Bone remodeling during fracture repair: The cellular picture

Gerstenfeld, 2003, Fracture healing as a post-natal developmental process: molecular, spatial, and temporal aspects of its regulation, J Cell Biochem, 88, 873, 10.1002/jcb.10435

Einhorn, 1998, The cell and molecular biology of fracture healing, Clin Orthop Relat Res, 355, S7, 10.1097/00003086-199810001-00003

Bolander, 1992, Regulation of fracture repair by growth factors, Proc Soc Exp Biol Med, 200, 165, 10.3181/00379727-200-43410A

AI-Aql, 2008, Molecular mechanisms controlling bone formation during fracture healing and distraction. Osteogenesis, J Dent Res, 87, 107, 10.1177/154405910808700215

Giannoudis, 2007, Fracture healing: the diamond concept, Injury, 38, S3, 10.1016/S0020-1383(08)70003-2

Stone, 1997, A molecular approach to bone regeneration, Br J Plast Surg, 1997, 369, 10.1016/S0007-1226(97)90547-6

Barnes, 1999, Growth factor regulation of fracture repair, J Bone Miner Res, 14, 1805, 10.1359/jbmr.1999.14.11.1805

Iwaki, 1997, Localization and quantification of proliferating cells during rat fracture repair: detection of proliferating cell nuclear antigen by immunohistochemistry, J Bone Miner Res, 12, 96, 10.1359/jbmr.1997.12.1.96

Shapiro, 2008, Bone development and its relation to fracture repair. The role of mesenchymal osteoblasts and surface osteoblasts, Eur Cells Mater, 15, 53, 10.22203/eCM.v015a05

Malizos, 2005, The healing potential of the periosteum. Molecular aspects, Injury, 36, S13, 10.1016/j.injury.2005.07.030

Baksh, 2004, Adult mesenchymal stem cells: characterization, differentiation, and application in cell and gene therapy, J Cell Mol Med, 8, 301, 10.1111/j.1582-4934.2004.tb00320.x

Colnot, 2006, Analyzing the cellular contribution of bone marrow to fracture healing using bone marrow transplantation in mice, Biochem Biophys Res Commun, 350, 557, 10.1016/j.bbrc.2006.09.079

Eghbali-Fatourechi, 2005, Circulating osteoblast-lineage cells in humans, N Engl J Med, 352, 1959, 10.1056/NEJMoa044264

Rumi, 2005, The origin of osteoprogenitor cells responsible for heterotopic ossification following hip surgery: an animal model in the rabbit, J Orthop Res, 23, 34, 10.1016/j.orthres.2004.05.003

Kalfas, 2001, Principles of bone healing, Neurosurg Focus, 10, E1, 10.3171/foc.2001.10.4.2

Deckers, 2002, Bone morphogenetic proteins stimulate angiogenesis through osteoblast-derived vascular endothelial growth factor A, Endocrinology, 143, 1545, 10.1210/en.143.4.1545

Zelzer, 2001, Tissue specific regulation of VEGF expression during bone development requires Cbfa1/Runx2, Mech Dev, 106, 97, 10.1016/S0925-4773(01)00428-2

Pufe, 2002, Quantitative measurement of the splice variants 120 and 164 of the angiogenic peptide vascular endothelial growth factor in the time flow of fracture healing: a study in the rat, Cell Tissue Res, 309, 387, 10.1007/s00441-002-0605-0

Chen, 2004, Bone morphogenetic proteins, Growth Factors, 22, 233, 10.1080/08977190412331279890

Nakase, 2006, Potential roles of bone morphogenetic proteins (BMPs) in skeletal repair and regeneration, J Bone Miner Metab, 24, 425, 10.1007/s00774-006-0718-8

Hutmacher, 2003, Periosteal cells in bone tissue engineering, Tissue Eng., 9, S45, 10.1089/10763270360696978

Collett, 2005, Angiogenesis and pericytes in the initiation of ectopic calcification, Circ Res, 96, 930, 10.1161/01.RES.0000163634.51301.0d

Peng, 2005, VEGF improves, whereas sFlt1 inhibits, BMP2-induced bone formation and bone healing through modulation of angiogenesis, J Bone Miner Res, 20, 2017, 10.1359/JBMR.050708

Tarkka, 2003, Adenoviral VEGF-A gene transfer induces angiogenesis and promotes bone formation in healing osseous tissues, J Gene Med, 5, 560, 10.1002/jgm.392

Brownlow, 2002, The vascularity of atrophic non-unions, Injury, 33, 145, 10.1016/S0020-1383(01)00153-X

Reed, 2002, Human atrophic fracture non-unions are not avascular, J Orthop Res, 20, 593, 10.1016/S0736-0266(01)00142-5

Harry, 2004, The effect of muscle and fasciocutaneous flaps on angiogenesis in open tibial fractures, J Orthop Trauma, 18, S15

Mulari, 2004, Osteoblast-like cells complete osteoclastic bone resorption and form new mineralized bone matrix in vitro, Calcif Tissue Int, 75, 253, 10.1007/s00223-004-0172-3

Teitelbaum, 2000, Bone resorption by osteoclasts, Science, 289, 1504, 10.1126/science.289.5484.1504

Fan, 1997, Macrophage colony stimulating factor down-regulates MCSF-receptor expression and entry of progenitors into the osteoclast lineage, J Bone Miner Res, 12, 1387, 10.1359/jbmr.1997.12.9.1387

Kong, 1999, OPGL is a key regulator of osteoclastogenesis, lymphocyte development and lymph-node organogenesis, Nature, 397, 315, 10.1038/16852

Blair, 2006, Mechanisms of disease: roles of OPG, RANKL and RANK in the pathophysiology of skeletal metastasis, Nat Clin Pract Oncol, 3, 41, 10.1038/ncponc0381

Quinn, 2005, Modulation of osteoclast formation, Biochem Biophys Res Commun, 328, 739, 10.1016/j.bbrc.2004.11.076

Lee, 2006, Cytokines regulating osteoclast formation and function, Curr Opin Rheumatol, 18, 411, 10.1097/01.bor.0000231911.42666.78

Kaneko, 2000, Direct stimulation of osteoclastic bone resorption by bone morphogenetic protein (BMP)-2 and expression of BMP receptors in mature osteoclasts, Bone, 27, 479, 10.1016/S8756-3282(00)00358-6

Little, 2007, The anabolic and catabolic responses in bone repair, J Bone Joint Surg [Br], 89, 425, 10.1302/0301-620X.89B4.18301

Einhorn, 2005, The science of fracture healing. 1, J Orthop Trauma, 19, S4, 10.1097/00005131-200511101-00002

Schell, 2006, Osteoclastic activity begins early and increases over the course of bone healing, Bone, 38, 547, 10.1016/j.bone.2005.09.018

Sawae, 2003, Osteoclast differentiation at growth plate cartilage-trabecular bone junction in newborn rat femur, J Electron Microsc (Tokyo), 52, 493, 10.1093/jmicro/52.6.493

Takahara, 2004, Matrix metalloproteinase-9 expression, tartrate-resistant acid phosphatase activity, and DNA fragmentation in vascular and cellular invasion into cartilage preceding primary endochondral ossification in long bones, J Orthop Res, 22, 1050, 10.1016/j.orthres.2004.01.011

Cole, 1987, Tartrate-resistant acid phosphatase in bone and cartilage following decalcification and cold-embedding in plastic, J Histochem Cytochem, 35, 203, 10.1177/35.2.3540104

McDonald, 2007, Bisphosphonate treatment and fracture repair, BoneKey Osteovision, 9, 236, 10.1138/20070271

Odvina, 2005, Severely suppressed bone turnover: a potential complication of alendronate therapy, J Clin Endocrinol Metab, 90, 1294, 10.1210/jc.2004-0952

McDonald MM, Dulai S, Godfrey C, Amanat N, Sztynda T, Little DG. Bolus or weekly zoledronic acid administration does not delay endochondral fracture repair but weekly dosing enhances delays in hard callus remodelling. Bone; 2008 Jun 3. [Epub ahead of print].

McDonald, 2007, Normal endochondral bone healing follows continuous bisphosphonate pre-treatment in a rat closed fracture model, J Bone Miner Res, 22, S253

Namkung-Matthai, 2001, Osteoporosis influences the early period of fracture healing in a rat osteoporotic model, Bone, 28, 80, 10.1016/S8756-3282(00)00414-2

Meyer, 2001, Age and ovariectomy impair both the normalization of mechanical properties and the accretion of mineral by the fracture callus in rats, J Orthop Res, 19, 428, 10.1016/S0736-0266(00)90034-2

Walsh, 1997, Fracture healing in a rat osteopenia model, Clin Orthop Relat Res, 342, 218, 10.1097/00003086-199709000-00029

Reinholt, 1991, Extensive clear zone and defective ruffled border formation in osteoclasts of osteopetrotic (ia/ia) rats: implications for secretory function, Exp Cell Res, 251, 477, 10.1006/excr.1999.4585

McDonald, 2007, The osteopetrotic Incisor absent rat exhibits reduced osteoclast activity, resulting in normal endochondral fracture union but delayed hard callus remodelling, J Bone Miner Res, 22, S253

Flick, 2003, Effects of receptor activator of NFkappaB (RANK) signaling blockade on fracture healing, J Orthop Res, 21, 676, 10.1016/S0736-0266(03)00011-1

Ulrich-Vinther, 2005, Osteoprotegerin treatment impairs remodeling and apparent material properties of callus tissue without influencing structural fracture strength, Calcif Tissue Int, 76, 280, 10.1007/s00223-004-0126-9

Ulrich-Vinther, 2005, Gene therapy with human osteoprotegerin decreases callus remodeling with limited effects on biomechanical properties, Bone, 37, 751, 10.1016/j.bone.2005.07.021

Vu, 1998, MMP-9/Gelatinase B is a key regulator of growth plate angiogenesis and apoptosis of hypertrophic chondrocytes, Cell, 93, 411, 10.1016/S0092-8674(00)81169-1

Colnot, 2003, Altered fracture repair in the absence of MMP9, Development, 130, 4123, 10.1242/dev.00559

Behonick, 2007, Role of Matrix metalloproteinase 13 in both endochondral and intramembranous ossification during skeletal regeneration, Plos One, 11, e1150, 10.1371/journal.pone.0001150

Malemud, 2006, Matrix metalloproteinases: role in skeletal development and growth plate disorders, Front Biosci, 11, 1702, 10.2741/1916

Tang, 2005, Extracellular matrix metalloproteinase inducer stimulates tumor angiogenesis by elevating vascular endothelial growth factor and matrix metalloproteinases, Cancer Res, 65, 3193, 10.1158/0008-5472.CAN-04-3605

Mark, 2004, Microvascular invasion during endochondral ossification in experimental fractures in rats, Bone, 35, 535, 10.1016/j.bone.2004.04.010

Hausman, 2001, Prevention of fracture healing in rats by an inhibitor of angiogenesis, Bone, 29, 560, 10.1016/S8756-3282(01)00608-1

Street, 2002, Vascular endothelial growth factor stimulates bone repair by promoting angiogenesis and bone turnover, Proc Natl Acad Sci USA, 99, 9656, 10.1073/pnas.152324099

Holsetin, 2008, Rapamycin Affects early fracture healing in mice, Br J Pharmacol, 10.1038/bjp.2008.167

Wan, 2008, Activation of the hypoxia-inducible factor-1alpha pathway accelerates bone regeneration, Proc Natl Acad Sci USA, 105, 686, 10.1073/pnas.0708474105

Tarvainen, 1994, Effect of clodronate on fracture healing in denervated rats, Bone, 15, 701, 10.1016/8756-3282(94)90320-4

Li, 1999, Effect of bisphosphonate (incadronate) on fracture healing of long bones in rats, J Bone Miner Res, 14, 969, 10.1359/jbmr.1999.14.6.969

Li, 2000, Concentration of bisphosphonate (incadronate) in callus area and its effects on fracture healing in rats, J Bone Miner Res, 15, 2042, 10.1359/jbmr.2000.15.10.2042

Choi, 2007, Inhibition of bone healing by pamidronate in calvarial bony defects, Oral Surg Oral Med Oral Pathol Oral Radiol Endod, 103, 321, 10.1016/j.tripleo.2006.06.057

Munns, 2005, Effects of intravenous pamidronate treatment in infants with osteogenesis imperfecta: clinical and histomorphometric outcome, J Bone Miner Res, 20, 1235, 10.1359/JBMR.050213

Amanat, 2005, A single systemic dose of pamidronate improves bone mineral content and accelerates restoration of strength in a rat model of fracture repair, J Orthop Res, 23, 1029, 10.1016/j.orthres.2005.03.004

Amanat, 2007, Optimal timing of a single dose of zoledronic acid to increase strength in rat fracture repair, J Bone Miner Res, 22, 867, 10.1359/jbmr.070318

Schmidt, 1977, A radiographic and histologic study of fracture healing in osteopetrotic rats, Radiology, 122, 517, 10.1148/122.2.517

Marks, 1978, Bone remodeling as an expression of altered phenotype: studies of fracture healing in untreated and cured osteopetrotic rats, Clin Orthop Relat Res, 137, 259

Nakamura, 2003, Osteoprotegerin regulates bone formation through a coupling mechanism with bone resorption, Endocrinology, 144, 5441, 10.1210/en.2003-0717

Grigoriadis, 1994, c-Fos: a key regulator of osteoclast-macrophage lineage determination and bone remodeling, Science, 266, 443, 10.1126/science.7939685

Nishino, 2001, Histochemical examination of osteoblastic activity in op/op mice with or without injection of recombinant M-CSF, J Bone Miner Metab, 19, 267, 10.1007/s007740170010

Martin, 2005, Osteoclast-derived activity in the coupling of bone formation to resorption, Trends Mol Med, 11, 76, 10.1016/j.molmed.2004.12.004

Mohan S, Baylink DJ. The role of IGF-II in the coupling of bone formation to resorption. In: Modern concepts of insulin-like growth factors (ed. Spencer, EM) 1991; p. 19–174.

Marzia, 2000, Decreased c-Src expression enhances osteoblast differentiation and bone formation, J Cell Biol, 151, 311, 10.1083/jcb.151.2.311

Schaller, 2004, The chloride channel inhibitor NS3736 prevents bone resorption in ovariectomized rats without changing bone formation, J Bone Miner Res, 19, 1144, 10.1359/JBMR.040302

Nakamura, 2007, Osteoclast inhibitory lectin (OCIL) inhibits osteoblast differentiation and function in vitro, Bone, 40, 305, 10.1016/j.bone.2006.09.001

Rubin, 2002, Skeletal adaptation to mechanical stimuli in the absence of formation or resorption of bone, J Musculoskelet Neuronal Interact, 2, 264

van Bezooijen, 2005, SOST/sclerostin, an osteocyte-derived negative regulator of bone formation, Cytokine Growth Factor Rev, 16, 319, 10.1016/j.cytogfr.2005.02.005

Winkler, 2003, Osteocyte control of bone formation via sclerostin, a novel BMP antagonist, EMBO J, 22, 6267, 10.1093/emboj/cdg599

Robling, 2008, Mechanical stimulation of bone in vivo reduces osteocyte expression of Sost/sclerostin, J Biol Chem, 283, 5866, 10.1074/jbc.M705092200

van Bezooijen, 2004, Sclerostin is an osteocyte-expressed negative regulator of bone formation, but not a classical BMP antagonist, J Exp Med, 199, 805, 10.1084/jem.20031454