Properties and function of extensible ligaments in the necks of turkeys (Meleagris gallopavo) and other birds

Highly extensible ligaments connect the neural spines of successive vertebrae in the proximal part of the neck in birds. Their elastic properties, and their lengths in different neck positions, have been investigated in experiments on turkeys. Their properties are very similar to those of the ligamentum nuchae of artiodactyls. They are stretched to large strains when the head is lowered but do not develop sufficient force to support the extended head and neck without the aid of muscles.     

Torsion and Bending in the Neck and Tail of Sauropod Dinosaurs and the Function of Cervical Ribs: Insights from Functional Morphology and Biomechanics

The long necks of sauropods have been subject to many studies regarding their posture and flexibility. Length of the neck varies among groups. Here, we investigate neck posture and morphology in several clades from a mechanical viewpoint. Emphasis is put on comparing sauropod necks and tails with structures in living archosaurs and mammals. Differences in the use made of necks and tails lead to clear-cut differences in the mechanical loads occurring in the same models. Ways of sustaining loads are identified by theoretical considerations. If the observed skeletal structures are suited to resist the estimated loading in a particular posture, this concordance is taken as an argument that this posture or movement was of importance during the life of the individual. Apart from the often-discussed bending in side view, we analyze the often overlooked torsion. Because torsional stresses in a homogenous element concentrate near the periphery, a cylindrical cross section gives greatest strength, and the direction of forces is oblique. In a vertebrate neck, during e.g. shaking the head and twisting the neck, oblique muscles, like the mm. scaleni, if activated unilaterally initiate movement, counterbalance the torsional moments and keep the joints between neck vertebrae in equilibrium. If activated bilaterally, these muscles keep the neck balanced in an energy-saving upright posture. The tendons of the mm. scaleni may have ossified as cervical ribs The long cervical ribs in brachiosaurids and mamenchisaurids seem to have limited flexibility, whereas the shorter cervical ribs in Diplodocidae allowed free movement. The tails of sauropods do not show pronounced adaptation to torsion, and seem to have been carried more or less in a horizontal, extended posture. In this respect, sauropod tails resemble the necks of herbivorous cursorial mammals. These analyses provide an improved understanding of neck use that will be extended to other sauropods in subsequent studies.     

The anatomical arrangement of muscle and tendon enhances limb versatility and locomotor performance

The arrangement of muscles and tendons has been studied in detail by anatomists, surgeons and biomechanists for over a century, and the energetics and mechanics of muscle contraction for almost as long. Investigation of how muscles function during locomotion and the relative length change in muscle fibres and the associated elastic tendon has, however, been more challenging. In recent years, novel in vivo measurement methods such as ultrasound and sonomicrometry have contributed to our understanding of the dynamics of the muscle tendon unit during locomotion. Here, we examine both published and new data to explore how muscles are arranged to deliver the wide repertoire of locomotor function and the trade-offs between performance and economy that result.     

Histology shows that elongated neck ribs in sauropod dinosaurs are ossified tendons

The histology of cervical ribs of Sauropoda reveals a primary bone tissue, which largely consists of longitudinally oriented mineralized collagen fibres, essentially the same tissue as found in ossified tendons. The absence of regular periosteal bone and the dominance of longitudinal fibres contradict the ventral bracing hypothesis (VBH) postulated for sauropod necks. The VBH predicts histologically primary periosteal bone with fibres oriented perpendicular to the rib long axis, indicative of connective tissue between overlapping hyperelongated cervical ribs. The transformation of the cervical ribs into ossified tendons makes the neck more flexible and implies that tension forces acted mainly along the length of the neck. This is contrary to the VBH, which requires compressive forces along the neck. Tension forces would allow important neck muscles to shift back to the trunk region, making the neck much lighter.     

Patterns of regional variation in the vertebral column of terrestrial salamanders

Regional variation in the vertebral column of several species of salamanders (families Ambystomatidae, Salamandridae and Plethodontidae) is analyzed. Measurements of three dimensions, centrum length, prezygapophyseal width, and transverse process length, provide the data. Ontogenetic, interspecific, intergeneric and interfamilial patterns of positional variation are diagrammed and discussed. Distinctive patterns of variation characterize the families, genera, and to a lesser extent, the species. The patterns of ambystomatid salamanders are the most generalized, and probably reflect derivation from a primitive ancestral stock. The most specialized conditions occur in the fully terrestrial plethodontids, a group generally considered to be highly derived. Data such as those presented here will aid in the identification of fossils. The patterns described have functional significance. For example, species which have an aquatic larval stage and which return to aquatic breeding sites have vertebrae which taper in length and width behind the pelvis. This is a feature associated with production of a traveling wave in the tail which is necessary for propulsion in water. Fully terrestrial species do not have a tapering column. In them, standing waves, such as occur in the trunk region of all species, typically occur in the tail. The caudal vertebrae of terrestrial species are rather uniform in dimensions for some distance, and the tail is cylindrical in form. Other functionally important features include the narrowing and shortening of some anterior vertebrae, associated with the development of a neck in some species with tongue feeding mechanisms. In contrast, species which use their heads as wedges during locomotion have broadened anterior vertebrae which serve as sites of origin for hypertrophied neck muscles.     

Reconstruction of the neck of Azhdarcho lancicollis and lifestyle of azhdarchids (Pterosauria, Azhdarchidae)

A computer reconstruction of isolated cervical vertebrae of Azhdarcho lancicollis from the Turonian of Uzbekistan allows three-dimensional model of the cervical region of the vertebral column of this animal. The relative length of cervical vertebrae (I + II < III < IV < V > VI > VII > VIII > IX) is the same as in pterodactyloids with short cervical vertebrae. An increase in neck length is provided mostly by the middle cervical vertebrae (IV–VI). In a neutral posture, the neck of azhdarchids was not straight, as often reconstructed, but S-shaped, with the maximum angles between the V–VI (20°), VI–VII (20°), and VIII–IX (17°) vertebrae. The feeding strategy of azhdarchids was probably similar to that of pelicans. In a search for prey, azhdarchids were soaring above the water surface of large inland or nearshore marine water bodies. Their prey (predominantly fish) was captured by the widely open mouth and fell into the throat sac, the presence of which is suggested by the spiral jaw joint. Prey was swallowed during the abrupt neck flexion in the posterior segment, which brought the head in an almost horizontal position. A storklike wading ecology for azhdarchids is less probable, because these clumsy on land animals were vulnerable to terrestrial predators.     

Morphological variations in bluegill, Lepomis macrochirus, with particular emphasis on growth-related changes

Morphological variations in bluegill, Lepomis macrochirus, including growth-related changes, sexual dimorphism and morphological differences between populations in different habitats, were examined in samples from three reservoirs in Kagawa, Japan. Body measurements demonstrated frequent growth-related proportional changes, particularly in body depth, body width, caudal peduncle length and head length, which all showed relative increases, whereas first dorsal fin length, caudal fin length and orbital diameter became relatively shorter. Body weight increased relatively with growth, with relative growth coefficients of 3.339–3.454 for regressions between total length and body weight. Such body weight increases were likely due to the relative increases in body depth and width, and caudal peduncle depth. Although counts of fin spines, fin rays and gill rakers did not change with growth, those of scales tended to increase. Males were significantly larger than females in body depth, caudal peduncle length, head length and body weight. Sexual dimorphism was also apparent in body coloration during the breeding season. In addition, a number of morphological differences were observed between individuals from different habitats.     

The effect of three-dimensional geometrical changes during adolescent growth on the biomechanics of a spinal motion segment

During adolescent growth, vertebrae and intervertebral discs undergo various geometrical changes. Although such changes in geometry are well known, their effects on spinal stiffness remains poorly understood. However, this understanding is essential in the treatment of spinal abnormalities during growth, such as scoliosis.A finite element model of an L3–L4 motion segment was developed, validated and applied to study the quantitative effects of changing geometry during adolescent growth on spinal stiffness in flexion, extension, lateral bending and axial rotation. Height, width and depth of the vertebrae and intervertebral disc were varied, as were the width of the transverse processes, the length of the spinous process, the size of the nucleus, facet joint areas and ligament size. These variations were based on average growth data for girls, as reported in literature.Overall, adolescent growth increases the stiffness with 36% (lateral bending and extension) to 44% (flexion). Two thirds of this increase occurs between 10 and 14 years of age and the last third between 14 years of age and maturity.Although the height is the largest geometrical change during adolescent growth, its effect on the biomechanics is small. The depth increase of the disc and vertebrae significantly affects the stiffness in all directions, while the width increase mainly affects the lateral bending stiffness. Hence, when analysing the biomechanics of the growing adolescent spine (for instance in scoliosis research), the inclusion of depth and width changes, in addition to the usually implemented height change, is essential.     

Interrelation between ischium,thigh extending muscles and locomotion in some primates

The relative length of the ischium varies according to the species' locomotory behavior, and is useful in estimation of the total relative weight of all the muscles that arise from it. There are three groups of muscles which are involved in thigh extension. Each of these muscles have other function(s) as well. When thigh extending muscles are well developed in a locomotory category or species, it is not only as a result of a response to their function of thigh extension, but also their other functions. In the macaques and guereza, the relative weight of the thigh extending muscles is high due to the massive hamstrings, particularly thebiceps femoris, and the ischium is relatively long. In the chimpanzee, because theadductor magnus muscle is well developed, the relative weight of the thigh extending muscles is high, and the ischium relatively long. In the gibbon, the relative weight of the thigh extending muscles is medium due to the feeble hamstrings and the heavyadductor magnus muscle, and the ischium is of medium relative length. The relative weight of the thigh extending muscles in the slow loris is close to medium with none of the muscles being more dominant than the others, and having ischium of somewhat medium relative length. In the grand and lesser bush babies, the relative weight of the thigh extending muscles and the relative length of the ischium is medium and small respectively, with none of the groups dominating the others, although they, particularly the lesser bush baby, have a massivesemimembranosus. Discussing the ischium in relation with the hamstring muscles alone is not satisfactory, since the ischium serves the thigh extending muscles, includingadductor magnus andquadratus femoris ventralis, attached on it as a lever. A long lever, in these cases the ischium, is an indication of the relatively heavy muscles that originate from it.     

Four forearm flexor muscles of the horse, Equus caballus: anatomy and histochemistry.

Two of the forearm flexors of the horse, the superficial and deep digital flexor muscles, are critical to support the digital and fetlock joints, exhibit differing insertions, and are passively supported by the proximal and distal check ligaments, respectively. These two muscles differ in histochemical composition and architecture. The differences are correlated with the different stress levels transmitted through their tendons, and the different frequencies of clinical breakdown that have been reported. Both muscles contain type I and type IIa fibers. A few type IIb fibers occurred in the deep digital flexor. The superficial digital flexor contained approximately 56% type I fibers, extremely short muscle fibers, and extensive connective tissue investment. In contrast, the deep digital flexor had three muscle heads: ulnar, radial, and "long" and "short" regions of the humeral head. The "long" and "short" regions of the humeral head contained 33% and 44% type I fibers, respectively, fiber lengths three to four times as long as those in the superficial digital flexor, and relatively less connective tissue investment. Flexor carpi radialis and flexor carpi ulnaris compared most closely with the humeral head of the deep digital flexor. These data suggest a correlation of the unique architecture of superficial digital flexor with its proposed elastic storage properties during locomotion in horses, and an explanation for the frequent breakdown of the superficial digital flexor in athletic horses.     

A three-dimensional model of the human cervical spine for impact simulation

A three-dimensional analytical model of the cervical spine is described. The cervical vertebrae and the head are modeled as rigid bodies which are interconnected by deformable elements representing the intervertebral disks, facet joints, ligaments and muscles. A special pentahedral continuum element for representing the articular facets is described which effectively maintains stability of the cervical spine in both lateral and frontal plane accelerations, which is very difficult with multi-spring models of the facets. A simplified representation is used for the spine and body below the level of T1. The neck musculature is modeled by over 100 muscle elements representing 22 major muscle groups in the neck. The model has been validated for frontal and sideways impact accelerations by simulating published experimental data. Results are also presented to show the effects of the stretch reflex response on the dynamics of the head and neck under moderate acceleration.     

The mechanics of jumping by a dog (Canis familiaris)

A force platform has been used to obtain records of the forces exerted on the ground by an Alsatian dog, during take-off for running long jumps and standing scale jumps. The records have been analysed in conjunction with cinematograph film, taken simultaneously, and anatomical data. Stresses in the principal muscles of the hind limb, and in the triceps, have been calculated and the values obtained are compared with the stresses found by other investigators in isometric experiments with excised mammal muscles. Stresses in certain tendons and bones have been calculated, and the values obtained are compared with published values for the strength of tendon and bone. Evidence is presented that the gastrocnemius and plantaris muscles behave, in take-off for a jump, essentially as passive elastic bodies. Most of the elastic energy is probably stored in their tendons. A tendency for distal limb muscles to be pinnate, with much shorter fibres than proximal limb muscles, is noted and discussed.     

棒花鱼形态特征的两性异形和雌性个体生育力

测定了棒花鱼(Abbottina rivularis)繁殖期形态特征包括体长、头长、头宽、头高、眼间距、鼻间距、背鳍基长、胸鳍长、胸鳍腹鳍间距、尾柄长、尾鳍长和体重的两性异形和雌性个体生育力。结果表明,雄性个体的数量显著多于雌性个体,雄性个体的体长显著大于雌性个体。特定体长的雌性个体的胸鳍腹鳍间距显著大于雄性个体,头长、头宽、头高、眼间距、鼻间距、背鳍基长、胸鳍长、尾柄长和尾鳍长显著小于雄性个体,雌雄两性体重不存在显著差异。棒花鱼的怀卵数量与体长和体重回归关系显著。偏相关分析显示,当控制第三者恒定时,怀卵数量与体长和体重呈正相关但不显著。棒花鱼存在个体大小和其他局部特征显著的两性异形,雌性个体主要通过腹腔容积的增加提高个体生育力。棒花鱼形态特征的两性异形是性选择和生育力选择共同作用的结果。     

Sagittal plane kinematics of the adult hyoid bone

The hyoid bone is a unique bone in the skeleton not articulated to any other bone. The hyoid muscles, which attach to the hyoid bone, may play a role in neck mechanics, but analysis of their function requires quantifying hyoid bone mechanics. The goal of this study was to obtain the detailed kinematics of the hyoid bone over a large range of flexion-extension motion using radiographs at 5 postures. The position of the hyoid bone in the sagittal plane was characterized with respect to head, jaw, and vertebral movements. Sex differences in hyoid kinematics were also investigated. We hypothesized that (1) the position of the hyoid bone in the sagittal plane is linearly correlated with motion of the head, jaw, and vertebrae, and (2) the hyoid position, size, and kinematics are sex-specific. We found that the hyoid bone X, Y, and angular position generally had strong linear correlations with the positions of the head, jaw, and the cervical vertebrae C1-C4. Hyoid X and angular position was also correlated to C5. Sex differences were found in some regressions of the hyoid bone with respect to C1-C5. The angular and linear measurements of the hyoid bone showed sex differences in absolute values, which were not evident after normalization by posture or neck size. Incorporating these results to neck models would enable accurate modeling of the hyoid muscles. This may have implications for analyzing the mechanics of the cervical spine, including loads on neck structures and implants.     

Architectural differences between the hamstring muscles

The purpose of this study was to understand the detailed architectural properties of the human hamstring muscles. The long (BFlh) and short (BFsh) head of biceps femoris, semimembranosus (SM) and semitendinosus (ST) muscles were dissected and removed from their origins in eight cadaveric specimens (age 67.8±4.3 years). Mean fiber length, sarcomere length, physiological cross-section area and pennation angle were measured. These data were then used to calculate a similarity index (δ) between pairs of muscles. The results indicated moderate similarity between BFlh and BFsh (δ=0.54) and between BFlh and SM (δ=0.35). In contrast, similarity was low between SM and ST (δ=0.98) and between BFlh and SM (δ=1.17). The fascicle length/muscle length ratio was higher for the ST (0.58) and BFsh (0.50) compared with the BFlh (0.27) and SM (0.22). There were, however, high inter-correlations between individual muscle architecture values, especially for muscle thickness and fascicle length data sets. Prediction of the whole hamstring architecture was achieved by combining data from all four muscles. These data show different designs of the hamstring muscles, especially between the SM and ST (medial) and BFlh and BFsh (lateral) muscles. Modeling the hamstrings as one muscle group by assuming uniform inter-muscular architecture yields less accurate representation of human hamstring muscle function.     

Size and shape in the phylum Phoronida

The lophophorate phylum Phoronida consists of about 13 species, which differ in body length and width, number of longitudinal muscles, lophophore geometry and number of lophophore tentacles. In absolute terms large species have a larger body width, more tentacles, more longitudinal muscles and greater coiling of the lophophore than small species. However, size and shape analyses suggest that with increasing size: (I) the body surface area to volume ratio increases because body length increases faster than body width; (2) the relative number longitudinal muscles decreases, and (3) the relative feeding surface area of the lophophore decreases because tentacle diameter is constant while tentacle number increases at the same rate as body length and tentacle length increases more slowly than tentacle number. Coiling and spiraling of the lophophore in large species may be an attempt to compensate for this last relationship. We suggest that the habits, mode of growth and feeding mechanism of phoronids constrain size-related changes in shape.     

Comparative morphometric study of the invasive pearl oyster <Emphasis Type="Italic">Pinctada radiata</Emphasis> along the Tunisian coastline

In order to study the relative growth of the pearl oyster Pinctada radiata in Tunisia, a total of 330 individuals of this species were collected from six sites along the Tunisian coastline. Quantitative measurements of collected oysters were conducted for shell height, shell length, shell width, hinge length, height and width of the nacreous part and wet weight. The size structure of the sampled populations was described and the relative growth between different morphometric characteristics was estimated as allometric growth lines for the six P. radiata samples. It appeared that the majority of examined samples were dominated by large individuals that exceed a shell height of 42 mm. The maximum size (100.5 mm), recorded in Bizerta lagoon, is bigger than that recorded elsewhere in particular in the Red Sea. Size distribution analysis also showed that the majority of P. radiata samples were dominated by two or more size groups. Differences of allometric regression were found between the examined samples for the tested relationships. Moreover, the Factorial Discriminant Analysis, coupled with Ascending Hierarchic Classification, classified the sub-populations according to geographic locations.     

黄颡鱼MSTN基因多态性及其与生长性状的相关性分析

肌肉生长抑制素基因(Myostatin,MSTN)属于转化生长因子β家族,其主要功能是负向调控肌肉的生长发育。采用PCR-SSCP技术对黄颡鱼MSTN基因进行单核苷酸多态性检测和分型,并与其生长性状进行关联分析。结果表明,在第一内含子部分检测到1个缺失位点和2个突变位点(T1003del、G1022A和T1063G),基因型分别为:AA、AB、CC、CD和DD;在第三外显子部分检测到1个突变位点(T132C),基因型分别为:EE和EF。关联性分析表明,AA基因型个体的全长、体长、体高、体厚、头长和体重显著大于CD和DD基因型(P<0.05),AA基因型雌性个体的全长、体长、体高、体厚、头长、尾柄高、尾柄厚和体重也显著大于DD基因型(P<0.05)。由此推断,AA基因型是影响雌性黄颡鱼生长性状的有利基因型,DD基因型是影响雌性黄颡鱼生长性状的不利基因型,可以尝试利用这两个位点对雌性黄颡鱼进行标记辅助选育。     

Sexual selection is not the origin of long necks in giraffes

The evolutionary origin of the long neck of giraffes is enigmatic. One theory (the 'sexual selection' theory) is that their shape evolved because males use their necks and heads to achieve sexual dominance. Support for this theory would be that males invest more in neck and head growth than do females. We have investigated this hypothesis in 17 male and 21 female giraffes with body masses ranging from juvenile to mature animals, by measuring head mass, neck mass, neck and leg length and the neck length to leg length ratio. We found no significant differences in any of these dimensions between males and females of the same mass, although mature males, whose body mass is significantly (50%) greater than that of mature females, do have significantly heavier (but not longer) necks and heavier heads than mature females. We conclude that morphological differences between males and females are minimal, that differences that do exist can be accounted for by the larger final mass of males and that sexual selection is not the origin of a long neck in giraffes.     

New Insights into Muscle Function during Pivot Feeding in Seahorses

Seahorses, pipefish and their syngnathiform relatives are considered unique amongst fishes in using elastic recoil of post-cranial tendons to pivot the head extremely quickly towards small crustacean prey. It is known that pipefish activate the epaxial muscles for a considerable time before striking, at which rotations of the head and the hyoid are temporarily prevented to allow energy storage in the epaxial tendons. Here, we studied the motor control of this system in seahorses using electromyographic recordings of the epaxial muscles and the sternohyoideus-hypaxial muscles with simultaneous high-speed video recordings of prey capture. In addition we present the results from a stimulation experiment including the muscle hypothesised to be responsible for the locking and triggering of pivot feeding in seahorses (m. adductor arcus palatini). Our data confirmed that the epaxial pre-activation pattern observed previously for pipefish also occurs in seahorses. Similar to the epaxials, the sternohyoideus-hypaxial muscle complex shows prolonged anticipatory activity. Although a considerable variation in displacements of the mouth via head rotation could be observed, it could not be demonstrated that seahorses have control over strike distance. In addition, we could not identify the source of the kinematic variability in the activation patterns of the associated muscles. Finally, the stimulation experiment supported the previously hypothesized role of the m. adductor arcus palatini as the trigger in this elastic recoil system. Our results show that pre-stressing of both the head elevators and the hyoid retractors is taking place. As pre-activation of the main muscles involved in pivot feeding has now been demonstrated for both seahorses and pipefish, this is probably a generalized trait of Syngnathidae.