The axial skeleton forms the central axis of the body and includes the bones of the skull, ossicles of the middle ear, hyoid bone of the throat, vertebral column, and the thoracic cage (ribcage) (Figure 38.5). The function of the axial skeleton is to provide support and protection for the brain, the spinal cord, and the organs in the ventral body cavity. It provides a surface for the attachment of muscles that move the head, neck, and trunk, performs respiratory movements, and stabilizes parts of the appendicular skeleton.

Figure 38.5 The axial skeleton consists of the bones of the skull, ossicles of the middle ear, hyoid bone, vertebral column, and rib cage. (credit: modification of work by Mariana Ruiz Villareal)

The Skull

The bones of the skull support the structures of the face and protect the brain. The skull consists of 22 bones, which are divided into two categories: cranial bones and facial bones. The cranial bones are eight bones that form the cranial cavity, which encloses the brain and serves as an attachment site for the muscles of the head and neck. The eight cranial bones are the frontal bone, two parietal bones, two temporal bones, occipital bone, sphenoid bone, and the ethmoid bone. Although the bones developed separately in the embryo and fetus, in the adult, they are tightly fused with connective tissue and adjoining bones do not move (Figure 38.6).

Figure 38.6 The bones of the skull support the structures of the face and protect the brain. (credit: modification of work by Mariana Ruiz Villareal)

The auditory ossicles of the middle ear transmit sounds from the air as vibrations to the fluid-filled cochlea. The auditory ossicles consist of three bones each: the malleus, incus, and stapes. These are the smallest bones in the body and are unique to mammals.

Fourteen facial bones form the face, provide cavities for the sense organs (eyes, mouth, and nose), protect the entrances to the digestive and respiratory tracts, and serve as attachment points for facial muscles. The 14 facial bones are the nasal bones, the maxillary bones, zygomatic bones, palatine, vomer, lacrimal bones, the inferior nasal conchae, and the mandible. All of these bones occur in pairs except for the mandible and the vomer (Figure 38.7).

Figure 38.7 The cranial bones, including the frontal, parietal, and sphenoid bones, cover the top of the head. The facial bones of the skull form the face and provide cavities for the eyes, nose, and mouth.

Although it is not found in the skull, the hyoid bone is considered a component of the axial skeleton. The hyoid bone lies below the mandible in the front of the neck. It acts as a movable base for the tongue and is connected to muscles of the jaw, larynx, and tongue. The mandible articulates with the base of the skull. The mandible controls the opening to the airway and gut. In animals with teeth, the mandible brings the surfaces of the teeth in contact with the maxillary teeth.

The Vertebral Column

The vertebral column, or spinal column, surrounds and protects the spinal cord, supports the head, and acts as an attachment point for the ribs and muscles of the back and neck. The adult vertebral column comprises 26 bones: the 24 vertebrae, the sacrum, and the coccyx bones. In the adult, the sacrum is typically composed of five vertebrae that fuse into one. The coccyx is typically 3–4 vertebrae that fuse into one. Around the age of 70, the sacrum and the coccyx may fuse together. We begin life with approximately 33 vertebrae, but as we grow, several vertebrae fuse together. The adult vertebrae are further divided into the 7 cervical vertebrae, 12 thoracic vertebrae, and 5 lumbar vertebrae (Figure 38.8).

Figure 38.8 (a) The vertebral column consists of seven cervical vertebrae (C1–7) twelve thoracic vertebrae (Th1–12), five lumbar vertebrae (L1–5), the os sacrum, and the coccyx. (b) Spinal curves increase the strength and flexibility of the spine. (credit a: modification of work by Uwe Gille based on original work by Gray's Anatomy; credit b: modification of work by NCI, NIH)

Each vertebral body has a large hole in the center through which the nerves of the spinal cord pass. There is also a notch on each side through which the spinal nerves, which serve the body at that level, can exit from the spinal cord. The vertebral column is approximately 71 cm (28 inches) in adult male humans and is curved, which can be seen from a side view. The names of the spinal curves correspond to the region of the spine in which they occur. The thoracic and sacral curves are concave (curve inwards relative to the front of the body) and the cervical and lumbar curves are convex (curve outwards relative to the front of the body). The arched curvature of the vertebral column increases its strength and flexibility, allowing it to absorb shocks like a spring (Figure 38.8).

Intervertebral discs composed of fibrous cartilage lie between adjacent vertebral bodies from the second cervical vertebra to the sacrum. Each disc is part of a joint that allows for some movement of the spine and acts as a cushion to absorb shocks from movements such as walking and running. Intervertebral discs also act as ligaments to bind vertebrae together. The inner part of discs, the nucleus pulposus, hardens as people age and becomes less elastic. This loss of elasticity diminishes its ability to absorb shocks.

The Thoracic Cage

The thoracic cage, also known as the ribcage, is the skeleton of the chest, and consists of the ribs, sternum, thoracic vertebrae, and costal cartilages (Figure 38.9). The thoracic cage encloses and protects the organs of the thoracic cavity, including the heart and lungs. It also provides support for the shoulder girdles and upper limbs, and serves as the attachment point for the diaphragm, muscles of the back, chest, neck, and shoulders. Changes in the volume of the thorax enable breathing.

The sternum, or breastbone, is a long, flat bone located at the anterior of the chest. It is formed from three bones that fuse in the adult. The ribs are 12 pairs of long, curved bones that attach to the thoracic vertebrae and curve toward the front of the body, forming the ribcage. Costal cartilages connect the anterior ends of the ribs to the sternum, with the exception of rib pairs 11 and 12, which are free-floating ribs.

Figure 38.9 The thoracic cage, or rib cage, protects the heart and the lungs. (credit: modification of work by NCI, NIH)

Axial Skeleton is the central, vertical axis of the body, including the skull, vertebral column, and thoracic cage.

Axial Skeleton (80 bones)

Skull (28)

Cranial Bones

• Parietal (2)
• Temporal (2)
• Frontal (1)
• Occipital (1)
• Ethmoid (1)
• Sphenoid (1)

Facial Bones

• Maxilla (2)
• Zygomatic (2)
• Mandible (1)
• Nasal (2)
• Platine (2)
• Inferior nasal concha (2)
• Lacrimal (2)
• Vomer (1)

Auditory Ossicles

• Malleus (2)
• Incus (2)
• Stapes (2)

Hyoid (1)

Vertebral Column

• Cervical vertebrae (7)
• Thoracic vertebrae (12)
• Lumbar vertebrae (5)
• Sacrum (1)
• Coccyx (1)

Thoracic Cage

• Sternum (1)
• Ribs (24)

The skeleton is subdivided into two major divisions—the axial and appendicular. The axial skeleton forms the vertical, central axis of the body and includes all bones of the head, neck, chest, and back (image). It serves to protect the brain, spinal cord, heart, and lungs. It also serves as the attachment site for muscles that move the head, neck, and back, and for muscles that act across the shoulder and hip joints to move their corresponding limbs.

The axial skeleton of the adult consists of 80 bones, including the skull, the vertebral column, and the thoracic cage. The skull is formed by 22 bones. Also associated with the head are an additional seven bones, including the hyoid bone and the ear ossicles (three small bones found in each middle ear). The vertebral column consists of 24 bones, each called a vertebra, plus the sacrum and coccyx. The thoracic cage includes the 12 pairs of ribs, and the sternum, the flattened bone of the anterior chest.

Axial and Appendicular Skeleton

The axial skeleton begins to form during early embryonic development. However, growth, remodeling, and ossification (bone formation) continue for several decades after birth before the adult skeleton is fully formed. Knowledge of the developmental processes that give rise to the skeleton is important for understanding the abnormalities that may arise in skeletal structures.

Development of the Skull

During the third week of embryonic development, a rod-like structure called the notochord develops dorsally along the length of the embryo. The tissue overlying the notochord enlarges and forms the neural tube, which will give rise to the brain and spinal cord. By the fourth week, mesoderm tissue located on either side of the notochord thickens and separates into a repeating series of block-like tissue structures, each of which is called a somite. As the somites enlarge, each one will split into several parts. The most medial of these parts is called a sclerotome. The sclerotomes consist of an embryonic tissue called mesenchyme, which will give rise to the fibrous connective tissues, cartilages, and bones of the body.

The bones of the skull arise from mesenchyme during embryonic development in two different ways. The first mechanism produces the bones that form the top and sides of the brain case. This involves the local accumulation of mesenchymal cells at the site of the future bone. These cells then differentiate directly into bone producing cells, which form the skull bones through the process of intramembranous ossification. As the brain case bones grow in the fetal skull, they remain separated from each other by large areas of dense connective tissue, each of which is called a fontanelle (image). The fontanelles are the soft spots on an infant’s head. They are important during birth because these areas allow the skull to change shape as it squeezes through the birth canal. After birth, the fontanelles allow for continued growth and expansion of the skull as the brain enlarges. The largest fontanelle is located on the anterior head, at the junction of the frontal and parietal bones. The fontanelles decrease in size and disappear by age 2. However, the skull bones remained separated from each other at the sutures, which contain dense fibrous connective tissue that unites the adjacent bones. The connective tissue of the sutures allows for continued growth of the skull bones as the brain enlarges during childhood growth.

The second mechanism for bone development in the skull produces the facial bones and floor of the brain case. This also begins with the localized accumulation of mesenchymal cells. However, these cells differentiate into cartilage cells, which produce a hyaline cartilage model of the future bone. As this cartilage model grows, it is gradually converted into bone through the process of endochondral ossification. This is a slow process and the cartilage is not completely converted to bone until the skull achieves its full adult size.

At birth, the brain case and orbits of the skull are disproportionally large compared to the bones of the jaws and lower face. This reflects the relative underdevelopment of the maxilla and mandible, which lack teeth, and the small sizes of the paranasal sinuses and nasal cavity. During early childhood, the mastoid process enlarges, the two halves of the mandible and frontal bone fuse together to form single bones, and the paranasal sinuses enlarge. The jaws also expand as the teeth begin to appear. These changes all contribute to the rapid growth and enlargement of the face during childhood.

Development of the Vertebral Column and Thoracic cage

Development of the vertebrae begins with the accumulation of mesenchyme cells from each sclerotome around the notochord. These cells differentiate into a hyaline cartilage model for each vertebra, which then grow and eventually ossify into bone through the process of endochondral ossification. As the developing vertebrae grow, the notochord largely disappears. However, small areas of notochord tissue persist between the adjacent vertebrae and this contributes to the formation of each intervertebral disc.

The ribs and sternum also develop from mesenchyme. The ribs initially develop as part of the cartilage model for each vertebra, but in the thorax region, the rib portion separates from the vertebra by the eighth week. The cartilage model of the rib then ossifies, except for the anterior portion, which remains as the costal cartilage. The sternum initially forms as paired hyaline cartilage models on either side of the anterior midline, beginning during the fifth week of development. The cartilage models of the ribs become attached to the lateral sides of the developing sternum. Eventually, the two halves of the cartilaginous sternum fuse together along the midline and then ossify into bone. The manubrium and body of the sternum are converted into bone first, with the xiphoid process remaining as cartilage until late in life.