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Muscular System

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Muscular System AccessScience from McGraw-Hill Education Page 1 of 29 www.accessscience.com Muscular system Contributed by: Warren F. Walker, Iain S. Young, John D. Altringham, Charles R. Noback Publication year: 2014 The muscular system consists of muscle cells, the contractile elements with the specialized property of exerting tension during contraction, and associated connective tissues. The three morphologic types of muscles are voluntary muscle, involuntary muscle, and cardiac muscle. The voluntary, striated, or skeletal muscles are involved with general posture and movements of the head, body, and limbs. The involuntary, nonstriated, or smooth muscles are the muscles of the walls of hollow organs of the digestive, circulatory, respiratory, and reproductive systems, and other visceral structures. Cardiac muscle is the intrinsic muscle tissue of the heart. See also: MUSCLE . In this article, the comparative embryology of the voluntary and involuntary muscles of the vertebrates will be outlined, followed by the comparative anatomy of the muscular system. Comparative Anatomy Phylogenetically speaking, muscles are very plastic organs. They have undergone considerable change during the evolution of vertebrates, correlated in large part with changes in the organisms’ environments and in their methods of support and locomotion. Establishment of homologies among muscles is not easy. Adult relationships can be misleading because muscles have subdivided during their evolution, and parts have migrated far from their original positions. Nerve supply is a more reliable criterion, because nerves have tended to follow the muscles through their evolutionary gymnastics, but often homologies cannot be established without recourse to embryonic development. Determination of the development of the thousands of individual muscles among the vertebrate classes is a monumental task. Comparison of muscles among vertebrates is greatly facilitated if the muscular system is subdivided into groups whose homology can be more easily established in the various classes. Muscle groups are particularly distinct in elasmobranchs and other primitive fishes, and they are generally defined on the basis of their embryonic origin in these animals. Two major groups of skeletal muscles are recognized, somatic (parietal) muscles, which develop from the myotomes, and branchiomeric muscles, which develop in the pharyngeal wall from lateral plate mesoderm. The somatic musculature is subdivided into axial muscles, which develop directly from the myotomes and lie along the longitudinal axis of the body, and appendicular muscles, which develop within the limb bud from mesoderm derived phylogenetically as buds from the myotomes. The vertebrate muscular system is the largest of the organ systems, making up 35–40% of the body weight in humans. The movement of vertebrates is accomplished exclusively by muscular action, and muscles play the AccessScience from McGraw-Hill Education Page 2 of 29 www.accessscience.com ImageFig. 1 Superficialof 1 muscles. ( a ) Cyclostome ( Petromyzon ). ( b ) Elasmobranch ( Squalus ). ( After H. W. Rand, The Chordates, Blakiston, 1950 ) major role in transporting materials within the body. Muscles also help to tie the bones of the skeleton together and supplement the skeleton in supporting the body against gravity. See also: SKELETAL SYSTEM . Axial musculature Most of the axial musculature is located along the back and flanks of the body, and this part is referred to as trunk musculature. But anteriorly the axial musculature is modified and assigned to other subgroups. Certain of the occipital and neck myotomes form the hypobranchial muscles, and the most anterior myotomes form the extrinsic ocular muscles. The trunk musculature of cyclostomes consists of a long series of segmental myomeres, each consisting of many longitudinal fibers attaching onto the myosepta ( Fig. 1 ). Each is folded in such a way as to appear approximately zigzag-shaped on the surface. The arrangement in jawed fishes is essentially the same, but the folding of the myomeres is more complex, and each is divided by a horizontal connective-tissue septum into dorsal (epaxial) and ventral (hypaxial) portions. A spinal nerve passes to each myomere, the dorsal ramus going to the epaxial portion and the ventral ramus to the hypaxial portion. This pattern of innervation persists in all higher vertebrates. Epaxial musculature. The epaxial musculature remains powerful in most cases. In amphibians, it consists of a group of medial and deep fibers that interlace the vertebrae, and a larger group of superficial fibers (dorsalis trunci). Segmentation is retained and undulations of the trunk and tail still play a role in the locomotion of many amphibians ( Fig. 2 ). AccessScience from McGraw-Hill Education Page 3 of 29 www.accessscience.com ImageFig. 2 Superficialof 2 muscles of three vertebrates, showing segmentation. ( a ) Amphibian ( Necturus ). ( b ) Reptile ( Sphenodon ). ( c ) Mammal ( Felis ). ( After H. W. Rand, The Chordates, Blakiston, 1950 ) In typical reptiles, the epaxial musculature is more complex. A medial and deep group of small, largely segmental muscles bind the vertebrae together and constitute the transversospinalis system; more laterally the musculature is arranged in two more extensive longitudinal groups, the longissimus dorsi, which lies dorsal to the transverse processes, and the iliocostalis, which is attached to the ribs. These three main divisions persist in mammals, but posteriorly there is a union of the iliocostalis, longissimus, and sometimes the more superficial part of the transversospinalis system to form a powerful erector spinae (sacropinalis) complex that helps to support the vertebral column. In mammals, the body is held off the ground by the limbs; thus the backbone is sometimes compared to a girder supported anteriorly and posteriorly by AccessScience from McGraw-Hill Education Page 4 of 29 www.accessscience.com pillars. Much of the epaxial musculature functions as tie members resisting tension stresses along this girder. Anteriorly there is a cleavage of the epaxial divisions into a host of muscles associated with the complex head and neck movements. In birds, the epaxial musculature in the trunk is greatly reduced, correlated with a fusion of many of the trunk vertebrae. Hypaxial musculature. The hypaxial musculature of tetrapods can be subdivided into three groups: (1) a subvertebral (hyposkeletal) group located ventral to the transverse processes and lateral to the centra of the vertebrae, (2) the flank muscles forming the lateral part of the body wall, and (3) the ventral abdominal muscles located on each side of the midventral line. The subvertebral musculature assists the epaxial muscles in the support and movement of the vertebral column. In mammals, it consists of longitudinal bundles—the longus colli in the neck and the anterior thorax, the quadratus lumborum, and psoas minor more posteriorly. Most of the flank musculature takes the form of broad, thin sheets of muscle that form much of the body wall and support the viscera. The ancestral, segmental nature of this musculature is retained throughout the trunk in salamanders, but is lost in higher tetrapods except in those parts of the trunk where ribs are well developed (Fig. 2). Three layers can be distinguished in the abdominal region of most tetrapods: a superficial external oblique, whose fibers extend caudally and ventrally; an internal oblique with fibers at right angles to the preceding; and a deep transversus abdominis. This pattern is much the same in the costal region, external intercostals, internal intercostals, and a reduced transversus thoracis being present in mammals. In reptiles, the pattern is more complex; the external layer is represented by supracostals, external intercostals, and sometimes a subcutaneous muscle. Respiratory movements of reptiles and birds are accomplished by the costal and abdominal muscles described above, but in mammals, which have a higher metabolic rate, additional respiratory muscles have evolved from the hypaxial muscles: the diaphragm (a derivative of cervical myotomes), serratus dorsalis, scalenes, and transversus costarum. See also: RESPIRATORY SYSTEM . Other parts of the hypaxial flank musculature have gained an attachment to the pectoral girdle where they help to transfer body weight from the vertebral column to the girdle and appendage and help to regulate the movement of the girdle. Only a few muscles of this type, the thoracoscapularis and levator scapulae, for example, are present in primitive tetrapods such as salamanders, and the body is not held far off the ground. In mammals, however, this group includes such large and powerful muscles as the serratus ventralis, rhomboideus, and levator scapulae ventralis. In the pelvic region of tetrapods, weight is transferred to the appendage directly across the sacroiliac joint and not by muscles. AccessScience from McGraw-Hill Education Page 5 of 29 www.accessscience.com The midventral hypaxial musculature in all tetrapods consists of the rectus abdominis, a longitudinal muscle on each side of the midline that extends from the pelvic region to the anterior part of the trunk (Fig. 2). It has evolved from the oblique flank muscles and in some salamanders remains closely associated with them. Transverse tendinous inscriptions are often present and are believed to represent persistent myosepta. Hypobranchial musculature. The hypobranchial musculature extends from the pectoral girdle forward along the ventral surface of the neck
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