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A bone is a rigid organ that constitutes part of the skeleton in most
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
animals. Bones protect the various other organs of the body, produce
red Red is the color at the long wavelength end of the visible spectrum of light, next to orange and opposite violet. It has a dominant wavelength of approximately 625–740 nanometres. It is a primary color in the RGB color model and a secondar ...
and white blood cells, store
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2 ...
s, provide structure and support for the body, and enable
mobility Mobility may refer to: Social sciences and humanities * Economic mobility, ability of individuals or families to improve their economic status * Geographic mobility, the measure of how populations and goods move over time * Mobilities, a conte ...
. Bones come in a variety of shapes and sizes and have complex internal and external structures. They are lightweight yet strong and hard and serve multiple functions. Bone tissue (osseous tissue), which is also called bone in the
uncountable In mathematics, an uncountable set (or uncountably infinite set) is an infinite set that contains too many elements to be countable. The uncountability of a set is closely related to its cardinal number: a set is uncountable if its cardinal num ...
sense of that word, is
hard tissue Hard tissue, refers to "normal" calcified tissue, is the tissue which is mineralized and has a firm intercellular matrix. The hard tissues of humans are bone, tooth enamel, dentin, and cementum. The term is in contrast to soft tissue. Bone Bon ...
, a type of specialized connective tissue. It has a
honeycomb A honeycomb is a mass of hexagonal prismatic wax cells built by honey bees in their nests to contain their larvae and stores of honey and pollen. Beekeepers may remove the entire honeycomb to harvest honey. Honey bees consume about of honey ...
-like
matrix Matrix most commonly refers to: * ''The Matrix'' (franchise), an American media franchise ** ''The Matrix'', a 1999 science-fiction action film ** "The Matrix", a fictional setting, a virtual reality environment, within ''The Matrix'' (franchis ...
internally, which helps to give the bone rigidity. Bone tissue is made up of different types of
bone cell An osteocyte, an oblate shaped type of bone cell with dendritic processes, is the most commonly found cell in mature bone. It can live as long as the organism itself. The adult human body has about 42 billion of them. Osteocytes do not divide and ...
s. Osteoblasts and
osteocyte An osteocyte, an oblate shaped type of bone cell with dendritic processes, is the most commonly found cell in mature bone. It can live as long as the organism itself. The adult human body has about 42 billion of them. Osteocytes do not divide an ...
s are involved in the formation and mineralization of bone;
osteoclast An osteoclast () is a type of bone cell that breaks down bone tissue. This function is critical in the maintenance, repair, and remodeling of bones of the vertebral skeleton. The osteoclast disassembles and digests the composite of hydrated pro ...
s are involved in the
resorption Resorption is the absorption of cells or tissue into the circulatory system, usually by osteoclasts. Types of resorption include: * Bone resorption * Herniated Disc Resorption * Tooth resorption * Fetal resorption * Blood resorption See also ...
of bone tissue. Modified (flattened) osteoblasts become the lining cells that form a protective layer on the bone surface. The mineralized matrix of bone tissue has an organic component of mainly collagen called ''
ossein Ossein is the organic extracellular matrix of bone, which is made of 95% collagen. It can be isolated by treating bones with hydrochloric acid, which dissolves the inorganic matrix (calcium phosphate and calcium carbonate). This substance is used i ...
'' and an inorganic component of
bone mineral Bone mineral (also called inorganic bone phase, bone salt, or bone apatite) is the inorganic component of bone tissue. It gives bones their compressive strength. Bone mineral is formed predominantly from carbonated hydroxyapatite with lower cry ...
made up of various salts. Bone tissue is
mineralized tissue Mineralized tissues are biological tissues that incorporate minerals into soft matrices. Typically these tissues form a protective shield or structural support. Bone, mollusc shells, deep sea sponge ''Euplectella'' species, radiolarians, diato ...
of two types, cortical bone and cancellous bone. Other types of tissue found in bones include bone marrow, endosteum,
periosteum The periosteum is a membrane that covers the outer surface of all bones, except at the articular surfaces (i.e. the parts within a joint space) of long bones. Endosteum lines the inner surface of the medullary cavity of all long bones. Structu ...
, nerves,
blood vessel The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away ...
s and cartilage. In the human body at birth, there are approximately 300 bones present; many of these fuse together during development, leaving a total of 206 separate bones in the adult, not counting numerous small sesamoid bones. The largest bone in the body is the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
or thigh-bone, and the smallest is the
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foo ...
in the
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
. The Greek word for bone is ὀστέον ("osteon"), hence the many terms that use it as a prefix—such as
osteopathy Osteopathy () is a type of alternative medicine that emphasizes physical manipulation of the body's muscle tissue and bones. Practitioners of osteopathy are referred to as osteopaths. Osteopathic manipulation is the core set of techniques in ...
. In
anatomical terminology Anatomical terminology is a form of scientific terminology used by anatomists, zoologists, and health professionals such as doctors. Anatomical terminology uses many unique terms, suffixes, and prefixes deriving from Ancient Greek and Latin. ...
, including the
Terminologia Anatomica ''Terminologia Anatomica'' is the international standard for human anatomical terminology. It is developed by the Federative International Programme on Anatomical Terminology, a program of the International Federation of Associations of Anatomi ...
international standard, the word for a bone is '' os'' (for example, os breve, os longum, os sesamoideum).


Structure

Bone is not uniformly solid, but consists of a flexible
matrix Matrix most commonly refers to: * ''The Matrix'' (franchise), an American media franchise ** ''The Matrix'', a 1999 science-fiction action film ** "The Matrix", a fictional setting, a virtual reality environment, within ''The Matrix'' (franchis ...
(about 30%) and bound minerals (about 70%) which are intricately woven and endlessly remodeled by a group of specialized bone cells. Their unique composition and design allows bones to be relatively
hard Hard may refer to: * Hardness, resistance of physical materials to deformation or fracture * Hard water, water with high mineral content Arts and entertainment * ''Hard'' (TV series), a French TV series * Hard (band), a Hungarian hard rock supe ...
and strong, while remaining lightweight. Bone matrix is 90 to 95% composed of elastic collagen fibers, also known as ossein, and the remainder is
ground substance Ground substance is an amorphous gel-like substance in the extracellular space that contains all components of the extracellular matrix (ECM) except for fibrous materials such as collagen and elastin. Ground substance is active in the development, m ...
. The elasticity of collagen improves fracture resistance. The matrix is hardened by the binding of inorganic mineral salt,
calcium phosphate The term calcium phosphate refers to a family of materials and minerals containing calcium ions (Ca2+) together with inorganic phosphate anions. Some so-called calcium phosphates contain oxide and hydroxide as well. Calcium phosphates are whi ...
, in a chemical arrangement known as
bone mineral Bone mineral (also called inorganic bone phase, bone salt, or bone apatite) is the inorganic component of bone tissue. It gives bones their compressive strength. Bone mineral is formed predominantly from carbonated hydroxyapatite with lower cry ...
, a form of calcium
hydroxylapatite Hydroxyapatite, also called hydroxylapatite (HA), is a naturally occurring mineral form of calcium apatite with the formula Ca5(PO4)3(OH), but it is usually written Ca10(PO4)6(OH)2 to denote that the crystal unit cell comprises two entities. ...
. It is the mineralization that gives bones rigidity. Bone is actively constructed and remodeled throughout life by special bone cells known as osteoblasts and osteoclasts. Within any single bone, the tissue is woven into two main patterns, known as cortical and cancellous bone, each with a different appearance and characteristics.


Cortex

The hard outer layer of bones is composed of cortical bone, which is also called compact bone as it is much denser than cancellous bone. It forms the hard exterior (cortex) of bones. The cortical bone gives bone its smooth, white, and solid appearance, and accounts for 80% of the total bone mass of an adult human skeleton. It facilitates bone's main functions—to support the whole body, to protect organs, to provide
lever A lever is a simple machine consisting of a beam or rigid rod pivoted at a fixed hinge, or '' fulcrum''. A lever is a rigid body capable of rotating on a point on itself. On the basis of the locations of fulcrum, load and effort, the lever is d ...
s for movement, and to store and release chemical elements, mainly calcium. It consists of multiple microscopic columns, each called an
osteon In osteology, the osteon or haversian system (; named for Clopton Havers) is the fundamental functional unit of much compact bone. Osteons are roughly cylindrical structures that are typically between 0.25 mm and 0.35 mm in diameter. Their ...
or Haversian system. Each column is multiple layers of osteoblasts and
osteocyte An osteocyte, an oblate shaped type of bone cell with dendritic processes, is the most commonly found cell in mature bone. It can live as long as the organism itself. The adult human body has about 42 billion of them. Osteocytes do not divide an ...
s around a central canal called the
haversian canal Haversian canals (sometimes canals of Havers) are a series of microscopic tubes in the outermost region of bone called cortical bone. They allow blood vessels and nerves to travel through them to supply the osteocytes. Structure Each Haversian ...
.
Volkmann's canal Volkmann's canals, also known as perforating holes or channels, are anatomic arrangements in cortical bones. Volkmann's canals are inside osteons. They interconnect the haversian canals with each other and the periosteum The periosteum is a mem ...
s 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. Cortical bone is covered by a
periosteum The periosteum is a membrane that covers the outer surface of all bones, except at the articular surfaces (i.e. the parts within a joint space) of long bones. Endosteum lines the inner surface of the medullary cavity of all long bones. Structu ...
on its outer surface, and an endosteum on its inner surface. The endosteum is the boundary between the cortical bone and the cancellous bone. The primary anatomical and functional unit of cortical bone is the
osteon In osteology, the osteon or haversian system (; named for Clopton Havers) is the fundamental functional unit of much compact bone. Osteons are roughly cylindrical structures that are typically between 0.25 mm and 0.35 mm in diameter. Their ...
.


Trabecules

Cancellous bone, also called trabecular or spongy bone, is the internal tissue of the skeletal bone and is an open cell
porous Porosity or void fraction is a measure of the void (i.e. "empty") spaces in a material, and is a fraction of the volume of voids over the total volume, between 0 and 1, or as a percentage between 0% and 100%. Strictly speaking, some tests measure ...
network that follows the material properties of
biofoams Biofoams are biological or biologically derived foams, making up lightweight and porous cellular solids. A relatively new term, its use in academia began in the 1980s in relation to the scum that formed on activated sludge plants. Biofoams is a br ...
. Cancellous bone has a higher
surface-area-to-volume ratio The surface-area-to-volume ratio, also called the surface-to-volume ratio and variously denoted sa/vol or SA:V, is the amount of surface area per unit volume of an object or collection of objects. SA:V is an important concept in science and engin ...
than cortical bone and it is less
dense Density (volumetric mass density or specific mass) is the substance's mass per unit of volume. The symbol most often used for density is ''ρ'' (the lower case Greek letter rho), although the Latin letter ''D'' can also be used. Mathematically ...
. This makes it weaker and more flexible. The greater surface area also makes it suitable for metabolic activities such as the exchange of calcium ions. Cancellous bone is typically found at the ends of long bones, near joints, and in the interior of vertebrae. Cancellous bone is highly
vascular The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
and often contains red bone marrow where
hematopoiesis Haematopoiesis (, from Greek , 'blood' and 'to make'; also hematopoiesis in American English; sometimes also h(a)emopoiesis) is the formation of blood cellular components. All cellular blood components are derived from haematopoietic stem cells ...
, the production of blood cells, occurs. The primary anatomical and functional unit of cancellous bone is the
trabecula A trabecula (plural trabeculae, from Latin for "small beam") is a small, often microscopic, tissue element in the form of a small beam, strut or rod that supports or anchors a framework of parts within a body or organ. A trabecula generally ha ...
. The trabeculae are aligned towards the mechanical load distribution that a bone experiences within long bones such as the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
. As far as short bones are concerned, trabecular alignment has been studied in the
vertebral The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordates ...
pedicle Pedicle or pedicel may refer to: Human anatomy *Pedicle of vertebral arch, the segment between the transverse process and the vertebral body, and is often used as a radiographic marker and entry point in vertebroplasty and kyphoplasty procedures ...
. Thin formations of osteoblasts covered in endosteum create an irregular network of spaces, known as trabeculae. Within these spaces are bone marrow and hematopoietic stem cells that give rise to
platelet Platelets, also called thrombocytes (from Greek θρόμβος, "clot" and κύτος, "cell"), are a component of blood whose function (along with the coagulation factors) is to react to bleeding from blood vessel injury by clumping, thereby i ...
s,
red blood cell Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek ''erythros'' for "red" and ''kytos'' for "holl ...
s 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. The words ''cancellous'' and ''trabecular'' refer to the tiny lattice-shaped units (trabeculae) that form the tissue. It was first illustrated accurately in the engravings of
Crisóstomo Martinez Crisóstomo Martínez (1638–1694) was a Spanish painter, engraver, anatomist and microscopist from Valencia, known for his atlas of anatomy. His work has been ascribed to the Spanish intellectual movement called "''Novator''" which refers to th ...
.


Marrow

Bone marrow, also known as
myeloid tissue Myeloid tissue, in the bone marrow sense of the word ''wikt:myeloid#Adjective, myeloid'' (''wikt:myelo-#Prefix, myelo-'' + ''wikt:-oid#Suffix, -oid''), is tissue (biology), tissue of bone marrow, of bone marrow cell lineage, or resembling bone ...
in red bone marrow, can be found in almost any bone that holds
cancellous tissue A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, an ...
. In
newborns An infant or baby is the very young offspring of human beings. ''Infant'' (from the Latin word ''infans'', meaning 'unable to speak' or 'speechless') is a formal or specialised synonym for the common term ''baby''. The terms may also be used t ...
, all such bones are filled exclusively with red marrow or hematopoietic marrow, but as the child ages the hematopoietic fraction decreases in quantity and the fatty/ yellow fraction called
marrow adipose tissue Bone marrow adipose tissue (BMAT), sometimes referred to as marrow adipose tissue (MAT), is a type of fat deposit in bone marrow. It increases in states of low bone density -osteoporosis, anorexia nervosa/ caloric restriction, skeletal unweigh ...
(MAT) increases in quantity. In adults, red marrow is mostly found in the bone marrow of the femur, the ribs, the vertebrae and
pelvic bones The hip bone (os coxae, innominate bone, pelvic bone or coxal bone) is a large flat bone, constricted in the center and expanded above and below. In some vertebrates (including humans before puberty) it is composed of three parts: the ilium, ischi ...
.


Vascular supply

Bone receives about 10% of cardiac output. Blood enters the endosteum, flows through the marrow, and exits through small vessels in the cortex. In humans, blood oxygen tension in bone marrow is about 6.6%, compared to about 12% in arterial blood, and 5% in venous and capillary blood.


Cells

Bone is metabolically active tissue composed of several types of cells. These cells include osteoblasts, which are involved in the creation and mineralization of bone tissue,
osteocyte An osteocyte, an oblate shaped type of bone cell with dendritic processes, is the most commonly found cell in mature bone. It can live as long as the organism itself. The adult human body has about 42 billion of them. Osteocytes do not divide an ...
s, and
osteoclast An osteoclast () is a type of bone cell that breaks down bone tissue. This function is critical in the maintenance, repair, and remodeling of bones of the vertebral skeleton. The osteoclast disassembles and digests the composite of hydrated pro ...
s, which are involved in the reabsorption of bone tissue. Osteoblasts and osteocytes are derived from
osteoprogenitor Osteoblasts (from the Greek language, Greek combining forms for "bone", ὀστέο-, ''osteo-'' and βλαστάνω, ''blastanō'' "germinate") are cell (biology), cells with a single Cell nucleus, nucleus that synthesize bone. However, in the p ...
cells, but
osteoclast An osteoclast () is a type of bone cell that breaks down bone tissue. This function is critical in the maintenance, repair, and remodeling of bones of the vertebral skeleton. The osteoclast disassembles and digests the composite of hydrated pro ...
s are derived from the same cells that differentiate to form macrophages and
monocyte Monocytes are a type of leukocyte or white blood cell. They are the largest type of leukocyte in blood and can differentiate into macrophages and conventional dendritic cells. As a part of the vertebrate innate immune system monocytes also ...
s. Within the marrow of the bone there are also hematopoietic stem cells. These cells give rise to other cells, including white blood cells,
red blood cell Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek ''erythros'' for "red" and ''kytos'' for "holl ...
s, and
platelet Platelets, also called thrombocytes (from Greek θρόμβος, "clot" and κύτος, "cell"), are a component of blood whose function (along with the coagulation factors) is to react to bleeding from blood vessel injury by clumping, thereby i ...
s.


Osteoblast

Osteoblasts are mononucleate bone-forming cells. They are located on the surface of osteon seams and make a
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
mixture known as
osteoid In histology, osteoid is the unmineralized, organic portion of the bone matrix that forms prior to the maturation of bone tissue. Osteoblasts begin the process of forming bone tissue by secreting the osteoid as several specific proteins. When ...
, which mineralizes to become bone. The osteoid seam is a narrow region of a 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
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are require ...
s, such as prostaglandins, to act on the bone itself. The osteoblast creates and repairs new bone by actually building around itself. First, the osteoblast puts up collagen fibers. These collagen fibers are used as a framework for the osteoblasts' work. The osteoblast then deposits calcium phosphate which is hardened by
hydroxide Hydroxide is a diatomic anion with chemical formula OH−. It consists of an oxygen and hydrogen atom held together by a single covalent bond, and carries a negative electric charge. It is an important but usually minor constituent of water. I ...
and
bicarbonate In inorganic chemistry, bicarbonate (IUPAC-recommended nomenclature: hydrogencarbonate) is an intermediate form in the deprotonation of carbonic acid. It is a polyatomic anion with the chemical formula . Bicarbonate serves a crucial biochem ...
ions. The brand-new bone created by the osteoblast is called
osteoid In histology, osteoid is the unmineralized, organic portion of the bone matrix that forms prior to the maturation of bone tissue. Osteoblasts begin the process of forming bone tissue by secreting the osteoid as several specific proteins. When ...
. Once the osteoblast is finished working it is actually trapped inside the bone once it hardens. When the osteoblast becomes trapped, it becomes known as an osteocyte. Other osteoblasts remain on the top of the new bone and are used to protect the underlying bone, these become known as lining cells.


Osteocyte

Osteocyte An osteocyte, an oblate shaped type of bone cell with dendritic processes, is the most commonly found cell in mature bone. It can live as long as the organism itself. The adult human body has about 42 billion of them. Osteocytes do not divide an ...
s are cells of mesenchymal origin and originate from osteoblasts that have migrated into and become trapped and surrounded by a bone matrix that they themselves produced. The spaces the cell body of osteocytes occupy within the mineralized collagen type I matrix are known as lacunae, while the osteocyte cell processes occupy channels called canaliculi. The many processes of osteocytes reach out to meet osteoblasts, osteoclasts, bone lining cells, and other osteocytes probably for the purposes of communication. Osteocytes remain in contact with other osteocytes in the bone through gap junctions—coupled cell processes which pass through the canalicular channels.


Osteoclast

Osteoclast An osteoclast () is a type of bone cell that breaks down bone tissue. This function is critical in the maintenance, repair, and remodeling of bones of the vertebral skeleton. The osteoclast disassembles and digests the composite of hydrated pro ...
s are very large
multinucleate Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordina ...
cells that are responsible for the breakdown of bones by the process of
bone resorption Bone resorption is resorption of bone tissue, that is, the process by which osteoclasts break down the tissue in bones and release the minerals, resulting in a transfer of calcium from bone tissue to the blood. The osteoclasts are multi-nuclea ...
. New bone is then formed by the osteoblasts. Bone is constantly remodeled 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 Monocytes are a type of leukocyte or white blood cell. They are the largest type of leukocyte in blood and can differentiate into macrophages and conventional dendritic cells. As a part of the vertebrate innate immune system monocytes also ...
stem-cell lineage, they are equipped with
phagocytic Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is c ...
-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 Tartrate-resistant acid phosphatase (TRAP or TRAPase), also called acid phosphatase 5, tartrate resistant (ACP5), is a glycosylated monomeric metalloprotein enzyme expressed in mammals. It has a molecular weight of approximately 35kDa, a basic iso ...
, are
secreted 440px Secretion is the movement of material from one point to another, such as a secreted chemical substance from a cell or gland. In contrast, excretion is the removal of certain substances or waste products from a cell or organism. The classical ...
against the mineral substrate. The reabsorption of bone by osteoclasts also plays a role in
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar t ...
homeostasis In biology, homeostasis (British also homoeostasis) (/hɒmɪə(ʊ)ˈsteɪsɪs/) is the state of steady internal, physical, and chemical conditions maintained by living systems. This is the condition of optimal functioning for the organism and ...
.


Composition

Bones consist of living cells (osteoblasts and osteocytes) embedded in a mineralized organic matrix. The primary inorganic component of human bone is hydroxyapatite, the dominant
bone mineral Bone mineral (also called inorganic bone phase, bone salt, or bone apatite) is the inorganic component of bone tissue. It gives bones their compressive strength. Bone mineral is formed predominantly from carbonated hydroxyapatite with lower cry ...
, having the nominal composition of Ca10(PO4)6(OH)2. The organic components of this matrix consist mainly of
type I collagen Type I collagen is the most abundant collagen of the human body. It forms large, eosinophilic fibers known as collagen fibers. It is present in scar tissue, the end product when tissue heals by repair, as well as tendons, ligaments, the endomy ...
—"organic" referring to materials produced as a result of the human body—and inorganic components, which alongside the dominant hydroxyapatite phase, include other compounds of
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar t ...
and
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phosph ...
including salts. Approximately 30% of the acellular component of bone consists of organic matter, while roughly 70% by mass is attributed to the inorganic phase. The collagen fibers give bone its
tensile strength Ultimate tensile strength (UTS), often shortened to tensile strength (TS), ultimate strength, or F_\text within equations, is the maximum stress that a material can withstand while being stretched or pulled before breaking. In brittle materials t ...
, and the interspersed crystals of hydroxyapatite give bone its compressive strength. These effects are synergistic. The exact composition of the matrix may be subject to change over time due to nutrition and
biomineralization Biomineralization, also written biomineralisation, is the process by which living organisms produce minerals, often to harden or stiffen existing tissues. Such tissues are called mineralized tissues. It is an extremely widespread phenomenon; ...
, with the ratio of
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar t ...
to
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phosph ...
varying between 1.3 and 2.0 (per weight), and trace minerals such as
magnesium Magnesium is a chemical element with the symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 of the periodic ta ...
,
sodium Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable ...
,
potassium Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosph ...
and
carbonate A carbonate is a salt of carbonic acid (H2CO3), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word ''carbonate'' may also refer to a carbonate ester, an organic compound containing the carbonate ...
also be found. Type I collagen composes 90–95% of the organic matrix, with the remainder of the matrix being a homogenous liquid called
ground substance Ground substance is an amorphous gel-like substance in the extracellular space that contains all components of the extracellular matrix (ECM) except for fibrous materials such as collagen and elastin. Ground substance is active in the development, m ...
consisting of proteoglycans such as hyaluronic acid and
chondroitin sulfate Chondroitin sulfate is a sulfated glycosaminoglycan (GAG) composed of a chain of alternating sugars ( N-acetylgalactosamine and glucuronic acid). It is usually found attached to proteins as part of a proteoglycan. A chondroitin chain can have ov ...
, as well as non-collagenous proteins such as
osteocalcin Osteocalcin, also known as bone gamma-carboxyglutamic acid-containing protein (BGLAP), is a small (49-amino-acid) noncollagenous protein hormone found in bone and dentin, first identified as a calcium-binding protein. Because osteocalcin has g ...
,
osteopontin Osteopontin (OPN), also known as bone /sialoprotein I (BSP-1 or BNSP), early T-lymphocyte activation (ETA-1), secreted phosphoprotein 1 (SPP1), 2ar and Rickettsia resistance (Ric), is a protein that in humans is encoded by the ''SPP1'' gene (secr ...
or
bone sialoprotein Bone sialoprotein (BSP) is a component of mineralized tissues such as bone, dentin, cementum and calcified cartilage. BSP is a significant component of the bone extracellular matrix and has been suggested to constitute approximately 8% of all ...
. Collagen consists of strands of repeating units, which give bone tensile strength, and are arranged in an overlapping fashion that prevents shear stress. The function of ground substance is not fully known. Two types of bone can be identified microscopically according to the arrangement of collagen: woven and lamellar. * Woven bone (also known as ''fibrous bone''), which is characterized by a haphazard organization of collagen fibers and is mechanically weak.Currey, John D. (2002)
"The Structure of Bone Tissue"
pp. 12–14 in ''Bones: Structure and Mechanics''. Princeton University Press. Princeton, NJ.
* 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 A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal develo ...
bones, but is later replaced by more resilient lamellar bone. In adults, woven bone is created after
fractures Fracture is the separation of an object or material into two or more pieces under the action of stress. The fracture of a solid usually occurs due to the development of certain displacement discontinuity surfaces within the solid. If a displa ...
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 humans in the
fetus A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal dev ...
during the third trimester,Salentijn, L. ''Biology of Mineralized Tissues: Cartilage and Bone'',
Columbia University College of Dental Medicine The Columbia University College of Dental Medicine, often abbreviated CDM, is one of the twenty graduate and professional schools of Columbia University. It is located at 630 West 168th Street in Manhattan, New York City. According to American ...
post-graduate dental lecture series, 2007
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 Torsion may refer to: Science * Torsion (mechanics), the twisting of an object due to an applied torque * Torsion of spacetime, the field used in Einstein–Cartan theory and ** Alternatives to general relativity * Torsion angle, in chemistry Bi ...
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.


Deposition

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 fibers rapidly
polymer A polymer (; Greek '' poly-'', "many" + ''-mer'', "part") is a substance or material consisting of very large molecules called macromolecules, composed of many repeating subunits. Due to their broad spectrum of properties, both synthetic a ...
ise to form collagen strands. At this stage, they are not yet mineralised, and are called "osteoid". Around the strands calcium and phosphate
precipitate In an aqueous solution, precipitation is the process of transforming a dissolved substance into an insoluble solid from a super-saturated solution. The solid formed is called the precipitate. In case of an inorganic chemical reaction leading ...
on the surface of these strands, within days to weeks becoming crystals of hydroxyapatite. In order to mineralise the bone, the osteoblasts secrete
vesicles Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features o ...
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.


Types

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 The diaphysis is the main or midsection (shaft) of a long bone. It is made up of cortical bone and usually contains bone marrow and adipose tissue (fat). It is a middle tubular part composed of compact bone which surrounds a central marrow cavit ...
, that is much longer than its width; and by an
epiphysis The epiphysis () is the rounded end of a long bone, at its joint with adjacent bone(s). Between the epiphysis and diaphysis (the long midsection of the long bone) lies the metaphysis, including the epiphyseal plate (growth plate). At the jo ...
, a rounded head at each end of the shaft. They are made up mostly of
compact bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, and ...
, with lesser amounts of marrow, located within the
medullary cavity The medullary cavity (''medulla'', innermost part) is the central cavity of bone shafts where red bone marrow and/or yellow bone marrow ( adipose tissue) is stored; hence, the medullary cavity is also known as the marrow cavity. Located in the m ...
, and areas of spongy, cancellous bone at the ends of the bones. Most bones of the limbs, including those of the
fingers A finger is a limb of the body and a type of digit, an organ of manipulation and sensation found in the hands of most of the Tetrapods, so also with humans and other primates. Most land vertebrates have five fingers (Pentadactyly). Chambers 1 ...
and
toes Toes are the digits (fingers) of the foot of a tetrapod. Animal species such as cats that walk on their toes are described as being '' digitigrade''. Humans, and other animals that walk on the soles of their feet, are described as being '' pl ...
, are long bones. The exceptions are the eight carpal bones of the
wrist In human anatomy, the wrist is variously defined as (1) the carpus or carpal bones, the complex of eight bones forming the proximal skeletal segment of the hand; "The wrist contains eight bones, roughly aligned in two rows, known as the carp ...
, the seven articulating
tarsal bone In the human body, the tarsus is a cluster of seven articulating bones in each foot situated between the lower end of the tibia and the fibula of the lower leg and the metatarsus. It is made up of the midfoot (cuboid, medial, intermediate, and la ...
s of the
ankle The ankle, or the talocrural region, or the jumping bone (informal) is the area where the foot and the leg meet. The ankle includes three joints: the ankle joint proper or talocrural joint, the subtalar joint, and the inferior tibiofibular joi ...
and the sesamoid bone of the
kneecap The patella, also known as the kneecap, is a flat, rounded triangular bone which articulates with the femur (thigh bone) and covers and protects the anterior articular surface of the knee joint. The patella is found in many tetrapods, such as m ...
. Long bones such as the clavicle, that have a differently shaped shaft or ends are also called ''modified long bones''. *
Short bone Short bones are designated as those bones that are as wide as they are long. Their primary function is to provide support and stability with little to no movement. They are one of five types of bones: short, long, flat, irregular and sesamoid ...
s 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. *
Flat bone Flat bones are bones whose principal function is either extensive protection or the provision of broad surfaces for muscular attachment. These bones are expanded into broad, flat plates,''Gray's Anatomy'' (1918). (See infobox) as in the cranium ...
s are thin and generally curved, with two parallel layers of compact bone sandwiching a layer of spongy bone. Most of the bones of the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
are flat bones, as is the
sternum The sternum or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major blood vessels from injury. Sha ...
. * 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 The patella, also known as the kneecap, is a flat, rounded triangular bone which articulates with the femur (thigh bone) and covers and protects the anterior articular surface of the knee joint. The patella is found in many tetrapods, such as ...
and the
pisiform The pisiform bone ( or ), also spelled pisiforme (from the Latin ''pisifomis'', pea-shaped), is a small knobbly, sesamoid bone that is found in the wrist. It forms the ulnar border of the carpal tunnel. Structure The pisiform is a sesamoid bone, ...
. *
Irregular bone The irregular bones are bones which, from their peculiar form, cannot be grouped as long, short, flat or sesamoid bones. Irregular bones serve various purposes in the body, such as protection of nervous tissue (such as the vertebrae protect th ...
s 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 The ethmoid bone (; from grc, ἡθμός, hēthmós, sieve) is an unpaired bone in the skull that separates the nasal cavity from the brain. It is located at the roof of the nose, between the two orbits. The cubical bone is lightweight due to a ...
and sphenoid bones.


Terminology

In the study of
anatomy Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
, 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 Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
and
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
. 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, they are said to "articulate". If the two bones have a fibrous connection and are relatively immobile, then the joint is called a "suture".


Development

The formation of bone is called
ossification Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in ...
. During the fetal stage of development this occurs by two processes: intramembranous ossification and endochondral ossification. Intramembranous ossification involves the formation of bone from connective tissue whereas endochondral ossification involves the formation of bone from cartilage. Intramembranous ossification mainly occurs during formation of the flat bones of the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
but also the mandible, maxilla, and clavicles; the bone is formed from connective tissue such as mesenchyme tissue rather than from cartilage. The process includes: the development of the
ossification center An ossification center is a point where ossification of the cartilage begins. The first step in ossification is that the cartilage cells at this point enlarge and arrange themselves in rows.Gray and Spitzka (1910), page 44. The matrix in which t ...
,
calcification Calcification is the accumulation of calcium salts in a body tissue. It normally occurs in the formation of bone, but calcium can be deposited abnormally in soft tissue,Miller, J. D. Cardiovascular calcification: Orbicular origins. ''Nature Ma ...
, trabeculae formation and the development of the periosteum. Endochondral ossification occurs in long bones and most other bones in the body; it involves the development of bone from cartilage. This process includes the development of a cartilage model, its growth and development, development of the primary and secondary
ossification center An ossification center is a point where ossification of the cartilage begins. The first step in ossification is that the cartilage cells at this point enlarge and arrange themselves in rows.Gray and Spitzka (1910), page 44. The matrix in which t ...
s, and the formation of articular cartilage and the
epiphyseal plate The epiphyseal plate (or epiphysial plate, physis, or growth plate) is a hyaline cartilage plate in the metaphysis at each end of a long bone. It is the part of a long bone where new bone growth takes place; that is, the whole bone is alive, wit ...
s. 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 Birth is the act or process of bearing or bringing forth offspring, also referred to in technical contexts as parturition. In mammals, the process is initiated by hormones which cause the muscular walls of the uterus to contract, expelling the f ...
. 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 The epiphysis () is the rounded end of a long bone, at its joint with adjacent bone(s). Between the epiphysis and diaphysis (the long midsection of the long bone) lies the metaphysis, including the epiphyseal plate (growth plate). At the jo ...
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 The epiphyseal plate (or epiphysial plate, physis, or growth plate) is a hyaline cartilage plate in the metaphysis at each end of a long bone. It is the part of a long bone where new bone growth takes place; that is, the whole bone is alive, wit ...
). At 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.


Functions

Bones have a variety of functions:


Mechanical

Bones serve a variety of mechanical functions. Together the bones in the body form the skeleton. They provide a frame to keep the body supported, and an attachment point for skeletal muscles,
tendon A tendon or sinew is a tough, high-tensile-strength band of dense fibrous connective tissue that connects muscle to bone. It is able to transmit the mechanical forces of muscle contraction to the skeletal system without sacrificing its ability ...
s, ligaments and
joint A joint or articulation (or articular surface) is the connection made between bones, ossicles, or other hard structures in the body which link an animal's skeletal system into a functional whole.Saladin, Ken. Anatomy & Physiology. 7th ed. McGraw- ...
s, which 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 Biomechanics is the study of the structure, function and motion of the mechanical aspects of biological systems, at any level from whole organisms to organs, cells and cell organelles, using the methods of mechanics. Biomechanics is a branch of ...
). Bones protect internal organs, such as the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
protecting the
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a ve ...
or the
ribs The rib cage, as an enclosure that comprises the ribs, vertebral column and sternum in the thorax of most vertebrates, protects vital organs such as the heart, lungs and great vessels. The sternum, together known as the thoracic cage, is a semi- ...
protecting the
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide to t ...
and
lungs The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side ...
. Because of the way that bone is formed, bone has a high compressive strength of about , poor
tensile strength Ultimate tensile strength (UTS), often shortened to tensile strength (TS), ultimate strength, or F_\text within equations, is the maximum stress that a material can withstand while being stretched or pulled before breaking. In brittle materials t ...
of 104–121 MPa, and a very low
shear stress Shear stress, often denoted by (Greek: tau), is the component of stress coplanar with a material cross section. It arises from the shear force, the component of force vector parallel to the material cross section. ''Normal stress'', on the ...
strength (51.6 MPa). This means that bone resists pushing (compressional) stress well, resist pulling (tensional) stress less well, but only poorly resists shear stress (such as due to torsional loads). While bone is essentially
brittle A material is brittle if, when subjected to stress, it fractures with little elastic deformation and without significant plastic deformation. Brittle materials absorb relatively little energy prior to fracture, even those of high strength. Br ...
, bone does have a significant degree of elasticity, contributed chiefly by collagen. Mechanically, bones also have a special role in
hearing Hearing, or auditory perception, is the ability to perceive sounds through an organ, such as an ear, by detecting vibrations as periodic changes in the pressure of a surrounding medium. The academic field concerned with hearing is audit ...
. The ossicles are three small bones in the
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
which are involved in sound transduction.


Synthetic

The cancellous part of bones contain bone marrow. Bone marrow produces blood cells in a process called
hematopoiesis Haematopoiesis (, from Greek , 'blood' and 'to make'; also hematopoiesis in American English; sometimes also h(a)emopoiesis) is the formation of blood cellular components. All cellular blood components are derived from haematopoietic stem cells ...
. Blood cells that are created in bone marrow include
red blood cell Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek ''erythros'' for "red" and ''kytos'' for "holl ...
s,
platelet Platelets, also called thrombocytes (from Greek θρόμβος, "clot" and κύτος, "cell"), are a component of blood whose function (along with the coagulation factors) is to react to bleeding from blood vessel injury by clumping, thereby i ...
s and white blood cells. Progenitor cells such as the hematopoietic stem cell divide in a process called mitosis to produce precursor cells. These include precursors which eventually give rise to
white blood cells White blood cells, also called leukocytes or leucocytes, are the cells of the immune system that are involved in protecting the body against both infectious disease and foreign invaders. All white blood cells are produced and derived from mult ...
, and
erythroblast A nucleated red blood cell (NRBC), also known by several other names, is a red blood cell that contains a cell nucleus. Almost all vertebrate organisms have hemoglobin-containing cells in their blood, and with the exception of mammals, all of t ...
s which give rise to red blood cells. Unlike red and white blood cells, created by mitosis, platelets are shed from very large cells called megakaryocytes. This process of progressive differentiation occurs within the bone marrow. After the cells are matured, they enter the circulatory system, circulation. Every day, over 2.5 billion red blood cells and platelets, and 50–100 billion granulocytes are produced in this way. As well as creating cells, bone marrow is also one of the major sites where defective or aged red blood cells are destroyed.


Metabolic

* Mineral storage – bones act as reserves of minerals important for the body, most notably
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar t ...
and phosphorus. Determined by the species, age, and the type of bone, bone cells make up to 15 percent of the bone. 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 –
marrow adipose tissue Bone marrow adipose tissue (BMAT), sometimes referred to as marrow adipose tissue (MAT), is a type of fat deposit in bone marrow. It increases in states of low bone density -osteoporosis, anorexia nervosa/ caloric restriction, skeletal unweigh ...
(MAT) acts as a storage reserve of fatty acids. * Acid-Base (chemistry), base balance – bone buffers the blood against excessive pH changes by absorbing or releasing Alkali salt, 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 system, Endocrine organ – bone controls
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phosph ...
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 Osteocalcin, also known as bone gamma-carboxyglutamic acid-containing protein (BGLAP), is a small (49-amino-acid) noncollagenous protein hormone found in bone and dentin, first identified as a calcium-binding protein. Because osteocalcin has g ...
, which contributes to the regulation of blood sugar (glucose) and Adipose tissue, fat deposition. Osteocalcin increases both the insulin secretion and sensitivity, in addition to boosting the number of beta cell, 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

Bone is constantly being created and replaced in a process known as Bone remodeling, remodeling. This ongoing turnover of bone is a process of resorption followed by replacement of bone with little change in shape. This is accomplished through osteoblasts and osteoclasts. Cells are stimulated by a variety of paracrine, signals, and together referred to as a 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 Microdamage in bone, microdamaged bones from everyday stress, and to shape 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 piezoelectricity, piezoelectric properties, which cause bone to generate small electrical potentials under stress. The action of osteoblasts and osteoclasts are controlled by a number of chemical enzymes 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. For example, 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. Osteoblasts can also be stimulated to increase bone mass through increased secretion of
osteoid In histology, osteoid is the unmineralized, organic portion of the bone matrix that forms prior to the maturation of bone tissue. Osteoblasts begin the process of forming bone tissue by secreting the osteoid as several specific proteins. When ...
and by Enzyme inhibitor, inhibiting the ability of osteoclasts to break down osseous tissue. 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 reabsorption 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.


Volume

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 cancellous bone volume in postmenopausal osteoporosis may be determined by the relationship between the total bone forming surface and the percent of surface resorption.


Clinical significance

A number of diseases can affect bone, including arthritis, fractures, infections, osteoporosis and tumors. Conditions relating to bone can be managed by a variety of doctors, including Rheumatology, rheumatologists for joints, and orthopedic surgeons, who may conduct surgery to fix broken bones. Other doctors, such as rehabilitation medicine, rehabilitation specialists may be involved in recovery, radiology, 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.


Fractures

In normal bone,
fractures Fracture is the separation of an object or material into two or more pieces under the action of stress. The fracture of a solid usually occurs due to the development of certain displacement discontinuity surfaces within the solid. If a displa ...
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. When serious, depending on the fractures type and location, complications may include flail chest, compartment syndromes or fat embolism. Compound fractures involve the bone's penetration through the skin. Some complex fractures can be treated by the use of bone grafting procedures that replace missing bone portions. Fractures and their underlying causes can be investigated by X-rays, CT scans and MRIs. Fractures are described by their location and shape, and several classification systems exist, depending on the location of the fracture. A common long bone fracture in children is a 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 Physical medicine and rehabilitation, rehabilitation.


Tumors

There are several types of tumor that can affect bone; examples of benign bone tumors include osteoma, osteoid osteoma, osteochondroma, osteoblastoma, enchondroma, giant-cell tumor of bone, and aneurysmal bone cyst.


Cancer

Cancer can arise in bone tissue, and bones are also a common site for other cancers to spread (metastasise) to. Cancers that arise in bone are called "primary" cancers, although such cancers are rare. Metastases within bone are "secondary" cancers, with the most common being breast cancer, lung cancer, prostate cancer, thyroid cancer, and kidney cancer. Secondary cancers that affect bone can either destroy bone (called a "lytic cycle, lytic" cancer) or create bone (a "sclerosis (medicine), sclerotic" cancer). Cancers of the bone marrow inside the bone can also affect bone tissue, examples including leukemia and multiple myeloma. Bone may also be affected by cancers in other parts of the body. Cancers in other parts of the body may release parathyroid hormone or parathyroid hormone-related peptide. This increases bone reabsorption, and can lead to bone fractures. Bone tissue that is destroyed or altered as a result of cancers is distorted, weakened, and more prone to fracture. This may lead to compression of the spinal cord, destruction of the marrow resulting in bruising, bleeding and immunosuppression, and is one cause of bone pain. If the cancer is metastatic, then there might be other symptoms depending on the site of the original cancer. Some bone cancers can also be felt. Cancers of the bone are managed according to their type, their cancer staging, stage, prognosis, and what symptoms they cause. Many primary cancers of bone are treated with radiotherapy. Cancers of bone marrow may be treated with chemotherapy, and other forms of targeted therapy such as immunotherapy may be used. Palliative care, which focuses on maximising a person's quality of life, may play a role in management, particularly if the likelihood of 5-year survival, survival within five years is poor.


Other painful conditions

* Osteomyelitis is inflammation of the bone or bone marrow due to bacterial infection. * Osteomalacia is a painful softening of adult bone caused by severe vitamin D deficiency. * Osteogenesis imperfecta * Osteochondritis dissecans * Ankylosing spondylitis * Skeletal fluorosis is a bone disease caused by an excessive accumulation of fluoride in the bones. In advanced cases, skeletal fluorosis damages bones and joints and is painful.


Osteoporosis

Osteoporosis is a disease of bone where there is reduced bone mineral density, increasing the likelihood of bone fracture, fractures. Osteoporosis is defined in women by the World Health Organization as a bone mineral density of 2.5 standard deviations below peak bone mass, relative to the age and sex-matched average. This density is measured using dual energy X-ray absorptiometry (DEXA), 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 (medicine), chronic diseases or as a result of tobacco smoking, 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 are at risk of fracture. One of the most important risk factors for osteoporosis is ageing, advanced age. Accumulation of oxidative DNA damage (naturally occurring), DNA damage in osteocyte, osteoblastic and
osteoclast An osteoclast () is a type of bone cell that breaks down bone tissue. This function is critical in the maintenance, repair, and remodeling of bones of the vertebral skeleton. The osteoclast disassembles and digests the composite of hydrated pro ...
ic cells appears to be a key factor in age-related osteoporosis. Osteoporosis treatment includes advice to stop smoking, decrease alcohol consumption, exercise regularly, and have a healthy diet. Calcium and trace mineral supplements may also be advised, as may Vitamin D. When medication is used, it may include bisphosphonates, Strontium ranelate, and hormone replacement therapy.


Osteopathic medicine

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. , over 77,000 physicians Osteopathic medicine in the United States, in the United States are trained in osteopathic medical schools.


Osteology

The study of bones and teeth is referred to as osteology. It is frequently used in anthropology, archeology and forensics, 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 the process of maceration (bone), maceration. 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 also be made from external bones such as antlers.


Other animals

Bird skeletons are very lightweight. Their bones are smaller and thinner, to aid flight. 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 them being hollow. A bird's beak is primarily made of bone as projections of the mandibles which are covered in keratin. Some bones, primarily formed separately in subcutaneous tissues, include headgears (such as bony core of horns, antlers, ossicones), osteoderm, and os penis/ os clitoris. A deer's antlers are composed of bone which is an unusual example of bone being outside the skin of the animal once the velvet is shed. The extinct predatory fish ''Dunkleosteus'' had sharp edges of hard exposed bone along its jaws. The proportion of cortical bone that is 80% in the human skeleton may be much lower in other animals, especially in marine mammals and marine turtles, or in various Mesozoic marine reptiles, such as ichthyosaurs, among others. This proportion can vary quickly in evolution; it often increases in early stages of returns to an aquatic lifestyle, as seen in early whales and pinnipeds, among others. It subsequently decreases in pelagic taxa, which typically acquire spongy bone, but aquatic taxa that live in shallow water can retain very thick, Pachyostosis, pachyostotic, Osteosclerosis, osteosclerotic, or pachyosteosclerotic bones, especially if they move slowly, like Sirenia, sea cows. In some cases, even marine taxa that had acquired spongy bone can revert to thicker, compact bones if they become adapted to live in shallow water, or in Hypersaline lake, hypersaline (denser) water. Many animals, particularly herbivores, practice osteophagy—the eating of bones. This is presumably carried out in order to replenish lacking
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phosph ...
. Many bone diseases that affect humans also affect other vertebrates—an example of one disorder is skeletal fluorosis.


Society and culture

Bones from slaughtered animals have a number of uses. In prehistoric times, they have been used for making bone tools. They have further been used in bone carving, already important in prehistoric art, and also in modern time as crafting materials for buttons, beads, handles, bobbins, Napier's bones, calculation aids, head nuts, dice, poker chips, pick-up sticks, arrows, scrimshaw, ornaments, etc. Bone glue can be made by prolonged boiling of ground or cracked bones, followed by filtering and evaporation to thicken the resulting fluid. Historically once important, bone glue and other animal glues today have only a few specialized uses, such as in antiques restoration. Essentially the same process, with further refinement, thickening and drying, is used to make gelatin. Broth is made by simmering several ingredients for a long time, traditionally including bones. Bone char, a porous, black, granular material primarily used for filtration and also as a black pigment, is produced by charring mammal bones. Oracle bone script was a writing system used in Ancient China based on inscriptions in bones. Its name originates from oracle bones, which were mainly ox clavicle. The Ancient Chinese (mainly in the Shang dynasty), would write their questions on the oracle bone, and burn the bone, and where the bone cracked would be the answer for the questions. To :wikt:point the bone, point the bone at someone is considered bad luck in some cultures, such as Australian aborigines, such as by the Kurdaitcha#Bone pointing, Kurdaitcha. The Furcula, wishbones of fowl have been used for divination, and are still customarily used in a tradition to determine which one of two people pulling on either prong of the bone may make a wish. Various cultures throughout history have adopted the custom of shaping an infant's head by the practice of artificial cranial deformation. A widely practised custom in China was that of foot binding to limit the normal growth of the foot.


Additional images

File:Gray72-en.svg, Cells in bone marrow File:Bertazzo S - SEM deproteined trabecular - wistar rat - x100.tif, Scanning electron microscope of bone at 100× magnification File:Bone structure marco photo.jpg, Structure detail of an animal bone


See also

* Artificial bone * Bone health * Distraction osteogenesis * National Bone Health Campaign * Skeleton


References


Further reading

* * * * – ''drawings by Philip J.'' * * – ''Anthony edits the current version; Harrison edited previous versions.''


External links


Educational resource materials (including animations) by the American Society for Bone and Mineral Research


* [http://www.scq.ubc.ca/?p=400 A good basic overview of bone biology from the Science Creative Quarterly] *
Bone histology photomicrographs
{{Authority control Bones Skeletal system Connective tissue