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By the end of this section, you will be able to:
  • Explain the function of cartilage
  • List the steps of endochondral ossification
  • Explain the growth activity at the epiphyseal plate
  • Compare and contrast the processes of modeling and remodeling

In the early stages of embryonic development, the embryo’s skeleton consists of fibrous membranes and hyaline cartilage. By the sixth or seventh week of embryonic life, the actual process of bone development, ossification    , begins.

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. 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.

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.

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.

Endochondral ossification

Image A shows a small piece of hyaline cartilage that looks like a bone but without the characteristic enlarged ends. The hyaline cartilage is surrounded by a thin perichondrium. In image B, the hyaline cartilage has increased in size and the ends have begun to bulge outwards. A group of dark granules form at the center of the cartilage. This is labeled the calcified matrix, as opposed to the rest of the cartilage, which is uncalcified matrix. In image C, the hyaline cartilage has again increased in size and spongy bone has formed at the calcified matrix. This is now called the primary ossification center. A nutrient artery has invaded the ossification center and is growing through the cavities of the new spongy bone. In image D, the cartilage now looks like a bone, as it has greatly increased in size and each end has two bulges. Only the proximal half of the bone is shown in all of the remaining images. In image D, spongy bone has completely developed in the medullary cavity, which is surrounded, on both sides, by compact bone. Now, the calcified matrix is located at the border between the proximal metaphysis and the proximal epiphysis. The epiphysis is still composed of uncalcified matrix. In image E, arteries and veins have now invaded the epiphysis, forming a calcified matrix at its center. This is called a secondary ossification center. In image F, the interior of the epiphysis is now completely calcified into bone. The outer edge of the epiphysis remains as cartilage, forming the articular cartilage at the joint. In addition, the border between the epiphysis and the metaphysis remains uncalcified, forming the epiphyseal plate.
Endochondral ossification follows five steps. (a) Mesenchymal cells differentiate into chondrocytes. (b) The cartilage model of the future bony skeleton. (c) Capillaries penetrate cartilage. 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 ( [link] c ).

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Source:  OpenStax, Human biology. OpenStax CNX. Dec 01, 2015 Download for free at http://legacy.cnx.org/content/col11903/1.3
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