A bone dating from the Pleistocene Ice Age of an extinct species of elephant.
A scanning electronic micrograph of bone at 10,000x magnification.
Bones are rigid organs that constitute part of the endoskeleton of vertebrates. They support and protect the various organs of the body, produce red and white blood cells and store minerals and also enable mobility. Bone tissue is a type of dense connective tissue. Bones come in a variety of shapes and sizes and have a complex internal and external structure, are lightweight yet strong and hard, and serve multiple functions. One of the types of tissue that makes up bone is the mineralized osseous tissue, also called bone tissue, that gives it rigidity and a coral-like three-dimensional internal structure. Other types of tissue found in bones include marrow, endosteum, periosteum, nerves, blood vessels and cartilage.
In the human at birth, there are over 270 bones, but many of these fuse together during development, leaving a total of 206 separate bones in the adult, not counting numerous small sesamoid bones and ossicles. The largest bone in the human body is the thigh-bone (femur) and the smallest bone of the 206 is the stapes.
- 1 Structure
- 2 Function
- 3 Clinical significance
- 4 Society and culture
- 5 Other animals
- 6 See also
- 7 References
- 8 Footnotes
- 9 External links
The primary tissue of bone, osseous tissue, is relatively hard and lightweight. It is mostly made up of a composite material incorporating the mineral calcium phosphate in the chemical arrangement termed calcium hydroxylapatite (this is the osseous tissue that gives bones their rigidity) and collagen, an elastic protein which improves fracture resistance.
Bone is not a uniformly solid material, but is rather a complex set of materials.
There are five types of bones in the human body: long, short, flat, irregular, and sesamoid.
- Long bones are characterized by a shaft, the diaphysis, that is much longer than it is wide. They are made up mostly of compact bone, with lesser amounts of marrow, located within the medullary cavity, and spongy bone. Most bones of the limbs, including those of the fingers and toes, are long bones. The exceptions are those of the wrist, ankle and kneecap.
- Short bones are roughly cube-shaped, and have only a thin layer of compact bone surrounding a spongy interior. The bones of the wrist and ankle are short bones, as are the sesamoid bones.
- Flat bones are thin and generally curved, with two parallel layers of compact bones sandwiching a layer of spongy bone. Most of the bones of the skull are flat bones, as is the sternum.
- Sesamoid bones are bones embedded in tendons. Since they act to hold the tendon further away from the joint, the angle of the tendon is increased and thus the leverage of the muscle is increased. Examples of sesamoid bones are the patella and the pisiform.
- Irregular bones do not fit into the above categories. They consist of thin layers of compact bone surrounding a spongy interior. As implied by the name, their shapes are irregular and complicated. Often this irregular shape is due to their many centers of ossification or because they contain bony sinuses. The bones of the spine, Pelvis, and some bones of the skull are irregular bones. Examples include the ethmoid and sphenoid bones.
The hard outer layer of bones is composed of compact bone tissue, so-called due to its minimal gaps and spaces. The hard outer layer gives bones its smooth, white, and solid appearance, and accounts for 80% of the total bone mass of an adult skeleton.
Compact bone consists of multiple microscopic columns, each called an osteon. Each column is multiple layers of osteoblasts and osteocytes around a central canal called the Haversian canal. Volkmann's canals at right angles connect the osteons together. The columns are metabolically active, and as bone is reabsorbed and created the nature and location of the cells within the osteon will change. Compact bone is covered by a periosteum on its outer surface, and an endosteum on its inner surface. The endosteum is the boundary between the compact band trabecular bone. 
Filling the interior of the bone is the trabecular bone tissue. It is an open cell porous network also called cancellous or spongy bone. Thin formations of osteoblasts covered in endosteum create an irregular network of spaces. Within these spaces are bone marrow and haematopoetic stem cells that give rise to platelets, red blood cells and white blood cells. Trabecular marrow is composed of a network of rod- and plate-like elements that make the overall organ lighter and allow room for blood vessels and marrow. Trabecular bone accounts for the remaining 20% of total bone mass but has nearly ten times the surface area of compact bone.
Bone marrow can be found in almost any bone that holds cancellous tissue. In newborns, all such bones are filled exclusively with red marrow, but as the child ages it is mostly replaced by yellow, or fatty marrow. In adults, red marrow is mostly found in the marrow bones of the femur, the ribs, the vertebrae and pelvic bones.
The majority of bone is made of the bone matrix. It is composed primarily of inorganic hydroxyapatite and organic collagen. Bone is formed by the hardening of this matrix around entrapped cells. When these cells become entrapped from osteoblasts they become osteocytes.
Bone is a metabolically active tissue composed of several types of cells. Cells that make up bone tissue include osteoblasts, which are involved in the creation and mineralisation of bone tissue, and osteoclasts, which are involved in the reabsorption of bone tissue. Osteoblasts and osteocytes are derived from osteoprotenitor cells, but osteoclasts are derived from the same cells that differentiate to form macrophages and monocytes. Within the marrow of the bone there are also haematopoetic stem cells. These cells give rise to other cells, including white blood cells, red blood cells, and platelets.
- Osteoblasts are mononucleate bone-forming cells. They are located on the surface of osteoid seams and make a protein mixture known as osteoid, which mineralizes to become bone. The osteoid seam is a narrow region of newly formed organic matrix, not yet mineralized, located on the surface of a bone. Osteoid is primarily composed of Type I collagen. Osteoblasts also manufacture hormones, such as prostaglandins, to act on the bone itself. They robustly produce alkaline phosphatase, an enzyme that has a role in the mineralisation of bone, as well as many matrix proteins.
- Osteocytes are mostly inactive osteoblasts. Osteocytes originate from osteoblasts that have migrated into and become trapped and surrounded by bone matrix that they themselves produced. The spaces they occupy are known as lacunae. Osteocytes have many processes that reach out to meet osteoblasts and other osteocytes probably for the purposes of communication.
- Osteoclasts are the cells responsible for bone resorption, thus they break down bone. New bone is then formed by the osteoblasts. Bone is constantly remodelled by the resorption of osteoclasts and created by osteoblasts. Osteoclasts are large cells with multiple nuclei located on bone surfaces in what are called Howship's lacunae (or resorption pits). These lacunae are the result of surrounding bone tissue that has been reabsorbed. Because the osteoclasts are derived from a monocyte stem-cell lineage, they are equipped with phagocytic-like mechanisms similar to circulating macrophages. Osteoclasts mature and/or migrate to discrete bone surfaces. Upon arrival, active enzymes, such as tartrate resistant acid phosphatase, are secreted against the mineral substrate. The reabsorption of bone by osteoclasts also plays a role in calcium homeostasis.
Bones consist of living cells embedded in a mineralized organic matrix. This matrix consists of organic components, mainly collagen - "organic" referring to materials produced as a result of the human body - and inorganic components, primarily hydroxyapatite and other salts of calcium and phosphate. Above 30% of the acellular part of bone consists of the organic components, and 70% of salts. The strands of collagen given bone its tensile strength, and the interspersed crystals of hydroxyapatite give bone its compressional strength. These effects are synergistic.
Hence, bone has a relatively high compressive strength of about 170 MPa (1800 kgf/cm²) but poor tensile strength of 104–121 MPa and very low shear stress strength (51.6 MPa), meaning it resists pushing forces well, but not pulling or torsional forces. While bone is essentially brittle, it does have a significant degree of elasticity, contributed chiefly by collagen.
The inorganic composition of bone (bone mineral) is primarily formed from salts of calcium and phosphate, the major salt being hydroxyapatite (Ca10(PO4)6(OH)2). The exact composition of the matrix may change over time and with nutrition, with the ratio of calcium to phosphate varying between 1.3-2 (per weight), and trace minerals such as magnesium, sodium, potassium and carbonate also being found.
The organic part of matrix is mainly composed of Type I collagen. Collagen composes about 90-95% of the organic matrix, with remainder of the matrix being a homogenous liquid called "ground substance" consisting of proteoglycans such as hyaluronic acid and chondroitin sulfate. Collagen consists of strands of repeating units, which give bone tensile strength, and are arranged in an overlapping fashion that prevents sheer stress. The function of ground substance is not fully known.
Two types of bone can be identified microscopically according to the pattern of collagen forming the osteoid (collagenous support tissue of type I collagen embedded in glycosaminoglycan gel):
- Woven bone, which is characterized by haphazard organization of collagen fibers and is mechanically weak
- Lamellar bone, which has a regular parallel alignment of collagen into sheets (lamellae) and is mechanically strong 
Woven bone is produced when osteoblasts produce osteoid rapidly, which occurs initially in all fetal bones (but is later replaced by more resilient lamellar bone). In adults woven bone is created after fractures or in Paget's disease. Woven bone is weaker, with a smaller number of randomly oriented collagen fibers, but forms quickly; it is for this appearance of the fibrous matrix that the bone is termed woven. It is soon replaced by lamellar bone, which is highly organized in concentric sheets with a much lower proportion of osteocytes to surrounding tissue. Lamellar bone, which makes its first appearance in the fetus during the third trimester, is stronger and filled with many collagen fibers parallel to other fibers in the same layer (these parallel columns are called osteons). In cross-section, the fibers run in opposite directions in alternating layers, much like in plywood, assisting in the bone's ability to resist torsion forces. After a fracture, woven bone forms initially and is gradually replaced by lamellar bone during a process known as "bony substitution." Compared to woven bone, lamellar bone formation takes place more slowly. The orderly deposition of collagen fibers restricts the formation of osteoid to about 1 to 2 µm per day. Lamellar bone also requires a relatively flat surface to lay the collagen fibers in parallel or concentric layers.
The extracellular matrix of bone is laid down by osteoblasts, which secrete both collagen and ground substance. These synthesise collagen within the cell, and then secrete collagen fibrils. The collagen fibres rapidly polymerise to form collagen strands. At this stage they are not get mineralised, and are called "osteoid". Around the strands calcium and phosphate precipitate on the surface of these strands, within a days to weeks becoming crystals of hydroxyapatite.
In order to mineralise the bone, the osteoblasts secrete vesicles containing alkaline phosphatase. This cleaves the phosphate groups and acts as the foci for calcium and phosphate deposition. The vesicles then rupture and act as a centre for crystals to grow on. More particularly, bone mineral is formed from globular and plate structures, distributed among the collagen fibrils of bone and forming yet larger structure.
Intramembranous ossification mainly occurs during formation of the flat bones of the skull but also the mandible, maxilla, and clavicles; the bone is formed from connective tissue such as mesenchyme tissue rather than from cartilage. The steps in intramembranous ossification are:
- Development of ossification center
- Formation of trabeculae
- Development of periosteum
Endochondral ossification, on the other hand, occurs in long bones and most of the rest of the bones in the body; it involves an initial hyaline cartilage that continues to grow. The steps in endochondral ossification are:
- Development of cartilage model
- Growth of cartilage model
- Development of the primary ossification center
- Development of the secondary ossification center
- Formation of articular cartilage and epiphyseal plate
Endochondral ossification begins with points in the cartilage called "primary ossification centers." They mostly appear during fetal development, though a few short bones begin their primary ossification after birth. They are responsible for the formation of the diaphyses of long bones, short bones and certain parts of irregular bones. Secondary ossification occurs after birth, and forms the epiphyses of long bones and the extremities of irregular and flat bones. The diaphysis and both epiphyses of a long bone are separated by a growing zone of cartilage (the epiphyseal plate). When the child reaches skeletal maturity (18 to 25 years of age), all of the cartilage is replaced by bone, fusing the diaphysis and both epiphyses together (epiphyseal closure).
In the upper limbs, only the diaphyses of the long bones and scapula are ossified. The epiphyses, carpal bones, coracoid process, medial border of the scapula, and acromion are still cartilaginous. The following steps are followed in the conversion of cartilage to bone:
- Zone of reserve cartilage. This region, farthest from the marrow cavity, consists of typical hyaline cartilage that as yet shows no sign of transforming into bone.
- Zone of cell proliferation. A little closer to the marrow cavity, chondrocytes multiply and arrange themselves into longitudinal columns of flattened lacunae.
- Zone of cell hypertrophy. Next, the chondrocytes cease to divide and begin to hypertrophy (enlarge), much like they do in the primary ossification center of the fetus. The walls of the matrix between lacunae become very thin.
- Zone of calcification. Minerals are deposited in the matrix between the columns of lacunae and calcify the cartilage. These are not the permanent mineral deposits of bone, but only a temporary support for the cartilage that would otherwise soon be weakened by the breakdown of the enlarged lacunae.
- Zone of bone deposition. Within each column, the walls between the lacunae break down and the chondrocytes die. This converts each column into a longitudinal channel, which is immediately invaded by blood vessels and marrow from the marrow cavity. Osteoblasts line up along the walls of these channels and begin depositing concentric lamellae of matrix, while osteoclasts dissolve the temporarily calcified cartilage.
Ossification in long bones
Several terms are also used to refer to specific features of long bones:
|diaphysis||The long, relatively straight main body of a long bone; region of primary ossification. Also known as the shaft.|
|epiphysis||The end regions of a long bone; regions of secondary ossification.|
|epiphyseal plate||Also known as the growth plate or physis. In a long bone it is a thin disc of hyaline cartilage that is positioned transversely between the epiphysis and metaphysis. In the long bones of humans, the epiphyseal plate disappears by twenty years of age.|
|metaphysis||The region of a long bone lying between the epiphysis and diaphysis.|
In the study of anatomy, anatomists use a number of anatomical terms to describe the appearance, shape and function of bones. Other anatomical terms are also used to describe the location of bones. Like other anatomical terms, many of these derive from Latin and Greek. Some anatomists still use Latin to refer to bones. The term "osseous", and the prefix "osteo-", referring to things related to bone, are still used commonly today.
Some examples of terms used to describe bones include the term "foramen" to describe a hole through which something passes, and a "canal" or "meatus" to describe a tunnel-like structure. A protrusion from a bone can be called a number of terms, including a "condyle", "crest", "spine", "eminence", "tubercle" or "tuberosity", depending on the protrusion's shape and location. In general, long bones are said to have a "head", "neck", and "body".
When two bones join together, they are said to "articulate". If the two bones have a fibrous connection and are relatively immobile, then the joint is called a "suture".
Bones have a variety of functions:
- Protection — bones can serve to protect internal organs, such as the skull protecting the brain or the ribs protecting the heart and lungs.
- Structure — bones provide a frame to keep the body supported.
- Movement — bones provide leverage system for, skeletal muscles, tendons, ligaments and joints function together to generate and transfer forces so that individual body parts or the whole body can be manipulated in three-dimensional space. The interaction between bone and muscle is studied in biomechanics.
- Sound transduction — bones are important in the mechanical aspect of overshadowed hearing.
- Blood production — the marrow, located within the medullary cavity of long bones and interstices of cancellous bone, produces blood cells in a process called hematopoiesis.
- Mineral storage — bones act as reserves of minerals important for the body, most notably calcium and phosphorus.
- Growth factor storage — mineralized bone matrix stores important growth factors such as insulin-like growth factors, transforming growth factor, bone morphogenetic proteins and others.
- Fat storage — the yellow bone marrow acts as a storage reserve of fatty acids.
- Acid-base balance — bone buffers the blood against excessive pH changes by absorbing or releasing alkaline salts.
- Detoxification — bone tissues can also store heavy metals and other foreign elements, removing them from the blood and reducing their effects on other tissues. These can later be gradually released for excretion.
- Endocrine organ — bone controls phosphate metabolism by releasing fibroblast growth factor – 23 (FGF-23), which acts on kidneys to reduce phosphate reabsorption. Bone cells also release a hormone called osteocalcin, which contributes to the regulation of blood sugar (glucose) and fat deposition. Osteocalcin increases both the insulin secretion and sensitivity, in addition to boosting the number of insulin-producing cells and reducing stores of fat.
- Calcium balance—The process of bone resorption by the osteoclasts releases stored calcium into the systemic circulation and is an important process in regulating calcium balance. As bone formation actively fixes circulating calcium in its mineral form, removing it from the bloodstream, resorption actively unfixes it thereby increasing circulating calcium levels. These processes occur in tandem at site-specific locations.
Remodeling or bone turnover is the process of resorption followed by replacement of bone with little change in shape and occurs throughout a person's life. Osteoblasts and osteoclasts, coupled together via paracrine cell signalling, are referred to as bone remodeling unit. Approximately 10% of the skeletal mass of an adult is remodelled each year. The purpose of remodeling is to regulate calcium homeostasis, repair micro-damaged bones (from everyday stress) but also to shape and sculpt the skeleton during growth.
Repeated stress, such as weight-bearing exercise or bone healing, results in the bone thickening at the points of maximum stress (Wolff's law). It has been hypothesized that this is a result of bone's piezoelectric properties, which cause bone to generate small electrical potentials under stress.
Paracrine cell signalling
The action of osteoblasts and osteoclasts are controlled by a number of chemical factors that either promote or inhibit the activity of the bone remodeling cells, controlling the rate at which bone is made, destroyed, or changed in shape. The cells also use paracrine signalling to control the activity of each other.
- Osteoblast stimulation
Bone building through increased secretion of osteoid is stimulated by the secretion of growth hormone by the pituitary, thyroid hormone and the sex hormones (estrogens and androgens). These hormones also promote increased secretion of osteoprotegerin. Osteoblasts can also be induced to secrete a number of cytokines that promote reabsorbtion of bone by stimulating osteoclast activity and differentiation from progenitor cells. Vitamin D, parathyroid hormone and stimulation from osteocytes induce osteoblasts to increase secretion of RANK-ligand and interleukin 6, which cytokines then stimulate increased reabsorption of bone by osteoclasts. These same compounds also increase secretion of macrophage colony-stimulating factor by osteoblasts, which promotes the differentiation of progenitor cells into osteoclasts, and decrease secretion of osteoprotegerin.
- Osteoclast inhibition
The rate at which osteoclasts resorb bone is inhibited by calcitonin and osteoprotegerin. Calcitonin is produced by parafollicular cells in the thyroid gland, and can bind to receptors on osteoclasts to directly inhibit osteoclast activity. Osteoprotegerin is secreted by osteoblasts and is able to bind RANK-L, inhibiting osteoclast stimulation.
Bone volume is determined by the rates of bone formation and bone resorption. Recent research has suggested that certain growth factors may work to locally alter bone formation by increasing osteoblast activity. Numerous bone-derived growth factors have been isolated and classified via bone cultures. These factors include insulin-like growth factors I and II, transforming growth factor-beta, fibroblast growth factor, platelet-derived growth factor, and bone morphogenetic proteins. Evidence suggests that bone cells produce growth factors for extracellular storage in the bone matrix. The release of these growth factors from the bone matrix could cause the proliferation of osteoblast precursors. Essentially, bone growth factors may act as potential determinants of local bone formation. Research has suggested that trabecular bone volume in postemenopausal osteoporosis may be determined by the relationship between the total bone forming surface and the percent of surface resorption.
A number of diseases can affect bone, including arthritis, fractures, infections, and osteoporosis. Conditions relating to bone can be managed by a variety of doctors, including rheumatologists for joints, and orthopedic surgeons, who may conduct surgery to fix broken bones. Other doctors, such as rehabilitation specialists may be involved in recovery, radiologists in interpreting the findings on imaging, and pathologists in investigating the cause of the disease, and family doctors may play a role in preventing complications of bone disease such as osteoporosis.
When a doctor sees a patient, a history and exam will be taken. Bones are then often imaged, called radiography. This might include ultrasound X-ray, CT scan, MRI scan and other imaging such as a Bone scan, which may be used to investigate cancer. Other tests such as a blood test for autoimmune markers may be taken, or a synovial fluid aspirate may be taken.
In normal bone, fractures occur when there is significant force applied, or repetitive trauma over a long time. Fractures can also occur when a bone is weakened, such as with osteoporosis, or when there is a structural problem, such as when the bone remodels excessively (such as Paget's disease) or is the site of the growth of cancer. Common fractures include wrist fractures and hip fractures, associated with osteoporosis, vertebral fractures associated with high-energy trauma and cancer, and fractures of long-bones. Not all fractures are painful, in fact some may not cause any pain at all. When serious, depending on the fractures type and location, complications may include flail chest, compartment syndromes or fat embolism.
Fractures are investigated using imaging such as X-ray of the joint or CT scans and MRI, and investigation of any underlying causes. Fractures are described by the location and shape of fracture, and several classification systems exist, depending on the location of the fracture. Fractures in children are described with the Salter–Harris fracture. When fractures are managed, pain relief is often given, and the fractured area is often immobilised. This is to promote bone healing. In addition, surgical measures such as internal fixation may be used. Because of the immobilisation, people with fractures are often advised to undergo rehabilitation.
|This section requires expansion. (November 2014)|
Osteoporosis is a disease of bone where there is reduced bone mineral density, increasing the likelihood of fractures. Osteoporosis is defined by the World Health Organization in women as a bone mineral density 2.5 standard deviations below peak bone mass, relative to the age and sex-matched average, as measured by Dual energy X-ray absorptiometry, with the term "established osteoporosis" including the presence of a fragility fracture. Osteoporosis is most common in women after menopause, when it is called "postmenopausal osteoporosis", but may develop in men and premenopausal women in the presence of particular hormonal disorders and other chronic diseases or as a result of smoking and medications, specifically glucocorticoids. Osteoporosis usually has no symptoms until a fracture occurs. For this reason, DEXA scans are often done in people with one or more risk factors, who have developed osteoporosis and be at risk of fracture.
Osteoporosis treatment includes advice to stop smoking, decrease alcohol consumption, exercise regularly, and have a healthy diet. Calcium supplements may also be advised, as may Vitamin D. When medication is used, it may include bisphosphonates, Strontium ranelate, and osteoporosis may be one factor considered when commencing Hormone replacement therapy.
Society and culture
Bones from slaughtered animals have a number of uses. They have been used as crafting materials for buttons, handles, ornaments etc. A special genre is scrimshaw. Ground bones are used as an organic phosphorus-nitrogen fertilizer and as additive in animal feed. Bones, in particular after calcination to bone ash is used as source of calcium phosphate for the production of bone china and previously also phosphorus chemicals.
Osteopathic medicine is a school of medical thought originally developed based on the idea of the link between the musculoskeletal system and overall health, but now very similar to mainstream medicine. As of 2012[update], over 77,000 physicians in the United States are trained in Osteopathic medicine colleges.
The study of bones and teeth is referred to as osteology. It is frequently used in anthropology, archeology and forensic science for a variety of tasks. This can include determining the nutritional, health, age or injury status of the individual the bones were taken from. Preparing fleshed bones for these types of studies can involve maceration – boiling fleshed bones to remove large particles, then hand-cleaning.
Typically anthropologists and archeologists study bone tools made by Homo sapiens and Homo neanderthalensis. Bones can serve a number of uses such as projectile points or artistic pigments, and can be made from endoskeletal or external bones such as antler or tusk.
It is a common misconception that bird skeletons are lightweight. While their bones are smaller and thinner, bird bone tissue is generally denser than mammalian bone tissue, allowing them to be stronger and offer more support with less volume and weight. Among mammals, bats come closest to birds in terms of bone density, suggesting that small dense bones are a flight adaptation. Many bird bones have little marrow due to being hollow, though not all bird bones are hollow.
Alternatives to bony endoskeletons
- Exoskeletons offer support, protection and levers for movement similar to endoskeletal bone. Different types of exoskeletons include shells, carapaces (consisting of calcium compounds or silica) and chitinous exoskeletons.
- A true endoskeleton (that is, protective tissue derived from mesoderm) is also present in Echinoderms. Porifera (sponges) possess simple endoskeletons that consist of calcareous or siliceous spicules and a spongin fiber network.
Bone penetrating the skin and being exposed to the outside can be both a natural process in some animals, and due to injury:
- A deer's antlers are composed of bone.
- Instead of teeth, the extinct predatory fish Dunkleosteus had sharp edges of hard exposed bone along its jaws.
- A compound fracture occurs when the edges of a broken bone puncture the skin.
- Though not strictly speaking exposed, a bird's beak is primarily bone covered in a layer of keratin over a vascular layer containing blood vessels and nerve endings.
- This article uses anatomical terminology; for an overview, see anatomical terminology.
|Wikimedia Commons has media related to Bones.|
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