38.2: Bone

38.2.1: Bone

Learning Objective

Distinguish between compact and spongy bone tissues

Key Points

• Compact bone is the hard external layer of all bones that protects, strengthens, and surrounds the medullary cavity filled with marrow.
• Cylindrical structures, called osteons, are aligned along lines of the greatest stress to the bone in order to resist bending or fracturing.
• Spongy or cancellous bone tissue consists of trabeculae that are arranged as rods or plates with red bone marrow in between.
• Spongy bone is prominent in regions where the bone is less dense and at the ends of long bones where the bone has to be more compressible due to stresses that arrive from many directions.

Key Terms

epiphysis

the rounded end of any long bone

trabecula

a small mineralized spicule that forms a network in spongy bone

osteocyte

a mature bone cell involved with the maintenance of bone

osteon

any of the central canals and surrounding bony layers found in compact bone

Bone Tissue

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.

Compact Bone Tissue

Compact bone (or cortical bone), forming the hard external layer of all bones, surrounds the medullary cavity (innermost part or bone marrow). It provides protection and strength to bones . Compact bone tissue consists of units called osteons or Haversian systems. Osteons are cylindrical structures that contain a mineral matrix and living osteocytes connected by canaliculi which transport blood. They are aligned parallel to the long axis of the bone. Each osteon consists of lamellae, layers of compact matrix that surround a central canal (the Haversian or osteonic canal), which contains the bone's blood vessels and nerve fibers. Osteons in compact bone tissue are aligned in the same direction along lines of stress, helping the bone resist bending or fracturing. Therefore, compact bone tissue is prominent in areas of bone at which stresses are applied in only a few directions.

Components of compact bone tissue

Compact bone tissue consists of osteons that are aligned parallel to the long axis of the bone and the Haversian canal that contains the bone's blood vessels and nerve fibers. The inner layer of bones consists of spongy bone tissue. The small dark ovals in the osteon represent the living osteocytes.

Spongy Bone Tissue

Compact bone tissue forms the outer layer of all bones while spongy or cancellous bone forms the inner layer of all bones. Spongy bone tissue does not contain osteons. Instead, it consists of trabeculae, which are lamellae that are arranged as rods or plates . Red bone marrow is found between the trabuculae. Blood vessels within this tissue deliver nutrients to osteocytes and remove waste. The red bone marrow of the femur and the interior of other large bones, such as the ileum, forms blood cells.

Arrangement of trabeculae in spongy bone

Trabeculae in spongy bone are arranged such that one side of the bone bears tension and the other withstands compression.

Spongy bone reduces the density of bone, allowing the ends of long bones to compress as the result of stresses applied to the bone. Spongy bone is prominent in areas of bones that are not heavily stressed or where stresses arrive from many directions . The epiphysis of a bone, such as the neck of the femur, is subject to stress from many directions. Imagine laying a heavy-framed picture flat on the floor. You could hold up one side of the picture with a toothpick if the toothpick were perpendicular to the floor and the picture. Now, drill a hole and stick the toothpick into the wall to hang up the picture. In this case, the function of the toothpick is to transmit the downward pressure of the picture to the wall. The force on the picture is straight down to the floor, but the force on the toothpick is both the picture wire pulling down and the bottom of the hole in the wall pushing up. The toothpick will break off right at the wall.

The neck of the femur is horizontal like the toothpick in the wall. The weight of the body pushes it down near the joint, but the vertical diaphysis of the femur pushes it up at the other end. The neck of the femur must be strong enough to transfer the downward force of the body weight horizontally to the vertical shaft of the femur.

38.2.2: Cell Types in Bones

Learning Objective

Distinguish among the four cell types in bone

Key Points

• Osteogenic cells are the only bone cells that divide.
• Osteogenic cells differentiate and develop into osteoblasts which, in turn, are responsible for forming new bones.
• Osteoblasts synthesize and secrete a collagen matrix and calcium salts.
• When the area surrounding an osteoblast calcifies, the osteoblast becomes trapped and transforms into an osteocyte, the most common and mature type of bone cell.
• Osteoclasts, the cells that break down and reabsorb bone, stem from monocytes and macrophages rather than osteogenic cells..
• There is a continual balance between osteoblasts generating new bone and osteoclasts breaking down bone.

Key Terms

osteoblast

a mononucleate cell from which bone develops

lacuna

a small opening; a small pit or depression; a small blank space; a gap or vacancy; a hiatus

endosteum

a membranous vascular layer of cells which line the medullary cavity of a bone

periosteum

a membrane surrounding a bone

canaliculus

any of many small canals or ducts in bone or in some plants

osteoprogenitor

a stem cell that is the precursor of an osteoblast

osteocyte

a mature bone cell involved with the maintenance of bone

osteoclast

a large multinuclear cell associated with the resorption of bone

Cell Types in Bones

Bone consists of four types of cells: osteoblasts, osteoclasts, osteocytes, and osteoprogenitor (or osteogenic) cells . Each cell type has a unique function and is found in different locations in bones . The osteoblast, the bone cell responsible for forming new bone, 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 becomes trapped within it. As a result, it changes in structure, becoming an osteocyte, the primary cell of mature bone and the most common type of bone cell. Each osteocyte is located in a space (lacuna) surrounded by bone tissue. Osteocytes maintain the mineral concentration of the matrix via the secretion of enzymes. As is the case with osteoblasts, osteocytes lack mitotic activity. They are able to communicate with each other and receive nutrients via long cytoplasmic processes that extend through canaliculi (singular = canaliculus), channels within the bone matrix.

Bone cell types

Table listing the function and location of the four types of bone cells.

Four types of 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; they become osteocytes. Osteoclasts develop from monocytes and macrophages and differ in appearance from other bone cells.

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; they are the only bone cells that divide. Immature osteogenic cells are found in the deep layers of the periosteum and the marrow. When they differentiate, they develop into osteoblasts. The dynamic nature of bone means that new tissue is constantly formed, while old, injured, or unnecessary bone is dissolved for repair or for calcium release. The cell responsible for bone resorption, or breakdown, is the osteoclast, which is found on bone surfaces, is multinucleated, and originates from monocytes and macrophages (two types of white blood cells) rather than from osteogenic cells. Osteoclasts continually break down old bone while osteoblasts continually form new bone. The ongoing balance between osteoblasts and osteoclasts is responsible for the constant, but subtle, reshaping of bone.

38.2.3: Bone Development

Learning Objective

Distinguish between intramembranous and endochondral ossification

Key Points

• The ossification of the flat bones of the skull, the mandible, and the clavicles begins with mesenchymal cells, which then differentiate into calcium-secreting and bone matrix-secreting osteoblasts.
• Osteoids form spongy bone around blood vessels, which is later remodeled into a thin layer of compact bone.
• During enchondral ossification, the cartilage template in long bones is calcified; dying chondrocytes provide space for the development of spongy bone and the bone marrow cavity in the interior of the long bones.
• The periosteum, an irregular connective tissue around bones, aids in the attachment of tissues, tendons, and ligaments to the bone.
• Until adolescence, lengthwise long bone growth occurs in secondary ossification centers at the epiphyseal plates (growth plates) near the ends of the bones.

Key Terms

osteoid

an organic matrix of protein and polysaccharides, secreted by osteoblasts, that becomes bone after mineralization

endochondral

within cartilage

chondrocyte

a cell that makes up the tissue of cartilage

diaphysis

the central shaft of any long bone

Development of Bone

Ossification, or osteogenesis, is the process of bone formation by osteoblasts. Ossification is distinct from the process of calcification; whereas calcification takes place during the ossification of bones, it can also occur in other tissues. Ossification begins approximately six weeks after fertilization in an embryo. Before this time, the embryonic skeleton consists entirely of fibrous membranes and hyaline cartilage. The development of bone from fibrous membranes is called intramembranous ossification; development from hyaline cartilage is called endochondral ossification. Bone growth continues until approximately age 25. Bones can grow in thickness throughout life, but after age 25, ossification functions primarily in bone remodeling and repair.

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.

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 .

Process of endochondral ossification

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.

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, while 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.

38.2.4: Growth of Bone

Learning Objective

Describe the processes of post-fetal bone growth and bone thickening

Key Points

• The epiphyseal plate, the area of growth composed of four zones, is where cartilage is formed on the epiphyseal side while cartilage is ossified on the diaphyseal side, thereby lengthening the bone.
• Each of the four zones has a role in the proliferation, maturation, and calcification of bone cells that are added to the diaphysis.
• The longitudinal growth of long bones continues until early adulthood at which time the chondrocytes in the epiphyseal plate stop proliferating and the epiphyseal plate transforms into the epiphyseal line as bone replaces the cartilage.
• Bones can increase in diameter even after longitudinal growth has stopped.
• Appositional growth is the process by which old bone that lines the medullary cavity is reabsorbed and new bone tissue is grown beneath the periosteum, increasing bone diameter.

Key Terms

hypertrophy

to increase in size

chondrocyte

a cell that makes up the tissue of cartilage

ossification

the normal process by which bone is formed

periosteum

a membrane surrounding a bone

metaphysis

the part of a long bone that grows during development

diaphysis

the central shaft of any long bone

epiphysis

the rounded end of any long bone

medullary

pertaining to, consisting of, or resembling, marrow or medulla

Growth of Bone

Long bones continue to lengthen (potentially throughout adolescence) through the addition of bone tissue at the epiphyseal plate. They also increase in width through appositional growth.

Lengthening of Long Bones

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, allowing the diaphysis to grow in length. The metaphysis is the wide portion of a long bone between the epiphysis and the narrow diaphysis. It is considered a part of the growth plate: the part of the bone that grows during childhood, which, as it grows, ossifies near the diaphysis and the epiphyses.

The epiphyseal plate is composed of four zones of cells and activity.

1. The reserve zone, the region closest to the epiphyseal end of the plate, contains small chondrocytes within the matrix . These chondrocytes do not participate in bone growth; instead, they secure the epiphyseal plate to the osseous tissue of the epiphysis.
2. The proliferative zone, the next layer toward the diaphysis, contains stacks of slightly-larger chondrocytes . It continually makes new chondrocytes via mitosis.
3. The zone of maturation and hypertrophy contains chondrocytes that are older and larger than those in the proliferative zone . The more mature cells are situated closer to the diaphyseal end of the plate. In this zone, lipids, glycogen, and alkaline phosphatase accumulate, causing the cartilaginous matrix to calcify. The longitudinal growth of bone is a result of cellular division in the proliferative zone along with the maturation of cells in the zone of maturation and hypertrophy.
4. The zone of calcified matrix, the zone closest to the diaphysis, contains chondrocytes that are dead because the matrix around them has calcified . Capillaries and osteoblasts from the diaphysis penetrate this zone. 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.

After the zone of calcified matrix, there is the zone of ossification, which is actually part of the metaphysis . Arteries from the metaphysis branch through the newly-formed trabeculae in this zone. The newly-deposited bone tissue at the top of the zone of ossification is called the primary spongiosa. The older bone at the bottom of the zone of ossification is called the secondary spongiosa.

Longitudinal bone growth

The epiphyseal plate is responsible for longitudinal bone growth. This illustration shows the zones bordering the epiphyseal plate of the epiphysis. The topmost layer of the epiphysis is the reserve zone. The second zone, the proliferative zone, is where chondrocytes are continually undergoing mitosis. The next zone is the zone of maturation and hypertrophy where lipids, glycogen, and alkaline phosphatase accumulate, causing the cartilaginous matrix to calcify. The following zone is the calcified matrix where the chondrocytes have hardened and die as the matrix around them has calcified. The bottom-most row is the zone of ossification which is part of the metaphysis. The newly-deposited bone tissue at the top of the zone of ossification is called the primary spongiosa, while the older bone is labeled the secondary spongiosa.

Bones continue to grow in length until early adulthood with the rate of growth controlled by hormones. 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 .

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.

Thickening of Long Bones

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, cells that work to break down bone, resorb old bone that lines the medullary cavity. At the same time, 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.

38.2.5: Bone Remodeling and Repair

Learning Objective

Outline the process of bone remodeling and repair

Key Points

• Bone replacement involves the osteoclasts which break down bone and the osteoblasts which make new bone.
• Bone turnover rates differ depending on the bone and the area within the bone.
• There are four stages in the repair of a broken bone: 1) the formation of hematoma at the break, 2) the formation of a fibrocartilaginous callus, 3) the formation of a bony callus, and 4) remodeling and addition of compact bone.
• Proper bone growth and maintenance requires many vitamins (D, C, and A), minerals (calcium, phosphorous, and magnesium), and hormones (parathyroid hormone, growth hormone, and calcitonin).

Key Terms

spicule

a sharp, needle-like piece

callus

the material of repair in fractures of bone which is at first soft or cartilaginous in consistency, but is ultimately converted into true bone and unites the fragments into a single piece

fibroblast

a cell found in connective tissue that produces fibers, such as collagen

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 or bone production done by osteoblasts and bone resorption done by osteoclasts, which break down old bone. 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, the rates at which old bone is replaced by new bone, are quite high, with five to seven percent of bone mass being recycled every week. Differences in turnover rates 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 everyday stress (for example, when a limb is in a cast) will begin to lose mass.

Stages of 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.

A fractured or broken bone undergoes repair through four stages:

1. Hematoma formation: Blood vessels in the broken bone tear and hemorrhage, resulting in the formation of clotted blood, or a hematoma, at the site of the break. The severed blood vessels at the broken ends of the bone are sealed by the clotting process. Bone cells deprived of nutrients begin to die.
2. Bone generation: Within days of the fracture, capillaries grow into the hematoma, while phagocytic cells begin to clear away the dead cells. Though fragments of the blood clot may remain, fibroblasts and osteoblasts enter the area and begin to reform bone. Fibroblasts produce collagen fibers that connect the broken bone ends, while osteoblasts start to form spongy bone. The repair tissue between the broken bone ends, the fibrocartilaginous callus, is composed of both hyaline and fibrocartilage . Some bone spicules may also appear at this point.
3. Bony callous formation: The fibrocartilaginous callus is converted into a bony callus of spongy bone. It takes about two months for the broken bone ends to be firmly joined together after the fracture. This is similar to the endochondral formation of bone when cartilage becomes ossified; osteoblasts, osteoclasts, and bone matrix are present.
4. Bone remodeling: The bony callus is then remodelled by osteoclasts and osteoblasts, with excess material on the exterior of the bone and within the medullary cavity being removed. Compact bone is added to create bone tissue that is similar to the original, unbroken bone. This remodeling can take many months; the bone may remain uneven for years.

Attributions

• Bone
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  • "osteon." http://en.wiktionary.org/wiki/osteon. Wiktionary CC BY-SA 3.0.
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  • "osteocyte." http://en.wiktionary.org/wiki/osteocyte. Wiktionary CC BY-SA 3.0.
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  • "OpenStax College, Biology. October 17, 2013." http://cnx.org/content/m44789/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "OpenStax College, Bone. October 17, 2013." http://cnx.org/content/m44789/latest/Figure_38_02_04.png. OpenStax CNX CC BY 3.0.
  • "OpenStax College, Bone. October 17, 2013." http://cnx.org/content/m44789/latest/Figure_38_02_05.jpg. OpenStax CNX CC BY 3.0.
• Cell Types in Bones
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• Bone Development
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  • "diaphysis." http://en.wiktionary.org/wiki/diaphysis. Wiktionary CC BY-SA 3.0.
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• Growth of Bone
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  • "hypertrophy." http://en.wiktionary.org/wiki/hypertrophy. Wiktionary CC BY-SA 3.0.
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• Bone Remodeling and Repair
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  • "fibroblast." http://en.wiktionary.org/wiki/fibroblast. Wiktionary CC BY-SA 3.0.
  • "callus." http://en.wiktionary.org/wiki/callus. Wiktionary CC BY-SA 3.0.
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  • "spicule." http://en.wiktionary.org/wiki/spicule. Wiktionary CC BY-SA 3.0.
  • "Stages of Fracture Repair." http://commons.wikimedia.org/wiki/File:613_Stages_of_Fracture_Repair.jpg. Wikimedia Commons CC BY-SA 3.0.