Bone tissue (osseous tissue) differs greatly from other tissues in the body. Bone is hard and many of its functions depend on that characteristic hardness.

Gross Anatomy of Bone

The structure of a long bone allows for the best visualization of all of the parts of a bone (Figure 1). A long bone has two parts: the diaphysis and the epiphysis. The diaphysis is the tubular shaft that runs between the proximal and distal ends of the bone. The hollow region in the diaphysis is called the medullary cavity, which is filled with yellow marrow. The walls of the diaphysis are composed of dense and hard compact bone.

The wider section at each end of the bone is called the epiphysis (plural = epiphyses), which is filled with spongy bone. Red marrow fills the spaces in the spongy bone. Each epiphysis meets the diaphysis at the metaphysis, the narrow area that contains the epiphyseal plate (growth plate), a layer of hyaline (transparent) cartilage in a growing bone. When the bone stops growing in early adulthood (approximately 18–21 years), the cartilage is replaced by osseous tissue and the epiphyseal plate becomes an epiphyseal line.

The medullary cavity has a delicate membranous lining called the endosteum (end– = “inside”; oste– = “bone”), where bone growth, repair, and remodeling occur. The outer surface of the bone is covered with a fibrous membrane called the periosteum (peri– = “around” or “surrounding”). The periosteum contains blood vessels, nerves, and lymphatic vessels that nourish compact bone. Tendons and ligaments also attach to bones at the periosteum. The periosteum covers the entire outer surface except where the epiphyses meet other bones to form joints (Figure 2). In this region, the epiphyses are covered with articular cartilage, a thin layer of cartilage that reduces friction and acts as a shock absorber.

Flat bones, like those of the cranium, consist of a layer of diploë (spongy bone), lined on either side by a layer of compact bone (Figure 2). The two layers of compact bone and the interior spongy bone work together to protect the internal organs. If the outer layer of a cranial bone fractures, the brain is still protected by the intact inner layer.

This illustration depicts an anterior view of the right femur, or thigh bone. The inferior end that connects to the knee is at the bottom of the diagram and the superior end that connects to the hip is at the top of the diagram. The bottom end of the bone contains a smaller lateral bulge and a larger medial bulge. A blue articular cartilage covers the inner half of each bulge as well as the small trench that runs between the bulges. This area of the inferior end of the bone is labeled the distal epiphysis. Above the distal epiphysis is the metaphysis, where the bone tapers from the wide epiphysis into the relatively thin shaft. The entire length of the shaft is the diaphysis. The superior half of the femur is cut away to show its internal contents. The bone is covered with an outer translucent sheet called the periosteum. At the midpoint of the diaphysis, a nutrient artery travels through the periosteum and into the inner layers of the bone. The periosteum surrounds a white cylinder of solid bone labeled compact bone. The cavity at the center of the compact bone is called the medullary cavity. The inner layer of the compact bone that lines the medullary cavity is called the endosteum. Within the diaphysis, the medullary cavity contains a cylinder of yellow bone marrow that is penetrated by the nutrient artery. The superior end of the femur is also connected to the diaphysis by a metaphysis. In this upper metaphysis, the bone gradually widens between the diaphysis and the proximal epiphysis. The proximal epiphysis of the femur is roughly hexagonal in shape. However, the upper right side of the hexagon has a large, protruding knob. The femur connects and rotates within the hip socket at this knob. The knob is covered with a blue colored articular cartilage. The internal anatomy of the upper metaphysis and proximal epiphysis are revealed. The medullary cavity in these regions is filled with the mesh like spongy bone. Red bone marrow occupies the many cavities within the spongy bone. There is a clear, white line separating the spongy bone of the upper metaphysis with that of the proximal epiphysis. This line is labeled the epiphyseal line.

Figure 1. Anatomy of a Long Bone. A typical long bone shows the gross anatomical characteristics of bone.

This illustration shows a cross section of a cranial bone, constructed somewhat like a sandwich. The topmost and bottommost layers are the thin, translucent, periosteum. The upper and lower periosteum cover an upper and lower layer of compact bone, respectively. The compact bone is solid, with each layer occupying about one tenth of the thickness of the cranial bone. The majority of the cross section is occupied by the spongy bone, or diploe, sandwiched between the upper and lower compact bone. The spongy bone contains many crisscrossing threads of bone. Dark air spaces occur between the threads, giving the bone a porous appearance, much like that of a sponge or Swiss cheese.

Figure 2. Anatomy of a Flat Bone. This cross-section of a flat bone shows the spongy bone (diploë) lined on either side by a layer of compact bone.

Bone Cells and Tissue

Bone contains a relatively small number of cells entrenched in a matrix of collagen fibers that provide a surface for inorganic salt crystals to adhere. These salt crystals form when calcium phosphate and calcium carbonate combine to create hydroxyapatite, which incorporates other inorganic salts like magnesium hydroxide, fluoride, and sulfate as it crystallizes, or calcifies, on the collagen fibers. The hydroxyapatite crystals give bones their hardness and strength, while the collagen fibers give them flexibility so that they are not brittle.

The top of this diagram shows the cross section of a generic bone with three zoom in boxes. The first box is on the periosteum. The second box is on the middle of the compact bone layer. The third box is on the inner edge of the compact bone where it transitions into the spongy bone. The callout in the periosteum points to two images. In the first image, four osteoblast cells are sitting end to end on the periosteum. The osteoblasts are roughly square shaped, except for one of the cells which is developing small, finger like projections. The caption says, “Osteoblasts form the matrix of the bone.” The second image called out from the periosteum shows a large, amorphous osteogenic cell sitting on the periosteum. The osteogenic cell is surrounded on both sides by a row of much smaller osteoblasts. The cell is shaped like a mushroom cap and also has finger like projections. The cell is a stem cell that develops into other bone cells. The box in the middle of the compact bone layer is pointing to an osteocyte. The osteocyte is a thin cell, roughly diamond shaped, with many branching, finger-like projections. The osteoctyes maintain bone tissue. The box at the inner edge of the compact bone is pointing to an osteoclast. The osteoclast is a large, round cell with multiple nuclei. It also has rows of fine finger like projections on its lower surface where it is sitting on the compact bone. The osteoclast reabsorbs bone.

Figure 3. Bone Cells. Four types of cells are found within bone tissue. Osteogenic cells are undifferentiated and develop into osteoblasts. When osteoblasts get trapped within the calcified matrix, their structure and function changes, and they become osteocytes. Osteoclasts develop from monocytes and macrophages and differ in appearance from other bone cells.

Although bone cells compose a small amount of the bone volume, they are crucial to the function of bones. Four types of cells are found within bone tissue: osteoblasts, osteocytes, osteogenic cells, and osteoclasts (Figure 3).

The osteoblast. is the bone cell responsible for forming new bone and is found in the growing portions of bone, including the periosteum and endosteum. Osteoblasts, which do not divide, synthesize and secrete the collagen matrix and calcium salts. As the secreted matrix surrounding the osteoblast calcifies, the osteoblast become trapped within it; as a result, it changes in structure and becomes an osteocyte, the primary cell of mature bone and the most common type of bone cell. Each osteocyte is located in a space called a lacuna and is surrounded by bone tissue. Osteocytes maintain the mineral concentration of the matrix via the secretion of enzymes. Like osteoblasts, osteocytes lack mitotic activity. They can communicate with each other and receive nutrients via long cytoplasmic processes that extend through canaliculi (singular = canaliculus), channels within the bone matrix.

If osteoblasts and osteocytes are incapable of mitosis, then how are they replenished when old ones die? The answer lies in the properties of a third category of bone cells—the osteogenic cell. These osteogenic cells are undifferentiated with high mitotic activity and they are the only bone cells that divide. Immature osteogenic cells are found in the deep layers of the periosteum and the marrow. They differentiate and develop into osteoblasts.

The dynamic nature of bone means that new tissue is constantly formed, and old, injured, or unnecessary bone is dissolved for repair or for calcium release. The cell responsible for bone resorption, or breakdown, is the osteoclast. They are found on bone surfaces, are multinucleated, and originate from monocytes and macrophages, two types of white blood cells, not from osteogenic cells. Osteoclasts are continually breaking down old bone while osteoblasts are continually forming new bone. The ongoing balance between osteoblasts and osteoclasts is responsible for the constant but subtle reshaping of bone. Table 2 reviews the bone cells, their functions, and locations.

Table 2. Bone Cells
Cell type	Function	Location
Osteogenic cells	Develop into osteoblasts	Deep layers of the periosteum and the marrow
Osteoblasts	Bone formation	Growing portions of bone, including periosteum and endosteum
Osteocytes	Maintain mineral concentration of matrix	Entrapped in matrix
Osteoclasts	Bone resorption	Bone surfaces and at sites of old, injured, or unneeded bone

Bone Tissue Types

Bones are considered organs because they contain various types of tissue, such as blood, connective tissue, nerves, and bone tissue. Osteocytes, the living cells of bone tissue, form the mineral matrix of bones. There are two types of bone tissue: compact and spongy.

The differences between compact and spongy bone are best explored via their histology. Most bones contain compact and spongy osseous tissue, but their distribution and concentration vary based on the bone’s overall function. Compact bone is dense so that it can withstand compressive forces, while spongy (cancellous) bone has open spaces and supports shifts in weight distribution.

Compact Bone: Compact bone is the denser, stronger of the two types of bone tissue (Figure 4). It can be found under the periosteum and in the diaphyses of long bones, where it provides support and protection.

A generic long bone is shown at the top of this illustration. The bone is split in half lengthwise to show its internal anatomy. The outer gray covering of the bone is labeled the periosteum. Within the periosteum is a thin layer of compact bone. The compact bone surrounds a central cavity called the medullary cavity. The medullary cavity is filled with spongy bone at the two epiphyses. A callout box shows that the main image is zooming in on the compact bone on the left side of the bone. On the main image, the periosteum is being peeled back to show its two layers. The outer layer of the periosteum is the outer fibrous layer. This layer has a periosteal artery and a periosteal vein running along its outside edge. The inner layer of the periosteum is labeled the inner osteogenic layer. The compact bone lies to the right of the periosteum and occupies the majority of the main image. Two flat layers of compact bone line the inner surface of the ostegenic periosteum. These sheets of compact bone are called the circumferential lamellae. The majority of the compact bone has lamellae running perpendicular to that of the circumferential lamellae. These concentric lamellae are arranged in a series of concentric tubes. There are small cavities between the layers of concentric lamellae called lacunae. The centermost concentric lamella surrounds a hollow central canal. A blue vein, a red artery, a yellow nerve and a green lymph vessel run vertically through the central canal. A set of concentric lamellae, its associated lacunae and the vessels and nerves of the central canal are collectively called an osteon. The front edge of the diagram shows a longitudinal cross section of one of the osteons. The vessels and nerve are visible running through the center of the osteon throughout its length. In addition, blood vessels can run from the periosteum through the sides of the osteons and connect with the vessels of the central canal. The blood vessels travel through the sides of the osteons via a perforating canal. The open areas between neighboring osteons are also filled with compact bone. This “filler” bone is referred to as the interstitial lamellae. At the far right of the compact bone, the edge of the spongy bone is visible. The spongy bone is a series of crisscrossing bony arches called trabeculae. There are many open spaces between the trabeculae, giving the spongy bone its sponge-like appearance.

Figure 4. Diagram of Compact Bone. (a) This cross-sectional view of compact bone shows the basic structural unit, the osteon. (b) In this micrograph of the osteon, you can clearly see the concentric lamellae and central canals. LM × 40. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)

The microscopic structural unit of compact bone is called an osteon, or Haversian system. Each osteon is composed of concentric rings of calcified matrix called lamellae (singular = lamella). Running down the center of each osteon is the central canal, or Haversian canal, which contains blood vessels, nerves, and lymphatic vessels. These vessels and nerves branch off at right angles through a perforating canal, also known as Volkmann’s canals, to extend to the periosteum and endosteum.

The osteocytes are located inside spaces called lacunae (singular = lacuna), found at the borders of adjacent lamellae. As described earlier, canaliculi connect with the canaliculi of other lacunae and eventually with the central canal. This system allows nutrients to be transported to the osteocytes and wastes to be removed from them.

This illustration shows the spongy bone within the proximal epiphysis of the femur in two successively magnified images. The lower-magnification image shows two layers of crisscrossing trabeculae. The surface of each is dotted with small black holes which are the openings of the canaliculi. One of the trabeculae is in a cross section to show its internal layers. The outermost covering of the lamellae is called the endosteum. This endosteum surrounds several layers of concentric lamellae. The higher-magnification image shows the cross section of the trabeculae more clearly. Three concentric lamellae are shown in this view, each possessing perpendicular black lines. These lines are the canaliculi and are oriented on the round lamellae similar to the spokes of a wheel. In between the lamellae are small cavities called lacunae which house cells called osteocytes. In addition, two large osteoclasts are seated on the outer edge of the outermost lamellae. The outermost lamellae are also surrounded by groups of small, white, osteoblasts.

Figure 5. Diagram of Spongy Bone. Spongy bone is composed of trabeculae that contain the osteocytes. Red marrow fills the spaces in some bones.

Spongy (Cancellous) Bone: Like compact bone, spongy bone, also known as cancellous bone, contains osteocytes housed in lacunae, but they are not arranged in concentric circles. Instead, the lacunae and osteocytes are found in a lattice-like network of matrix spikes called trabeculae (singular = trabecula) (Figure 5). The trabeculae may appear to be a random network, but each trabecula forms along lines of stress to provide strength to the bone. The spaces of the trabeculated network provide balance to the dense and heavy compact bone by making bones lighter so that muscles can move them more easily. In addition, the spaces in some spongy bones contain red marrow, protected by the trabeculae, where hematopoiesis occurs.

Aging and the Skeletal System: Paget’s Disease

Paget’s disease usually occurs in adults over age 40. It is a disorder of the bone remodeling process that begins with overactive osteoclasts. This means more bone is resorbed than is laid down. The osteoblasts try to compensate but the new bone they lay down is weak and brittle and therefore prone to fracture.

This illustration shows the normal skeletal structure of the legs from an anterior view. The flesh of the legs and feet are outlined around the skeleton for reference. A second illustration shows the legs of someone with Paget’s disease. The affected person’s left femur is curved outward, causing the left leg to be bowed and shorter than the right leg.

Figure 6. Paget’s Disease. Normal leg bones are relatively straight, but those affected by Paget’s disease are porous and curved.

While some people with Paget’s disease have no symptoms, others experience pain, bone fractures, and bone deformities (Figure 6). Bones of the pelvis, skull, spine, and legs are the most commonly affected. When occurring in the skull, Paget’s disease can cause headaches and hearing loss.

What causes the osteoclasts to become overactive? The answer is still unknown, but hereditary factors seem to play a role. Some scientists believe Paget’s disease is due to an as-yet-unidentified virus.

Paget’s disease is diagnosed via imaging studies and lab tests. X-rays may show bone deformities or areas of bone resorption. Bone scans are also useful. In these studies, a dye containing a radioactive ion is injected into the body. Areas of bone resorption have an affinity for the ion, so they will light up on the scan if the ions are absorbed. In addition, blood levels of an enzyme called alkaline phosphatase are typically elevated in people with Paget’s disease.

Bisphosphonates, drugs that decrease the activity of osteoclasts, are often used in the treatment of Paget’s disease. However, in a small percentage of cases, bisphosphonates themselves have been linked to an increased risk of fractures because the old bone that is left after bisphosphonates are administered becomes worn out and brittle. Still, most doctors feel that the benefits of bisphosphonates more than outweigh the risk; the medical professional has to weigh the benefits and risks on a case-by-case basis. Bisphosphonate treatment can reduce the overall risk of deformities or fractures, which in turn reduces the risk of surgical repair and its associated risks and complications.

This illustration shows an anterior view if the right femur. The femur is split in half lengthwise to show its internal anatomy. The outer covering of the femur is labeled the periosteum. Within it is a thin layer of compact bone that surrounds a central cavity called the medullary or marrow cavity. This cavity is filled with spongy bone at both epiphyses. A nutrient artery and vein travels through the periosteum and compact bone at the center of the diaphysis. After entering the bone, the nutrient arteries and veins spread throughout the marrow cavity in both directions. Some of the arteries and veins in the marrow cavity also spread into the spongy bone within the distal and proximal epiphyses. However, additional blood vessels called the metaphyseal arteries and the metaphyseal veins enter into the metaphysis from outside of the bone.

Figure 7. Diagram of Blood and Nerve Supply to Bone. Blood vessels and nerves enter the bone through the nutrient foramen.

Blood and Nerve Supply

The spongy bone and medullary cavity receive nourishment from arteries that pass through the compact bone. The arteries enter through the nutrient foramen (plural = foramina), small openings in the diaphysis (Figure 7). The osteocytes in spongy bone are nourished by blood vessels of the periosteum that penetrate spongy bone and blood that circulates in the marrow cavities. As the blood passes through the marrow cavities, it is collected by veins, which then pass out of the bone through the foramina.

In addition to the blood vessels, nerves follow the same paths into the bone where they tend to concentrate in the more metabolically active regions of the bone. The nerves sense pain, and it appears the nerves also play roles in regulating blood supplies and in bone growth, hence their concentrations in metabolically active sites of the bone.

Bone Formation and Development

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 (osteogenesis), begins. There are two osteogenic pathways—intramembranous ossification and endochondral ossification—but bone is the same regardless of the pathway that produces it.

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

Intramembranous ossification is the process of bone development from fibrous membranes. It is involved in the formation of the flat bones of the skull, the mandible, and the clavicles. Ossification begins as mesenchymal cells form a template of the future bone. They then differentiate into osteoblasts at the ossification center. Osteoblasts secrete the extracellular matrix and deposit calcium, which hardens the matrix. The non-mineralized portion of the bone or osteoid continues to form around blood vessels, forming spongy bone. Connective tissue in the matrix differentiates into red bone marrow in the fetus. The spongy bone is remodeled into a thin layer of compact bone on the surface of the spongy bone.

Image A shows seven osteoblasts, cells with small, finger like projections. They are surrounded by granules of osteoid. Both the cells and the osteoid are contained within a blue, circular, ossification center that is surrounded by a “socket” of dark, string-like collagen fibers and gray mesenchymal cells. The cells are generally amorphous, similar in appearance to an amoeba. In image B, the ossification center is no longer surrounded by a ring of osteoblasts. The osteoblasts have secreted bone into the ossification center, creating a new bone matrix. There are also five osteocytes embedded in the new bone matrix. The osteocytes are thin, oval-shaped cells with many fingerlike projections. Osteoid particles are still embedded in the bony matrix in image B. In image C, the ring of osteoblasts surrounding the ossification center has separated, forming an upper and lower layer of osteoblasts sandwiched between the two layers of mesenchyme cells. A label indicates that the mesenchyme cells and the surrounding collagen fibers form the periosteum. The osteoblasts secrete spongy bone into the space between the two osteoblast rows. Therefore, the accumulating spongy bone pushes the upper and lower rows of osteoblasts away from each other. In this image, most of the spongy bone has been secreted by the osteoblasts, as the trabeculae are visible. In addition, an artery has already broken through the periosteum and invaded the spongy bone. Image D looks similar to image C, except that the rows of osteoblasts are now secreting layers of compact bone between the spongy bone and the periosteum. The artery has now branched and spread throughout the spongy bone. A label indicates that the cavities between the trabeculae now contain red bone marrow.

Figure 8. 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

Endochondral ossification is the process of bone development from hyaline cartilage. All of the bones of the body, except for the flat bones of the skull, mandible, and clavicles, are formed through endochondral ossification.

In long bones, chondrocytes form a template of the hyaline cartilage diaphysis. Responding to complex developmental signals, the matrix begins to calcify. This calcification prevents diffusion of nutrients into the matrix, resulting in chondrocytes dying and the opening up of cavities in the diaphysis cartilage. Blood vessels invade the cavities, and osteoblasts and osteoclasts modify the calcified cartilage matrix into spongy bone. Osteoclasts then break down some of the spongy bone to create a marrow, or medullary, cavity in the center of the diaphysis. Dense, irregular connective tissue forms a sheath (periosteum) around the bones. The periosteum assists in attaching the bone to surrounding tissues, tendons, and ligaments. The bone continues to grow and elongate as the cartilage cells at the epiphyses divide.

In the last stage of prenatal bone development, the centers of the epiphyses begin to calcify. Secondary ossification centers form in the epiphyses as blood vessels and osteoblasts enter these areas and convert hyaline cartilage into spongy bone. Until adolescence, hyaline cartilage persists at the epiphyseal plate (growth plate), which is the region between the diaphysis and epiphysis that is responsible for the lengthwise growth of long bones.

Illustration shows bone growth, which begins with a hyaline cartilage model that has the appearance of a small bone. A primary ossification center forms in the center of the narrow part of the bone, and a bone collar forms around the outside. The periosteum forms around the outside of the bone. Next, blood vessels begin to form in the bone and secondary ossification centers form in the epiphyses. The primary ossification center hollows out to form the medullary cavity, and an epiphyseal plate grows, separating the epiphyses from the diaphysis.

Figure 9. Endochondral ossification is the process of bone development from hyaline cartilage. The periosteum is the connective tissue on the outside of bone that acts as the interface between bone, blood vessels, tendons, and ligaments.

How Bones Grow in Length

This illustration shows anterior views of a right and left femur. The left femur possesses a growth plate at the border of its distal metaphysis and distal epiphysis. The proximal epiphysis has two growth plates. The first is located below the head of the femur while the second is located below the trochanter, which is the bump on the lateral side of the femur. The right femur has the same plates as the left femur. However, the left femur represents a mature long bone. Here, growth is completed and the epiphyseal plate has degraded to a thin, faint, epiphyseal line.

Figure 10. 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.

The epiphyseal plate (growth 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.

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 10).

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.

Diseases of the Skeletal System

Osteogenesis imperfecta (OI) is a genetic disease in which bones do not form properly and therefore are fragile and break easily. It is also called brittle bone disease. The disease is present from birth and affects a person throughout life.

The genetic mutation that causes OI affects the body’s production of collagen, one of the critical components of bone matrix. The severity of the disease can range from mild to severe. Those with the most severe forms of the disease sustain many more fractures than those with a mild form. Frequent and multiple fractures typically lead to bone deformities and short stature. Bowing of the long bones and curvature of the spine are also common in people afflicted with OI. Curvature of the spine makes breathing difficult because the lungs are compressed.

Because collagen is such an important structural protein in many parts of the body, people with OI may also experience fragile skin, weak muscles, loose joints, easy bruising, frequent nosebleeds, brittle teeth, blue sclera, and hearing loss. There is no known cure for OI. Treatment focuses on helping the person retain as much independence as possible while minimizing fractures and maximizing mobility. Toward that end, safe exercises, like swimming, in which the body is less likely to experience collisions or compressive forces, are recommended. Braces to support legs, ankles, knees, and wrists are used as needed. Canes, walkers, or wheelchairs can also help compensate for weaknesses.

When bones do break, casts, splints, or wraps are used. In some cases, metal rods may be surgically implanted into the long bones of the arms and legs. Research is currently being conducted on using bisphosphonates to treat OI. Smoking and being overweight are especially risky in people with OI, since smoking is known to weaken bones, and extra body weight puts additional stress on the bones.

Bone Remodeling and Repair

Bone renewal continues after birth into adulthood. Bone remodeling is the replacement of old bone tissue by new bone tissue. It involves the processes of bone deposition by osteoblasts and bone resorption by osteoclasts. Normal bone growth requires vitamins D, C, and A, plus minerals such as calcium, phosphorous, and magnesium. Hormones such as parathyroid hormone, growth hormone, and calcitonin are also required for proper bone growth and maintenance.

Bone turnover rates are quite high, with five to seven percent of bone mass being recycled every week. Differences in turnover rate exist in different areas of the skeleton and in different areas of a bone. For example, the bone in the head of the femur may be fully replaced every six months, whereas the bone along the shaft is altered much more slowly.

Bone remodeling allows bones to adapt to stresses by becoming thicker and stronger when subjected to stress. Bones that are not subject to normal stress, for example when a limb is in a cast, will begin to lose mass. A fractured or broken bone undergoes repair through four stages

Fractures & Bone Repair

A fracture is a broken bone. It will heal whether or not a physician resets it in its anatomical position. If the bone is not reset correctly, the healing process will keep the bone in its deformed position.

When a broken bone is manipulated and set into its natural position without surgery, the procedure is called a closed reduction. Open reduction requires surgery to expose the fracture and reset the bone. While some fractures can be minor, others are quite severe and result in grave complications. For example, a fractured diaphysis of the femur has the potential to release fat globules into the bloodstream. These can become lodged in the capillary beds of the lungs, leading to respiratory distress and if not treated quickly, death.

Types of Fractures

Fractures are classified by their complexity, location, and other features. Table 1 outlines common types of fractures. Some fractures may be described using more than one term because it may have the features of more than one type (e.g., an open transverse fracture).

$In this illustration, each type of fracture is shown on the right femur from an anterior view. In the closed fracture, the femur is broken in the middle of the shaft with the upper and lower halves of the bone completely separated. However, the two halves of the bones are still aligned in that the broken edges are still facing each other. In an open fracture, the femur is broken in the middle of the shaft with the upper and lower halves of the bone completely separated. Unlike the closed fracture, in the open fracture, the two bone halves are misaligned. The lower half is turned laterally and it has protruded through the skin of the thigh. The broken ends no longer line up with each other. In a transverse fracture, the bone has a crack entirely through its width, however, the broken ends are not separated. The crack is perpendicular to the long axis of the bone. Arrows indicate that this is usually caused by compression of the bone in a superior-inferior direction. A spiral fracture travels diagonally through the diameter of the bone. In a comminuted fracture, the bone has several connecting cracks at its middle. It is possible that the bone could splinter into several small pieces at the site of the comminuted fracture. In an impacted fracture, the crack zig zags throughout the width of the bone like a lightning bolt. An arrow indicates that these are usually caused by an impact that pushes the femur up into the body. A greenstick fracture is a small crack that does not extend through the entire width of the bone. The oblique fracture shown here is travelling diagonally through the shaft of the femur at about a thirty degree angle.$

Figure 11. Types of Fractures. Compare healthy bone with different types of fractures: (a) closed fracture, (b) open fracture, (c) transverse fracture, (d) spiral fracture, (e) comminuted fracture, (f) impacted fracture, (g) greenstick fracture, and (h) oblique fracture.

Table 1. Types of Fractures
Type of fracture	Description
Transverse	Occurs straight across the long axis of the bone
Oblique	Occurs at an angle that is not 90 degrees
Spiral	Bone segments are pulled apart as a result of a twisting motion
Comminuted	Several breaks result in many small pieces between two large segments
Impacted	One fragment is driven into the other, usually as a result of compression
Greenstick	A partial fracture in which only one side of the bone is broken
Open (or compound)	A fracture in which at least one end of the broken bone tears through the skin; carries a high risk of infection
Closed (or simple)	A fracture in which the skin remains intact

Bone Repair

When a bone breaks, blood flows from any vessel torn by the fracture. These vessels could be in the periosteum, osteons, and/or medullary cavity. The blood begins to clot, and about six to eight hours after the fracture, the clotting blood has formed a fracture hematoma (Figure 12).

The disruption of blood flow to the bone results in the death of bone cells around the fracture.

$This illustration shows a left to right progression of bone repair. The break is shown in the leftmost image, where the femur has an oblique, closed fracture in the middle of its shaft. The next image magnifies the break, showing that blood has filled the area between the broken bones. Blood has also filled in around the lateral and medial sides of the break. The influx of blood causes the broken area to swell, creating a hematoma. In the next image, the hematoma has been replaced with an external callus between the two broken ends. Within the internal callus, the blood vessels have reconnected and some spongy bone has regenerated in the gap between the two bone halves. In the next image, spongy bone has completely regenerated, connecting the two broken ends, referred to as the bony callus. The external callus still remains on the lateral and medial sides of the break, as the compact bone has not yet regenerated. In the final image, the compact bone has fully regenerated, encapsulating the bony callus and completely reconnecting the two bone halves. The bone has a slight bulge at the location of the healed fracture, which is clearly shown in the final image, which shows a zoomed out image of the completely healed femur.$

Figure 12 Stages in Fracture Repair. The healing of a bone fracture follows a series of progressive steps: (a) A fracture hematoma forms. (b) Internal and external calli form. (c) Cartilage of the calli is replaced by trabecular bone. (d) Remodeling occurs.

Within about 48 hours after the fracture, chondrocytes from the endosteum have created an internal callus (plural = calli) by secreting a fibrocartilaginous matrix between the two ends of the broken bone, while the periosteal chondrocytes and osteoblasts create an external callus of hyaline cartilage and bone, respectively, around the outside of the break (Figure 12b). This stabilizes the fracture.

Over the next several weeks, osteoclasts resorb the dead bone; osteogenic cells become active, divide, and differentiate into osteoblasts. The cartilage in the calli is replaced by trabecular bone via endochondral ossification (Figure 12c).

Eventually, the internal and external calli unite, compact bone replaces spongy bone at the outer margins of the fracture, and healing is complete. A slight swelling may remain on the outer surface of the bone, but quite often, that region undergoes remodeling (Figure 12d), and no external evidence of the fracture remains.

Scientific Method Connection

Decalcification of Bones
Question: What effect does the removal of calcium and collagen have on bone structure?

Background: Conduct a literature search on the role of calcium and collagen in maintaining bone structure. Conduct a literature search on diseases in which bone structure is compromised.

Hypothesis: Develop a hypothesis that states predictions of the flexibility, strength, and mass of bones that have had the calcium and collagen components removed. Develop a hypothesis regarding the attempt to add calcium back to decalcified bones.

Test the hypothesis: Test the prediction by removing calcium from chicken bones by placing them in a jar of vinegar for seven days. Test the hypothesis regarding adding calcium back to decalcified bone by placing the decalcified chicken bones into a jar of water with calcium supplements added. Test the prediction by denaturing the collagen from the bones by baking them at 250°C for three hours.

Analyze the data: Create a table showing the changes in bone flexibility, strength, and mass in the three different environments.

Report the results: Under which conditions was the bone most flexible? Under which conditions was the bone the strongest?

Draw a conclusion: Did the results support or refute the hypothesis? How do the results observed in this experiment correspond to diseases that destroy bone tissue?

Chapter 7- The Skeletal System

Bone Structure, Formation, Development And Bone Repair