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5

The Skeletal System:

Osseous Tissue and

Skeletal Structure

Introduction

The skeletal system is made of:

• Skeletal bones

• Cartilage

• Ligaments

• Connective tissue to stabilize the skeleton

Bones are dynamic organs, which consist of several tissue types

Introduction

Functions of the skeletal system

• Support

• Provides the framework for the attachment of other organs

• Storage of minerals

• Calcium ions: 98% of the body’s calcium ions are in the bones

• Phosphate ions

• Blood cell production

• Bone marrow produces erythrocytes, leukocytes, and platelets

Introduction

Functions of the skeletal system (continued)

• Leverage

• Muscles pull on the bones to produce movement

• Protection

• Ribs protect heart and lungs

• Skull protects the brain

• Vertebrae protect the spinal cord

• Pelvic bones protect the reproductive organs

Anatomy of Skeletal Elements

There are seven broad categories of bones according to their shapes

• Sutural bones

• Irregular bones

• Short bones

• Pneumatized bones

• Flat bones

• Long bones

• Sesamoid bones

Classification of Bones

Long Bones: longer than they are wide, upper and lower limbs.

Short Bones: Broad as they are long, cube shaped, round, i.e. tarsals & carpals

Flat Bones: Thin, flat and curved, skull (parietal), sternum, scapulae.

Irregular Bones: Don’t fit in the others, face, vertebrae

Sesamoid: Sesame seed, patella, small and flat

Sutural bones: Wormian bones, borders like a jigsaw puzzle.

Figure 5.11 Shapes of Bones

Sutural Bones

Sutures

Sutural bone

Pneumatized Bones

Ethmoid

Air cells

Irregular Bones

Vertebra

Short Bones

Carpal bones

Flat Bones

Parietal bone External table

Internal table

Diploë

(spongy bone)

Long Bones

Sesamoid Bones

Humerus

Patella

Structure of Bone

Bones (osseous tissue)

• Supporting connective tissue

• Specialized cells

• Solid matrix

• Outer lining

• Called the periosteum

• Inner lining

• Called the endosteum

Structure of Bone

The Histological Organization of Mature Bone

• The matrix

• Calcium phosphate eventually converts to

hydroxyapatite crystals

• Hydroxyapatite crystals resist compression

Structure of Bone

The Histological Organization of Mature Bone

• Collagen fibers

• Make up 2/3 of the bone matrix

• Contribute to the tensile strength of bones

• Collagen and hydroxyapatite make bone tissue extremely strong

• Bone cells

• Contribute only 2% of the bone mass

Structure of Bone

The Cells of Mature Bone

• Osteocytes

• Mature bone cells

• Maintain the protein and mineral content of the matrix

• Osteoblasts

• Immature bone cells

• Found on the inner and outer surfaces of bones

• Produce osteoid, which is involved in making the matrix

• Osteoblasts are involved in making new bone. This is a process called osteogenesis

• Osteoblasts can convert to osteocytes

Structure of Bone

The Cells of Mature Bone (continued)

• Osteoprogenitor cells

• Found on the inner and outer surfaces of bones

• Differentiate to form new osteoblasts

• Heavily involved in the repair of bones after a break

• Osteoclasts

• Secrete acids, which dissolve the bones thereby causing the release of stored calcium ions and phosphate ions into the blood

• This process is called osteolysis

Figure 5.1a Histological Structure of a Typical Bone

Canaliculi Osteocyte Matrix

Osteocyte: Mature bone cell that maintains the bone matrix

Osteoblast Osteoid Matrix

Osteoblast: Immature bone cell that secretes organic components of matrix

The cells of bone

Endosteum Osteoprogenitor cell

Medullary cavity

Osteoprogenitor cell:

Stem cell whose divisions produce osteoblasts

Osteoclast Matrix

Medullary cavity

Osteoclast: Multinucleate cell that secretes acids and enzymes to dissolve bone matrix

Structure of Bone

The Osteon

• It is the basic unit of skeletal bones

• Consists of:

• Central canal

• Canaliculi

• Osteocytes

• Lacunae

• Lamellae

Figure 5.1c Histological Structure of a Typical Bone

Osteons LM

220

A thin section through compact bone; in this procedure the intact matrix and central canals appear white, and the lacunae and canaliculi are shown in black.

Canaliculi

Concentric lamellae

Central canal

Osteon

Lacunae

Figure 5.1d Histological Structure of a Typical Bone

Canaliculi

Concentric lamellae

Central canal

Osteon

Lacunae

Osteon

A single osteon at higher magnification

LM

343

Figure 5.1b Histological Structure of a Typical Bone

Osteon

Lacunae

Central canals

Osteons SEM

182

A scanning electron micrograph of several osteons in compact bone

Lamellae

Structure of Bone

Two types of osseous tissue

Compact bone (dense bone)

• Compact bones are dense and solid

• Forms the walls of bone outlining the medullary cavity

• Medullary cavity consists of bone marrow

Spongy bone (trabecular bone)

• Open network of plates

Figure 5.2a-c The Internal Organization in Representative Bones

Spongy bone

Blood vessels

Compact bone

Medullary cavity

Endosteum

Collagen fiber orientation

Concentric lamellae

The organization of collagen fibers within concentric lamellae

Periosteum

Gross anatomy of the humerus

Compact bone

Spongy bone

Medullary cavity

Concentric lamellae

Interstitial lamellae

Capillary

Small vein

Central canal

Endosteum

Circumferential lamellae

Osteons

Periosteum

Trabeculae of spongy bone

Artery Vein

Perforating canal

Central canal

Diagrammatic view of the histological organization of compact and spongy bone

Figure 5.2d The Internal Organization in Representative Bones

Trabeculae of spongy bone

Canaliculi opening on surface

Location and structure of spongy bone. The photo shows a sectional view of the proximal end of the femur.

Endosteum

Lamellae

Structure of Bone

Structural Differences

• Compact bone

• Consists of osteons

• Makes up the dense, solid portion of bone

• Spongy bone

• Trabeculae are arranged in parallel struts

• Trabeculae form branching plates

• Trabeculae form an open network

• Creates the lightweight nature of bones

Structure of Bone

Functional Differences

• Compact bone

• Conducts stress from one area of the body to another area of the body

• Generates tremendous strength from end to end

• Weak strength when stress is applied to the side

• Spongy bone

• Trabeculae create strength to deal with stress from the side

Figure 5.3a Anatomy of a Representative Bone

Epiphysis

Metaphysis Articular surface of head of femur

Spongy bone

Compact bone

Diaphysis

(shaft)

Metaphysis

Epiphysis

Posterior view Sectional view

The femur, or thigh bone, in superficial and sectional views. The femur has a diaphysis (shaft) with walls of compact bone and epiphyses (ends) filled with spongy bone. A metaphysis separates the diaphysis and epiphysis at each end of the shaft. The body weight is transferred to the femur at the hip joint.

Because the hip joint is off center relative to the axis of the shaft, the body weight is distributed along the bone so that the medial portion of the shaft is compressed and the lateral portion is stretched.

Medullary cavity

Figure 5.3b Anatomy of a Representative Bone

An intact femur chemically cleared to show the orientation of the trabeculae in the epiphysis

Figure 5.3c Anatomy of a Representative Bone

Articular surface of head of femur

Trabeculae of spongy bone

Cortex

Medullary cavity

Compact bone

A photograph showing the epiphysis after sectioning

Structure of Bone

Organization of Compact and Spongy Bone

• Epiphysis

• Each end of the long bones

• Diaphysis

• Shaft of the long bones

• Metaphysis

• Narrow growth zone between the epiphysis and the diaphysis

Figure 5.3a Anatomy of a Representative Bone

Epiphysis

Metaphysis Articular surface of head of femur

Spongy bone

Compact bone

Diaphysis

(shaft)

Metaphysis

Epiphysis

Posterior view Sectional view

The femur, or thigh bone, in superficial and sectional views. The femur has a diaphysis (shaft) with walls of compact bone and epiphyses (ends) filled with spongy bone. A metaphysis separates the diaphysis and epiphysis at each end of the shaft. The body weight is transferred to the femur at the hip joint.

Because the hip joint is off center relative to the axis of the shaft, the body weight is distributed along the bone so that the medial portion of the shaft is compressed and the lateral portion is stretched.

Medullary cavity

Structure of Bone

The Periosteum and Endosteum

• Periosteum

• Outer surface of the bone

• Isolates and protects the bone from surrounding tissue

• Provides a route and a place for attachment for circulatory and nervous supply

• Actively participates in bone growth and repair

• Attaches the bone to the connective tissue network of the deep fascia

Structure of Bone

The Periosteum and Endosteum

• Periosteum and Tendons

• Tendons are cemented into the lamellae by osteoblasts

• Therefore, tendons are actually a part of the bone

Figure 5.4c Anatomy and Histology of the Periosteum and Endosteum

LM

100

Zone of tendon  bone attachment

Tendon

Periosteum

Medullary cavity

Endosteum

Spongy bone of epiphysis

Epiphyseal cartilage

A tendon

 bone junction

Structure of Bone

The Periosteum and Endosteum

• Endosteum

• Inner surface of bone

• Lines the medullary cavity

• Consists of osteoprogenitor cells

• Actively involved in repair and growth

Figure 5.4ab Anatomy and Histology of the Periosteum and Endosteum

Endosteum

Joint capsule

Cellular layer of periosteum

Fibrous layer of periosteum

Compact bone

Circumferential lamellae

Cellular layer of periosteum

Fibrous layer of periosteum

Canaliculi

Lacuna

Osteocyte

Perforating fibers

The periosteum contains outer

(fibrous) and inner (cellular) layers.

Collagen fibers of the periosteum are continuous with those of the bone, adjacent joint capsules, and attached tendons and ligaments.

Bone matrix

Giant multinucleate osteoclast

Endosteum

Osteoprogenitor cell

Osteocyte

Osteoid

Osteoblasts

The endosteum is an incomplete cellular layer containing osteoblasts, osteoprogenitor cells, and osteoclasts.

Bone Development and Growth

Before six weeks of development, the skeleton is cartilage

• Cartilage cells will be replaced by bone cells

• This is called ossification

Osteogenesis

• Bone formation

Calcification

• The deposition of calcium ions into the bone tissue

Bone Development and Growth

There are two types of ossification

• Intramembranous ossification

• Involved in the development of clavicle, mandible, skull, and face

• Endochondral ossification

• Involved in the development of limbs, vertebrae, and hips

Bone Development and Growth

Intramembranous ossification

• Mesenchymal cells differentiate to form osteoblasts

• Osteoblasts begin secreting a matrix

• Osteoblasts become trapped in the matrix

• Osteoblasts differentiate and form osteocytes

• More osteoblasts are produced, thus move outward

• Eventually, compact bone is formed

Figure 5.5 Histology of Intramembranous Ossification

Mesenchymal cells aggregate, differentiate into osteoblasts, and begin the ossification process. The bone expands as a series of spicules that spread into surrounding tissues.

Osteocyte in lacuna

Bone matrix

Osteoblast

Osteoid

Embryonic connective tissue

Mesenchymal cell

Blood vessel

As the spicules interconnect, they trap blood vessels within the bone.

Osteocytes in lacunae

Blood vessels

Osteoblast layer

Over time, the bone assumes the structure of spongy bone. Areas of spongy bone may later be removed, creating medullary cavities.

Through remodeling, spongy bone formed in this way can be converted to compact bone.

Blood vessel

Blood vessel Osteoblasts Spicules

LM  32 LM  32

Bone Development and Growth

Endochondral ossification

• The developing bone begins as cartilage cells

• Cartilage matrix grows inward

• Interstitial growth

• Cartilage matrix grows outward

• Appositional growth

• Blood vessels grow around the cartilage

Bone Development and Growth

Endochondral ossification (continued)

• Perichondrial cells convert to osteoblasts

• Osteoblasts develop a superficial layer of bone around the cartilage

• Blood vessels penetrate the cartilage

• Osteoblasts begin to develop spongy bone in the diaphysis

• This becomes the primary center of ossification

Figure 5.7a Anatomical and Histological Organization of Endochondral Ossification (Part 1 of 2)

As the cartilage enlarges, chondrocytes near the center of the shaft increase greatly in size.

The matrix is reduced to a series of small struts that soon begin to calcify. The enlarged chondrocytes then die and disintegrate, leaving cavities within the cartilage.

Enlarging chondrocytes within calcifying matrix

Blood vessels grow around the edges of the cartilage, and the cells of the perichondrium convert to osteoblasts.

The shaft of the cartilage then becomes ensheathed in a superficial layer of bone.

Epiphysis

Diaphysis

Bone formation

Hyaline cartilage

Steps in the formation of a long bone from a hyaline cartilage model

Figure 5.7a Anatomical and Histological Organization of Endochondral Ossification (Part 2 of 2)

Blood vessels penetrate the cartilage and invade the central region. Fibroblasts migrating with the blood vessels differentiate into osteoblasts and begin producing spongy bone at a primary center of ossification. Bone formation then spreads along the shaft toward both ends.

Remodeling occurs as growth continues, creating a medullary cavity. The bone of the shaft becomes thicker, and the cartilage near each epiphysis is replaced by shafts of bone. Further growth involves increases in length (Steps 5 and 6) and diameter (see Figure 5.9).

Blood vessel

Medullary cavity

Primary ossification center

Superficial bone

Spongy bone

Medullary cavity

See

Figure 5.9

Metaphysis

Steps in the formation of a long bone from a hyaline cartilage model

Bone Development and Growth

Endochondral ossification (continued)

• The cartilage near the epiphysis converts to bone

• Blood vessels penetrate the epiphysis

• Osteoblasts begin to develop spongy bone in the epiphysis

• Epiphysis becomes the secondary center of ossification

Figure 5.7b Anatomical and Histological Organization of Endochondral Ossification (Part 1 of 2)

Capillaries and osteoblasts migrate into the epiphyses, creating secondary ossification centers.

Hyaline cartilage

Epiphysis

Soon the epiphyses are filled with spongy bone. An articular cartilage remains exposed to the joint cavity; over time it will be reduced to a thin superficial layer. At each metaphysis, an epiphyseal cartilage separates the epiphysis from the diaphysis.

Articular cartilage

Spongy bone

Metaphysis

Periosteum

Compact bone

Epiphyseal cartilage

Diaphysis

Secondary ossification center

Bone Development and Growth

Epiphyseal plate

• Area of cartilage in the metaphysis

• Also called the epiphyseal cartilage

• Cartilage near the diaphysis is converted to bone

• The width of this zone gets narrower as we age

Figure 5.7b Anatomical and Histological Organization of Endochondral Ossification (Part 2 of 2)

Epiphyseal cartilage matrix

Cartilage cells undergoing division

Zone of proliferation

Zone of hypertrophy

Medullary cavity

Osteoblasts Osteoid

Epiphyseal cartilage LM

250

Light micrograph showing the zones of cartilage and the advancing osteoblasts at an epiphyseal cartilage

Figure 5.7 Anatomical and Histological Organization of Endochondral Ossification

As the cartilage enlarges, chondrocytes near the center of the shaft increase greatly in size.

The matrix is reduced to a series of small struts that soon begin to calcify. The enlarged chondrocytes then die and disintegrate, leaving cavities within the cartilage.

Enlarging chondrocytes within calcifying matrix

Hyaline cartilage

Blood vessels grow around the edges of the cartilage, and the cells of the perichondrium convert to osteoblasts. The shaft of the cartilage then becomes ensheathed in a superficial layer of bone.

Bone formation

Epiphysis

Diaphysis

Blood vessels penetrate the cartilage and invade the central region. Fibroblasts migrating with the blood vessels differentiate into osteoblasts and begin producing spongy bone at a primary center of ossification. Bone formation then spreads along the shaft toward both ends.

Blood vessel

Medullary cavity

Primary ossification center

Superficial bone

Spongy bone

Remodeling occurs as growth continues, creating a medullary cavity. The bone of the shaft becomes thicker, and the cartilage near each epiphysis is replaced by shafts of bone. Further growth involves increases in length (Steps 5 and 6) and diameter (see Figure 5.9).

Medullary cavity

Capillaries and osteoblasts migrate into the epiphyses, creating secondary ossification centers.

Hyaline cartilage

Epiphysis

Metaphysis

Periosteum

Compact bone

See

Figure 5.9

Metaphysis

Secondary ossification center

Steps in the formation of a long bone from a hyaline cartilage model

Soon the epiphyses are filled with spongy bone. An articular cartilage remains exposed to the joint cavity; over time it will be reduced to a thin superficial layer. At each metaphysis, an epiphyseal cartilage separates the epiphysis from the diaphysis.

Articular cartilage

Spongy bone

Epiphyseal cartilage

Diaphysis

Epiphyseal cartilage matrix

Cartilage cells undergoing division

Medullary cavity

Osteoblasts

Epiphyseal cartilage

Osteoid

LM

250

Light micrograph showing the zones of cartilage and the advancing osteoblasts at an epiphyseal cartilage

Zone of proliferation

Zone of hypertrophy

Figure 5.8 Epiphyseal Cartilages and Lines

X-ray of the hand of a young child. The arrows indicate the locations of the epiphyseal cartilages.

X-ray of the hand of an adult. The arrows indicate the locations of epiphyseal lines.

Figure 5.6 Fetal Intramembranous and Endochondral Ossification

Intramembranous ossification produces the roofing bones of the skull

At 10 weeks the fetal skull clearly shows both membrane and cartilaginous bone, but the boundaries that indicate the limits of future skull bones have yet to be established.

Endochondral ossification replaces cartilages of embryonic skull

Primary ossification centers of the diaphyses (bones of the lower limb)

Future hip bone

Temporal bone

Mandible

Clavicle

Scapula

Humerus

Ribs

Vertebrae

Hip bone

(ilium)

Femur

Parietal bone

Frontal bone

Metacarpal bones

Phalanges

Radius

Ulna

Cartilage

Fibula

Tibia

Phalanx

Metatarsal bones

At 16 weeks the fetal skull shows the irregular margins of the future skull bones. Most elements of the appendicular skeleton form through endochondral ossification. Note the appearance of the wrist and ankle bones at 16 weeks versus at 10 weeks.

Bone Development and Growth

Enlarging the diameter of bone

• Called appositional growth

• Blood vessels that run parallel to the bone becomes surrounded by bone cells

• “Tunnels” begin to form

• Each “tunnel” has a blood vessel in it

Bone Development and Growth

Enlarging the diameter of bone

• Osteoblasts begin to produce matrix, thus creating concentric rings

• As osteoblasts are laying down more bone material, osteoclasts are dissolving the inner bone, thus creating the marrow cavity

Figure 5.9a Appositional Bone Growth (Part 1 of 2)

Bone formation at the surface of the bone produces ridges that parallel a blood vessel.

Ridge Periosteum

The ridges enlarge and create a deep pocket.

Perforating canal

Artery

Three-dimensional diagrams illustrate the mechanism responsible for increasing the diameter of a growing bone.

The ridges meet and fuse, trapping the vessel inside the bone.

Figure 5.9a Appositional Bone Growth (Part 2 of 2)

Bone deposition proceeds inward toward the vessel, beginning the creation of a typical osteon.

Additional circumferential lamellae are deposited and the bone continues to increase in diameter.

Circumferential lamellae

Osteon is complete with new central canal around blood vessel. Second blood vessel becomes enclosed.

Periosteum

Central canal of new osteon

Three-dimensional diagrams illustrate the mechanism responsible for increasing the diameter of a growing bone.

Figure 5.9b Appositional Bone Growth

Bone resorbed by osteoclasts

Infant

Bone deposited by osteoblasts

Child

A bone grows in diameter as new bone is added to the outer surface. At the same time, osteoclasts resorb bone on the inside, enlarging the medullary cavity.

Young adult Adult

Bone Development and Growth

There are four major sets of blood vessels associated with the long bones

• Nutrient vessels

• Enter the diaphysis and branch toward the epiphysis

• Re-enter the compact bone, leading to the central canals of the osteons

• Metaphyseal vessels

• Supply nutrients to the diaphyseal edge of the epiphysis

Bone Development and Growth

Four major sets of blood vessels (continued)

• Epiphyseal vessels

• Supply nutrients to the medullary cavities of the epiphysis

• Periosteal vessels

• Supply nutrients to the superficial osteons

Figure 5.10 Circulatory Supply to a Mature Bone

Branches of nutrient artery and vein

Periosteum

Periosteal arteries and veins

Connections to superficial osteons

Nutrient artery and vein

Nutrient foramen

Metaphyseal artery and vein

Articular cartilage

Epiphyseal artery and vein

Metaphyseal artery and vein

Periosteum

Compact bone

Medullary cavity

Metaphysis

Epiphyseal line

Bone Development and Growth

Factors Regulating Bone Growth

• Nutrition

• Calcium ions

• Phosphate ions

• Magnesium ions

• Citrate

• Carbonate ions

• Sodium ions

• Vitamins A, C, D (calcitriol)

Bone Development and Growth

Factors Regulating Bone Growth (continued)

• Hormones: Parathyroid gland

• Releases parathyroid hormone

• Stimulates osteoclasts

• Stimulates osteoblasts

• Increases calcium ion absorption from the small intestine to the blood

Bone Development and Growth

Factors Regulating Bone Growth (continued)

• Hormones: Thyroid gland

Releases calcitonin

• Inhibits osteoclasts

• Removes calcium ions from blood and adds it to bone

Bone Development and Growth

Factors Regulating Bone Growth (continued)

• Hormones: Thyroid gland

• Releases thyroxine (T

4

)

• Maintains normal activity of the epiphyseal cartilage

Bone Development and Growth

Factors Regulating Bone Growth (continued)

• Hormones: Pituitary gland

• Releases growth hormone (somatotropin)

• Maintains normal activity of the epiphyseal cartilage

Bone Maintenance, Remodeling, and Repair

Aging and the Skeletal System

• When we’re young, osteoblast activity balances with osteoclast activity

• When we get older, osteoblast activity slows faster than osteoclast activity

• When osteoclast activity is faster than osteoblast activity, bones become porous

• Estrogen keeps osteoclast activity under control

Bone Maintenance, Remodeling, and Repair

Aging and the Skeletal System

• As women age, estrogen levels drop

• Osteoclast control is lost

• Osteoclasts are overactive

• Bones become porous

• This is osteoporosis

Bone Maintenance, Remodeling, and Repair

Injury and Repair

• When a bone is broken, bleeding occurs

• A network of spongy bone forms

• Osteoblasts are overly activated, thus resulting in enlarged callused area

• This area is now stronger and thicker than normal bone

Clinical Note 5.3 Fractures and Their Repair (Part 3 of 4)

Repair of a fracture

Fracture hematoma

Dead bone

Bone fragments

Immediately after the fracture, extensive bleeding occurs.

Over a period of several hours, a large blood clot, or fracture hematoma, develops.

Spongy bone of external callus

Periosteum

An internal callus forms as a network of spongy bone units the inner edges, and an external callus of cartilage and bone stabilizes the outer edges.

Clinical Note 5.3 Fractures and Their Repair (Part 4 of 4)

Internal callus

External callus

The cartilage of the external callus has been replaced by bone, and struts of spongy bone now unite the broken ends. Fragments of dead bone and the areas of bone closest to the break have been removed and replaced.

External callus

A swelling initially marks the location of the fracture.

Over time, this region will be remodeled, and little evidence of the fracture will remain.

Anatomy of Skeletal Elements

Bone markings include:

• Projections

• Depressions

• Fissures

• Foramina

• Canals (meatuses)

Figure 5.12a Examples of Bone Markings (Surface Features)

Trochanter

Head

Neck

Facet

Tubercle

Femur

Condyle

Figure 5.12b Examples of Bone Markings (Surface Features)

Fissure

Ramus

Skull, anterior view

Process

Foramen

Figure 5.12c Examples of Bone Markings (Surface Features)

Canal

Sinuses

Meatus

Skull, sagittal section

Figure 5.12d Examples of Bone Markings (Surface Features)

Tubercle

Head

Sulcus

Neck

Tuberosity

Condyle

Humerus

Fossa

Trochlea

Figure 5.12e Examples of Bone Markings (Surface Features)

Spine

Line

Foramen

Pelvis

Ramus

Crest

Fossa

Table 5.1 Common Bone Marking Terminology

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