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Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by
cells Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
named
osteoblast 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 ...
s. It is synonymous with bone tissue formation. There are two processes resulting in the formation of normal, healthy bone tissue:
Intramembranous ossification Intramembranous ossification is one of the two essential processes during fetal development of the gnathostome (excluding chondrichthyans such as sharks) skeletal system by which rudimentary bone tissue is created. Intramembranous ossification is a ...
is the direct laying down of bone into the primitive connective tissue (
mesenchyme Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every o ...
), while
endochondral ossification Endochondral ossification is one of the two essential processes during fetal development of the mammalian skeletal system by which bone tissue is produced. Unlike intramembranous ossification, the other process by which bone tissue is produced, c ...
involves
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck an ...
as a precursor. In
fracture healing Bone healing, or fracture healing, is a proliferative physiological process in which the body facilitates the repair of a bone fracture. Generally, bone fracture treatment consists of a doctor reducing (pushing) displaced bones back into place ...
, endochondral osteogenesis is the most commonly occurring process, for example in fractures of long bones treated by plaster of Paris, whereas fractures treated by
open reduction and internal fixation Internal fixation is an operation in orthopedics that involves the surgical implementation of implants for the purpose of repairing a bone, a concept that dates to the mid-nineteenth century and was made applicable for routine treatment in the m ...
with metal plates,
screws A screw and a bolt (see '' Differentiation between bolt and screw'' below) are similar types of fastener typically made of metal and characterized by a helical ridge, called a ''male thread'' (external thread). Screws and bolts are used to fa ...
, pins, rods and nails may heal by intramembranous osteogenesis.
Heterotopic ossification Heterotopic ossification (HO) is the process by which bone tissue forms outside of the skeleton in muscles and soft tissue. Symptoms In traumatic heterotopic ossification (traumatic myositis ossificans), the patient may complain of a warm, tend ...
is a process resulting in the formation of bone tissue that is often atypical, at an extraskeletal location. Calcification is often confused with ossification. Calcification is synonymous with the formation of calcium-based salts and crystals within
cells Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
and tissue. It is a process that occurs during ossification, but not necessarily ''vice versa''. The exact mechanisms by which bone development is triggered remains unclear, but growth factors and
cytokines Cytokines are a broad and loose category of small proteins (~5–25 kDa) important in cell signaling. Cytokines are peptides and cannot cross the lipid bilayer of cells to enter the cytoplasm. Cytokines have been shown to be involved in autocrin ...
appear to play a role.


Intramembranous ossification

Intramembranous ossification Intramembranous ossification is one of the two essential processes during fetal development of the gnathostome (excluding chondrichthyans such as sharks) skeletal system by which rudimentary bone tissue is created. Intramembranous ossification is a ...
forms the flat bones of the skull, mandible and hip bone.


Endochondral ossification

Endochondral ossification Endochondral ossification is one of the two essential processes during fetal development of the mammalian skeletal system by which bone tissue is produced. Unlike intramembranous ossification, the other process by which bone tissue is produced, c ...
is the formation of long bones and other bones. This requires a
hyaline cartilage Hyaline cartilage is the glass-like (hyaline) and translucent cartilage found on many joint surfaces. It is also most commonly found in the ribs, nose, larynx, and trachea. Hyaline cartilage is pearl-gray in color, with a firm consistency and has ...
precursor. There are two centers of ossification for
endochondral ossification Endochondral ossification is one of the two essential processes during fetal development of the mammalian skeletal system by which bone tissue is produced. Unlike intramembranous ossification, the other process by which bone tissue is produced, c ...
. The primary center In long bones, bone tissue first appears in the diaphysis (middle of shaft).
Chondrocyte Chondrocytes (, from Greek χόνδρος, ''chondros'' = cartilage + κύτος, ''kytos'' = cell) are the only cells found in healthy cartilage. They produce and maintain the cartilaginous matrix, which consists mainly of collagen and proteog ...
s multiply and form trebeculae. Cartilage is progressively eroded and replaced by hardened bone, extending towards the epiphysis. A perichondrium layer surrounding the cartilage forms the periosteum, which generates sperm cells that then go on to make a collar that encircles the outside of the bone and remodels the medullary cavity on the inside. The nutrient artery enters via the
nutrient foramen All bones possess larger or smaller foramina (openings) for the entrance of blood-vessels; these are known as the nutrient foramina, and are particularly large in the shafts of the larger long bones, where they lead into a nutrient canal, which ext ...
from a small opening in the diaphysis. It invades the primary center of ossification, bringing osteogenic cells (
osteoblast 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 ...
s on the outside, osteoclasts on the inside.) The canal of the
nutrient foramen All bones possess larger or smaller foramina (openings) for the entrance of blood-vessels; these are known as the nutrient foramina, and are particularly large in the shafts of the larger long bones, where they lead into a nutrient canal, which ext ...
is directed away from more active end of bone when one end grows more than the other. When bone grows at same rate at both ends, the nutrient artery is perpendicular to the bone. Most other bones (e.g. vertebrae) also have primary ossification centers, and bone is laid down in a similar manner. Secondary centers The secondary centers generally appear at the epiphysis. Secondary ossification mostly occurs after birth (except for distal femur and proximal tibia which occurs during 9th month of fetal development). The epiphyseal arteries and osteogenic cells invade the epiphysis, depositing
osteoclasts 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 prote ...
and
osteoblasts 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 ...
which erode the cartilage and build bone, respectively. This occurs at both ends of long bones but only one end of digits and ribs.


Evolution

Several hypotheses have been proposed for how bone evolved as a structural element in vertebrates. One hypothesis is that bone developed from tissues that evolved to store minerals. Specifically, calcium-based minerals were stored in cartilage and bone was an
exaptation Exaptation and the related term co-option describe a shift in the function of a trait during evolution. For example, a trait can evolve because it served one particular function, but subsequently it may come to serve another. Exaptations are common ...
development from this calcified cartilage. However, other possibilities include bony tissue evolving as an osmotic barrier, or as a protective structure.


See also

* Dystrophic calcification * Mechanostat, a model describing ossification and bone loss * Ossicone, the horn-like (or antler-like) protuberances on the heads of giraffes and related species * Osteogenesis imperfecta, a juvenile bone disease * Fibrodysplasia ossificans progressiva, an extremely rare genetic disease which causes fibrous tissue (muscle, tendon, ligament etc.) to ossify when damaged * Primrose syndrome, a rare genetic disease in which cartilage becomes ossified.


References

{{Ossification Animal physiology Skeletal system *Ossification