69:
996:
40:
947:
981:
876:
H drives precipitation under a wide variety of extracellular conditions, as long as calcium and phosphate are available in the matrix compartment. The mechanism by which acid transits the barrier layer remains uncertain. Osteoblasts have capacity for Na/H exchange via the redundant Na/H exchangers, NHE1 and NHE6. This H exchange is a major element in acid removal, although the mechanism by which H is transported from the matrix space into the barrier osteoblast is not known.
966:
804:
935:
675:, are secreted in bone's organic matrix. Osteocalcin is not expressed at significant concentrations except in bone, and thus osteocalcin is a specific marker for bone matrix synthesis. These proteins link organic and mineral component of bone matrix. The proteins are necessary for maximal matrix strength due to their intermediate localization between mineral and collagen.
725:. Cartilage mineralizes by massive expression of phosphate-producing enzymes, which cause high local concentrations of calcium and phosphate that precipitate. This mineralized cartilage is not dense or strong. In the air breathing vertebrates it is used as a scaffold for formation of cellular bone made by osteoblasts, and then it is removed by
875:
and stopping further precipitation. Cartilage presents no barrier to diffusion and acid therefore diffuses away, allowing precipitation to continue. In the osteon, where matrix is separated from extracellular fluid by tight junctions, this cannot occur. In the controlled, sealed compartment, removing
1020:
In 1974 Jones et al. found that osteoblasts moved laterally in vivo and in vitro under different experimental conditions and described the migration method in detail. The osteoblasts were, however, contaminated by cells migrating from the vascular openings, which might include endothelial cells and
602:
hormones, is not fully understood, although it is likely that ACTH is bifunctional, like PTH, supporting bone formation with periodic spikes of ACTH, but causing bone destruction in large concentrations. In mice, mutations that reduce the efficiency of ACTH-induced glucocorticoid production in the
414:
Osteoclasts are multinucleated cells that derive from hematopoietic progenitors in the bone marrow which also give rise to monocytes in peripheral blood. Osteoclasts break down bone tissue, and along with osteoblasts and osteocytes form the structural components of bone. In the hollow within bones
1012:
The first isolation technique by microdissection method was originally described by Fell et al. using chick limb bones which were separated into periosteum and remaining parts. She obtained cells which possessed osteogenic characteristics from cultured tissue using chick limb bones which were
558:
gland under the control of serum calcium activity. PTH also has important systemic functions, including to keep serum calcium concentrations nearly constant regardless of calcium intake. Increasing dietary calcium results in minor increases in blood calcium. However, this is not a significant
655:
type I, which forms dense crosslinked ropes that give bone its tensile strength. By mechanisms still unclear, osteoblasts secrete layers of oriented collagen, with the layers parallel to the long axis of the bone alternating with layers at right angles to the long axis of the bone every few
48:(purple) rimming a bony spicule (pink - on diagonal of image). In this routinely fixed and decalcified (bone mineral removed) tissue, the osteoblasts have retracted and are separated from each other and from their underlying matrix. In living bone, the cells are linked by
423:
that supply oxygen and nutrients for bone formation. Bone is a highly vascular tissue, and active formation of blood vessel cells, also from mesenchymal stem cells, is essential to support the metabolic activity of bone. The balance of bone formation and
911:. The active osteoblast produces substantial collagen type I. About 10% of the bone matrix is collagen with the balance mineral. The osteoblast's nucleus is spherical and large. An active osteoblast is characterized morphologically by a prominent
642:
into osteoblasts and showing that the dye diffused to surrounding and deeper cells in the bone-forming unit. Bone is composed of many of these units, which are separated by impermeable zones with no cellular connections, called cement lines.
559:
mechanism supporting osteoblast bone formation, except in the condition of low dietary calcium; further, abnormally high dietary calcium raises the risk of serious health consequences not directly related to bone mass including
1493:
Sun L, Peng Y, Sharrow AC, Iqbal J, Zhang Z, Papachristou DJ, Zaidi S, Zhu LL, Yaroslavskiy BB, Zhou H, Zallone A, Sairam MR, Kumar TR, Bo W, Braun J, Cardoso-Landa L, Schaffler MB, Moonga BS, Blair HC, Zaidi M (April 2006).
946:
752:
bind strongly to bone mineral, when administered for short periods. They then accumulate in narrow bands in the new bone. These bands run across the contiguous group of bone-forming osteoblasts. They occur at a narrow
915:
that appears histologically as a clear zone adjacent to the nucleus. The products of the cell are mostly for transport into the osteoid, the non-mineralized matrix. Active osteoblasts can be labeled by antibodies to
887:
Feedback from physical activity maintains bone mass, while feedback from osteocytes limits the size of the bone-forming unit. An important additional mechanism is secretion by osteocytes, buried in the matrix, of
736:, packed around the collagen ropes. This is a strong composite material that allows the skeleton to be shaped mainly as hollow tubes. Reducing the long bones to tubes reduces weight while maintaining strength.
768:
older process by which cartilage is mineralized: tetracycline does not label mineralized cartilage at narrow bands or in specific sites, but diffusely, in keeping with a passive mineralization mechanism.
1016:
Enzymatic digestion is one of the most advanced techniques for isolating bone cell populations and obtaining osteoblasts. Peck et al. (1964) described the original method that is now often used by many
980:
788:(that is, by passive transporters, which do not pump calcium against a gradient). In contrast, phosphate is actively produced by a combination of secretion of phosphate-containing compounds, including
1949:
Roach HI (June 1994). "Why does bone matrix contain non-collagenous proteins? The possible roles of osteocalcin, osteonectin, osteopontin and bone sialoprotein in bone mineralisation and resorption".
586:, which are important in reproduction and energy metabolism regulation. Bone turnover involves major expenditures of energy for synthesis and degradation, involving many additional signals including
522:(BMPs) that determine to a major extent where chondrocyte differentiation occurs and where spaces are left between bones. The system of cartilage replacement by bone has a complex regulatory system.
288:
throughout life in the air-breathing vertebrates. The skeleton, often referred to as the skeletal system, is important both as a supporting structure and for maintenance of calcium, phosphate, and
761:, coordinated across the bone-forming group, is involved in bone formation, and that only cell-mediated mineral formation occurs. That is, dietary calcium does not create mineral by mass action.
1058:
Pittenger MF, Mackay AM, Beck SC, Jaiswal RK, Douglas R, Mosca JD, Moorman MA, Simonetti DW, Craig S, Marshak DR (April 1999). "Multilineage potential of adult human mesenchymal stem cells".
210:, osteoblasts function in groups of connected cells. Individual cells cannot make bone. A group of organized osteoblasts together with the bone made by a unit of cells is usually called the
965:
1031:
1036:
631:, small pores that connect osteoblasts, allowing the cells in one cohort to function as a unit. The gap junctions also connect deeper layers of cells to the surface layer (
1434:
Zaidi M, Sun L, Robinson LJ, Tourkova IL, Liu L, Wang Y, Zhu LL, Liu X, Li J, Peng Y, Yang G, Shi X, Levine A, Iqbal J, Yaroslavskiy BB, Isales C, Blair HC (May 2010).
550:
Many other regulatory systems are involved in the transition of cartilage to bone and in bone maintenance. A particularly important bone-targeted hormonal regulator is
757:) mineralization front. Most bone surfaces express no new bone formation, no tetracycline uptake and no mineral formation. This strongly suggests that facilitated or
1145:
Blair HC, Zaidi M, Huang CL, Sun L (November 2008). "The developmental basis of skeletal cell differentiation and the molecular basis of major skeletal defects".
151:
406:. Osteocytes remain alive and are connected by cell processes to a surface layer of osteoblasts. Osteocytes have important functions in skeletal maintenance.
2790:
995:
2263:"Concerning pH gradients between the extracellular compartment and fluids bathing the bone mineral surface and their relation to calcium ion distribution"
567:. Intermittent PTH stimulation increases osteoblast activity, although PTH is bifunctional and mediates bone matrix degradation at higher concentrations.
428:
tends to be negative with age, particularly in post-menopausal women, often leading to a loss of bone serious enough to cause fractures, which is called
892:, a protein that inhibits a pathway that maintains osteoblast activity. Thus, when the osteon reaches a limiting size, it deactivates bone synthesis.
538:
differentiation, which generally precedes bone formation for endochondral ossification. An additional family of essential regulatory factors is the
1188:
Blair HC, Sun L, Kohanski RA (November 2007). "Balanced regulation of proliferation, growth, differentiation, and degradation in skeletal cells".
1994:"Fourier transform infrared microspectroscopic analysis of bones of osteocalcin-deficient mice provides insight into the function of osteocalcin"
800:
is a membrane-anchored protein that is a characteristic marker expressed in large amounts at the apical (secretory) face of active osteoblasts.
682:), accumulation of mineral was not notably affected, indicating that organization of matrix is not significantly related to mineral transport.
2570:
1545:"Absence of HDL cholesteryl ester uptake in mice via SR-BI impairs an adequate adrenal glucocorticoid-mediated stress response to fasting"
1648:"Junctions between early developing osteoblasts of rat calvaria as revealed by freeze-fracture and ultrathin section electron microscopy"
2635:
Peck, W. A.; Birge, S. J.; Fedak, S. A. (11 December 1964). "Bone Cells: Biochemical and
Biological Studies after Enzymatic Isolation".
2446:
2757:
2739:
2245:
1334:
879:
In bone removal, a reverse transport mechanism uses acid delivered to the mineralized matrix to drive hydroxyapatite into solution.
698:, occur. Chondrocytes do not have intercellular connections and are not coordinated in units. Cartilage is composed of a network of
324:
with excellent tensile and compressive strength, which can bend under a strain and recover its shape without damage. This is called
316:. The matrix is mineralized by deposition of hydroxyapatite (alternative name, hydroxylapatite). This mineral is hard, and provides
368:
among other cell types. Osteoblast quantity is understood to be inversely proportional to that of marrow adipocytes which comprise
1013:
separated into periosteum and remaining parts. She obtained cells which possessed osteogenic characteristics from cultured tissue.
3168:
2783:
158:
2359:
Blair HC, Teitelbaum SL, Ghiselli R, Gluck S (August 1989). "Osteoclastic bone resorption by a polarized vacuolar proton pump".
627:
passage and thus create a bone compartment separate from the general extracellular fluid. The osteoblasts are also connected by
498:, although isolated mesenchymal stem cells in tissue culture may also form osteoblasts under permissive conditions that include
2596:"The Osteogenic Capacity in vitro of Periosteum and Endosteum Isolated from the Limb Skeleton of Fowl Embryos and Young Chicks"
678:
However, in mice where expression of osteocalcin or osteopontin was eliminated by targeted disruption of the respective genes (
527:
460:, which is then removed and replaced by bone, made by osteoblasts. Intramembranous ossification is the direct ossification of
68:
146:
2896:
772:
Osteoblasts separate bone from the extracellular fluid by tight junctions by regulated transport. Unlike in cartilage,
595:
523:
445:
419:. Components that are essential for osteoblast bone formation include mesenchymal stem cells (osteoblast precursor) and
2776:
934:
591:
491:
476:
448:. Endochondral ossification is the process of forming bone from cartilage and this is the usual method. This form of
39:
531:
519:
402:
When the bone-forming unit is not actively synthesizing bone, the surface osteoblasts are flattened and are called
289:
261:
2901:
2734:
and
Margaret W. Neuman. (1958). The Chemical Dynamics of Bone Mineral. Chicago: The University of Chicago Press.
441:
530:(TGF-Ξ²), is part of a superfamily of proteins that include BMPs, which possess common signaling elements in the
3163:
744:
The mechanisms of mineralization are not fully understood. Fluorescent, low-molecular weight compounds such as
539:
487:
472:
134:
122:
2033:
Thurner PJ, Chen CG, Ionova-Martin S, Sun L, Harman A, Porter A, Ager JW, Ritchie RO, Alliston T (June 2010).
694:, a solid avascular (without blood vessels) tissue in which individual cartilage-matrix secreting cells, or
661:
2553:
Baron, Roland; Rawadi, Georges; Roman-Roman, Sergio (2006). "WNT Signaling: A Key
Regulator of Bone Mass".
721:
In air-breathing vertebrates, cartilage is replaced by cellular bone. A transitional tissue is mineralized
789:
1543:
Hoekstra M, Meurs I, Koenders M, Out R, Hildebrand RB, Kruijt JK, Van Eck M, Van Berkel TJ (April 2008).
2824:
908:
785:
560:
369:
357:
218:
989:
of undecalcified tissue displaying osteoblasts actively synthesizing rudimentary bone tissue (center).
139:
3173:
3050:
2644:
2368:
1792:
1447:
1391:
Reid IR, Bristow SM, Bolland MJ (April 2015). "Cardiovascular complications of calcium supplements".
1197:
1067:
956:
797:
503:
480:
317:
244:, that is deposited in a highly regulated manner, into the organic matrix forming a strong and dense
3072:
1887:
Aubin JE, Liu F, Malaval L, Gupta AK (August 1995). "Osteoblast and chondroblast differentiation".
624:
616:
551:
325:
245:
3082:
2711:
2668:
2427:
2213:
1974:
1869:
1761:
1712:
1525:
1416:
1299:
1221:
1170:
921:
321:
1781:"Transitions in collagen types during matrix-induced cartilage, bone, and bone marrow formation"
959:
cancellous bone displaying osteoblasts actively synthesizing osteoid, containing two osteocytes.
387:, is mineralized by the osteoblasts. Before the organic matrix is mineralized, it is called the
1326:
732:
Osteoblasts produce an advanced type of bone matrix consisting of dense, irregular crystals of
2857:
2852:
2753:
2735:
2731:
2703:
2660:
2617:
2576:
2566:
2528:
2510:
2419:
2384:
2341:
2292:
2241:
2205:
2170:
2116:
2064:
2015:
1966:
1931:
1904:
1861:
1820:
1753:
1704:
1669:
1625:
1576:
1517:
1475:
1408:
1373:
1330:
1291:
1256:
1213:
1162:
1124:
1083:
395:. During bone formation, the surface layer of osteoblasts consists of cuboidal cells, called
328:. Forces that exceed the capacity of bone to behave elastically may cause failure, typically
3097:
3087:
3000:
2695:
2652:
2607:
2558:
2518:
2502:
2494:
2454:
2411:
2376:
2331:
2323:
2282:
2274:
2197:
2160:
2152:
2106:
2098:
2054:
2046:
2005:
1958:
1896:
1851:
1810:
1800:
1743:
1696:
1659:
1615:
1607:
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1556:
1507:
1465:
1455:
1400:
1363:
1283:
1248:
1205:
1154:
1114:
1075:
796:
that cleave phosphate to create a high phosphate concentration at the mineralization front.
758:
636:
313:
308:
It is synthesized and then the mineral is added. The vast majority of the organic matrix is
3045:
3004:
2745:
2686:
Jones, S.J.; Boyde, A. (December 1977). "Some morphological observations on osteoclasts".
2498:
1368:
1351:
917:
912:
819:
715:
620:
425:
341:
285:
49:
2648:
2482:
2372:
1796:
1451:
1201:
1071:
803:
570:
The skeleton is also modified for reproduction and in response to nutritional and other
3133:
2862:
2612:
2595:
2523:
2336:
2311:
2235:
2165:
2140:
2111:
2086:
2059:
2034:
1620:
1595:
1470:
1435:
781:
733:
707:
604:
583:
344:, which produce and secrete matrix proteins and transport mineral into the matrix, and
237:
207:
195:
179:
2562:
2287:
2262:
2010:
1993:
1815:
1780:
1319:
1252:
974:
of undecalcified tissue displaying osteoblasts actively synthesizing osteoid (center).
3157:
3138:
2819:
2139:
Blair HC, Robinson LJ, Huang CL, Sun L, Friedman PA, Schlesinger PH, Zaidi M (2011).
1900:
1158:
815:
679:
495:
465:
420:
329:
110:
53:
2715:
2672:
2402:
Klein-Nulend J, Nijweide PJ, Burger EH (June 2003). "Osteocyte and bone structure".
1978:
1873:
1716:
1420:
1303:
1225:
1174:
660:. Defects in collagen type I cause the commonest inherited disorder of bone, called
2943:
2891:
2883:
2837:
2431:
2217:
1765:
1529:
871:
In a closed system as mineral precipitates, acid accumulates, rapidly lowering the
765:
745:
703:
628:
599:
429:
241:
199:
2656:
900:
57:
1992:
Boskey AL, Gadaleta S, Gundberg C, Doty SB, Ducy P, Karsenty G (September 1998).
1079:
127:
3102:
2842:
2188:
Frost HM (1969). "Tetracycline-based histological analysis of bone remodeling".
1840:"Mutations in collagen genes: causes of rare and some common diseases in humans"
793:
726:
695:
672:
668:
639:
555:
457:
416:
345:
257:
253:
230:
226:
98:
2050:
1856:
1839:
1748:
1731:
1611:
1561:
1544:
1512:
1495:
784:
isolate the bone formation space. Calcium is transported across osteoblasts by
510:
that provides high concentrations of phosphate at the mineral deposition site.
376:, the thin connective tissue layer on the outside surface of bones, and in the
3123:
3065:
3024:
3009:
2927:
2829:
2459:
2415:
1687:
Doty SB (1981). "Morphological evidence of gap junctions between bone cells".
1287:
986:
971:
952:
904:
903:
staining (H&E) shows that the cytoplasm of active osteoblasts is slightly
889:
754:
657:
461:
392:
384:
373:
2514:
356:
Osteoblasts are the major cellular component of bone. Osteoblasts arise from
3128:
3118:
3092:
3060:
3014:
2995:
2990:
2922:
2811:
2803:
2380:
1805:
1460:
1274:
Nicks KM, Fowler TW, Gaddy D (June 2010). "Reproductive hormones and bone".
1209:
773:
722:
691:
587:
542:(FGFs) that determine where skeletal elements occur in relation to the skin
535:
499:
453:
377:
361:
269:
17:
2664:
2621:
2580:
2532:
2423:
2345:
2296:
2174:
2120:
2068:
1962:
1757:
1629:
1580:
1521:
1479:
1412:
1377:
1295:
1217:
1166:
1128:
1087:
2388:
2209:
2019:
1970:
1935:
1908:
1865:
1708:
1673:
1596:"The atherogenic Scarb1 null mouse model shows a high bone mass phenotype"
1260:
635:
when surrounded by bone). This was demonstrated directly by injecting low
452:
is the more complex form: it follows the formation of a first skeleton of
3077:
2707:
2035:"Osteopontin deficiency increases bone fragility but preserves bone mass"
1922:
Delmas PD, Demiaux B, Malaval L, Chapuy MC, Meunier PJ (April 1986). "".
1824:
699:
652:
579:
309:
281:
249:
222:
1664:
1647:
1594:
Martineau C, Martin-Falstrault L, Brissette L, Moreau R (January 2014).
1239:
Loutit, J.F.; Nisbet, N.W. (January 1982). "The Origin of
Osteoclasts".
822:
precipitating from phosphate, calcium, and water at a slightly alkaline
764:
The mechanism of mineral formation in bone is clearly distinct from the
3019:
2953:
2699:
2201:
2102:
1700:
1119:
1102:
777:
749:
575:
571:
388:
365:
301:
297:
265:
2506:
2327:
2278:
1571:
1404:
164:
3040:
2948:
2768:
2156:
925:
564:
507:
211:
2750:
Musculoskeletal system: anatomy, physiology, and metabolic disorders
2312:"High capacity Na+/H+ exchange activity in mineralizing osteoblasts"
2310:
Liu L, Schlesinger PH, Slack NM, Friedman PA, Blair HC (June 2011).
1732:"Functional gap junctions between osteocytic and osteoblastic cells"
1436:"ACTH protects against glucocorticoid-induced osteonecrosis of bone"
518:
Key growth factors in endochondral skeletal differentiation include
1730:
Yellowley CE, Li Z, Zhou Z, Jacobs CR, Donahue HJ (February 2000).
651:
Almost all of the organic (non-mineral) component of bone is dense
598:. The physiological role for responses to these, and several other
217:
Osteoblasts are specialized, terminally differentiated products of
1325:(3. ed.). Philadelphia, Pa.: Churchill Livingstone. pp.
802:
711:
483:
702:
type II held in tension by water-absorbing proteins, hydrophilic
2974:
2969:
2874:
2799:
2087:"Mechanisms balancing skeletal matrix synthesis and degradation"
449:
203:
183:
87:
2772:
814:
At least one more regulated transport process is involved. The
236:
In organized groups of disconnected cells, osteoblasts produce
225:
and specialized proteins in much smaller quantities, including
2481:
Robling, Alexander G.; Bonewald, Lynda F. (10 February 2020).
383:
Normally, almost all of the bone matrix, in the air breathing
256:. It is also an important store of minerals for physiological
872:
823:
667:
Minor, but important, amounts of small proteins, including
1646:
Arana-Chavez VE, Soares AM, Katchburian E (August 1995).
615:
In well-preserved bone studied at high magnification via
296:, is entirely extracellular. The bone matrix consists of
780:
cannot move in or out by passive diffusion, because the
292:
in the whole organism. The functional part of bone, the
1103:"Regulation of bone cell function by acid-base balance"
714:. It develops as the initial skeleton in more advanced
56:, and integrated with underlying osteocytes and matrix
619:, individual osteoblasts are shown to be connected by
1032:
List of human cell types derived from the germ layers
810:, the osteon, composed of osteoblasts and osteocytes.
729:, which specialize in degrading mineralized tissue.
554:(PTH). Parathyroid hormone is a protein made by the
252:
is the main support for the bodies of air breathing
3111:
3033:
2983:
2962:
2936:
2910:
2882:
2873:
2810:
2447:"Fun facts about bones: More than just scaffolding"
1779:Reddi AH, Gay R, Gay S, Miller EJ (December 1977).
1037:
List of distinct cell types in the adult human body
145:
133:
121:
109:
104:
93:
83:
78:
32:
1318:
924:dye fast blue to demonstrate alkaline phosphatase
340:Bone is a dynamic tissue that is constantly being
1838:Kuivaniemi H, Tromp G, Prockop DJ (April 1991).
372:. Osteoblasts are found in large numbers in the
320:. Thus, the collagen and mineral together are a
2134:
2132:
2130:
2085:Blair HC, Zaidi M, Schlesinger PH (June 2002).
2080:
2078:
1641:
1639:
1140:
1138:
1053:
1051:
391:. Osteoblasts buried in the matrix are called
2784:
2752:. Summit, New Jersey: Ciba-Geigy Corporation
2229:
2227:
603:adrenals cause the skeleton to become dense (
8:
486:. A second required transcription factor is
233:, which compose the organic matrix of bone.
1352:"Transcriptional control of skeletogenesis"
2879:
2791:
2777:
2769:
808:Major features of the bone-forming complex
526:also regulates early skeletal patterning.
248:, the mineralized matrix. The mineralized
2611:
2522:
2458:
2335:
2286:
2164:
2110:
2058:
2009:
1855:
1814:
1804:
1747:
1663:
1619:
1570:
1560:
1511:
1469:
1459:
1367:
1118:
1001:Osteoblasts lining bone (H&E stain).
860:
856:
852:
848:
844:
440:Bone is formed by one of two processes:
73:Illustration showing a single osteoblast
2555:Current Topics in Developmental Biology
1047:
940:Osteoblast (Wright Giemsa stain, 100x)
930:
464:as happens during the formation of the
920:, or using naphthol phosphate and the
162:
29:
2499:10.1146/annurev-physiol-021119-034332
2237:The Chemical Dynamics of Bone Mineral
1369:10.1146/annurev.genom.9.081307.164437
818:of bone mineral basically is that of
534:. TGF-Ξ² is particularly important in
494:differentiate under the influence of
360:(MSC). MSC give rise to osteoblasts,
221:. They synthesize dense, crosslinked
7:
896:Morphology and histological staining
27:Cells secreting extracellular matrix
2234:Neuman WF, Neuman MW (1958-01-01).
907:due to the substantial presence of
2557:. Vol. 76. pp. 103β127.
2261:Schartum S, Nichols G (May 1962).
1496:"FSH directly regulates bone mass"
25:
2445:Dance, Amber (23 February 2022).
1600:Am. J. Physiol. Endocrinol. Metab
415:are many other cell types of the
1159:10.1111/j.1469-185X.2008.00048.x
994:
979:
964:
945:
933:
706:. This is the adult skeleton in
348:, which break down the tissues.
159:Anatomical terms of microanatomy
67:
38:
2240:. University of Chicago Press.
647:Collagen and accessory proteins
611:Organization and ultrastructure
528:Transforming growth factor beta
477:the developing progenitor cells
1:
2657:10.1126/science.146.3650.1476
2563:10.1016/S0070-2153(06)76004-5
2483:"The Osteocyte: New Insights"
2011:10.1016/s8756-3282(98)00092-1
1253:10.1016/S0171-2985(82)80074-0
206:. However, in the process of
1901:10.1016/8756-3282(95)00183-E
1785:Proc. Natl. Acad. Sci. U.S.A
1440:Proc. Natl. Acad. Sci. U.S.A
1080:10.1126/science.284.5411.143
596:follicle stimulating hormone
546:Steroid and protein hormones
492:Osteochondroprogenitor cells
446:intramembranous ossification
2487:Annual Review of Physiology
1317:Larsen, William J. (2001).
909:rough endoplasmic reticulum
592:adrenocorticotropic hormone
590:hormones. Two of these are
520:bone morphogenetic proteins
514:Bone morphogenetic proteins
370:marrow adipose tissue (MAT)
3190:
2141:"Calcium and bone disease"
2051:10.1016/j.bone.2010.02.014
1857:10.1096/fasebj.5.7.2010058
1749:10.1359/jbmr.2000.15.2.209
1612:10.1152/ajpendo.00421.2013
1562:10.1194/jlr.M700475-JLR200
1513:10.1016/j.cell.2006.01.051
782:tight osteoblast junctions
690:The primitive skeleton is
532:TGF beta signaling pathway
2594:Fell, HB (January 1932).
2460:10.1146/knowable-022222-1
2416:10.1007/s11914-003-0002-y
1288:10.1007/s11914-010-0014-3
574:stresses; it responds to
540:fibroblast growth factors
468:of the skull and others.
442:endochondral ossification
284:is a large organ that is
157:
66:
37:
2688:Cell and Tissue Research
1356:Annu Rev Genom Hum Genet
1147:Biol Rev Camb Philos Soc
1007:Isolation of Osteoblasts
488:Sp7 transcription factor
304:. The protein forms the
3169:Connective tissue cells
2381:10.1126/science.2528207
1806:10.1073/pnas.74.12.5589
1461:10.1073/pnas.0912176107
1210:10.1196/annals.1402.029
662:osteogenesis imperfecta
479:express the regulatory
342:reshaped by osteoblasts
1963:10.1006/cbir.1994.1088
811:
740:Mineralization of bone
358:mesenchymal stem cells
219:mesenchymal stem cells
2825:fibrocartilage callus
901:Hematoxylin and eosin
806:
786:facilitated transport
686:Bone versus cartilage
182:combining forms for "
2606:(Pt 2): 157β180.11.
1895:(2 Suppl): 77Sβ83S.
1190:Ann. N. Y. Acad. Sci
798:Alkaline phosphatase
708:cartilaginous fishes
504:alkaline phosphatase
481:transcription factor
404:inactive osteoblasts
318:compressive strength
2649:1964Sci...146.1476P
2643:(3650): 1476β1477.
2404:Curr Osteoporos Rep
2373:1989Sci...245..855B
1797:1977PNAS...74.5589R
1665:10.1679/aohc.58.285
1652:Arch. Histol. Cytol
1452:2010PNAS..107.8782Z
1350:Karsenty G (2008).
1276:Curr Osteoporos Rep
1202:2007NYASA1116..165B
1072:1999Sci...284..143P
928:activity directly.
625:extracellular fluid
617:electron microscopy
552:parathyroid hormone
502:and substrates for
326:elastic deformation
286:formed and degraded
2700:10.1007/bf00220298
2600:Journal of Anatomy
2202:10.1007/BF02058664
2103:10.1042/BJ20020165
1736:J. Bone Miner. Res
1701:10.1007/BF02409482
1689:Calcif. Tissue Int
1120:10.1079/pns2003268
883:Osteocyte feedback
812:
471:During osteoblast
397:active osteoblasts
322:composite material
246:mineralized tissue
140:H2.00.03.7.00002
3151:
3150:
3147:
3146:
3056:connective tissue
3051:Volkmann's canals
2732:William F. Neuman
2572:978-0-12-153176-8
2451:Knowable Magazine
2328:10.1002/jcp.22501
2279:10.1172/JCI104569
2190:Calcif Tissue Res
1405:10.1002/jcb.25028
1101:Arnett T (2003).
312:, which provides
262:acid-base balance
194:"germinate") are
173:
172:
168:
16:(Redirected from
3181:
3098:medullary cavity
3073:Sharpey's fibres
3046:Haversian canals
3001:epiphyseal plate
2991:subchondral bone
2880:
2793:
2786:
2779:
2770:
2746:Netter, Frank H.
2720:
2719:
2683:
2677:
2676:
2632:
2626:
2625:
2615:
2591:
2585:
2584:
2550:
2544:
2543:
2541:
2539:
2526:
2478:
2472:
2471:
2469:
2467:
2462:
2442:
2436:
2435:
2399:
2393:
2392:
2356:
2350:
2349:
2339:
2316:J. Cell. Physiol
2307:
2301:
2300:
2290:
2258:
2252:
2251:
2231:
2222:
2221:
2185:
2179:
2178:
2168:
2157:10.1002/biof.143
2136:
2125:
2124:
2114:
2097:(Pt 2): 329β41.
2082:
2073:
2072:
2062:
2030:
2024:
2023:
2013:
1989:
1983:
1982:
1946:
1940:
1939:
1919:
1913:
1912:
1884:
1878:
1877:
1859:
1835:
1829:
1828:
1818:
1808:
1776:
1770:
1769:
1751:
1727:
1721:
1720:
1684:
1678:
1677:
1667:
1643:
1634:
1633:
1623:
1591:
1585:
1584:
1574:
1564:
1540:
1534:
1533:
1515:
1490:
1484:
1483:
1473:
1463:
1431:
1425:
1424:
1393:J. Cell. Biochem
1388:
1382:
1381:
1371:
1347:
1341:
1340:
1324:
1321:Human embryology
1314:
1308:
1307:
1271:
1265:
1264:
1247:(3β4): 193β203.
1236:
1230:
1229:
1185:
1179:
1178:
1142:
1133:
1132:
1122:
1098:
1092:
1091:
1055:
998:
987:Light micrograph
983:
972:Light micrograph
968:
953:Light micrograph
949:
937:
864:
847:O + 10 Ca β Ca
842:
841:
838:
766:phylogenetically
759:active transport
640:fluorescent dyes
637:molecular weight
623:, which prevent
450:bone development
314:tensile strength
290:acid-base status
202:that synthesize
165:edit on Wikidata
71:
42:
30:
21:
3189:
3188:
3184:
3183:
3182:
3180:
3179:
3178:
3164:Skeletal system
3154:
3153:
3152:
3143:
3107:
3029:
3005:epiphyseal line
2979:
2958:
2932:
2906:
2897:intramembranous
2869:
2806:
2797:
2767:
2728:
2726:Further reading
2723:
2685:
2684:
2680:
2634:
2633:
2629:
2593:
2592:
2588:
2573:
2552:
2551:
2547:
2537:
2535:
2480:
2479:
2475:
2465:
2463:
2444:
2443:
2439:
2401:
2400:
2396:
2367:(4920): 855β7.
2358:
2357:
2353:
2309:
2308:
2304:
2267:J. Clin. Invest
2260:
2259:
2255:
2248:
2233:
2232:
2225:
2187:
2186:
2182:
2138:
2137:
2128:
2084:
2083:
2076:
2032:
2031:
2027:
1991:
1990:
1986:
1948:
1947:
1943:
1921:
1920:
1916:
1886:
1885:
1881:
1837:
1836:
1832:
1791:(12): 5589β92.
1778:
1777:
1773:
1729:
1728:
1724:
1686:
1685:
1681:
1645:
1644:
1637:
1593:
1592:
1588:
1542:
1541:
1537:
1492:
1491:
1487:
1433:
1432:
1428:
1390:
1389:
1385:
1349:
1348:
1344:
1337:
1316:
1315:
1311:
1273:
1272:
1268:
1238:
1237:
1233:
1187:
1186:
1182:
1144:
1143:
1136:
1100:
1099:
1095:
1066:(5411): 143β7.
1057:
1056:
1049:
1045:
1028:
1009:
1002:
999:
990:
984:
975:
969:
960:
950:
941:
938:
918:Type-I collagen
913:Golgi apparatus
898:
885:
862:
858:
854:
850:
846:
839:
836:
835:
833:
742:
688:
649:
621:tight junctions
613:
584:glucocorticoids
548:
516:
473:differentiation
438:
426:bone resorption
412:
354:
338:
336:Bone remodeling
306:organic matrix.
278:
260:including both
169:
74:
62:
61:
50:tight junctions
28:
23:
22:
15:
12:
11:
5:
3187:
3185:
3177:
3176:
3171:
3166:
3156:
3155:
3149:
3148:
3145:
3144:
3142:
3141:
3136:
3131:
3126:
3121:
3115:
3113:
3109:
3108:
3106:
3105:
3100:
3095:
3090:
3085:
3080:
3075:
3070:
3069:
3068:
3063:
3053:
3048:
3043:
3037:
3035:
3031:
3030:
3028:
3027:
3022:
3017:
3012:
3007:
2998:
2993:
2987:
2985:
2981:
2980:
2978:
2977:
2972:
2966:
2964:
2960:
2959:
2957:
2956:
2951:
2946:
2940:
2938:
2934:
2933:
2931:
2930:
2925:
2920:
2914:
2912:
2908:
2907:
2905:
2904:
2899:
2894:
2888:
2886:
2877:
2871:
2870:
2868:
2867:
2866:
2865:
2860:
2855:
2847:
2846:
2845:
2840:
2832:
2827:
2822:
2816:
2814:
2808:
2807:
2798:
2796:
2795:
2788:
2781:
2773:
2766:
2765:External links
2763:
2762:
2761:
2743:
2727:
2724:
2722:
2721:
2678:
2627:
2586:
2571:
2545:
2493:(1): 485β506.
2473:
2437:
2394:
2351:
2322:(6): 1702β12.
2302:
2253:
2246:
2223:
2180:
2126:
2074:
2045:(6): 1564β73.
2025:
1984:
1951:Cell Biol. Int
1941:
1914:
1879:
1850:(7): 2052β60.
1830:
1771:
1722:
1679:
1635:
1586:
1535:
1485:
1446:(19): 8782β7.
1426:
1399:(4): 494β501.
1383:
1342:
1335:
1309:
1266:
1231:
1180:
1134:
1093:
1046:
1044:
1041:
1040:
1039:
1034:
1027:
1024:
1023:
1022:
1018:
1014:
1008:
1005:
1004:
1003:
1000:
993:
991:
985:
978:
976:
970:
963:
961:
951:
944:
942:
939:
932:
897:
894:
884:
881:
869:
868:
867:
866:
820:hydroxyapatite
741:
738:
734:hydroxyapatite
687:
684:
648:
645:
612:
609:
605:osteosclerotic
547:
544:
515:
512:
496:growth factors
466:membrane bones
437:
434:
411:
408:
353:
350:
337:
334:
330:bone fractures
277:
276:Bone structure
274:
238:hydroxyapatite
208:bone formation
198:with a single
190:and βλαΟΟάνΟ,
171:
170:
161:
155:
154:
149:
143:
142:
137:
131:
130:
125:
119:
118:
113:
107:
106:
102:
101:
95:
91:
90:
85:
81:
80:
76:
75:
72:
64:
63:
44:
43:
35:
34:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
3186:
3175:
3172:
3170:
3167:
3165:
3162:
3161:
3159:
3140:
3137:
3135:
3132:
3130:
3127:
3125:
3122:
3120:
3117:
3116:
3114:
3110:
3104:
3101:
3099:
3096:
3094:
3091:
3089:
3086:
3084:
3081:
3079:
3076:
3074:
3071:
3067:
3064:
3062:
3059:
3058:
3057:
3054:
3052:
3049:
3047:
3044:
3042:
3039:
3038:
3036:
3032:
3026:
3023:
3021:
3018:
3016:
3013:
3011:
3008:
3006:
3002:
2999:
2997:
2994:
2992:
2989:
2988:
2986:
2982:
2976:
2973:
2971:
2968:
2967:
2965:
2961:
2955:
2952:
2950:
2947:
2945:
2942:
2941:
2939:
2935:
2929:
2926:
2924:
2921:
2919:
2916:
2915:
2913:
2909:
2903:
2900:
2898:
2895:
2893:
2890:
2889:
2887:
2885:
2881:
2878:
2876:
2872:
2864:
2861:
2859:
2856:
2854:
2851:
2850:
2848:
2844:
2841:
2839:
2836:
2835:
2833:
2831:
2828:
2826:
2823:
2821:
2820:perichondrium
2818:
2817:
2815:
2813:
2809:
2805:
2801:
2794:
2789:
2787:
2782:
2780:
2775:
2774:
2771:
2764:
2759:
2758:0-914168-88-6
2755:
2751:
2747:
2744:
2741:
2740:0-226-57512-8
2737:
2733:
2730:
2729:
2725:
2717:
2713:
2709:
2705:
2701:
2697:
2694:(3): 387β97.
2693:
2689:
2682:
2679:
2674:
2670:
2666:
2662:
2658:
2654:
2650:
2646:
2642:
2638:
2631:
2628:
2623:
2619:
2614:
2609:
2605:
2601:
2597:
2590:
2587:
2582:
2578:
2574:
2568:
2564:
2560:
2556:
2549:
2546:
2534:
2530:
2525:
2520:
2516:
2512:
2508:
2504:
2500:
2496:
2492:
2488:
2484:
2477:
2474:
2461:
2456:
2452:
2448:
2441:
2438:
2433:
2429:
2425:
2421:
2417:
2413:
2409:
2405:
2398:
2395:
2390:
2386:
2382:
2378:
2374:
2370:
2366:
2362:
2355:
2352:
2347:
2343:
2338:
2333:
2329:
2325:
2321:
2317:
2313:
2306:
2303:
2298:
2294:
2289:
2284:
2280:
2276:
2273:(5): 1163β8.
2272:
2268:
2264:
2257:
2254:
2249:
2247:0-226-57512-8
2243:
2239:
2238:
2230:
2228:
2224:
2219:
2215:
2211:
2207:
2203:
2199:
2196:(1): 211β37.
2195:
2191:
2184:
2181:
2176:
2172:
2167:
2162:
2158:
2154:
2151:(3): 159β67.
2150:
2146:
2142:
2135:
2133:
2131:
2127:
2122:
2118:
2113:
2108:
2104:
2100:
2096:
2092:
2088:
2081:
2079:
2075:
2070:
2066:
2061:
2056:
2052:
2048:
2044:
2040:
2036:
2029:
2026:
2021:
2017:
2012:
2007:
2004:(3): 187β96.
2003:
1999:
1995:
1988:
1985:
1980:
1976:
1972:
1968:
1964:
1960:
1957:(6): 617β28.
1956:
1952:
1945:
1942:
1937:
1933:
1930:(14): 643β6.
1929:
1926:(in French).
1925:
1918:
1915:
1910:
1906:
1902:
1898:
1894:
1890:
1883:
1880:
1875:
1871:
1867:
1863:
1858:
1853:
1849:
1845:
1841:
1834:
1831:
1826:
1822:
1817:
1812:
1807:
1802:
1798:
1794:
1790:
1786:
1782:
1775:
1772:
1767:
1763:
1759:
1755:
1750:
1745:
1742:(2): 209β17.
1741:
1737:
1733:
1726:
1723:
1718:
1714:
1710:
1706:
1702:
1698:
1695:(5): 509β12.
1694:
1690:
1683:
1680:
1675:
1671:
1666:
1661:
1658:(3): 285β92.
1657:
1653:
1649:
1642:
1640:
1636:
1631:
1627:
1622:
1617:
1613:
1609:
1606:(1): E48β57.
1605:
1601:
1597:
1590:
1587:
1582:
1578:
1573:
1568:
1563:
1558:
1555:(4): 738β45.
1554:
1550:
1546:
1539:
1536:
1531:
1527:
1523:
1519:
1514:
1509:
1506:(2): 247β60.
1505:
1501:
1497:
1489:
1486:
1481:
1477:
1472:
1467:
1462:
1457:
1453:
1449:
1445:
1441:
1437:
1430:
1427:
1422:
1418:
1414:
1410:
1406:
1402:
1398:
1394:
1387:
1384:
1379:
1375:
1370:
1365:
1361:
1357:
1353:
1346:
1343:
1338:
1336:0-443-06583-7
1332:
1328:
1323:
1322:
1313:
1310:
1305:
1301:
1297:
1293:
1289:
1285:
1281:
1277:
1270:
1267:
1262:
1258:
1254:
1250:
1246:
1242:
1241:Immunobiology
1235:
1232:
1227:
1223:
1219:
1215:
1211:
1207:
1203:
1199:
1196:(1): 165β73.
1195:
1191:
1184:
1181:
1176:
1172:
1168:
1164:
1160:
1156:
1153:(4): 401β15.
1152:
1148:
1141:
1139:
1135:
1130:
1126:
1121:
1116:
1113:(2): 511β20.
1112:
1108:
1107:Proc Nutr Soc
1104:
1097:
1094:
1089:
1085:
1081:
1077:
1073:
1069:
1065:
1061:
1054:
1052:
1048:
1042:
1038:
1035:
1033:
1030:
1029:
1025:
1019:
1015:
1011:
1010:
1006:
997:
992:
988:
982:
977:
973:
967:
962:
958:
954:
948:
943:
936:
931:
929:
927:
923:
919:
914:
910:
906:
902:
895:
893:
891:
882:
880:
877:
874:
865:
831:
830:
829:
828:
827:
825:
821:
817:
816:stoichiometry
809:
805:
801:
799:
795:
791:
787:
783:
779:
775:
770:
767:
762:
760:
756:
751:
747:
739:
737:
735:
730:
728:
724:
719:
717:
713:
709:
705:
704:proteoglycans
701:
697:
693:
685:
683:
681:
680:knockout mice
676:
674:
670:
665:
663:
659:
654:
646:
644:
641:
638:
634:
630:
629:gap junctions
626:
622:
618:
610:
608:
606:
601:
597:
593:
589:
585:
581:
577:
573:
568:
566:
562:
557:
553:
545:
543:
541:
537:
533:
529:
525:
521:
513:
511:
509:
505:
501:
497:
493:
489:
485:
482:
478:
474:
469:
467:
463:
459:
455:
451:
447:
443:
435:
433:
431:
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746:tetracycline
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696:chondrocytes
689:
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600:glycoprotein
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242:bone mineral
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116:osteoblastus
115:
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3174:Human cells
3103:bone marrow
2843:chondrocyte
2410:(1): 5β10.
1282:(2): 60β7.
957:decalcified
727:osteoclasts
673:osteopontin
669:osteocalcin
658:micrometers
594:(ACTH) and
556:parathyroid
484:Cbfa1/Runx2
417:bone marrow
410:Osteoclasts
385:vertebrates
352:Osteoblasts
346:osteoclasts
294:bone matrix
258:homeostasis
254:vertebrates
231:osteopontin
227:osteocalcin
186:", α½ΟΟΞΞΏ-,
176:Osteoblasts
105:Identifiers
99:bone tissue
46:Osteoblasts
3158:Categories
3093:trabeculae
3088:canaliculi
3066:periosteum
3025:epicondyle
3010:metaphysis
2970:cancellous
2928:osteoclast
2918:osteoblast
2830:metaphysis
2507:1805/30982
2145:BioFactors
2091:Biochem. J
1924:Presse Med
1572:2066/69489
1362:: 183β96.
1043:References
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890:sclerostin
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633:osteocytes
462:mesenchyme
393:osteocytes
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362:adipocytes
178:(from the
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2515:0066-4278
922:diazonium
792:, and by
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692:cartilage
588:pituitary
536:cartilage
500:vitamin C
454:cartilage
378:endosteum
270:phosphate
3139:sesamoid
3078:enthesis
2975:cortical
2748:(1987).
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2665:14208576
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1167:18710437
1129:14506899
1088:10102814
1026:See also
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580:estrogen
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2858:elastic
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2645:Bibcode
2637:Science
2613:1248877
2538:8 March
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2369:Bibcode
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2020:9737340
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1068:Bibcode
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