The Skeletal System
The supporting tissues of animals which often serve to protect the body, or parts of it, and play an important role in the animal's physiology.
Skeletons can be divided into two main types based on the relative position of the skeletal tissues. When these tissues are located external to the soft parts, the animal is said to have an exoskeleton. If they occur deep within the body, they form an endoskeleton. All vertebrate animals possess an endoskeleton, but most also have components that are exoskeletal in origin. Invertebrate skeletons, however, show far more variation in position, morphology, and materials used to construct them.
The vertebrate endoskeleton is usually constructed of bone and cartilage; only certain fishes have skeletons that lack bone. In addition to an endoskeleton, many species possess distinct exoskeletal structures made of bone or horny materials. This dermal skeleton provides support and protection at the body surface.
Various structural components make up the human skeleton, including collagen, three different types of cartilage (hyaline, fibrocartilage, and elastic), and a variety of bone types (woven, lamellar, trabecular, and plexiform). See also Bone; Collagen; Connective tissue.
The vertebrate skeleton consists of the axial skeleton (skull, vertebral column, and associated structures) and the appendicular skeleton (limbs or appendages). The basic plan for vertebrates is similar, although large variations occur in relation to functional demands placed on the skeleton.
The axial skeleton supports and protects the organs of the head, neck, and torso, and in humans it comprises the skull, ear ossicles, hyoid bone, vertebral column, and rib cage.
The adult human skull consists of eight bones which form the cranium, or braincase, and 13 facial bones that support the eyes, nose, and jaws. There are also three small, paired ear ossicles—the malleus, incus, and stapes—within a cavity in the temporal bone. The total of 27 bones represents a large reduction in skull elements during the course of vertebrate evolution. The three components of the skull are the neurocranium, dermatocranium, and visceral cranium. See also Ear (vertebrate).
The brain and certain sense organs are protected by the neurocranium. All vertebrate neurocrania develop similarly, starting as ethmoid and basal cartilages beneath the brain, and as capsules partially enclosing the tissues that eventually form the olfactory, otic, and optic sense organs. Further development produces cartilaginous walls around the brain. Passages (foramina) through the cartilages are left open for cranial nerves and blood vessels. Endochondral ossification from four major centers follows in all vertebrates, except the cartilaginous fishes.
The visceral skeleton, the skeleton of the pharyngeal arches, is demonstrated in a general form by the elasmobranch fishes, where all the elements are cartilaginous and support the jaws and the gills. The mandibular (first) arch consists of two elements on each side of the body: the palatoquadrates dorsally, which form the upper jaw, and Meckel's cartilages, which join ventrally to form the lower jaw. The hyoid (second) arch has paired dorsal hyomandibular cartilages and lateral, gill-bearing ceratohyals. This jaw mechanism attaches to the neurocranium for support. In all jawed vertebrates except mammals, an articulation between the posterior ends of the palatoquadrate and Meckel's cartilages occurs between the upper and lower jaws. The bony fishes have elaborated on the primitive condition, where the upper jaw was fused to the skull and the lower jaw or mandible could move only in the manner of a simple hinge. Teleosts are able to protrude the upper and lower jaws. In the course of mammalian evolution, the dentary of the lower jaw enlarged and a ramus expanded upward in the temporal fossa. This eventually formed an articulation with the squamosal of the skull. With the freeing of the articular bone and the quadrate from their function in jaw articulation, they became ear ossicles in conjunction with the columella, that is, a skeletal rod that formed the first ear ossicle. The remaining visceral skeleton has evolved from jaw and gill structures in the fishes to become an attachment site for tongue muscles and to support the vocal cords in tetrapods. See also Mammalia.
The vertebral column is an endoskeletal segmented rod of mesodermal origin. It provides protection to the spinal cord, sites for muscle attachment, flexibility, and support, particularly in land-based tetrapods where it has to support the weight of the body. Hard, spool-shaped bony vertebrae alternate with tough but pliable intervertebral discs. Each typical vertebral body (centrum) has a bony neural arch extending dorsally. The spinal cord runs through these arches, and spinal nerves emerge through spaces. Bony processes and spines project from the vertebrae for the attachment of muscles and ligaments. Synovial articulations between adjacent vertebrae effectively limit and define the range of vertebral motion.
Vertebral morphology differs along the length of the column. There are two recognized regions in fishes (trunk and caudal) and five in mammals (cervical, thoracic, lumbar, sacral, and caudal), reflecting regional specializations linked to function. Humans have seven cervical, twelve thoracic, five lumbar, five (fused) sacral, and four coccygeal vertebrae. Most amphibians, reptiles, and mammals have seven cervical vertebrae regardless of neck length, whereas the number is variable in birds. Specific modification to the first two cervical vertebrae in most reptiles, birds, and mammals gives the head extra mobility. The presence of large ribs in the thoracic region often limits spinal flexibility. In typical tetrapods, the sacral region is usually modified for support of the pelvic girdle, while the number of caudal vertebrae varies greatly (from 0 to 50) between and within animal groups. See also Vertebra.
Sternum and ribs
Jawed fishes have ribs that help maintain the rigidity and support of the coelomic cavity. These ribs typically follow the connective tissue septa that divide successive muscle groups. In the caudal region, they are often small paired ventral ribs, fused on the midline to form the haemal arches. Ancestral tetrapods had ribs on all vertebrae, and their lengths varied between the vertebral regions. Modern amphibia (frogs and toads) have few thoracic ribs, and these are much reduced and never meet ventrally. Reptiles have varied rib arrangements, ranging from snakes with ribs on each vertebra (important for locomotor requirements) to turtles with only eight ribs which are fused to the inside of the carapace. Flying birds and penguins have a greatly enlarged sternum that links the ribs ventrally. In humans there are twelve pairs of ribs which form a strong but movable cage encompassing the heart and lungs.
This section of the skeletal system comprises the pectoral and pelvic limb girdles and bones of the free appendages. The girdles provide a supporting base onto which the usually mobile limbs attach.
The pectoral girdle has both endoskeletal and dermal components. The dermal components are derived from postopercular dermal armor of primitive fishes, and are represented by the clavicles and interclavicles in modern vertebrates, except where they are secondarily lost. Endochondral bone forms the scapula. In fishes, the main component of the girdle (the cleithrum) is anchored to the skull by other bony elements. Increased mobility of the girdle is seen in amphibia as it becomes independent of the skull. Further development and skeletal reduction have resulted in a wide range of morphologies, culminating in the paired clavicles and scapulae of mammals.
Birds have fused their paired clavicles and single interclavicle to form the wishbone or furcula. Clavicles have disappeared in certain groups of bounding mammals to allow greater movement of the scapula. Although humans, and most other mammals, have a coracoid process on the scapula, other tetrapods typically have a separate coracoid bracing the scapula against the sternum and forming part of the glenoid fossa.
The pelvic girdle forms by endochondral ossification, that is, the conversion of cartilage into bone. In the fishes, it is a small structure embedded in the body wall musculature just anterior to the cloaca. Each half of the girdle provides an anchor and articulation point for the pelvic fins. In tetrapods, the girdle attaches to the vertebral column to increase its stability and assist in the support of body weight and locomotor forces. Humans, like all other tetrapods, have a bilaterally symmetrical pelvic girdle, each half of which is formed from three fused bones: the ischium, ilium, and pubis. A part of each of these elements forms the acetabulum, the socket-shaped component of the hip joint, that articulates with the femoral head.
All urogenital and digestive products have to pass through the pelvic outlet. This accounts for the pelvic sexual dimorphism seen in most mammals, where the pelvic opening is broader in females, because of the physical demands of pregnancy and parturition. In birds (with the exception of the ostrich and the rhea), both sexes have an open pelvic girdle, a condition also found in female megachiropteran bats (flying foxes), gophers, and mole-rats.
Paired fins and tetrapod limbs
Paired fins in fishes come in different forms, but all are involved in locomotion. In the simplest form they are fairly rigid and extend from the body, functioning as stabilizers, but they are also capable of acting like a wing to produce lift as in sharks. In many fishes, the pectoral fins have narrow bases and are highly maneuverable as steering fins for low-speed locomotion. In addition, some fishes use their pectoral and pelvic fins to walk on the river bed, while others have greatly enlarged pectoral fins that take over as the main propulsive structures.
The basic mammalian pectoral limb consists of the humerus, radius, ulna, carpals, five metacarpals, and fourteen phalanges; and the pelvic limb consists of the femur, tibia, fibula, tarsal, five metatarsals, and fourteen phalanges. A typical bird pelvic limb consists of a femur, tibiotarsus (formed by fusion of the tibia with the proximal row of tarsal bones), fibula, and tarsometatarsus (formed by fusion of metatarsals II–IV), metatarsal I, and four digits (each consisting of two to five phalanges).
In biology, the skeleton or skeletal system is the biological system providing physical support in living organisms. (By extension, non-biological outline structures such as gantries or buildings may also acquire skeletons.)
Types and Classification
Skeletal systems are commonly divided into three types—external (an exoskeleton), internal (an endoskeleton), and fluid based (a hydrostatic skeleton), although hydrostatic skeletal systems may be classified separately from the other two, because they lack hardened support structures. An internal skeletal system consists of rigid or semi-rigid structures, within the body, moved by the muscular system. If the structures are mineralized or ossified, as they are in humans and other mammals, they are referred to as bones. Cartilage is another common component of skeletal systems, supporting and supplementing the skeleton. The human ear and nose are shaped by cartilage. Some organisms have a skeleton consisting entirely of cartilage and without any calcified bones at all, for example sharks. The bones or other rigid structures are connected by ligaments and connected to the muscular system via tendons.
Hydrostatic skeletons are similar to a water-filled balloon. Located internally in cnidarians (coral, jellyfish etc.) and annelids (leeches, earthworms etc.), among others, these animals can move by contracting the muscles surrounding the fluid-filled pouch, creating pressure within the pouch that causes movement. Animals such as earthworms use their hydrostatic skeletons to change their body shape, as they move forward, from long and thin to shorter and wider.