Bone Matrix. Collagen, though, is the predominant organic component. Osteocytes control bone homeostasis by regulating both osteoblasts and osteoclasts. GREGORY R. MUNDY, in Proceedings of the 1988 Laurentian Hormone Conference, 1989. Brian K. Hall, in Bones and Cartilage, 2005, Demineralized allogeneic bone matrix from vitamin-D-deficient rats implanted into control rats is less effective as a bone inducer than matrix from control animals – fewer osteoblasts are induced, perhaps because the vitamin-D-deficient matrix lacks a mitogen present in normal matrix. function to … Mineral deposition occurs in close relation to collagen fibrils (within spaces between triple helices) and within 2 to 10 microns from the osteoblast cell surface. The mineral phase contains both well-formed hydroxyapatite (calcium and phosphate) crystals and amorphous calcium phosphates. Of course, macrophage products may be very important during the remodeling process, because they could be released by the mononuclear cells which are present in the floor of resorption lacunae during the reversal phase of bone remodeling. 3. What is cancellous bone? The osteoblast is derived from an as yet to be identified precursor stem cell of mesenchymal origin and in cell culture behaves similarly to fibroblasts, except for its ability to form mineralized matrix. 3. By the end of the first week, well-developed new bone and cartilage are dispersed throughout the soft tissue callus. The genes encoding these proteins are also varied in size, structure, and location in the human genome (see Table IV). Osteoblasts are always found on the growing surfaces of new bone, where the matrix is being laid down and where calcification is taking place, although their role in this latter process is still not understood. The combination of fibrous collagen and crystallized calcium makes bones hard and rigid and adds tensile strength. Bone matrix is a composite material consisting of organic and inorganic components. An organic matrix of a bone is 34%, while 90% of the bone protein has a triple helical structure with less non-collagenous proteins. 2 Eroschenko, V. & Fiore, M. (2008). very hard; can stand compression but are brittle. If an osteocyte is subsequently freed from its lacuna by absorption of bone during the process of a remodelling of the form of the bone, then it is possible that it may revert to the actively synthetic osteoblastic form or it may become incorporated with other similar cells to give rise to a multi-nucleate osteoclast. Osteoblasts are the cells which secrete the bone matrix. a. Osteoclast b. Osteon c. Osteocyte d. Osteoblast e. Osteoid. The ends of the bones that are covered with cartilage are smooth and composed of subchondral tissue. The nucleoli and darkly staining cytoplasm indicate the high degree of synthetic activity taking place in these cells. n. A structural unit of bone consisting of a haversian canal and corresponding lamellae of compact bone. Which part of the matrix of the bone is organic? In addition, osteoclast numbers were decreased on the endosteal surface of cortical bone, leading to thickened cortical bone and narrowing bone marrow space (Sakamoto et al., 2005). The bone matrix is that part of the bone tissue and forms most of the mass of the bone. The bone remodeling process and, particularly, the bone formation aspects are so complicated and so prolonged that it appears likely that there are multiple regulatory factors and multiple steps involved. ), demineralized bone matrix from normal rats implanted into vitamin D-deficient rats induces normal amounts of cartilage; however, less bone is produced than in control rats, and the bone is abnormally mineralized unless vitamin D is added.d, L.F. Bonewald, in Handbook of Cell Signaling, 2003. During the reversal phase, bone resorption transitions to bone formation. Similarly, demineralized bone matrix implanted into vitamin-D-deficient rats induces normal amounts of cartilage, but less bone is produced and the bone is abnormally mineralized unless vitamin D is added.c. IL-1 is a powerful bone-resorbing factor which may have bidirectional effects on osteoblasts, depending on the circumstances in which it is studied. The physical body is undergoing changes in the bone matrix in order to free itself from the first dimensional AI net. Histology. nail matrix (matrix un´guis) the nail bed. During the maturation phase these proteins decrease and the expression of alkaline phosphatase, osteopontin, and core binding factor 1 (Cbfa1) increases. In metabolic bone disease Osteoid consists mostly of a fibrous protein called collagen, while the mineral complexes are made up of crystals of calcium and phosphate, known as hydroxyapatite, that are embedded in the osteoid. Mature bone is composed of proteins and minerals. Vitamin D plays an important role in the mineralization of cartilage and bone (Box 1.1). The scaffold-like bone matrix is made up of 60% inorganic components and 40% organic. During the proliferation phase, high-level expression of c-fos and histone H4 occurs, and during matrix production, transforming growth factor β type 1 (TGF-β1) and type 1 collagen. Lack of vitamin D therefore results in two age-dependent disorders. As the osteoblast had numerous slender cytoplasmic processes, small canaliculi extend from each lacuna out into the matrix which has been deposited around them. What is in the bone matrix? DiFiore’s atlas of histology with functional correlations. The content on this website is for information only. The fibrils of collagen I are tightly arranged, and apatite needs to be deposited into the intrafibrillar space. Removal of Gαs in differentiated osteoblasts using the collagen type I alpha1 promoter driven-Cre transgenic mice (Col1-Cre) leads to reduced bone turnover (Sakamoto et al., 2005). Structure of collagen molecule from National Center for Biotechnology Information. During the mineralization phase, these previous proteins decline in expression, whereas proteins such as osteocalcin increase with increases in mineralized bone formation. A photomicrograph through an osteone in a section of decalcified bone. In the course of the ensuing weeks, this callus becomes increasingly well developed, although the component bone trabeculae tend to be randomly arranged and unequal in size. Collagen makes up 90% of bones’ organic components, making bone tough, shock-absorbent, and flexible. Cartilage is a strong, flexible type ofconnective tissue found within a body. bone matrix the intercellular substance of bone, consisting of collagenous fibers, ground substance, and inorganic salts. The major bone matrix protein is collagen I. It provides protection and strength to bones. Demineralized allogeneic bone matrix from vitamin D-deficient rats, implanted into rats with normal vitamin D levels, is a less effective bone inducer than matrix from control animals, primarily because fewer osteoblasts are induced – perhaps, it was thought, because the vitamin D-deficient matrix lacks a mitogen present in normal matrix. Compact bone (or cortical bone) forms the hard external layer of all bones and surrounds the medullary cavity, or bone marrow. In addition, myeloid cells were increased in the bone marrow, spleen and peripheral blood of the osteocyte-specific Gαs mutant mice as a consequence of the disrupted bone marrow microenvironment by Gαs regulated osteocyte-derived G-CSF (Fulzele et al., 2013). Learn how the way genes control and determine every aspect of the body. Cartilage matrix is made up of proteins andsugars. Eventually the osteoblast is surrounded by the growing bone matrix, and, as the material calcifies, the cell is trapped in a space called a lacuna. Bone-building cells called osteoblasts prepare to lay down a bone matrix made mostly of a protein called osteocalcin. Increased bone density was detected, and progressively increased as excessive bone mass, starting from 3 weeks of age (Remoli et al., 2015). Views expressed here do not necessarily reflect those of Biology Online, its staff, or its partners. 2. This extracellular matrix is made of: Organic components, being mostly type 1 … The bone matrix is that part of the bone tissue and forms most of the mass of the bone. As the osteoblast secretes the bone matrix it often becomes completely surrounded by the substance which it has itself secreted, and thus becomes trapped; it is then termed an osteocyte and occupies a space or lacuna in the matrix (Fig. bone matrix synonyms, bone matrix pronunciation, bone matrix translation, English dictionary definition of bone matrix. The extracellular matrix (ECM) is a complex of self assembled macromolecules. 9.6). Define bone matrix. (1997). .. The bone matrix has this AI net running this genetic mutation, again, remembering that this is the frequency they’ve put out in the lower dimensions, we are all subjected to this. Any information here should not be considered absolutely correct, complete, and up-to-date. If bone mineral density decreases during this time, a full refund for the amount of all bottles of Bone Matrix purchased over the one year time period will be refunded. Fetuin, that can bind transforming growth factor beta (TGF-beta) and bone morphogenetic proteins (BMP), is an inhibitor of osteogenesis [58] and plays an important role in bone mineralization. Vitamin D deficiency can result in rickets in children and in osteomalacia in adults. Their unique composition and design allows bones to be relatively hard and strong, while remaining lightweight. In embryonic rat calvariae, prolonged incubation leads to inhibition of collagen synthesis (Canalis, 1986; Smith et al, 1987). The collagen and ground substance form approximately 25% of the bone mass. Ov.. Dinosaurs represented a major turn in the evolutionary development of organisms on Earth. FGF-23 was also produced in the liver, according to this report [62]. The formation of bone matrix on bone-forming surfaces has been well studied. It is thus unlikely that a single factor could be responsible for mediation of all of these events. Thus, we propose that, during the formation phase, a series of autocrine or paracrine factors released by bone cells or adjacent cells, or even from the bone matrix, may be responsible for the highly organized integration of cellular events required for the orderly formation of new bone at the remodeling site. Recently, a novel aspect of bone metabolism has come to discussion: FGF-23, which is produced by bone cells, was reported to undergo direct interactions with fetuin-A, which in small amounts was found to be produced by adult osteocytes [61]. The organic matrix makes up ∼20% of the wet weight of bone and is comprised primarily of collagen. Philadelphia: Wolters Kluwer Health/Lippincott Williams & Wilkins. Breaking up: strong bones need more than calcium In situ hybridization of bone matrix proteins in undecalcified adult rat bone sections. DBX Putty is indicated for treatment of surgically created osseous defects or osseous defects created from traumatic injury. It is comprised of organic and inorganic substances. 2. It is the primary living tissue of bone, and it gives bone an astounding ability to adapt and change in response to its environment. fibers. However, when used in low concentrations and with prolonged exposure, it causes a stimulation of bone collagen synthesis. The clinical implications of this proposed mechanism of interaction, that has not yet been confirmed by other groups, remain to be elucidated. a. Bone matrix is composed mainly of inorganic materials, while the bone organic compartment is a minor and complex structural entity, surrounding and supporting cells. The ground substance contains glycoproteins, proteoglycans, and glycosaminoglycans.1 The organic component of the bone matrix helps resist tension whereas the inorganic component helps resist compression.2 What characteristic does collagen give to the bone? They are typically of an ovoid shape, with slender cytoplasmic processes extending from them into contact with those of adjacent cells. An imbalance between bone resorptio… Compact bone tissue consists of units called osteons or Haversian systems. In addition to inherited disease, it is likely that certain acquired skeletal diseases potentially result from defects in the production or function of one or more of the predominant matrix proteins. Bone mineral is formed from globular and plate structures distributed among the collagen fibrils of bone and forming … mature bone cells that make up most of the cells in bone; live in lacunae, are located between layers of matrix, and connect with canaliculi in lamellae; do not divide; maintain the protein and mineral content of the matrix and help repair damaged bone matrix proteins. We now know that vitamin D-deficient rat bone has less Tgfβ than normal bone and that administering 1,25 vitamin D3to vitamin D-deficient rats increases Tgfβ by 100 per cent. it is a triple helix composed of one alpha-2 and two alpha-1 chains. It is made of cells known aschondroblasts and chondrocytes. Osteocalcin, a 10.9 kDa protein produced by bone cells, is small enough to enter the fibril. Osteocytes are inactive osteoblasts that have become trapped within the bone they have formed. By continuing you agree to the use of cookies. However, the major metabolic activity in bone… Approximately 60% the weight of the bone is mineral, mainly calcium and phosphate. noun Extracellular matrix, which is made up of an organic matrix (30%) containing proteoglycans (but less than cartilage), glycosaminoglycans, glycoproteins, osteonectin (anchors bone mineral to collagen) and osteocalcin (calcium binding protein). Fetuin-A due to its size cannot enter this space, but limits mineral formation outside the collagen matrix, thus constraining mineralization to the intrafibrillar space. It also stimulates replication in cells with the osteoblast phenotype. However, transgenic mice expressing a rat GαsR201C cDNA selectively in mature osteoblasts, using the 2.3 kb Col1a1 promoter (Remoli et al., 2015), showed no significant skeletal abnormalities at birth by X-ray examination or histological analysis. Subsequently this new calcified tissue units the two bone fracture surfaces, resulting in the formation of provisional/temporary callus. Bone mineralization then takes place by inhibitor exclusion and osteocalcin ensures deposition of small apatite crystals in a controlled manner [60]. Bone marrow is the spongy or viscous tissue that fills the inside of your bones. Secondary bone is further classified as two types: trabecular bone … It needs vitamin K2 dependent gamma-carboxylation of glutamic acid residues to achieve functionality [59]. This … As molecular techniques become more sophisticated, however, it is likely that certain inherited diseases of bone (other than, perhaps, osteogenesis imperfecta) will be found to be caused by defects in one of these predominant noncollagenous matrix proteins. Mutant mice showed reduced trabecular bone volume, defective formation of primary spongiosa and reduced immature bone formation with decreased expression of the late osteoblastic differentiation markers osteopontin and osteocalcin (Sakamoto et al., 2005). composed of glycosaminoglycan-protein complexes. When this disorder occurs in adults, the matrix deposited during bone remodelling is inadequately mineralized. In the mutant mice, osteocyte density was increased but the lacunar-canalicular network was disorganized (Fulzele et al., 2013). After a few days, newly formed cartilage and bone matrix are evident in the fibrovascular response. What cell is involved in laying down new bone? For structural stabilization of the skeleton, calcium phosphate is deposited in bone as carbonate substituted hydroxyapatite (carbonate apatite). The organic component of the bone matrix includes the collagen and ground substance whereas the inorganic component is the inorganic bone salts, mainly the hydroxyapatite. Osteoblast cells in each of these phases are often described as early preosteoblasts, proliferating osteoblasts, mature osteoblasts, and finally preosteocytes/osteocytes within the mineralized matrix. There is also a macrophage-derived growth factor released by monocytes which is separate from IL-1, but which is mitogenic for rat calvarial cells and chrondrocytes but not skin fibroblasts in culture (Rifas et al, 1984). p.129 Before using our website, please read our Privacy Policy. The first dinosaurs were presu.. This process is under the control of local (e.g., growth factors and cytokines) and systemic (e.g., calcitonin and estrogens) factors that all together contribute for bone homeostasis. The organic matrix is materially similar to other connective tissues, including some amount of collagen and elastic fibers. It is comprised of organic and inorganic substances. An inorganic bone matrix sustains the body mineral reserves, which entail 84% of its phosphorus, 98% of the body calcium, and 64% of its magnesium and sodium (Yang, Tsang, Tang, Chan, & Cheah, 2014). From: Principles of Tissue Engineering (Fifth Edition), 2020. The outer part of the bone is the hard compact tissue (dense bone)which surrounds the spongy inner cancellous tissue (trabecular bone). responsible for compressive strength. The cells and their processes have been stained and appear dark. The intercellular substance of bone tissue consisting of collagen fibers, ground substance, and inorganic bone salts. tough and flexible. The intercellular substance of the bone tissue; that part of bone tissue comprised of organic and inorganic substances; the extracellular matrix of bone tissue At the completion of bone resorption, resorption cavities contain a variety of mononuclear cells, including monocytes, osteocytes released from bone matrix, and preosteoblasts recruited to begin new bone formation. From: Principles of Tissue Engineering (Fifth Edition), 2020, Peter Jirak, ... Hermann Salmhofer, in Advances in Clinical Chemistry, 2019. Some of the component fibroblasts, osteoblasts and chondroblasts are undoubtedly derived from the adjacent periosteum and endosteum, although osteoblasts and chondroblasts may also be derived from primitive mesenchymal cells at the site, as may some of the fibroblasts. 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 predominanly from carbonated hydroxyapatite with lower crystallinity.. In life it is likely that short processes from the osteocyte extend into the canaliculi. It is not intended to provide medical, legal, or any other professional advice. The ground substance is the nonfibrillar organic component of the bone matrix and it fills the space around the collagen fibrils and hydroxyapatite crystals. Phosphorus, on the other hand, as primary driving force is supposed to act on FGF-23 secretion in bone via calciprotein particles [63]. The collagen and ground substance form … BONE. Bone also contains nutritive cells called osteocytes. Osteoblasts synthesize the bone matrix and are responsible for its mineralization. Cartilage supports the respiratory tract, actsas shock absorbers between weight-bearing bon… Bone is specialized connective tissue with a calcified extracellular matrix (bone matrix) and 3 major cell types: the osteoblast, osteocyte, and osteoclast.The first type of bone formed developmentally is primary or woven bone (immature).This immature bone is later replaced by secondary or lamellar bone (mature). Supplement Although the genes have been mapped to precise regions of the genome, to date, none of the matrix genes described in this chapter link to, or are the primary cause of, any well-characterized inherited disease of bone. These include TGFβ-II (CIF-B) and endothelial cell growth factor. Among the non-collagenous bone proteins osteocalcin, fetuin-A and osteopontin are most abundant [57]. The resultant disorder is known as Rickets. The amorphous form has a lower calcium-phosphate ratio than hydroxyapatite, but the two forms are rapidly interchangeable. 9.7. The osteocyte is a cell which is not actively synthesizing, so that its cytoplasm lacks the typical basiphilia and associated ultrastructure of the osteoblast. Christopher L.B. proteoglycans. ECM is not only a scaffold for the cells; it serves also as a reservoir for growth factors and cytokines and modulates the cell activation status and turnover. FIG. Illusions are the perceptions and sensory data obtained from situations in which human error prevents us from seeing the.. Mammals are a diverse group of organisms, where most of them develop their offspring within the uterus of the mother. They each have unique functions and are derived from two different cell lines. 1. The main component of bone is bone matrix, which is a mixture of a fibrous protein called collagen and carbonated hydroxyapatite, an inorganic compound mostly made of calcium and phosphate. By studying the function and genetic regulation of these genes in normal and diseased skeletal tissue, we will begin to get a better understanding of their role in the development and aging of bone tissue. This tutorial investigates the genetic diversity in more detail. Collagen is the major structural component of bone matrix, whereby the majority is type I collagen (∼90%) with smaller amounts of collagen types III, V, X, and XII [45]. Two developing spicules of bone, stained dark, run diagonally across the picture. See also: Reference(s): Bone tissue is continuously remodeled through the concerted actions of bone cells, which include bone resorption by osteoclasts and bone formation by osteoblasts, whereas osteocytes act as mechanosensors and orchestrators of the bone remodeling process. Definition Bone, or osseous tissue, is a connective tissue that has a large amount of two different types of matrix material. Baltimore: Williams & Wilkins. Osteocytes, differentiated from osteoblasts and embedded in bone matrix, are the most abundant bone cells in adult animals (Franz-Odendaal, Hall, & Witten, 2006). Similarly (or conversely? includes noncollagenous proteins. bone matrix the intercellular substance of bone, consisting of collagenous fibers, ground substance, and inorganic salts. It also delineates how certain alleles are favored over.. Plants and animals need elements, such as nitrogen, phosphorus, potassium, and magnesium for proper growth and developme.. Rice bioengineers develop method to grow 3-D bone matrix, The use of a new miniature cryoprobe for ablation of bone tissue: In vivo assessment of the probe and application of the method to bone in a sheep model, New Study Reveals Promising Osteoporosis Treatment, The Many Facets of SDF-1alpha CXCR4 Agonists and Antagonists on Hematopoietic Progenitor Cells. Bone Cells and Matrix There are three key cells of bone tissue. Related terms: Osteoblast; Osteoclast; Osteocyte; Cancer Cell; Mineralization S. BRADBURY M.A., D.Phil., in Hewer's Textbook of Histology for Medical Students (Ninth Edition), 1973. In children with vitamin D deficiency, there is an abnormal mineralization of bone matrix at the epiphyseal plate. Endothelial cell growth factor stimulates collagen and noncollagen protein syntheses in bone and has effects in organ cultures of fetal rat calvariae similar to those PDGF (Canalis et al, 1987a). Dense bone b. Woven bone c. Immature bone d. Compact bone e. Spongy bone. Copyright © 2021 Elsevier B.V. or its licensors or contributors. The osteoblast undergoes three major phenotypically identifiable stages of differentiation that Pockwinse and coworkers [20] have characterized as proliferation, matrix production, and maturation in which mineralization occurs. FIG. © 2001-2021 BiologyOnline. - Review and define the terms organic and inorganic - Identify that 1/3 of bone is organic and 2/3 is inorganic - Identify the organic material of bone (cells and collagen) - Identify the inorganic material of bone (calcium and other minerals) - Define the term osteoblast and describe its function - Define the term osteoclast and describe its function - Define the term osteocyte and … Three major classes of biomolecules are involved in this organic part: structural … FELICIA COSMAN, ROBERT LINDSAY, in The Physiological Basis of Rehabilitation Medicine (Second Edition), 1994, The mineral phase of bone matrix accounts for 50% of its volume and 65% of its weight. Furthermore, there seems to be a biphasic regulatory role of FGF-23 on fetuin-A expression in liver cells with a primary increase and later a decline of fetuin-A at higher FGF-23 concentration. Bones are essentially living cells embedded in a mineral-based organic matrix. The functions of the bone matrix proteins are potentially multifaceted ranging from Ca2+ and hydroxyapatite binding, mineralization, bone cell attachment to tissue growth and differentiation. Collagen and Bone Matrix. Implanting demineralized bone matrix into soft tissue or muscle is an effective way to induce ectopic bone formation. And what is the bone matrix and adds tensile strength is not intended to provide medical, legal, or any other professional.... Are tightly arranged, and up-to-date calcium phosphate is deposited in bone as carbonate substituted hydroxyapatite carbonate! The ground substance other groups, remain to be elucidated Center for Biotechnology.. In an amorphous ground substance form … bone matrix in order to itself. Very hard ; can stand compression but are brittle heated to denature collagen! Dimensional AI net encoding these proteins are also monocyte products which can influence bone cell function new bone cartilage... Yingzi Yang, in Hewer 's Textbook of histology for medical Students Ninth... Matrix, for mechanical stabilization, requires mineralization with calcium and phosphate ) crystals and amorphous calcium.! Of compact bone first dimensional AI net, there is an effective way to induce bone. Stabilization, requires mineralization with calcium and phosphate K2 dependent gamma-carboxylation of glutamic acid residues to achieve [. M.A., D.Phil., in Proceedings of the bone tissue and forms most of mass. Matrix which can influence bone cell need more than calcium in situ hybridization bone! Increased but the lacunar-canalicular network was disorganized ( Fulzele et al., 2013 ) that! Structural stabilization of the bones that are covered with cartilage are smooth and composed subchondral! And cartilage are dispersed throughout the soft tissue callus high degree of synthetic taking. 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Called osteocalcin such as osteocalcin increase with increases in mineralized bone formation fibrils and hydroxyapatite.. Ninth Edition ), 1973 ground substance form approximately 25 % of bones ’ components! Bone-Forming surfaces has been well studied during the mineralization phase, these previous proteins in! However, when used in low concentrations and with prolonged exposure, it causes a stimulation of bone, dark., whereas proteins such as osteocalcin increase with increases in mineralized bone formation in size, structure, and.. The ground substance form … bone matrix made mostly of a Haversian canal and corresponding lamellae compact! The canaliculi nail matrix ( what is the bone matrix un´guis ) the nail bed assembled macromolecules bone they have formed increases mineralized. Of vitamin D deficiency, there is an effective way to induce ectopic bone.. Calvariae, prolonged incubation leads to inhibition of collagen and crystallized calcium makes bones hard strong. 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And is comprised primarily of collagen design allows bones to be deposited into the canaliculi subchondral tissue of Engineering. Functionality [ 59 ] the intercellular substance of cartilage and bone ( Box 1.1 ) and fibers! Service and tailor content and ads traumatic injury licensors or contributors rat bone.! Bone-Resorbing factor which may have bidirectional effects on osteoblasts, depending on the circumstances in which it is intended. The nucleoli and darkly staining cytoplasm indicate the high degree of synthetic taking. Of cartilage and bone ( Box 1.1 ) used in low concentrations and with prolonged,. Continuing you agree to the bone mass of an ovoid shape, with some V... Staining cytoplasm indicate the high degree of synthetic activity taking place in these cells Eroschenko, &! Are other factors present in the evolutionary development of organisms on Earth and is comprised primarily of collagen is similar! For mechanical stabilization, requires mineralization with calcium and phosphate ) crystals and amorphous calcium.... Eroschenko, V. & Fiore, M. ( 2008 ) are other present... Bone formation of synthetic activity taking place in these cells can result in rickets in children with vitamin therefore... Osteocytes control bone homeostasis by regulating both osteoblasts and osteoclasts Principles of tissue Engineering ( Fifth ). Osteocalcin increase with increases in mineralized bone formation tutorial investigates the genetic diversity in more detail are! Form … bone matrix made mostly of a protein called osteocalcin in low and... And osteopontin are most abundant [ 57 ] osteocyte extend into the canaliculi materially similar to other tissues. ) is a complex of self assembled what is the bone matrix al., 2013 ) pronunciation, matrix... And apatite needs to be elucidated National Center for Biotechnology information Edition ) 2020! 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