Calcium is not only the most abundant mineral in bone, it is also the most abundant mineral in the human body. Calcium ions are needed not only for bone mineralization but for tooth health, regulation of the heart rate and strength of contraction, blood coagulation, contraction of smooth and skeletal muscle cells, and regulation of nerve impulse conduction. The normal level of calcium in the blood is about 10 mg/dL. When the body cannot maintain this level, a person will experience hypo- or hypercalcemia.
Hypocalcemia, a condition characterized by abnormally low levels of calcium, can have an adverse effect on a number of different body systems including circulation, muscles, nerves, and bone. Without adequate calcium, blood has difficulty coagulating, the heart may skip beats or stop beating altogether, muscles may have difficulty contracting, nerves may have difficulty functioning, and bones may become brittle. The causes of hypocalcemia can range from hormonal imbalances to an improper diet. Treatments vary according to the cause, but prognoses are generally good.
Conversely, in hypercalcemia, a condition characterized by abnormally high levels of calcium, the nervous system is underactive, which results in lethargy, sluggish reflexes, constipation and loss of appetite, confusion, and in severe cases, coma.
Obviously, calcium homeostasis is critical. The skeletal, endocrine, and digestive systems play a role in this, but the kidneys do, too. These body systems work together to maintain a normal calcium level in the blood (Figure 6.24).
Figure 6.24Pathways in Calcium Homeostasis The body regulates calcium homeostasis with two pathways; one is signaled to turn on when blood calcium levels drop below normal and one is the pathway that is signaled to turn on when blood calcium levels are elevated.
Calcium is a chemical element that cannot be produced by any biological processes. The only way it can enter the body is through the diet. The bones act as a storage site for calcium: The body deposits calcium in the bones when blood levels get too high, and it releases calcium when blood levels drop too low. This process is regulated by PTH, vitamin D, and calcitonin.
Cells of the parathyroid gland have plasma membrane receptors for calcium. When calcium is not binding to these receptors, the cells release PTH, which stimulates osteoclast proliferation and resorption of bone by osteoclasts. This demineralization process releases calcium into the blood. PTH promotes reabsorption of calcium from the urine by the kidneys, so that the calcium returns to the blood. Finally, PTH stimulates the synthesis of vitamin D, which in turn, stimulates calcium absorption from any digested food in the small intestine.
When all these processes return blood calcium levels to normal, there is enough calcium to bind with the receptors on the surface of the cells of the parathyroid glands, and this cycle of events is turned off (Figure 6.24).
When blood levels of calcium get too high, the thyroid gland is stimulated to release calcitonin (Figure 6.24), which inhibits osteoclast activity and stimulates calcium uptake by the bones, but also decreases reabsorption of calcium by the kidneys. All of these actions lower blood levels of calcium. When blood calcium levels return to normal, the thyroid gland stops secreting calcitonin.
All of the organ systems of your body are interdependent, and the skeletal system is no exception. The food you take in via your digestive system and the hormones secreted by your endocrine system affect your bones. Even using your muscles to engage in exercise has an impact on your bones.
Exercise and Bone Tissue:
During long space missions, astronauts can lose approximately 1 to 2 percent of their bone mass per month. This loss of bone mass is thought to be caused by the lack of mechanical stress on astronauts’ bones due to the low gravitational forces in space. Lack of mechanical stress causes bones to lose mineral salts and collagen fibers, and thus strength. Similarly, mechanical stress stimulates the deposition of mineral salts and collagen fibers. The internal and external structure of a bone will change as stress increases or decreases so that the bone is an ideal size and weight for the amount of activity it endures. That is why people who exercise regularly have thicker bones than people who are more sedentary. It is also why a broken bone in a cast atrophies while its contralateral mate maintains its concentration of mineral salts and collagen fibers. The bones undergo remodeling as a result of forces (or lack of forces) placed on them.
Numerous, controlled studies have demonstrated that people who exercise regularly have greater bone density than those who are more sedentary. Any type of exercise will stimulate the deposition of more bone tissue, but resistance training has a greater effect than cardiovascular activities. Resistance training is especially important to slow down the eventual bone loss due to aging and for preventing osteoporosis.
Nutrition and Bone Tissue:
The vitamins and minerals contained in all of the food we consume are important for all of our organ systems. However, there are certain nutrients that affect bone health.
Calcium and Vitamin D:
You already know that calcium is a critical component of bone, especially in the form of calcium phosphate and calcium carbonate. Since the body cannot make calcium, it must be obtained from the diet. However, calcium cannot be absorbed from the small intestine without vitamin D. Therefore, intake of vitamin D is also critical to bone health. In addition to vitamin D’s role in calcium absorption, it also plays a role, though not as clearly understood, in bone remodeling.
Milk and other dairy foods are not the only sources of calcium. This important nutrient is also found in green leafy vegetables, broccoli, and intact salmon and canned sardines with their soft bones. Nuts, beans, seeds, and shellfish provide calcium in smaller quantities.
Except for fatty fish like salmon and tuna, or fortified milk or cereal, vitamin D is not found naturally in many foods. The action of sunlight on the skin triggers the body to produce its own vitamin D (Figure 6.22), but many people, especially those of darker complexion and those living in northern latitudes where the sun’s rays are not as strong, are deficient in vitamin D. In cases of deficiency, a doctor can prescribe a vitamin D supplement.
Figure 6.22 Synthesis of Vitamin D Sunlight is one source of vitamin D.
Vitamin K also supports bone mineralization and may have a synergistic role with vitamin D in the regulation of bone growth. Green leafy vegetables are a good source of vitamin K.
The minerals magnesium and fluoride may also play a role in supporting bone health. While magnesium is only found in trace amounts in the human body, more than 60 percent of it is in the skeleton, suggesting it plays a role in the structure of bone. Fluoride can displace the hydroxyl group in bone’s hydroxyapatite crystals and form fluorapatite. Similar to its effect on dental enamel, fluorapatite helps stabilize and strengthen bone mineral. Fluoride can also enter spaces within hydroxyapatite crystals, thus increasing their density.
Omega-3 fatty acids have long been known to reduce inflammation in various parts of the body. Inflammation can interfere with the function of osteoblasts, so consuming omega-3 fatty acids, in the diet or in supplements, may also help enhance production of new osseous tissue. Table 6.5 summarizes the role of nutrients in bone health.Nutrients and Bone Health
Role in bone health
Needed to make calcium phosphate and calcium carbonate, which form the hydroxyapatite crystals that give bone its hardness
Needed for calcium absorption
Supports bone mineralization; may have synergistic effect with vitamin D
Structural component of bone
Structural component of bone
Omega-3 fatty acids
Reduces inflammation that may interfere with osteoblast function
Hormones and Bone Tissue:
The endocrine system produces and secretes hormones, many of which interact with the skeletal system. These hormones are involved in controlling bone growth, maintaining bone once it is formed, and remodeling it.
Hormones That Influence Osteoblasts and/or Maintain the Matrix:
Several hormones are necessary for controlling bone growth and maintaining the bone matrix. The pituitary gland secretes growth hormone (GH), which, as its name implies, controls bone growth in several ways. It triggers chondrocyte proliferation in epiphyseal plates, resulting in the increasing length of long bones. GH also increases calcium retention, which enhances mineralization, and stimulates osteoblastic activity, which improves bone density.
GH is not alone in stimulating bone growth and maintaining osseous tissue. Thyroxine, a hormone secreted by the thyroid gland promotes osteoblastic activity and the synthesis of bone matrix. During puberty, the sex hormones (estrogen in girls, testosterone in boys) also come into play. They too promote osteoblastic activity and production of bone matrix, and in addition, are responsible for the growth spurt that often occurs during adolescence. They also promote the conversion of the epiphyseal plate to the epiphyseal line (i.e., cartilage to its bony remnant), thus bringing an end to the longitudinal growth of bones. Additionally, calcitriol, the active form of vitamin D, is produced by the kidneys and stimulates the absorption of calcium and phosphate from the digestive tract.
Hormones That Influence Osteoclasts:
Bone modeling and remodeling require osteoclasts to resorb unneeded, damaged, or old bone, and osteoblasts to lay down new bone. Two hormones that affect the osteoclasts are parathyroid hormone (PTH) and calcitonin.
PTH stimulates osteoclast proliferation and activity. As a result, calcium is released from the bones into the circulation, thus increasing the calcium ion concentration in the blood. PTH also promotes the reabsorption of calcium by the kidney tubules, which can affect calcium homeostasis (see below).
The small intestine is also affected by PTH, albeit indirectly. Because another function of PTH is to stimulate the synthesis of vitamin D, and because vitamin D promotes intestinal absorption of calcium, PTH indirectly increases calcium uptake by the small intestine. Calcitonin, a hormone secreted by the thyroid gland, has some effects that counteract those of PTH. Calcitonin inhibits osteoclast activity and stimulates calcium uptake by the bones, thus reducing the concentration of calcium ions in the blood. As evidenced by their opposing functions in maintaining calcium homeostasis, PTH and calcitonin are generally not secreted at the same time. Table 6.6 summarizes the hormones that influence the skeletal system. Hormones That Affect the Skeletal System
Increases length of long bones, enhances mineralization, and improves bone density
Stimulates bone growth and promotes synthesis of bone matrix
Promote osteoblastic activity and production of bone matrix; responsible for adolescent growth spurt; promote conversion of epiphyseal plate to epiphyseal line
Stimulates absorption of calcium and phosphate from digestive tract
Stimulates osteoclast proliferation and resorption of bone by osteoclasts; promotes reabsorption of calcium by kidney tubules; indirectly increases calcium absorption by small intestine
Inhibits osteoclast activity and stimulates calcium uptake by bones
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 (Figure 6.20). Table 6.4 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).
Figure 6.20Types 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.Types of Fractures
Type of fracture
Occurs straight across the long axis of the bone
Occurs at an angle that is not 90 degrees
Bone segments are pulled apart as a result of a twisting motion
Several breaks result in many small pieces between two large segments
One fragment is driven into the other, usually as a result of compression
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
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 6.21a). The disruption of blood flow to the bone results in the death of bone cells around the fracture.
Figure 6.21Stages 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 6.21b). 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 6.21c).
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 6.21d), and no external evidence of the fracture remains.
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.
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.
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 6.16Intramembranous 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.
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.
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.
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. Later discussions in this chapter will show that bone is also dynamic in that its shape adjusts to accommodate stresses. This section will examine the gross anatomy of bone first and then move on to its histology.
Gross Anatomy of Bone:
The structure of a long bone allows for the best visualization of all of the parts of a bone (Figure 6.7). 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.
Figure 6.7 Anatomy of a Long Bone A typical long bone shows the gross anatomical characteristics of 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 6.8). In this region, the epiphyses are covered with articular cartilage, a thin layer of cartilage that reduces friction and acts as a shock absorber.
Figure 6.8Periosteum and Endosteum The periosteum forms the outer surface of bone, and the endosteum lines the medullary cavity.
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 6.9). 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.
Figure 6.9Anatomy 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.
The surface features of bones vary considerably, depending on the function and location in the body. Table 6.2 describes the bone markings, which are illustrated in (Figure 6.10). There are three general classes of bone markings: (1) articulations, (2) projections, and (3) holes. As the name implies, an articulation is where two bone surfaces come together (articulus = “joint”). These surfaces tend to conform to one another, such as one being rounded and the other cupped, to facilitate the function of the articulation. A projection is an area of a bone that projects above the surface of the bone. These are the attachment points for tendons and ligaments. In general, their size and shape is an indication of the forces exerted through the attachment to the bone. A hole is an opening or groove in the bone that allows blood vessels and nerves to enter the bone. As with the other markings, their size and shape reflect the size of the vessels and nerves that penetrate the bone at these points.Bone Markings
Where two bones meet
Prominent rounded surface
Head of femur
Spinous process of the vertebrae
Transverse process of vertebra
Small, rounded process
Tubercle of humerus
Slight, elongated ridge
Temporal lines of the parietal bones
Holes and depressions
Foramen (holes through which blood vessels can pass through)
Fovea capitis on the head of the femur
Sigmoid sulcus of the temporal bones
Passage in bone
Slit through bone
Hole through bone
Foramen magnum in the occipital bone
Opening into canal
External auditory meatus
Air-filled space in bone
Figure 6.10Bone Features The surface features of bones depend on their function, location, attachment of ligaments and tendons, or the penetration of blood vessels and nerves.
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.
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 6.11).
Figure 6.11Bone 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.
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 6.3 reviews the bone cells, their functions, and locations.Bone Cells
Develop into osteoblasts
Deep layers of the periosteum and the marrow
Growing portions of bone, including periosteum and endosteum
Maintain mineral concentration of matrix
Entrapped in matrix
Bone surfaces and at sites of old, injured, or unneeded bone
Compact and Spongy Bone:
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 is the denser, stronger of the two types of bone tissue (Figure 6.12). It can be found under the periosteum and in the diaphyses of long bones, where it provides support and protection.
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.
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 6.13). 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.
Figure 6.13 Diagram of Spongy Bone Spongy bone is composed of trabeculae that contain the osteocytes. Red marrow fills the spaces in some bones.
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 6.15). 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.
Figure 6.15Diagram of Blood and Nerve Supply to Bone Blood vessels and nerves enter the bone through the nutrient foramen.
The 206 bones that compose the adult skeleton are divided into five categories based on their shapes (Figure 6.6). Their shapes and their functions are related such that each categorical shape of bone has a distinct function.
Figure 6.6 Classifications of Bones Bones are classified according to their shape.
A long bone is one that is cylindrical in shape, being longer than it is wide. Keep in mind, however, that the term describes the shape of a bone, not its size. Long bones are found in the arms (humerus, ulna, radius) and legs (femur, tibia, fibula), as well as in the fingers (metacarpals, phalanges) and toes (metatarsals, phalanges). Long bones function as levers; they move when muscles contract.
A short bone is one that is cube-like in shape, being approximately equal in length, width, and thickness. The only short bones in the human skeleton are in the carpals of the wrists and the tarsals of the ankles. Short bones provide stability and support as well as some limited motion.
The term “flat bone” is somewhat of a misnomer because, although a flat bone is typically thin, it is also often curved. Examples include the cranial (skull) bones, the scapulae (shoulder blades), the sternum (breastbone), and the ribs. Flat bones serve as points of attachment for muscles and often protect internal organs.
An irregular bone is one that does not have any easily characterized shape and therefore does not fit any other classification. These bones tend to have more complex shapes, like the vertebrae that support the spinal cord and protect it from compressive forces. Many facial bones, particularly the ones containing sinuses, are classified as irregular bones.
A sesamoid bone is a small, round bone that, as the name suggests, is shaped like a sesame seed. These bones form in tendons (the sheaths of tissue that connect bones to muscles) where a great deal of pressure is generated in a joint. The sesamoid bones protect tendons by helping them overcome compressive forces. Sesamoid bones vary in number and placement from person to person but are typically found in tendons associated with the feet, hands, and knees. The patellae (singular = patella) are the only sesamoid bones found in common with every person. Table 6.1 reviews bone classifications with their associated features, functions, and examples.Bone Classifications
Bone, or osseous tissue, is a hard, dense connective tissue that forms most of the adult skeleton, the support structure of the body. In the areas of the skeleton where bones move (for example, the ribcage and joints), cartilage, a semi-rigid form of connective tissue, provides flexibility and smooth surfaces for movement. The skeletal system is the body system composed of bones and cartilage and performs the following critical functions for the human body:
supports the body
protects internal organs
produces blood cells
stores and releases minerals and fat
Support, Movement, and Protection
The most apparent functions of the skeletal system are the gross functions—those visible by observation. Simply by looking at a person, you can see how the bones support, facilitate movement, and protect the human body.
Just as the steel beams of a building provide a scaffold to support its weight, the bones and cartilage of your skeletal system compose the scaffold that supports the rest of your body. Without the skeletal system, you would be a limp mass of organs, muscle, and skin.
Bones also facilitate movement by serving as points of attachment for your muscles. While some bones only serve as a support for the muscles, others also transmit the forces produced when your muscles contract. From a mechanical point of view, bones act as levers and joints serve as fulcrums (Figure 6.2). Unless a muscle spans a joint and contracts, a bone is not going to move. For information on the interaction of the skeletal and muscular systems, that is, the musculoskeletal system, seek additional content.
Figure 6.2Bones Support Movement Bones act as levers when muscles span a joint and contract.
Bones also protect internal organs from injury by covering or surrounding them. For example, your ribs protect your lungs and heart, the bones of your vertebral column (spine) protect your spinal cord, and the bones of your cranium (skull) protect your brain (Figure 6.3).
Figure 6.3 Bones Protect Brain The cranium completely surrounds and protects the brain from non-traumatic injury.
Mineral Storage, Energy Storage, and Hematopoiesis
On a metabolic level, bone tissue performs several critical functions. For one, the bone matrix acts as a reservoir for a number of minerals important to the functioning of the body, especially calcium, and phosphorus. These minerals, incorporated into bone tissue, can be released back into the bloodstream to maintain levels needed to support physiological processes. Calcium ions, for example, are essential for muscle contractions and controlling the flow of other ions involved in the transmission of nerve impulses.
Bone also serves as a site for fat storage and blood cell production. The softer connective tissue that fills the interior of most bone is referred to as bone marrow (Figure 6.5). There are two types of bone marrow: yellow marrow and red marrow. Yellow marrow contains adipose tissue; the triglycerides stored in the adipocytes of the tissue can serve as a source of energy. Red marrow is where hematopoiesis—the production of blood cells—takes place. Red blood cells, white blood cells, and platelets are all produced in the red marrow.
Figure 6.5 Head of Femur Showing Red and Yellow Marrow The head of the femur contains both yellow and red marrow. Yellow marrow stores fat. Red marrow is responsible for hematopoiesis.
Bones make good fossils. While the soft tissue of a once living organism will decay and fall away over time, bone tissue will, under the right conditions, undergo a process of mineralization, effectively turning the bone to stone. A well-preserved fossil skeleton can give us a good sense of the size and shape of an organism, just as your skeleton helps to define your size and shape. Unlike a fossil skeleton, however, your skeleton is a structure of living tissue that grows, repairs, and renews itself. The bones within it are dynamic and complex organs that serve a number of important functions, including some necessary to maintain homeostasis.