This is a supportive connective tissue with a flexible rubbery matrix. The cells that secrete the matrix are called chondroblasts. The chondroblasts surround themselves with the matrix until they become trapped in little cavities called lacunae. Then they are called chondrocytes.

Cartilages do not have blood vessels except when transforming into bone. Nutrition and waste removal therefore depend on slow diffusion through the matrix from/to blood vessels in the dense irregular connective tissue membrane (perichondrium) surrounding it. Chondrocytes thus have slow rates of metabolism and cell division. Injured cartilages therefore heal slowly.

The matrix contains collagen fibres which give cartilages their flexibility and strength.

Cartilages are divided into three types: hyaline cartilage, elastic cartilage and fibrocartilage.

HYALINE CARTILAGE

Hyaline cartilage has a clear glassy blue-white appearance, arising from the invisible fineness of its collagen fibres. It is the most abundant type of cartilage. It covers the ends of bones where bones form joints (articular surfaces) and forms the cartilage rings of the respiratory tract (larynx, trachea and bronchi). It also forms the costal cartilages which attach the ribs to the sternum and the epiphyseal plates (growth plates) of bones and the embryonic skeleton. The tip of the nose is also supported by hyaline cartilage.

Hyaline cartilage provides support and some flexibility.

ELASTIC CARTILAGE

Elastic cartilage contains conspicuous elastic fibres among bundles of collagen fibres. It provides rigidity with even more flexibility than hyaline cartilage. It is found in the external ear, epiglottis and auditory tube.

FIBROCARTILAGE

Fibrocartilage contains coarse, readily visible bundles of collagen. It has more collagen fibres than hyaline cartilage. It is found in the discs between vertebrae, the symphysis pubis (joint between the public bones). It is somewhat flexible and capable of withstanding considerable pressure. There is no perichondrium around the fibrocartilage.

Endochondral Ossification

Endochondral ossification involves the replacement of hyaline cartilage with bony tissue. Most of the bones of the skeleton are formed in this manner. These bones are called endochondral bones. In this process, the future bones are first formed as hyaline cartilage models. During the third month after conception, the perichondrium that surrounds the hyaline cartilage "models" becomes infiltrated with blood vessels and osteoblasts and changes into a periosteum. The osteoblasts form a collar of compact bone around the diaphysis. At the same time, the cartilage in the center of the diaphysis begins to disintegrate. Osteoblasts penetrate the disintegrating cartilage and replace it with spongy bone. This forms a primary ossification center. Ossification continues from this center toward the ends of the bones. After spongy bone is formed in the diaphysis, osteoclasts break down the newly formed bone to open up the medullary cavity.

The cartilage in the epiphyses continues to grow so the developing bone increases in length. Later, usually after birth, secondary ossification centers form in the epiphyses. Ossification in the epiphyses is similar to that in the diaphysis except that the spongy bone is retained instead of being broken down to form a medullary cavity. When secondary ossification is complete, the hyaline cartilage is totally replaced by bone except in two areas. A region of hyaline cartilage remains over the surface of the epiphysis as the articular cartilage and another area of cartilage remains between the epiphysis and diaphysis. This is the epiphyseal plate or growth region.

Bone Growth

Bones grow in length at the epiphyseal plate by a process that is similar to endochondral ossification. The cartilage in the region of the epiphyseal plate next to the epiphysis continues to grow by mitosis. The chondrocytes, in the region next to the diaphysis, age and degenerate. Osteoblasts move in and ossify the matrix to form bone. This process continues throughout childhood and the adolescent years until the cartilage growth slows and finally stops. When cartilage growth ceases, usually in the early twenties, the epiphyseal plate completely ossifies so that only a thin epiphyseal line remains and the bones can no longer grow in length. Bone growth is under the influence of growth hormone from the anterior pituitary gland and sex hormones from the ovaries and testes.

Even though bones stop growing in length in early adulthood, they can continue to increase in thickness or diameter throughout life in response to stress from increased muscle activity or to weight. The increase in diameter is called appositional growth. Osteoblasts in the periosteum form compact bone around the external bone surface. At the same time, osteoclasts in the endosteum break down bone on the internal bone surface, around the medullary cavity. These two processes together increase the diameter of the bone and, at the same time, keep the bone from becoming excessively heavy and bulky.

As the name indicates, at a cartilaginous joint, the adjacent bones are united by cartilage, a tough but flexible type of connective tissue. These types of joints lack a joint cavity and involve bones that are joined together by either hyaline cartilage or fibrocartilage (Figure). There are two types of cartilaginous joints. A synchondrosis is a cartilaginous joint where the bones are joined by hyaline cartilage. Also classified as a synchondrosis are places where bone is united to a cartilage structure, such as between the anterior end of a rib and the costal cartilage of the thoracic cage. The second type of cartilaginous joint is a symphysis, where the bones are joined by fibrocartilage.

Synchondrosis

A synchondrosis (“joined by cartilage”) is a cartilaginous joint where bones are joined together by hyaline cartilage, or where bone is united to hyaline cartilage. A synchondrosis may be temporary or permanent. A temporary synchondrosis is the epiphyseal plate (growth plate) of a growing long bone. The epiphyseal plate is the region of growing hyaline cartilage that unites the diaphysis (shaft) of the bone to the epiphysis (end of the bone). Bone lengthening involves growth of the epiphyseal plate cartilage and its replacement by bone, which adds to the diaphysis. For many years during childhood growth, the rates of cartilage growth and bone formation are equal and thus the epiphyseal plate does not change in overall thickness as the bone lengthens. During the late teens and early 20s, growth of the cartilage slows and eventually stops. The epiphyseal plate is then completely replaced by bone, and the diaphysis and epiphysis portions of the bone fuse together to form a single adult bone. This fusion of the diaphysis and epiphysis is a synostosis. Once this occurs, bone lengthening ceases. For this reason, the epiphyseal plate is considered to be a temporary synchondrosis. Because cartilage is softer than bone tissue, injury to a growing long bone can damage the epiphyseal plate cartilage, thus stopping bone growth and preventing additional bone lengthening.

Growing layers of cartilage also form synchondroses that join together the ilium, ischium, and pubic portions of the hip bone during childhood and adolescence. When body growth stops, the cartilage disappears and is replaced by bone, forming synostoses and fusing the bony components together into the single hip bone of the adult. Similarly, synostoses unite the sacral vertebrae that fuse together to form the adult sacrum.

Examples of permanent synchondroses are found in the thoracic cage. One example is the first sternocostal joint, where the first rib is anchored to the manubrium by its costal cartilage. (The articulations of the remaining costal cartilages to the sternum are all synovial joints.) Additional synchondroses are formed where the anterior end of the other 11 ribs is joined to its costal cartilage. Unlike the temporary synchondroses of the epiphyseal plate, these permanent synchondroses retain their hyaline cartilage and thus do not ossify with age. Due to the lack of movement between the bone and cartilage, both temporary and permanent synchondroses are functionally classified as a synarthrosis.

Symphysis

A cartilaginous joint where the bones are joined by fibrocartilage is called a symphysis (“growing together”). Fibrocartilage is very strong because it contains numerous bundles of thick collagen fibers, thus giving it a much greater ability to resist pulling and bending forces when compared with hyaline cartilage. This gives symphyses the ability to strongly unite the adjacent bones, but can still allow for limited movement to occur. Thus, a symphysis is functionally classified as an amphiarthrosis.

The gap separating the bones at a symphysis may be narrow or wide. Examples in which the gap between the bones is narrow include the pubic symphysis and the manubriosternal joint. At the pubic symphysis, the pubic portions of the right and left hip bones of the pelvis are joined together by fibrocartilage across a narrow gap. Similarly, at the manubriosternal joint, fibrocartilage unites the manubrium and body portions of the sternum.

The intervertebral symphysis is a wide symphysis located between the bodies of adjacent vertebrae of the vertebral column. Here a thick pad of fibrocartilage called an intervertebral disc strongly unites the adjacent vertebrae by filling the gap between them. The width of the intervertebral symphysis is important because it allows for small movements between the adjacent vertebrae. In addition, the thick intervertebral disc provides cushioning between the vertebrae, which is important when carrying heavy objects or during high-impact activities such as running or jumping.

Review

There are two types of cartilaginous joints. A synchondrosis is formed when the adjacent bones are united by hyaline cartilage. A temporary synchondrosis is formed by the epiphyseal plate of a growing long bone, which is lost when the epiphyseal plate ossifies as the bone reaches maturity. The synchondrosis is thus replaced by a synostosis. Permanent synchondroses that do not ossify are found at the first sternocostal joint and between the anterior ends of the bony ribs and the junction with their costal cartilage. A symphysis is where the bones are joined by fibrocartilage and the gap between the bones may be narrow or wide. A narrow symphysis is found at the manubriosternal joint and at the pubic symphysis. A wide symphysis is the intervertebral symphysis in which the bodies of adjacent vertebrae are united by an intervertebral disc.

Cartilage is the tough but flexible tissue that covers the ends of your bones at a joint. It also gives shape and support to other parts of your body, such as your ears, nose and windpipe. Healthy cartilage helps you move by allowing your bones to glide over each other. It also protects bones by preventing them from rubbing against each other.

Injured, inflamed, or damaged cartilage can cause symptoms such as pain and limited movement. It can also lead to joint damage and deformity. Causes of cartilage problems include

• Tears and injuries, such as sports injuries
• Genetic factors
• Other disorders, such as some types of arthritis

Osteoarthritis results from breakdown of cartilage.

Cartilage Templates

Bone is a replacement tissue; that is, it uses a model tissue on which to lay down its mineral matrix. For skeletal development, the most common template is cartilage. During fetal development, a framework is laid down that determines where bones will form. This framework is a flexible, semi-solid matrix produced by chondroblasts and consists of hyaluronic acid, chondroitin sulfate, collagen fibers, and water. As the matrix surrounds and isolates chondroblasts, they are called chondrocytes. Unlike most connective tissues, cartilage is avascular, meaning that it has no blood vessels supplying nutrients and removing metabolic wastes. All of these functions are carried on by diffusion through the matrix. This is why damaged cartilage does not repair itself as readily as most tissues do.

Throughout fetal development and into childhood growth and development, bone forms on the cartilaginous matrix. By the time a fetus is born, most of the cartilage has been replaced with bone. Some additional cartilage will be replaced throughout childhood, and some cartilage remains in the adult skeleton.

Intramembranous Ossification

During intramembranous ossification, compact and spongy bone develops directly from sheets of mesenchymal (undifferentiated) connective tissue. The flat bones of the face, most of the cranial bones, and the clavicles (collarbones) are formed via intramembranous ossification.

The process begins when mesenchymal cells in the embryonic skeleton gather together and begin to differentiate into specialized cells (Figure 6.16a). Some of these cells will differentiate into capillaries, while others will become osteogenic cells and then osteoblasts. Although they will ultimately be spread out by the formation of bone tissue, early osteoblasts appear in a cluster called an ossification center.

The osteoblasts secrete osteoid, uncalcified matrix, which calcifies (hardens) within a few days as mineral salts are deposited on it, thereby entrapping the osteoblasts within. Once entrapped, the osteoblasts become osteocytes (Figure 6.16b). As osteoblasts transform into osteocytes, osteogenic cells in the surrounding connective tissue differentiate into new osteoblasts.

Osteoid (unmineralized bone matrix) secreted around the capillaries results in a trabecular matrix, while osteoblasts on the surface of the spongy bone become the periosteum (Figure 6.16c). The periosteum then creates a protective layer of compact bone superficial to the trabecular bone. The trabecular bone crowds nearby blood vessels, which eventually condense into red marrow (Figure 6.16d).

Figure 6.16 Intramembranous Ossification Intramembranous ossification follows four steps. (a) Mesenchymal cells group into clusters, and ossification centers form. (b) Secreted osteoid traps osteoblasts, which then become osteocytes. (c) Trabecular matrix and periosteum form. (d) Compact bone develops superficial to the trabecular bone, and crowded blood vessels condense into red marrow.

Intramembranous ossification begins in utero during fetal development and continues on into adolescence. At birth, the skull and clavicles are not fully ossified nor are the sutures of the skull closed. This allows the skull and shoulders to deform during passage through the birth canal. The last bones to ossify via intramembranous ossification are the flat bones of the face, which reach their adult size at the end of the adolescent growth spurt.

Endochondral Ossification

In endochondral ossification, bone develops by replacing hyaline cartilage. Cartilage does not become bone. Instead, cartilage serves as a template to be completely replaced by new bone. Endochondral ossification takes much longer than intramembranous ossification. Bones at the base of the skull and long bones form via endochondral ossification.

In a long bone, for example, at about 6 to 8 weeks after conception, some of the mesenchymal cells differentiate into chondrocytes (cartilage cells) that form the cartilaginous skeletal precursor of the bones (Figure 6.17a). Soon after, the perichondrium, a membrane that covers the cartilage, appears Figure 6.17b).

Figure 6.17 Endochondral Ossification Endochondral ossification follows five steps. (a) Mesenchymal cells differentiate into chondrocytes. (b) The cartilage model of the future bony skeleton and the perichondrium form. (c) Capillaries penetrate cartilage. Perichondrium transforms into periosteum. Periosteal collar develops. Primary ossification center develops. (d) Cartilage and chondrocytes continue to grow at ends of the bone. (e) Secondary ossification centers develop. (f) Cartilage remains at epiphyseal (growth) plate and at joint surface as articular cartilage.

As more matrix is produced, the chondrocytes in the center of the cartilaginous model grow in size. As the matrix calcifies, nutrients can no longer reach the chondrocytes. This results in their death and the disintegration of the surrounding cartilage. Blood vessels invade the resulting spaces, not only enlarging the cavities but also carrying osteogenic cells with them, many of which will become osteoblasts. These enlarging spaces eventually combine to become the medullary cavity.

As the cartilage grows, capillaries penetrate it. This penetration initiates the transformation of the perichondrium into the bone-producing periosteum. Here, the osteoblasts form a periosteal collar of compact bone around the cartilage of the diaphysis. By the second or third month of fetal life, bone cell development and ossification ramps up and creates the primary ossification center, a region deep in the periosteal collar where ossification begins (Figure 6.17c).

While these deep changes are occurring, chondrocytes and cartilage continue to grow at the ends of the bone (the future epiphyses), which increases the bone’s length at the same time bone is replacing cartilage in the diaphyses. By the time the fetal skeleton is fully formed, cartilage only remains at the joint surface as articular cartilage and between the diaphysis and epiphysis as the epiphyseal plate, the latter of which is responsible for the longitudinal growth of bones. After birth, this same sequence of events (matrix mineralization, death of chondrocytes, invasion of blood vessels from the periosteum, and seeding with osteogenic cells that become osteoblasts) occurs in the epiphyseal regions, and each of these centers of activity is referred to as a secondary ossification center (Figure 6.17e).

How Bones Grow in Length

The epiphyseal plate is the area of growth in a long bone. It is a layer of hyaline cartilage where ossification occurs in immature bones. On the epiphyseal side of the epiphyseal plate, cartilage is formed. On the diaphyseal side, cartilage is ossified, and the diaphysis grows in length. The epiphyseal plate is composed of four zones of cells and activity (Figure 6.18). The reserve zone is the region closest to the epiphyseal end of the plate and contains small chondrocytes within the matrix. These chondrocytes do not participate in bone growth but secure the epiphyseal plate to the osseous tissue of the epiphysis.

Figure 6.18 Longitudinal Bone Growth The epiphyseal plate is responsible for longitudinal bone growth.

The proliferative zone is the next layer toward the diaphysis and contains stacks of slightly larger chondrocytes. It makes new chondrocytes (via mitosis) to replace those that die at the diaphyseal end of the plate. Chondrocytes in the next layer, the zone of maturation and hypertrophy, are older and larger than those in the proliferative zone. The more mature cells are situated closer to the diaphyseal end of the plate. The longitudinal growth of bone is a result of cellular division in the proliferative zone and the maturation of cells in the zone of maturation and hypertrophy.

Most of the chondrocytes in the zone of calcified matrix, the zone closest to the diaphysis, are dead because the matrix around them has calcified. Capillaries and osteoblasts from the diaphysis penetrate this zone, and the osteoblasts secrete bone tissue on the remaining calcified cartilage. Thus, the zone of calcified matrix connects the epiphyseal plate to the diaphysis. A bone grows in length when osseous tissue is added to the diaphysis.

Bones continue to grow in length until early adulthood. The rate of growth is controlled by hormones, which will be discussed later. When the chondrocytes in the epiphyseal plate cease their proliferation and bone replaces the cartilage, longitudinal growth stops. All that remains of the epiphyseal plate is the epiphyseal line (Figure 6.19).

Figure 6.19 Progression from Epiphyseal Plate to Epiphyseal Line As a bone matures, the epiphyseal plate progresses to an epiphyseal line. (a) Epiphyseal plates are visible in a growing bone. (b) Epiphyseal lines are the remnants of epiphyseal plates in a mature bone.

How Bones Grow in Diameter

While bones are increasing in length, they are also increasing in diameter; growth in diameter can continue even after longitudinal growth ceases. This is called appositional growth. Osteoclasts resorb old bone that lines the medullary cavity, while osteoblasts, via intramembranous ossification, produce new bone tissue beneath the periosteum. The erosion of old bone along the medullary cavity and the deposition of new bone beneath the periosteum not only increase the diameter of the diaphysis but also increase the diameter of the medullary cavity. This process is called modeling.

Bone Remodeling

The process in which matrix is resorbed on one surface of a bone and deposited on another is known as bone modeling. Modeling primarily takes place during a bone’s growth. However, in adult life, bone undergoes remodeling, in which resorption of old or damaged bone takes place on the same surface where osteoblasts lay new bone to replace that which is resorbed. Injury, exercise, and other activities lead to remodeling. Those influences are discussed later in the chapter, but even without injury or exercise, about 5 to 10 percent of the skeleton is remodeled annually just by destroying old bone and renewing it with fresh bone.