Vertebrate muscle systems
Major types of vertebrate muscles
In terms of its microscopic structure, the musculature of vertebrates is usually divided into three types: striated, cardiac, and smooth muscle. Smooth and cardiac muscle are under the control of the involuntary, or autonomic, nervous system. Striated muscle, on the other hand, is mainly under the control of the voluntary, or central, nervous system. Smooth and cardiac muscle are also similar in their development, being generally associated with the yolk sac. Striated muscle develops directly from the middle of the three embryonic layers, arising largely from the mesodermal somites (see below). In the adult, smooth and cardiac muscle are associated with organs or tubes (viscera), and striated (skeletal) muscle is associated with the bony or cartilaginous skeleton.
The two major divisions of the vertebrate musculature are the visceral musculature and the somatic musculature (the striated muscles of the body wall). Somatic musculature may be divided into appendicular, or limb, muscles and axial muscles. The axial muscles include the muscles of the tail, trunk, and eyeballs as well as a group of muscles called hypobranchial muscles, which separate and migrate from the others during development.
Basic pattern of development
The gastrula is the stage of embryonic development at which the embryo appears as three distinct layers of cells (the germ layers): the exterior ectoderm, the middle mesoderm, and the interior endoderm. The mesoderm differentiates to form most of the tissues, structures, and organs of the body. As the embryo lengthens, the mesoderm lying along the midline differentiates to form the notochord, a hollow cartilaginous nerve tube. In the adult the notochord contributes only to the structure of the vertebrae. The mesoderm lateral to this midline then divides into three parts that ultimately form the somites (which subsequently form the vertebrae, the somatic muscles, and the skin), the structures of the urogenital system, and the coelom and its associated structures.
Comparative anatomy
One problem in discussing the differences in arrangement of muscles between the various vertebrate groups is in deciding which muscles in each species are homologous—that is, which have the same evolutionary and developmental origin. The problem arises because the position and attachment of muscles change during evolution; a muscle lying in the same position and attached to the same bone or cartilage in one vertebrate may have different origins from those of another vertebrate species. Comparison of the development of muscles in the embryo of each species and of their nerve supply would probably give the best clues. No single method may be relied upon in all cases, and many different types of evidence are considered before the homology is decided upon.
Vertebrate muscles are given names derived from Latin according to their attachments. In this system the Latin names of the bony points of attachment are either joined, as in sternocleidomastoid, naming the human muscle that runs from the sternum and clavicle to the mastoid region of the skull, or they may be named for their form or their gross function. There are several standard terms that describe form and function. A muscle may have more than one point of origin; thus, it may be described as having, for example, two “heads,” as in biceps femoris (bi- for two, -ceps for heads, femoris meaning “of the femur”). It may be long, longus, or short, brevis. It may run transversely across a body segment, transversus, or obliquely, obliquus. It may lie close to the surface, superficialis, or deep, profundus. In describing function, flexors are muscles that tend to close the angle made by the two bones to which they are attached; extensors tend to increase the angle. Adductors pull a bone or cartilage closer to the axis of the body, or limb, while abductors pull away from the axis. Rotators turn one bone or cartilage with respect to another or with respect to the midline. Pronators turn the sole of the foot or the palm of the hand to face the ground, while the opposite function is performed by supinators. Constrictors and sphincters diminish the volume of spaces or the area of structures, and dilators increase them. The names of muscles in humans often have been applied to grossly equivalent muscles in animals, a situation that often causes confusion.
Jawless fishes
The earliest known vertebrates were jawless fishes of the class Agnatha, and their only living representatives are the cyclostomes—the lampreys and the hagfishes. The modern agnathans retain much of the general organization of the ancestral vertebrates, and, therefore, much of their musculature is relevant to an understanding of the evolution of muscles in more-advanced vertebrates.
The cyclostomes are free-swimming animals with prominent axial somatic musculature, which during contraction produces undulating waves that propagate from head to tail to produce thrust. The axial muscles form a single segmented mass running vertically down the side of the fish. These muscle segments, known as myomeres, consist of relatively short fibres that insert into septa of connective tissue, the myocommata, between the adjacent myomeres. There is only a rudimentary axial skeleton and no appendicular skeleton, so there are no limb muscles. The eyes of cyclostomes are degenerate structures, and the six axially derived muscles normally found associated with vertebrate eyes are diminished or absent. The branchiomeric muscles in cyclostomes are represented by a sheet of constrictors that compresses the gill pouches and helps the pumping mechanism draw water through the pharynx to the gills. Other muscles of the branchiomeric series have been modified for specialized feeding functions. The branchiomeric musculature of more primitive jawless fishes would probably have been similar for each of the gill arches.