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Formation and remodelling of bone

Bone formation is an essential process in the development of the human body. It starts during the development of the foetus, and continues throughout childhood and adolescence as the skeleton grows. Bone remodelling meanwhile is a life-long process, consisting of resorption (the breaking down of old bone) and ossification (formation of new bone), and is key to shaping the skeleton and to the repair of bone fractures.

There are three types of cell present in bone that are of particular interest – osteoblasts, osteocytes and osteoclasts, which are respectively responsible for the production, maintenance and resorption of bone.

  • Osteoblasts
    Mononucleated “bone-forming” cells found near the surface of bones. They are responsible for making osteoid, which consists mainly of collagen. The osteoblasts then secrete alkaline phosphatase to create sites for calcium and phosphate deposition, which allows crystals of bone mineral to grow at these sites. The osteoid becomes mineralised, thus forming bone.
  • Osteocytes
    These are osteoblasts that are no longer on the surface of the bone, but are instead found in lacunae between the lamellae in bone. Their main role is homeostasis – maintaining the correct oxygen and mineral levels in the bone.
  • Osteoclasts
     Multinucleated cells responsible for bone resorption. They travel to specific sites on the surface of bone and secrete acid phosphatase, which unfixes the calcium in mineralised bone to break it down.

During foetal development there are two mechanisms for creating bone tissue:

  • Endochondral ossification
  • Intramembranous ossification

Intramembranous ossification occurs in the formation of flat bones such as those in the skull, and will not be covered further here. More information can be found through the Going Further page.

Endochondral ossification
This involves bone growth from an underlying cartilage model, and is seen in the formation and growth of long bones such as the femur.

The initial step involves the development of a cartilage model, which has the rough shape of the bone being formed. In the middle of the shaft is the primary ossification centre, where osteoblasts lay down osteoid on the shaft to form a bone collar.

The osteoid calcifies, and blood vessels grow into cavities within the matrix. Osteoblasts then use the calcified matrix as a support structure to lay down more osteoid and form trabeculae within the bone. Meanwhile osteoclasts break down spongy bone to create the medullary cavity, which contains bone marrow.

Initially the bone material is deposited with the collagen fibres in random directions, meaning the strength is much lower than at the final stage in which the fibres are aligned. The primary structure is called woven bone because the collagen fibres are woven together randomly. This is then converted into lamellar bone over time, which is much stronger due to the aligned fibres. The osteoid deposited by the osteoblasts calcifies to initially produce primitive cancellous bone. At sites where cortical bone is required, further deposition of osteoid occurs to increase the density of the structure.

At birth secondary ossification centres appear at either end of long bones. Between the primary and secondary centres is the epiphyseal plate, made of cartilage, which continues to form new cartilage and be replaced by bone such that the bone increases in length. This continues until a person is in their mid-twenties, when the plate is finally replaced by bone and no further growth occurs.

Note: This animation requires Adobe Flash Player 8 and later, which can be downloaded here.

Remodelling of bone

Ossification is also essential in the remodelling of bone. This occurs throughout a person’s lifetime, with ossification and resorption (removal of bone tissue) working together to reshape the skeleton during growth, maintain calcium levels in the body, and repair micro-fractures caused by everyday stress.

Note: This animation requires Adobe Flash Player 8 and later, which can be downloaded here.

The remodelling of cortical bone follows the same process as shown above, but with a different geometry in order to form the concentric lamellae seen in osteons.

Responsive material

Bone is considered to be a responsive material. The formation and resorption of bone occur continuously: the body responds to stress levels in different areas of bone to ensure the right amount of healthy bone tissue is maintained and the bone can be continually reshaped.

A stress of 25–40 MPa is sufficient to maintain the correct levels of bone. If the bone is under-stressed for prolonged periods of time, bone wastage will set in, and the bones will become thinner. This can be an issue if a patient is bed-ridden for a long time, and is also observed in astronauts after long periods in space. A similar effect occurs during osteoporosis, in which the activity of osteoblasts decreases with age. This results in an imbalance of resorption and formation, causing bones to become thinner and weaker.

The opposite effect can be seen when bones are suddenly subjected to higher levels of stress than normal. Studies have been conducted that show an increase in bone mass in new recruits to the army as they begin intensive training.

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