Bones are rigid organs that form part of the endoskeleton of vertebrates. They function to move, support, and protect the various organs of the body, produce red and white blood cells and store minerals. Because bones come in a variety of shapes and have a complex internal and external structure, they are lightweight, yet strong and hard, in addition to fulfilling their many other functions. One of the types of tissues that makes up bone is the mineralized osseous tissue, also called bone tissue, that gives it rigidity and honeycomb-like three-dimensional internal structure. Other types of tissue found in bones include marrow, endosteum and periosteum, nerves, blood vessels and cartilage. There are 206 bones in the adult human body and about 270 in an infant.
Bones have ten main functions:
- Protection — Bones can serve to protect internal organs, such as the skull protecting the brain or the ribs protecting the heart and lungs.
- Shape — Bones provide a frame to keep the body supported.
- Blood production — The marrow, located within the medullary cavity of long bones and interstices of cancellous bone, produces blood cells in a process called haematopoiesis.
- Mineral storage — Bones act as reserves of minerals important for the body, most notably calcium and phosphorus.
- Fat Storage — The yellow bone marrow acts as a storage reserve of fatty acids
- Movement — Bones, 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.
- 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.
- Sound transduction — Bones are important in the mechanical aspect of hearing.
The primary tissue of bone, osseous tissue
, is a relatively hard
and lightweight composite material
, formed mostly of calcium phosphate
in the chemical arrangement termed calcium hydroxylapatite
(this is the osseous tissue
that gives bones their rigidity). It has relatively high compressive strength
but poor tensile strength
, meaning it resists pushing forces well, but not pulling forces. While bone is essentially brittle, it does have a significant degree of elasticity
, contributed chiefly by collagen
. All bones consist of living cells
embedded in the mineralized organic matrix
that makes up the osseous tissue.
Bone is not a uniformly solid material, but rather has some spaces between its hard elements.
Compact bone or (Cortical bone)
The hard outer layer of bones is composed of compact bone
tissue, so-called due to its minimal gaps and spaces. This tissue gives bones their smooth, white, and solid appearance, and accounts for 80% of the total bone mass of an adult skeleton. Compact bone may also be referred to as dense bone.
Filling the interior of the organ is the trabecular bone
tissue (an open cell porous
network also called cancellous or spongy bone) which is composed of a network of rod- and plate-like elements that make the overall organ lighter and allowing 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.
There are several types of cells constituting the bone;
- Osteoblasts are mononucleate bone-forming cells which descend from osteoprogenitor cells. They are located on the surface of osteoid seams and make a protein mixture known as osteoid, which mineralizes to become 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. Osteoblasts are the immature bone cells.
- Bone lining cells are essentially inactive osteoblasts. They cover all of the available bone surface and function as a barrier for certain ions.
- Osteocytes originate from osteoblasts which have migrated into and become trapped and surrounded by bone matrix which they themselves produce. The spaces which they occupy are known as lacunae. Osteocytes have many processes which reach out to meet osteoblasts and other osteocytes probably for the purposes of communication. Their functions include to varying degrees: formation of bone, matrix maintenance and calcium homeostasis. They have also been shown to act as mechano-sensory receptors—regulating the bone's response to stress and mechanical load. They are mature bone cells.
- Osteoclasts are the cells responsible for bone resorption (remodeling of bone to reduce its volume). Osteoclasts are large, multinucleated cells located on bone surfaces in what are called Howship's lacunae or resorption pits. These lacunae, or resorption pits, are left behind after the breakdown of the bone surface. 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 matrix is the major constituent of bone, surrounding the cells. It has inorganic and organic parts.
The inorganic is mainly crystalline mineral salts and calcium, which is present in the form of hydroxyapatite
. The matrix is initially laid down as unmineralized osteoid (manufactured by osteoblasts). Mineralisation involves osteoblasts secreting 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.
The organic part of matrix is mainly composed of Type I collagen
. This is synthesised intracellularly as tropocollagen and then exported, forming fibrils
. The organic part is also composed of various growth factors, the functions of which are not fully known. These factors present include glycosaminoglycans
, bone sialo protein
and Cell Attachment Factor. One of the main things that distinguishes the matrix of a bone from that of another cell is that the matrix in bone is hard.
Woven or lamellar
Bone is first deposited as woven bone, in a disorganized structure with a high proportion of osteocytes in young and in healing injuries. Woven bone is weaker, with a small number of randomly oriented collagen fibers, but forms quickly. It is replaced by lamellar bone, which is highly organized in concentric sheets with a low proportion of osteocytes. Lamellar bone is stronger and filled with many collagen fibers parallel to other fibers in the same layer (these parallel columns are called osteons). The fibers run in opposite directions in alternating layers, much like plywood, assisting in the bone's ability to resist torsion forces. After a break, woven bone quickly forms and is gradually replaced by slow-growing lamellar bone on pre-existing calcified hyaline cartilage through a process known as "bony substitution."
Five types of bones
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 greater in length than width. They are comprised mostly of compact bone and lesser amounts of marrow, which is 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.
- 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. The bones of the spine and hips are irregular bones.
- 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 force of the muscle is increased. Examples of sesamoid bones are the patella and the pisiform
The formation of bone during the fetal stage of development occurs by two processes: intramembranous
and endochondral ossification
Intramembranous ossification mainly occurs during formation of the flat bones of the skull; the bone is formed from mesenchyme tissue. 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, such as limbs; the bone is formed from cartilage. 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).
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 flat bones of the skull, the ribs, the vertebrae and pelvic bones.
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 units.
The purpose of remodeling is to regulate calcium homeostasis
, repair micro-damaged bones (from everyday stress) but also to shape and sculpture the skeleton during growth.
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.
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
are controlled by a number of chemical factors
which either promote or inhibit the activity of the bone remodelling 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.
Osteoblasts can be stimulated to increase bone mass through increased secretion of osteoid
and by inhibiting
the ability of osteoclasts to break down osseous tissue
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 reabsorbtion 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.
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.
There are many disorders of the skeleton. One of the more prominent is osteoporosis
Osteoporosis is a disease
of bone - leading to an increased risk of fracture
. In osteoporosis, the bone mineral density
(BMD) is reduced, bone microarchitecture is disrupted, and the amount and variety of non-collagenous
proteins in bone is altered. Osteoporosis is defined by the World Health Organization
(WHO) in women as a bone mineral density 2.5 standard deviations
below peak bone mass (20-year-old sex-matched healthy person average) as measured by DXA
; the term "established osteoporosis" includes the presence of a fragility fracture
. Osteoporosis is most common in women after the 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
, specifically glucocorticoids
, when the disease is called steroid-
or glucocorticoid-induced osteoporosis
(SIOP or GIOP).
Osteoporosis can be prevented with lifestyle advice and medication, and preventing falls in people with known or suspected osteoporosis is an established way to prevent fractures. Osteoporosis can be treated with bisphosphonates
and various other medical treatments.
Other disorders of bone include:
The study of bones and teeth is referred to as osteology
. It is frequently used in anthropology
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.
Alternatives to bony endoskeletons
There are several evolutionary
alternatives to mammilary bone; though they have some similar functions, they are not completely functionally analogous to bone.
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.
Several terms are used to refer to features and components of bones throughout the body:
|| Definition |
| articular process
|| A projection that contacts an adjacent bone. |
|| The region where adjacent bones contact each other—a joint. |
|| A long, tunnel-like foramen, usually a passage for notable nerves or blood vessels. |
|| A large, rounded articular process. |
|| A prominent ridge. |
|| A relatively small projection or bump. |
|| A projection near to a condyle but not part of the joint. |
|| A small, flattened articular surface. |
|| An opening through a bone. |
|| A broad, shallow depressed area. |
|| A small pit on the head of a bone. |
|| A cavity within a bone. |
|| A long, thin projection, often with a rough surface. Also known as a ridge. |
|| One of two specific protuberances of bones in the ankle. |
|| A short canal. |
|| A relatively large projection or prominent bump.(gen.) |
|| An arm-like branch off the body of a bone. |
|| A cavity within a cranial bone. |
|| A relatively long, thin projection or bump. |
|| Articulation between cranial bones. |
|| One of two specific tuberosities located on the femur. |
|| A projection or bump with a roughened surface, generally smaller than a tuberosity. |
|| A projection or bump with a roughened surface. |
Several terms are used to refer to specific features of long bones:
| Bone feature
|| Definition |
|| The long, relatively straight main body of a long bone; region of primary ossification. Also known as the shaft. |
|| 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. |
|| The proximal articular end of the bone. |
|| The region of a long bone lying between the epiphysis and diaphysis. |
|| The region of bone between the head and the shaft. |