FIGURE 8.14 • Electron micrograph of an osteoclast. This micrograph shows a segment of bone surface (B) and a portion of an osteoclast that is in apposition to the partially digested bone. The resorption front (RF) of the osteoclast possesses numerous infoldings of the plasma membrane. When viewed in the light microscope, these infoldings are evident as the ruffled border. When the plane of section is parallel to the infoldings (asterisks), a broad, nonspecialized expanse of cytoplasm is seen. The cytoplasm of the osteoclast contains numerous mitochondria (M), lysosomes, and Golgi apparatus, all of which are functionally linked with the resorption and degradation of the bone matrix. In the upper part of the figure, some collagen fibrils are evident; the arrows indicate where 68-nm cross-banding is visible. 10,000.
the resorption site (see Fig. 8.12). At the electron microscopic level, hydroxyapatite crystals from the bone substance are observed between the processes of the ruffled border (Fig. 8.14). Internal to the ruffled border and in close proximity are numerous mitochondria and lysosomes. The nuclei are typically located in the part of the cell more removed from the bone surface. In this same region are profiles of rER, multiple stacks of Golgi apparatus, and many vesicles.
 The clear zone (sealing zone) is a ringlike perimeter of cytoplasm adjacent to the ruffled border that demarcates the bone area being resorbed. Essentially, the clear zone is a compartment at the site of the ruffled border where resorption and degradation of the matrix occurs. It contains abundant actin filaments but essentially lacks other organelles. The actin filaments are arranged in a ringlike structure surrounded on both sides by actin-binding proteins such as vinculin and talin (Fig. 8.15). The plasma membrane at the site of the clear zone contains cell and extracellular matrix adhesion molecules that are responsible for providing a tight seal between the plasma membrane and mineralized matrix of the bone. Several classes of integrin extracellular receptors (i.e., v 3 vitronectin receptor, 2 1 collagen receptor, or v 1 receptor) help maintain the seal.
 The basolateral region functions in the exocytosis of digested material (see Fig. 8.15). Transport vesicles containing degraded bone material endocytosed at the ruffled border fuse here with the cell membrane to release their contents. TRAP has been found within these vesicles, suggesting its role in the fragmentation of endocytosed material.
Osteoclasts resorb bone tissue by releasing protons and lysosomal hydrolases into the constricted microenvironment of the extracellular space.
Some, if not most, of the vesicles in the osteoclast are lysosomes. Their contents are released into the extracellular space in the clefts between the cytoplasmic processes of the ruffled border, a clear example of lysosomal enzymes functioning outside the cell. Once liberated, these hydrolytic enzymes, which include cathepsin K (a cysteine protease) and matrix metalloproteinases, degrade collagen and other proteins of the bone matrix.