Qiao T  Lu J 《PloS one》2012,7(4):e35103

Background

The pectoral and pelvic girdles support paired fins and limbs, and have transformed significantly in the diversification of gnathostomes or jawed vertebrates (including osteichthyans, chondrichthyans, acanthodians and placoderms). For instance, changes in the pectoral and pelvic girdles accompanied the transition of fins to limbs as some osteichthyans (a clade that contains the vast majority of vertebrates – bony fishes and tetrapods) ventured from aquatic to terrestrial environments. The fossil record shows that the pectoral girdles of early osteichthyans (e.g., Lophosteus, Andreolepis, Psarolepis and Guiyu) retained part of the primitive gnathostome pectoral girdle condition with spines and/or other dermal components. However, very little is known about the condition of the pelvic girdle in the earliest osteichthyans. Living osteichthyans, like chondrichthyans (cartilaginous fishes), have exclusively endoskeletal pelvic girdles, while dermal pelvic girdle components (plates and/or spines) have so far been found only in some extinct placoderms and acanthodians. Consequently, whether the pectoral and pelvic girdles are primitively similar in osteichthyans cannot be adequately evaluated, and phylogeny-based inferences regarding the primitive pelvic girdle condition in osteichthyans cannot be tested against available fossil evidence.

Methodology/Principal Findings

Here we report the first discovery of spine-bearing dermal pelvic girdles in early osteichthyans, based on a new articulated specimen of Guiyu oneiros from the Late Ludlow (Silurian) Kuanti Formation, Yunnan, as well as a re-examination of the previously described holotype. We also describe disarticulated pelvic girdles of Psarolepis romeri from the Lochkovian (Early Devonian) Xitun Formation, Yunnan, which resemble the previously reported pectoral girdles in having integrated dermal and endoskeletal components with polybasal fin articulation.

Conclusions/Significance

The new findings reveal hitherto unknown similarity in pectoral and pelvic girdles among early osteichthyans, and provide critical information for studying the evolution of pelvic girdles in osteichthyans and other gnathostomes.  相似文献   

16.
Albian sedimentary successions of northwestern Canada have yielded a diverse assemblage of Mesozoic marine vertebrates, and ichthyosaurs form an important component of these faunas. Here, we describe a partial postcranial skeleton of a small (estimated at less than 3 m total body length) ichthyosaur from the Wabiskaw Member of the Clearwater Formation (lowermost Albian). The semi-articulated specimen includes much of the presacral vertebral column, dorsal ribs and gastralia. Most significantly, it possesses an articulated pectoral girdle and humerus, and also preserves the pelvic girdle, allowing new insights into girdle evolution in ichthyosaurs. Whereas both sets of girdles are thought to exhibit large amounts of intraspecific variation, the pectoral girdle of ophthalmosaurids appears to evolve very slowly, remaining essentially unchanged from the Middle Jurassic onwards. In contrast, the pelvic girdle shows taxonomically informative changes within Ophthalmosauridae. The variable and poorly known nature of girdle morphology in Cretaceous ichthyosaurs precludes generic referral of the specimen.  相似文献   

17.
Walking of Chrysemys has been studied by cinephotography and x-rays. The lateral sequence, diagonal couplet gait, limb support sequence, and wide track provide great stability, yet a slight pitch and roll cause some plastral drag. Velocity ranges from 28 mm to 51 mm/second, and fluctuates within a stride. Limb movements and structure resemble those of other ectotherms, but incorporate modifications reflecting the animal's short, broad trunk encased in a shell and carried close to the ground. The triradiate pectoral girdle so articulates with the shell as to act as a truss for weight transfer to the ground. Girdle rotation increases the efficiency of the girdle as a truss, and contributes to locomotor efficiency. The glenoid cavities are more than twice as far apart as the acetabula, so a thrust from the pectoral girdle has less propulsive efficiency on the center of gravity than one from the acetabulum. The humerus and femur are protracted to a greater extent than in other ectotherms and their horizontal arcs of retraction are less. Rotation of these elements about their longitudinal axes contributes to the length of a stride and to foot placement and withdrawal. Differences in the movements of comparable segments of front and hind limbs correlate with differences in the width of the girdles, a crus longer than the antebrachium, and different capacities for joint rotation.  相似文献   

18.
The excursions of wing elements and the activity of eleven shoulder muscles were studied by cineradiography and electromyography in European starlings (Sturnus vulgaris) flying in a wind tunnel at speeds of 9–20 m s?1. At the beginning of downstroke the humerus is elevated 80–90° above horizontal, and both elbow and wrist are extended to 90° or less. During downstroke, protraction of the humerus (55°) remains constant; elbow and wrist are maximally extended (120° and 160°, respectively) as the humerus passes through a horizontal orientation. During the downstroke-upstroke transition humeral depression ceases (at about 20° below horizontal) and the humerus begins to retract. However, depression of the distal wing continues by rotation of the humerus and adduction of the carpometacarpus. Humeral retraction (to within about 30° of the body axis) is completed early in upstroke, accompanied by flexion of the elbow and carpometacarpus. Thereafter the humerus begins to protract as elevation continues. At mid-upstroke a rapid counterrotation of the humerus reorients the ventral surface of the wing to face laterad; extension of the elbow and carpometacarpus are initiated sequentially. The upstroke-downstroke transition is characterized by further extension of the elbow and carpometacarpus, and the completion of humeral protraction. Patterns of electromyographic activity primarily coincide with the transitional phases of the wingbeat cycle rather than being confined to downstroke or upstroke. Thus, the major downstroke muscles (pectoralis, coracobrachialis caudalis, sternocoracoideus, subscapularis, and humerotriceps) are activated in late upstroke to decelerate, extend, and reaccelerate the wing for the subsequent downstroke; electromyographic activity ends well before the downstroke is completed. Similarly, the upstroke muscles (supracoracoideus, deltoideus major) are activated in late downstroke to decelerate and then reaccelerate the wing into the upstroke; these muscles are deactivated by mid-upstroke. Only two muscles (scapulohumeralis caudalis, scapulotriceps) exhibit electromyographic activity exclusively during the downstroke. Starlings exhibit a functional partitioning of the two heads of the triceps (the humerotriceps acts with the pectoralis group, and does not overlap with the scapulotriceps). The biphasic pattern of the biceps brachii appears to correspond to this partitioning.  相似文献   

19.
The pectoral girdle is a unique skeletal element that underwent drastic morphological changes during its evolution, especially in association with the fin-to-limb transition. Comparative studies of its development are needed to gain a deeper understanding of its evolution. Transplantation experiments using the quail-chick chimeric system have revealed that not only lateral plate mesoderm but also somites contribute to the pectoral girdle in birds. Studies in mice and turtles also document somitic contributions to the pectoral girdle, but extirpation experiments in a salamander did not affect shoulder girdle development. Somitic contributions to the pectoral girdle therefore have been interpreted as a feature unique to amniotes. Here, we present a long-term fate map of single somites in the Mexican axolotl, based on transplantations of somites two to six from GFP-transgenic donors into wild-type hosts, as well as injections of fluorescein dextran into single somites. The results show a somitic derivation of the dorsal region of the suprascapula, demonstrating that somitic contributions to the pectoral girdle are not restricted to amniotes. Comparison with the few other species studied so far leads us to suggest a position-dependent origin of the pectoral girdle. We propose that embryonic origin is determined by the proximity of the developing pectoral girdle to the somites or to the lateral plate mesoderm, respectively. This position-dependent origin and the diversity of the anatomy of the pectoral girdle among vertebrates implies that the embryonic origin of the pectoral girdle is too variable to be useful for defining homologies or for phylogenetic analysis.  相似文献   

20.
The configuration of the pectoral girdle bones and muscles of numerous catfishes was studied in detail and compared with that of other siluriforms, as well as of other teleosts, described in the literature. The pectoral girdle of catfishes is composed of only three bones, which probably correspond to the posttemporo-supracleithrum (posttemporal + supracleithrum), scapulo-coracoid (scapula + coracoid), and cleithrum of other teleosts. These latter two bones constitute the place of origin of the pectoral girdle muscles. Two of these muscles are related to the movements of the pectoral fin. These two muscles correspond, very likely, to the abductor superficialis and to the adductor superficialis of other teleostean fishes. In relation to the pectoral spine (thickened first pectoral fin ray), it is usually moved by three well-developed muscles, which are probably homologous with the arrector ventralis, arrector dorsalis, and abductor profundus of nonsiluriform teleosts. The morphological diversity and the plesiomorphic configuration of these muscles, as well as of the other catfish pectoral girdle structures, are discussed.  相似文献   

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