The combined application of molecular, biochemical, histochemical, and ultrastructural approaches has defined a temporal sequence of gene expression associated with development of the bone cell phenotype in primary osteoblast cultures. The peak levels of expressed genes reflect a developmental sequence of bone cell differentiation characterized by three principal periods: proliferation, extracellular matrix maturation and mineralization, and two restriction points to which the cells can progress but cannot pass without further signals. The regulation of cell growth and bone-specific gene expression has been examined during this developmental sequence and is discussed within the context of several unique concepts. These are (1) that oncogene expression in proliferating osteoblasts contributes to the suppression of genes expressed postproliferatively, (2) that hormone modulation of a gene is dependent upon the maturational state of the osteoblast, and (3) that chromatin structure and the presence of nucleosomes contribute to three-dimensional organization of gene promoters that support synergistic and/or antagonistic activities of physiologic mediators of bone cell growth and differentiation.
References
1.
Abe, E., Y. Ishimi, N. Takahashi, T. Akatsu, H. Ozawa, H. Yamana, S. Yoshiki, and T. Suda: A differentiation-inducing factor produced by the osteoblastic cell line MC3T3E1 stimulates bone resorption by promoting osteoclast formation. J. Bone Min. Res. 3:635-645 (1988).
2.
Aronow, M.A., L.C. Gerstenfeld, T.A. Owen, M.S. Tassinari, G.S. Stein, and J.B. Lian: Factors that promote progressive development of the osteoblast phenotype in cultured fetal rat calvaria cells. J. Cell Physiol. 143:213-221 (1990a).
3.
Aronow, M.A., T.A. Owen, G.S. Stein, and J.B. Lian: Estrogen inhibition of osteocalcin mRNA expression occurs in cultured rat osteoblasts only after formation of a mineralized extracellular matrix. J. Bone Min. Res., 5:S273 (1990b).
4.
Barone, L.M., T.A. Owen, M.S. Tassinari, R. Bortell, G.S. Stein and J.B. Lian: Developmental expression and hormonal regulation of the rat matrix gla protein (MGP) gene in chondrogenesis and osteogenesis. J. Cell. Biochem. 46:351-365 (1991).
5.
Barrack, E.R. and D.S. Coffey: Hormone receptors and the nuclear matrix. Gene Regulation by Steroid Hormones, II, A. K. Roy and J. H. Clark, Eds. Springer-Verlag, New York, 1983, 239-266.
6.
Baumbach, L.L., G.S. Stein, and J.L. Stein: Regulation of human histone gene expression: Transcriptional and post-transcriptional control in the coupling of histone messenger RNA stability with DNA replication. Biochemistry26:6178-6187 (1987 ).
7.
Bellows, C.G., J.E. Aubin, J.N.M. Heersche, and M.E. Antosz: Mineralized bone nodules formed in vitro from enzymatically released rat calvaria cell populations. Calcif. Tissue Int. 38:143-154 (1986).
8.
Bellows, C.G., J.N.M. Heersche, and J.E. Aubin: Determination of the capacity for proliferation and differentiation of osteoprogenitor cells in the presence and absence of dexamethasone. Dev. Biol. 140:132-138 (1990).
9.
Benayhahu, D., Y. Kletter, D. Zipori, and S. Weintraub: Bone marrow-derived stromal cell line expressing osteoblastic phenotype in vitro and osteogenic capacity in vivo. J. Cell. Physiol. 140:1-7 (1989).
10.
Bentz, H., R.M. Nathan, D.M. Rosen, R.M. Armstrong, A.Y. Thompson, P.R. Segarini, M.C. Mathews, J.R. Dasch, K.A. Piez, and S.M. Seyedin: Purification and characterization of a unique osteoinductive factor from bovine bone. J. Biol. Chem. 264:20805-20810 (1989 ).
11.
Beresford, J.N., J.A. Gallagher, J.W. Poser, and R.G.G. Russell: Production of osteocalcin by human bone cells in vitro. Effects of 1,25(OH)2D3, 24,25(OH)2D3, parathyroid hormone, and glucocorticoids. Metab. Bone Dis. Relat. Res.5:229-234 (1984).
12.
Bemier, S.M., J. Desjardina, A.K. Sullivan, and D. Goltzman: Establishment of an osseous cell line from fetal rat calvaria using an immunocytolytic method of cell selection: Characterization of the cell line and of derived clones. J. Cell. Physiol145:274-285 (1990).
13.
Betsholtz, C., A. Johnsson, C.-H. Heldin, B. Westermark, P. Lind, M.S. Urdea, R. Eddy, T.B. Shows, K. Philpott, A.L. Mellor, T.J. Knott, and J. Scott: cDNA sequence and chromosomal localization of human platelet-derived growth factor A-chain and its expression in tumor cell lines. Nature, 320:695-699 (1986).
14.
Bhargava, U., M. Bar-Lev, C.G. Bellows, andJ.E. Aubin: Ultrastructural analysis of bone nodules formed in vitro by isolated fetal rat calvaria cells. Bone. 9:155-163 (1988).
15.
Binderman, I., E. Berger, N. Fine, Z. Shimshoni, A. Harell, and D. Somjen: Calvaria derived osteogenic cells: phenotypic expression in culture. Connect. Tissue Res. 20:41-47 (1989).
16.
Blau, H.M. and D. Baltimore: Differentiation requires continuous regulation. J. Cell Biol. 112:781-783 (1991).
17.
Bortell, R., L.M. Barone, M.S. Tassinari, J.B. Lian, and G.S. Stein: Gene expression during endochondral bone development: Evidence for coordinate expression of transforming growth factor β and collagen type I. J. Cell. Biochem. 44:81-91 (1990).
18.
Bortell, R.T., A. Owen, A.J. van Wijnen, J.P. Bidwell, P. Gavazzo, E. Breen, H. DeLuca, G.S. Stein, and J.B. Lian: Do multiple vitamin D receptor binding elements synergistically contribute to the level of osteocalcin gene promoter activity? Proc. Natl. Acad. Sci. submitted.
19.
Boskey, A.L., F.H. Wians Jr., and P.V. Hauschka: The effect of osteocalcin on in vitro lipid-induced hydroxyapatite formation and seeded hydroxyapatite growth. Calcif. Tiss. Int. 37:57-62 (1985).
20.
Boskey, A.L.: Noncollagenous matrix proteins and their role in mineralization. Bone Miner. 6:111-123 (1989).
21.
Bruder, S.P. and A. Caplan: First bone formation and the dissection of an osteogenic lineage in the embryonic chick tibia is revealed by monoclonal antibodies against osteoblasts. Bone. 11:359-375 (1989).
22.
Buckely, M.J., A.J. Banes, and R.D. Jordan: The effects of mechanical strain on osteoblasts in vitro. J. Oral Maxillofacial Surg. 48:276-282 (1990).
23.
Canalis, E.: The hormonal and local regulation of bone formation. Endocrinology. 112:931-939 (1983).
24.
Canalis, E. and J.B. Lian: 1,25(OH)2D3 effects on collagen and DNA synthesis in periosteum and periosteum-free calvaria. Bone. 6:457-460 (1985).
25.
Canalis, E., T. McCarthy, and M. Centrella: Growth factors and the regulation of bone remodeling. J. Clin. Invest. 81:277-281 (1988).
26.
Capco, D.G., K.M. Wan, and S. Penman: The nuclear matrix: three-dimensional architecture and protein composition. Cell. 29:847-858 (1982).
27.
Carrington, J.L., A.B. Roberts, K.C. Flanders, N.S. Roche, and A.H. Reddi: Accumulation, localization and compartmentation of transforming growth factor β during endochondral bone development. J. Cell Biol. 107:1969-1975 (1988).
28.
Celeste, A.J., V. Rosen, J.L. Buecker, R. Kriz, E.A. Wang, and J.M. Wozney: Isolation of the human gene for bone Gla protein utilizing mouse and rat cDNA clones. EMBO J.5:1885-1890 (1986 ).
29.
Centrella, M., T.L. McCarthy, and E. Canalis: Skeletal tissue and transforming growth factor β. FASEB J. 2:3066-3073 (1988).
30.
Centrella, M.,. T.L. McCarthy, and E. Canalis: β2 microglobulin enhances insulin-like growth factor I receptor levels and synthesis in bone cell cultures. J. Biol. Chem. 264:18268-18271 (1989).
31.
Collart, D., A. Ramsey-Ewing, R. Bortell, J. Lian, J. Stein, and G. Stein: Isolation and characterization of a cDNA from a human histone H2B gene which is reciprocally expressed in relation to replication-dependent H2B histone genes during HL60 cell differentiation. Biochemistry. 30:1610-1617 (1991).
32.
Craig, A.M., J.H. Smith, and D.T. Denhardt: Osteopontin, a transformation-associated cell adhesion phosphoprotein, is induced by 12-0-tetradecanoylphorbol 13-acetate in mouse epidermis. J. Biol. Chem. 264:9682-9689 (1989).
33.
DeLuca, H.F.: The vitamin D story: a collaborative effort of basic science and clinical medicine. FASEB J. 2:224-236(1988).
34.
Demay, M.B., D.A. Roth, and H.M. Kronenberg: Regions of the rat osteocalcin gene which mediate the effect of 1,25-dihydroxyvitamin D3 on gene transcription. J. Biol. Chem.264:2279-2282 (1989).
35.
Demay, M.B., J.M. Gerardi, H.F. DeLuca, and H.M. Kronenberg: DNA sequences in the rat osteocalcin gene that bind the 1,25-dihydroxyvitamin D3 receptor and confer responsive to 1,25-dihydroxyvitamin D3. Proc. Natl. Acad. Sci. U.S.A.87:369-373 (1990).
36.
Diamond, M.I., J.N. Miner, S.K. Yoshinaga, and K.R. Yamamoto: Transcription factor interactions: Selectors of positive or negative regulation from a single DNA element. Science. 249:1266-1272 (1990).
37.
Dworetzky, S.I., E.G. Fey, S. Penman, J.B. Lian, J.L. Stein, and G.S. Stein: Progressive changes in the protein composition of the nuclear matrix during rat osteoblast differentiation. Proc. Natl. Acad. Sci. U.S.A. 87:4605-4607 (1990).
38.
Dworetsky, S.I., K.L. Wright, E.G. Fey, S. Penman, J.B. Lian, J.L. Stein, and G.S. Stein: Sequence-specific DNA binding proteins are components of a nuclear matrix attachment site. Proc. Natl. Acad. Sci. U.S.A., in press.
39.
Ecarot-Charrier, B., F.H. Glorieux, M. van der Rest, and G. Pereira: Osteoblasts isolated from mouse calvaria initiate matrix mineralization in culture. J. Cell Biol. 96:639-643 (1983).
40.
Ecarot-Charrier, B., N. Shepard, G. Charette, M. Grynpas, and F.H. Glorieux: Mineralization in osteoblast cultures: A light and electron microscopic study. Bone. 9:147-154 (1988).
41.
Fell, H.B.: Skeletal development in tissue culture, In The Biochemistry and Physiology of Bone, G. H. Bourne, Ed., Academic Press, New York, 1956, 401-441.
42.
Fey, E.G. and S. Penman: Nuclear matrix proteins reflect cell type of origin in cultured human cells. Proc. Natl. Acad. Sci. U.S.A. 85:121-125 (1988).
43.
Fisher, L.W., G.R. Hawins, N. Turcoss, and J.D. Termine: Purification and partial characterization of small proteoglycans I and II, bone sialoproteins I and II, and osteonectin from the mineral compartment of developing human bone. J. Biol. Chem. 262:9702-9708 (1987).
44.
Fitton-Jackson, S.: The influence of tissue interactions and extracellular macromolecules on the control of phenotypic expression and synthetic capacity of bone and cartilage. Calcif. Tissue Res. Suppl. 23:489-495 (1978).
45.
Franceschi, R.T. and J. Young: Regulation of alkaline phosphatase by 1,25-dihydroxyvitamin D3 and ascorbic acid in bone-derived cells. J. Bone Min. Res.5:1157-1167 (1990).
46.
Franzen, A. and D. Heinegard: Isolation and characterization of two sialoproteins present only in bone calcified matrix. Biochem. J. 232:715-724 (1985).
47.
Fraser, J.D. and P.A. Price: Lung, heart, and kidney express high levels of mRNA for the vitamin K-dependent matrix Gla protein: implications for the possible functions of matrix Gla protein and for the tissue distribution of the γ-carboxylase. J. Biol. Chem. 263:11033-11036 (1988).
48.
Friedenstein, A.J. and R.K. Chailakhyan: Bone marrow osteogenic stem cells: In vitro cultivation and transplantation in diffusion chambers. Cell Tissue Kinet. 20:263-272 (1987).
49.
Gehron-Robey, P., M. Young, L. Fischer, and T. McClain: Thrombospondin is an osteoblast-derived component of mineralized extracellular matrix. J. Cell. Biol. 108:719-727 (1989).
50.
Gerstenfeld, L.C., S.D. Chipman, J. Glowacki, and J.B. Lian: Expression of differentiated function by mineralizing cultures of chicken osteoblasts. Dev. Biol. 122:49-60 (1987).
51.
Gerstenfeld, L.C., S.D. Chipman, C.M. Kelly, K.J. Hodgens, D.D. Lee, and W.J. Landis: Collagen expression illustrated assembly and mineralization in cultures of chicken calvaria osteoblasts. J. Cell Biol. 106:979-989 (1988).
52.
Gerstenfeld, L.C., C.M. Kelly, M. Von Deck, and J.B. Lian: Effect of 1,25-dihydroxyvitamin D3 on induction of chondrocyte maturation in culture: extracellular matrix gene expression and morphology. Endocrinology. 126:1599-1609 (1990).
53.
Glimcher, M.J.: Mechanism of calcification: Role of collagen fibrils and collagen-phosphoprotein complexes in vitro and in vivo. Anat. Rec. 224:139-153 (1989).
54.
Glowacki, J. and J. Lian: Impaired recruitment of osteoclast progenitors in deficient osteocalcin-depleted bone implants. Cell Differ. 21:247-254 (1987).
55.
Goldring, M.B. and S.R. Goldring: Basic science and pathology skeletal tissue response to cytokines. Clin. Orthop. Relat. Res. 258:245-278 (1990).
56.
Gorski, J.P. and K. Shimizu: Isolation of new phosphorylated glycoprotein from mineralized phase of bone that exhibits limited homology to adhesive protein osteopontin. J. Biol. Chem. 263:15938-15945 (1988).
57.
Grange, T., J. Roux, G. Rigeaud, and R. Pictet: Two remote glucocorticoid responsive units interact cooperatively to promote glucocorticoid induction of rat tyrosine aminotransferase gene expression. Nucleic Acids Res. 17:8695-8709 (1989 ).
58.
Grigoriadis, A.E., J.N.M. Heersche, and J.E. Aubin: Differentiation of muscle, fat, cartilage, and bone from progenitor cells present in a bone-derived clonal cell population: Effect of dexamethasone. J. Cell Biol. 106:2139-2151 (1988 ).
59.
Gronowicz, G., F.N. Woodiel, M-B. McCarthy, andL.G. Raisz: In vitro mineralization of fetal rat parietal bones in defined serum-free medium: effect of P-glycerol phosphate. J. Bone Min. Res. 4:313-324 (1989).
60.
Gundberg, C.M., P.V. Hauschka, J.B. Lian, and P.M. Gallop: Osteocalcin isolation, characterization, and detection. Methods Enzymol. 107:516-544 (1984).
61.
Hamilton, J.A., S. Lingelbach, N.C. Partridge, and T.J. Martin: Regulation of plasminogen activator production by bone-resorbing hormones in normal and malignant osteoblasts. Endocrinology116:2186-2191 (1985).
62.
Harrison, J.R., D.N. Petersen, A.C. Lichtler, A.T. Mador, D.W. Rowe, and B.E. Kream: 1,25-Dihydroxyvitamin D3 inhibits transcription of type I collagen genes in the rat osteosarcoma cell line ROS 17/ 2.8. Endocrinology125:327-333 (1989).
63.
Hauschka, P.V., A.E. Mavrakos, M.D. lafrati, S.E. Doleman, and M. Klagsbrun: Growth factors in bone matrix. Isolation of multiple types by affinity chromatography on heparin-sepharose. J. Biol. Chem. 261:12665-12675 (1986).
64.
Hauschka, P.V., J.B. Lian, D.E.C. Cole, and C.M. Gundberg: Osteocalcin and matrix Gla protein: vitamin K-dependent proteins in bone, Physiol. Rev. 69:990-1047 (1989).
65.
Heath, J.K., S.J. Atkinson, M.C. Meikle, and J.J. Reynolds: Mouse osteoblasts synthesize collagenase in response to bone resorbing agents. Biochim. Biophys. Acta. 802:151-154 (1984).
66.
Heath, J.K., S.B. Rodan, K. Yoon, and G.A. Rodan: Rat calvarial cell lines immortalized with SV-40 large T antigen: Constitutive and retinoic acid-inducible expression of osteoblastic features. Endocrinology. 124:3060-3068 (1989).
67.
Heine, U.I., E.F. Munoz, K.C. Flanders, L.R. Ellingsworth, H.-Y.P., Lam, N.L. Thompson, A.B. Roberts, and M.B. Sporn: Role of transforming growth factor-β in the development of the mouse embryo. J. Cell Biol. 105:2861-2876 (1987 ).
68.
Heinegård, D. and A. Oldberg: Structure and biology of cartilage and bone matrix noncollagenous macromolecules. FASEB J. 3:2042-2051 (1989 ).
69.
Hock, J.M., M. Gunnes-Hey, J. Poser, H. Olson, N.H. Bell, and L.G. Raisz: Stimulation of undermineralized matrix formation by 1,25 dihydroxyvitamin D3 in long bones of rats. Calcif. Tissue Int.28:79-86 (1986).
70.
Holthuis, J., T.A. Owen, A.J. van Wijnen, K.L. Wright, A. Ramsey-Ewing, M.B. Kennedy, R. Carter, S.C. Cosenza, K.J. Soprano, J.B. Lian, J.L. Stein, and G.S. Stein: Tumor cells exhibit deregulation of the cell cycle histone gene promoter factor HiNF-D. Science. 247:1454-1457 (1990 ).
71.
Hourdry, J.: Evolution of the amphibian regenerative capacity throughout metamorphosis, In The Proceedings of the 6th International Marcus Singer Symposium: Regeneration and Development, S. Inoue, Ed., Okada Printing and Publishing Company, Maebashi, Japan, 1988, 13-22.
72.
Howlett, C.R., J. Cave, M. Williamson, J. Farmer, S.Y. Ali, L. Bab, and M.E. Owen: Mineralization in in vitro cultures of rabbit marrow stromal cells. Clin. Orthop. 213:251-263 (1986).
73.
Ibaraki, K., J.D. Termine, M.F. Young, and S.W. Whitson: Differential bone matrix gene expression in a novel mineralizing cell culture system. J. Bone Min. Res. Suppl.5:S232 (1990 ).
74.
Jingushi, S., A. Heydemann, M. Joyce, R. Soltero, and M.E. Bolander: Acidic fibroblast growth factor is synthesized in fracture callus by inflammatory cells around the blood clot and regulated cartilage formation during fracture healing. J. Bone Min. Res.4:S285 (1989 ).
75.
Jingushi, S., A. Heydemann, S.K. Kana, L.R. Macey, and M.E. Bolander: Acidic fibroblast growth factor (aFGF) injection stimulates cartilage enlargement and inhibits cartilage gene expression in rat fracture healing. J. Orthop. Res. Unit.8:364-371 (1990).
76.
Kerner, S.A., R.A. Scott, and J.W. Pike: Sequence elements in the human osteocalcin gene confer basal activation and inducible response to hormonal vitamin D,. Proc. Natl. Acad. Sci. U.S.A. 86:4455-4459 (1989).
77.
Kiefer, M.C., A.C.S. Saphire, D.M. Bauer, and P.J. Barr: The cDNA and derived amino acid sequences of human and bovine bone Gla protein. Nucleic Acids Res. 18:1909 (1990).
78.
Kinne, R.W. and L.W. Fisher: Keratin sulfate proteoglycan in rabbit compact bone is sialoprotein II. J. Biol. Chem. 262:10206-10211 (1987).
79.
Kroeger, P., C. Stewart, T. Schaap, A. van Wijnen, J. Hirshman, S. Helms, G. Stein, and J. Stein: Proximal and distal regulatory elements that influence in vivo expression of a cell cycle-dependent human H4 histone gene. Proc. Natl. Acad. Sci. U.S.A. 84:3982-3986 (1987 ).
80.
Lane, T.F. and E.H. Sage: Functional mapping of SPARC: peptides from two distinct Ca+ (+)-binding sites modulate cell shape. J. Cell Biol. 111:3065-3076 (1990).
81.
Lawrence, J.B., R.H. Singer, C.A. Villnave, J.L. Stein, and G.S. Stein: Intracellular distribution of histone mRNAs in human fibroblasts studied by in situ hybridization. Proc. Natl. Acad. Sci. U.S.A.85:263-267 (1988).
82.
Leboy, P.S., L. Vaias, B. Uschmann, E. Golub, S.L. Adams, M. Pacifici: Ascorbic acid induces alkaline phosphatase, type X collagen, and calcium deposition in cultures chick chondrocytes. J. Biol. Chem. 264:17281-17286 (1989).
83.
Leboy, P.S., J.N. Beresford, C. Devlin, and M.E. Owen: Dexamethasone induction of osteoblast mRNAs in rat marrow stromal cell cultures. J. Cell. Physiol. 146:370-378 (1991).
84.
Lee, W., A. Haslinger, M. Karin, and R. Tjian: Activation of transcription by two factors that bind promoter and enhancer sequences of the human metallothionein gene and SV40. Nature (London). 325:368-372 (1987).
Lian, J.B. and P.A. Friedman: The vitamin K-dependent synthesis of gamma-carboxyglutamic acid by bone microsomes. J. Biol. Chem. 253:6623-6626 (1978 ).
87.
Lian, J.B., M.C. Couttes, and E. Canalis: Studies of hormonal regulation of osteocalcin synthesis in cultured fetal rat calvariae. J. Biol. Chem. 60:8706-8710 (1985).
88.
Lian, J.B., A.H. Roufosse, B. Reit, and M.J. Glimcher: Concentration of osteocalcin and phosphoprotein as a function of mineral content and age in cortical bone. Calcif. Tissue Int. 345:84 (1982).
89.
Lian, J.B.: Osteocalcin: Functional studies and postulated role in bone resorption, In Current Advances in Vitamin K Research J. W. Suttie, Ed., Elsevier, New York (1987).
90.
Lian, J.B., C. Stewart, E. Puchacz, S. Mackowiak, V. Shalhoub, D. Collart, G. Zambetti, and G. Stein: Structure of the rat osteocalcin gene and regulation of vitamin D-dependent expression. Proc. Natl. Acad. Sci. U.S.A. 86:1143-1147 (1989).
91.
Lian, J.B. and S.C. Marks, Jr.: Osteopetrosis in the rat: Coexistence of reductions in osteocalcin and bone resorption. Endocrinology. 126:955-962 (1990).
92.
Lian, J.B., G.S. Stein, R. Bortell, and T.A. Owen: Phenotype suppression: a postulated molecular mechanism for mediating the relationship of proliferation and differentiation by fos/jun interactions at AP-1 sites in steroid responsive promoter elements of tissue-specific genes. J. Cell. Biochem. 45:9-14 (1991).
93.
Lian, J. and G. Stein: Vitamin D regulation of osteocalcin, collagen and alkaline phosphatase gene expression is controlled by osteoblast growth and differentiation, in Eighth Workshop on Vitamin D-1991, A. W. Norman, Ed., Walter de Gruyter, Berlin, in press.
94.
Lian, J.B. and C.M. Gundberg: Clinical significance of serum osteocalcin and urine Gla measurements. Clin. Orthop. Rel. Res. 266:267-291 (1988).
95.
Liao, J., K. Ozono, T. Sone, D.P. McDonnel, and J.W. Pike: Vitamin D receptor interaction with specific DNA requires a nuclear protein and 1,25-dihydroxyvitamin D,. Proc. Natl. Acad. Sci. U.S.A.87:9751-9755 (1990).
96.
Lichtler, A., M.L. Stover, J. Angilly, B. Kream, and J. Rowe: Isolation and characterization of the rat α-1 (I) collagen promoter. J. Biol. Chem. 264:3072-3077 (1989).
97.
Long, M.S., J.L. Williams, and K.G. Mann: Expression of human bone-related proteins in the hematopoietic microenvironment. J. Clin. Invest. 86:1387-1395 (1990 ).
98.
Luyten, F.P., N.S. Cunningham, S. Ma, N. Muthukumaran, R.G. Hammonds, W.B. Nevins, W.I. Wood, and A.H. Reddi: Purification and partial amino acid sequence of osteogenin, a protein initiating bone differentiation. J. Biol. Chem.264:13377-13380 (1989).
99.
Lyons, K.M., R.W. Pelton, and B.L.M. Hogan: Patterns of expression of murine Vgr-1 and BMP-2a RNA suggest that transforming growth factor-β-like genes coordinately regulate aspects of embryonic development. Genes Dev. 1:1657-1668 (1989 ).
100.
Majeska, R.J., S.B. Rodan, andG.A. Rodan: Parathyroid-responsive clonal cell lines from rat osteosarcoma. Endocrinology. 107:1494-1501 (1980).
101.
Majeska, R.J. and G.A. Rodan: The effect of 1,25(OH)2D3 on alkaline phosphatase in osteoblastic osteosarcoma cells. J. Biol. Chem.257:3362-3365 (1982 ).
102.
Majeska, R.J., B.C. Nair, and G.A. Rodan: Glucocorticoid regulation of alkaline phosphatase in the osteoblastic osteosarcoma cell line ROS 17/2.8 Endocrinology. 116:170-179 (1985).
103.
Mardon, H. and J.T. Triffit: A tissue-specific protein in rat osteogenic tissues. J. Bone Min. Res. 2:191-199 (1987).
104.
Markose, E.R., J.L. Stein, G.S. Stein, and J.B. Lian: Vitamin D-mediated modifications in protein-DNA interactions at two promoter elements of the osteocalcin gene. Proc. Natl. Acad. Sci. U.S.A. 87:1701-1705 (1990).
105.
Minghetti, P.P. and A.W. Norman: 1,25(OH)2 vitamin D, receptors: gene regulation and genetic circuitry. FASEB J.2:3043-3053 (1989).
106.
Mohan, S. and D.J. Baylink: Bone growth factors. Clin. Orthop. Rel. Res. 263:30-48 (1991).
107.
Morris, T.D., L.A. Weber, E. Hickey, G.S. Stein, and J.L. Stein: Changes in the stability of a human H3 histone mRNA during the HeLa cell cycle. Mol. Cell. Biol. 11:544-553 (1991).
108.
Morrison, N.A., J. Shine, J.-C. Fragonas, V. Verkest, L. McMenemy, and J.A. Eisman: 1,25-dihydroxyvitamin D-responsive element and glucocorticoid repression in the osteocalcin gene. Science. 246:1158-1161 (1989).
109.
Mulliken, J.B. and J. Glowacki: Induced osteogenesis for repair and construction in the craniofacial region. Plast. Reconstr. Surg. 65:553-559 (1980).
110.
Nagata, T., C.G. Bellows, S. Kasugai, W.T. Butler, and J. Sodek: Biosynthesis of bone proteins [SPP-1 (secreted phosphoprotein-1, osteopontin), BSP (bone sialoprotein) and SPARC (osteonectin)] in association with mineralized-tissue formation by fetal-rat calvarial cells in culture. Biochem. J.274:513-520 (1991).
111.
Nanes, M.S., J. Rubin, L. Titus, G.N. Hendy, and B.D. Catherwood: Interferon-γ inhibits 1,25-dihydroxyvitamin D3-stimulated synthesis of bone GLA protein in rat osteosarcoma cells by a pretranslational mechanism. Endocrinology. 127:588-594 (1990).
112.
Nijweide, P.J. and R.J.P. Mulder: Identification of osteocytes in osteoblast-like cell cultures using a monoclonal antibody specifically directed against osteocytes. Histochemistry. 84:342-347 (1986).
113.
Nishimoto, S.K., W.F. Stryker, and M.E. Nimni: Calcification of osteoblastlike rat osteosarcoma cells in agarose suspension cultures. Calcif. Tissue Int. 41:274-280 (1987a).
114.
Nishimoto, S.K., C. Salka, and M.E. Nimni: Retinoic acid and glucocorticoids enhance the effect of 1,25-dihydroxyvitamin D3 on bone γ-carboxyglutamic acid protein synthesis by rat osteosarcoma cells. J. Bone Min. Res.2:571-577 (1987b).
115.
Noda, M. and G.A. Rodan: Type β transforming growth factor (TGFβ) regulation of alkaline phosphatase expression and other phenotype-related mRNAs in osteoblastic rat osteosarcoma cells. J. Cell. Physiol. 133:426-437 (1987).
116.
Noda, M., K. Yoon, and G.A. Rodan: Cyclic AMP-mediated stabilization of osteocalcin mRNA in rat osteoblast-like cells treated with parathyroid hormone. J. Biol. Chem. 263:18574-18577 (1988).
117.
Noda, M.: Transcriptional regulation of osteocalcin production by transforming growth factor-β in rat osteoblast-like cells. Endocrinology. 124:612-617 (1989).
118.
Noda, M., R.L. Vogel, D.M. Hasson, and G.A. Rodan: Leukemia inhibitory factor suppresses proliferation, alkaline phosphatase activity, and type I collagen messenger ribonucleic acid level and enhances osteopontin mRNA level in murine osteoblast-like (MC3T3E1) cells. Endocrinology. 127:18-190 (1990a).
119.
Noda, M., R.L. Vogel, A.M. Craig, J. Prahl, H.F. DeLuca, and D.T. Denhardt: Identification of a DNA sequence responsible for binding of the 1,25-dihydroxyvitamin D3 receptor and 1,25-dihydroxyvitamin D3 enhancement of mouse secreted phosphoprotein 1 (Spp-1 or osteopontin) gene expression. Proc. Natl. Acad. Sci. U.S.A.87:9995-9999 (1990b).
120.
Nomura, S., A.J. Wills, D.R. Edwards, J.K. Heath, and B.L.M. Hogan: Developmental expression of 2ar (osteopontin) and SPARC (osteonectin) RNA as revealed by in situ hybridization. J. Cell Biol. 106:441-450 (1988).
121.
Oldberg, A., A. Franzén, and D. Heinegard: Cloning and sequence analysis of rat bone sialoprotein (osteopontin) cDNA reveals an Arg-Gly-Asp cell-binding sequence. Proc. Natl. Acad. Sci. U.S.A. 83:8819-8823 (1986 ).
122.
Oldberg, A., B. Jirskog-Hed, S. Axelsson, and D. Heinegard: Regulation of bone sialoprotein mRNA by steroid hormones. J. Cell Biol. 109:3183-3186 (1989).
123.
Otsuka, K., J. Sodek, and H. Limeback: Synthesis of collagenase and collagenase inhibitors by osteoblast-like cells in culture. Eur. J. Biochem. 145:123-129 (1984).
124.
Owen, M. and A.J. Friedenstein: Stromal stem cells: marrow-derived osteogenic precursors, In Cell and Molecular Biology of Vertebrate Hard Tissues D. Evered and S. Harnett, Eds., John Wiley & Sons, Chichester, 198, 42-60, 1988.
125.
Owen, T.A., M. Aronow, V. Shalhoub, L.M. Barone, L. Wilming, M.S. Tassinari, M.B. Kennedy, S. Pockwinse, J.B. Lian, and G.S. Stein: Progressive development of the rat osteoblast phenotype in vitro: reciprocal relationships in expression of genes associated with osteoblast proliferation and differentiation during formation of the bone extracellular matrix. J. Cell. Physiol. 143:420-430 (1990a).
126.
Owen, T.A., J. Holthuis, E. Markose, A.J. van Wijnen, S.A. Wolfe, S. Grimes, J.B. Lian, and G.S. Stein: Modifications of protein-DNA interactions in the proximal promoter of a cell growth-regulated histone gene during the onset and progression of osteoblast differentiation. Proc. Natl. Acad. Sci. U.S.A.87:5129-5133 (1990b).
127.
Owen, T.A., R. Bortell, S.A. Yocum, S.L. Smock, M. Zhang, C. Abate, V. Shalhoub, N. Aronin, K.L. Wright, A.J. van Wijnen, J.L. Stein, T. Curran, J.B. Lian, and G.S. Stein: Coordinate occupancy of AP-1 sites in the vitamin D responsive and CCAAT box elements by fos-jun in the osteocalcin gene: a model for phenotype suppression of transcription. Proc. Natl. Acad. Sci. U.S.A.87:9990-9994 (1990c).
128.
Owen, T.A., M.A. Aronow, L.M. Barone, B. Bettencourt, G. Stein, and J.B. Lian: Pleiotropic effects of vitamin D on osteoblast gene expression are related to the proliferative and differentiated state of the bone cell phenotype: dependency upon basal levels of gene expression, duration of exposure and bone matrix competency in normal rat osteoblast cultures. Endocrinology. 128:1496-1504 (1991).
129.
Ozawa, H., K. Imamura, E. Abe, N. Takahashi, T. Hiraide, Y. Shibasaki, T. Fukuhara, and T. Suda: Effect of a continuously applied compressive pressure on mouse osteoblast-like cells (MC3T3-E1) in vitro. J. Cell. Physiol. 142:177-185 (1990).
130.
Ozkaynak, E., D.C. Rueger, E.A. Drier, C. Corbett, R.J. Ridge, T.K. Sampath, and H. Oppermann: OP-1 cDNA encodes an osteogenic protein in the TGF-β family. EMBO J. 9:2085-2093 (1990).
131.
Partridge, N.C., D. Alcorn, V.P. Michelangeli, G. Ryan, and T.J. Martin: Morphological and biochemical characterization of four clonal osteogenic sarcoma cell lines of rat origin. Cancer Res. 43:4308-4314 (1983).
132.
Pauli, U., S. Chrysogelos, G. Stein, J. Stein, and H. Nick: Protein-DNA interactions in vivo upstream of a cell cycle regulated human H4 histone gene. Science. 236:1308-1311 (1987).
133.
Pauli, U., S. Chrysogelos, J. Stein, and G. Stein: Native genomic blotting: high-resolution mapping of DNase I-hypersensitive sites and protein-DNA interactions. Proc. Natl. Acad. Sci. U.S.A. 85:16-20 (1988).
134.
Pauli, U., S. Chrysogelos, H. Nick, G. Stein, and J. Stein: In vivo protein binding sites and nuclease hypersensitivity in the promoter region of a cell cycle regulated human H3 histone gene. Nucleic Acids Res. 17:2333-2350 (1989).
135.
Pechak, D.G., J.J. Kujawa, and A.I. Caplan: Morphological and histochemical events during first bone formation in embryonic chick limbs. Bone. 7:441-458 (1986).
136.
Peck, W.A., S.J. Birge, and S.A. Fedax: Bone cells: biochemical and biological studies after enzymatic isolation. Science146:1476-1477 (1964).
137.
Plumb, M., J. Stein, and G. Stein: Coordinate regulation of multiple histone mRNAs during the cell cycle in HeLa cells. Nucleic Acids Res. 11:2391-2410 (1983).
138.
Price, P.A.: Role of vitamin K-dependent proteins in bone metabolism. Annu. Rev. Nutr. 8:865-883 (1988).
139.
Prince, C.W. and W.T. Butler: 1,25-Dihydroxyvitamin D2 regulated the biosynthesis of osteopontin, a bone-derived cell attachment protein, in clonal osteoblast-like osteosarcoma cells. Collagen Rel. Res.7:305-313 (1987).
140.
Puchacz, E., J.B. Lian, G.S. Stein, J. Wozney, K. Huebner, and C. Croce: Chromosomal localization of the human osteocalcin gene. Endocrinology.24:2648-2650 (1989).
141.
Rauscher, F.J., III, P.J. Voulalas, B.R. Franza, Jr., and T. Curran: Fos and jun bind cooperative to the AP-1 site: reconstitution in vitro. Genes Dev. 2:1687-1699 (1988).
142.
Reddi, A.H.: Cell biology and biochemistry of endochondral bone development. Collagen Res. 1:209-226 (1981).
143.
Reich, K.M., C.V. Gay, and J.A. Frangos: Fluid shear stress as a mediator of osteoblast cyclic adenosine monophosphate production. J. Cell. Physiol. 143:100-104 (1990).
144.
Reinholt, F.P., K. Hultenby, A. Oldberg, and D. HeinegArd: Osteopontin-a possible anchor of osteoclasts to bone. Proc. Natl. Acad. Sci. U.S.A. 87:4473-4475 (1990 ).
145.
Robey, P.G. and L.J. Termine: Human bone cells in vitro. Calcif. Tissue Int. 37:453-460 (1985).
146.
Rodan, G.A. and S.B. Rodan: Expression of the osteoblastic phenotype, In Bone and Mineral Research, Vol. 2, W. A. Peck, Ed., Elsevier Science, Amsterdam, 1984, 244-285.
147.
Rodan, S.B., Y. Imai, M.A. Thiede, G. Wesolowski, D. Thompson, Z. Bar-Shavit, S. Shull, K. Mann, and G.A. Rodan: Characterization of a human cell line (SAOS-2) with osteoblastic properties. Cancer Res. 47:4961 (1987).
148.
Rodan, S.B., G. Wesolowski, K. Yoon, and G.A. Rodan: Opposing effects of fibroblast growth factor and pertussis toxin on alkaline phosphatase, osteopontin, osteocalcin, and type I collagen mRNA levels in ROS 17/2.8 cells. J. Biol. Chem. 264:19934-19941 (1989).
149.
Rodan, G.A. and M. Noda: Gene expression in osteoblastic cells. Crit. Rev. Eukaryotic Gene Expression1:85-98 (1991).
150.
Romberg, R.W., P.G. Wemess, B.L. Riggs, and K.G. Mann: Inhibition of hydroxyapatite crystal growth by bone-specific and other calcium-binding proteins. Biochemistry. 25:1176-1180 (1986 ).
151.
Rosen, D.M., S.A. Stempien, A.Y. Thompson, and S.M. Seyedin: Transforming growth factor-β modulates the expression of osteoblast and chondrocyte phenotypes in vivo. J. Cell Physiol. 134:337-346 (1988).
152.
Ruoslahti, E.: Structure and biology of proteoglycans. Annu. Rev. Cell. Biol. 4:229-255 (1988).
153.
Ruther, U., C. Garber, D. Komitowski, R. Muller, and E.F. Wagner: Deregulated c-fos expression interferes with normal bone development in transgenic mice. Nature. 325:412-416 (1987).
154.
Sampath, T.K., N. Muthukumaran, and A.H. Reddi: Isolation of osteogenin, an extracellular matrix-associated, bone inductive protein, by heparin affinity chromatography. Proc. Natl. Acad. Sci. U.S.A. 84:7109-7113 (1987).
155.
Sandberg, M., H. Autio-Harmainen, and E. Vuorio: Localization of the expression of types I, III, and IV collagen, TGP-β1 and c-fos genes in developing human calvarial bones. Dev. Biol. 130:324-334 (1988).
156.
Schepmoes, G., E. Breen, T.A. Owen, M.A. Aronow, G.S.Stein, and J.B. Lian: Influence of dexamethasone on the vitamin D mediated regulation of osteocalcin gene expression. J. Cell. Biochem., 47:184-196, 1991.
157.
Schüle, R., M. Muller, C. Kaltschmidt, and R. Renkawitz: Many transcription factors interact synergistically with steroid receptors. Science. 241:1418-1420 (1988 ).
158.
Schüle, R., U. Kazuhiko, D.J. Mangelsdorf, J. Bolado, J.W. Pike, and R.M. Evans: Jun-Fos and receptors for vitamins A and D recognize a common response element in the human osteocalcin gene. Cell. 61:497-504 (1990a).
159.
Schüle, R., P. Rangarajan, S. Kliewer, L.J. Ransome, J. Bolado, N. Yang, I.M. Verma, and R.M. Evans: Functional antagonism between oncoprotein c-Jun and the glucocorticoid receptor. Cell. 62:1217-1226 (1990 b).
160.
Shalhoub, V., L.C. Gerstenfeld, J.B. Lian, J.L. Stein, and G.S. Stein: Down-regulation of cell growth and cell cycle regulated genes during chick osteoblast differentiation with the reciprocal expression of histone gene variants. Biochemistry. 28:5318-5322 (1989 ).
161.
Shalhoub, V., M.E. Jackson, J.B. Lian, G.S. Stein. and S.C. Marks, Jr.: Gene expression during skeletal development in three osteopetrotic rat mutations: evidence for osteoblast abnormalities. J. Biol. Chem. 255:9847-9856 (1991).
162.
Sierra, F., A. Lichtler, F. Marashi, R. Rickles, T. Van Dyke, S. Clark, J. Wells, G. Stein, and J. Stein: Organization of human histone genes. Proc. Natl. Acad. Sci. U.S.A. 79:1795-1799 (1982).
163.
Silbermann, M., D. Lewinson, H. Gonen, M.A. Lizarbe, and K. von der Mark: In vitro transformation of chondroprogenitor cells into osteoblasts and the formation of new membrane bone. Anat. Rec. 206:373-383 (1983).
164.
Smith, J.H. and D.T. Denhardt: Molecular cloning of a tumor promoter-inducible mRNA found in JB6 mouse epidermal cells: induction is stable at high, but not at low, cell densities. J. Cell. Biochem. 34:13-22 (1987).
165.
Smith, J.H. and D.T. Denhardt: Evidence for two pathways of protein kinase C induction of 2AR expression: correlation with mitogenesis. J. Cell. Physiol. 139:189-195 (1989).
166.
Smith, M.M. and B.K. Hall: Development and evolutionary origins of vertebrate skeletogenic and odontogenic tissue. Biol. Rev. 65:277-373 (1990).
167.
Soriano, P., C. Montgomery, R. Geske, and A. Bradley: Targeted disruption of the c-src proto-oncogene leads to osteopetrosis in mice. Cell. 64:693-702 (1991).
168.
Speiss, Y.H., P.A. Price, J.L. Deftos, and S.C. Manolagas: Phenotype-associated changes in the effects of 1,25-dihydroxyvitamin D3 on alkaline phosphatase and bone Gla-protein of rat osteoblastic cells. Endocrinology. 118:1340-1346 (1986).
169.
Sporn, M.B., A.B. Roberts, J.H. Shull, J.M. Smith, J.M. Ward, and J. Sodek: Polypeptide transforming growth factors isolated from bovine sources and used for wound healing in vivo. Science. 219:1329-1331 (1983).
170.
Stein, G., W. Park, C. Thrall, R. Mans, and J. Stein: Regulation of cell cycle stage-specific transcription of histone genes from chromatin by non-histone chromosomal proteins. Nature. 257:764-767 (1975).
171.
Stein, G., J. Lian, J. Stein, R. Briggs, V. Shalhoub, K. Wright, U. Pauli, and A.J. van Wijnen: Altered binding of human histone gene transcription factors during the shutdown of proliferation and onset of differentiation in HL-60 cells. Proc. Natl. Acad. Sci. U.S.A. 86:1865-1869 (1989a).
172.
Stein, G.S., J.B. Lian, L.C. Gerstenfeld, V. Shalhoub, M. Aronow, T. Owen, and E. Markose: The onset and progression of osteoblast differentiation is functionally related to cellular proliferation. Connect. Tissue Res. 20:3-13 (1989b).
173.
Stein, G.S., J.B. Lian, and T.A. Owen: Bone cell differentiation: a functionally coupled relationship between expression of cell-growth- and tissue-specific genes. Current Opin. Cell Biol. 2:1018-1027 (1990).
174.
Stein, G.S., J.B. Lian, and T.A. Owen: Relationship of cell growth to the regulation of tissue-specific gene expression during osteoblast differentiation. FASEB J. 4:3111-3123 (1990).
175.
Stein, G.S., J.B. Lian, R.A. Bortell, T.A. Owen, A. van Wijnen, S.I. Dworetsky, and J.P. Bidwell: Regulation of transcription factor activity during growth and differentiation: involvement of the nuclear matrix in concentration and localization of promoter binding proteins. J. Cell. Biochem. 47:300-305 (1991).
176.
Strauss, P.G., E.I. Closs, J. Schmidt, and V. Erfle: Gene expression during osteogenic differentiation in mandibular condyles in vitro. J. Cell. Biol. 110:1369-1378 (1990 ).
177.
Stromstedt, P.E., L. Poelinger, J.A. Gustafsson, and J. Cralstedt-Duke: The glucocorticoid receptor binds to a sequence overlapping the TATA box of the human osteocalcin promoter: a potential mechanism for negative regulation. Mol. Cell. Biol. 11:3379-3383 (1991).
178.
Sudo, H., H. Kodama, Y. Amagai, S. Yamamoto, and H. Kasai: In vitro differentiation and calcification in a new clone osteogenic cell line derived from newborn mouse calvaria. J. Cell Biol. 96:191 (1983).
179.
Termine, J.D., H.K. Kleinman, S.W. Whitson, K.M. Conn, M.L. McGarvey, and G.R. Martin: Osteonectin, a bone-specific protein linking mineral to collagen. Cell. 26:99-105 (1981).
180.
Thaller, C. and G. Eichele: Identification and spatial distribution of retinoids in the developing chick limb bud. Nature. 327:625-628 (1987).
181.
Theofan, G. and P.A. Price: Bone gla protein messenger ribonucleic acid is regulated by both 1,25-dihydroxyvitamin D3 and 3',5'-cyclic adenosine monophosphate in rat osteosarcomacells. Mol. Endocrinol.3:36 (1989).
182.
van Wijnen, A.J., J.L. Stein, and G.S. Stein: A nuclear protein with affinity for the 5' flanking region of a cell cycle dependent human H4 histone gene in vitro. Nucleic Acids Res. 15:1679-1698 (1987).
183.
van Wijnen, A.J., K.L. Wright, R.F. Massung, M. Gerretsen, J.L. Stein, and G.S. Stein: Two target sites for protein binding in the promoter region of a cell cycle regulated human H1 histone gene. Nucleic Acids Res. 16:571-592 (1988).
184.
Van Wijnen, A.J., K.L. Wright, J.B. Lian, J.L. Stein, and G.S. Stein: Human H4 histone gene transcription requires the proliferation-specific nuclear factor HiNF-D: Auxilliary roles for HiNF-C (Spl-Like) and HiNF-A (high mobility group-like). J. Biol. Chem.264:15034-15042 (1989).
185.
Van Wijnen, A.J., T.K. Choi, T.A. Owen, K.L. Wright, J.B. Lian, R. Jaenisch, J.L. Stein, and G.S. Stein: Involvement of the cell cycle-regulated nuclear factor HiNF-D in cell growth control of a human H4 histone gene during hepatic development in transgenic mice. Proc. Natl. Acad. Sci. U.S.A.88:2573-2577 (1991).
186.
Wagner, M., C. Thaller, T. Jessell, and G. Eichele: Polarizing activity and retinoid synthesis in the floor plate of the neural tube. Nature. 345:819-822 (1990).
187.
Wang, E.A., V. Rosen, J.S. D'Allesandro, M. Bauduy, P. Cordes, T. Harada, D.I. Israel, R.M. Hewick, K.M. Kerns, P. LaPan, D.P. Luxenberg, D. McQuaid, I.K. Moutsatsos, J. Nove, and J.M. Wozney: Recombinant human bone morphogenetic protein induces bone formation. Proc. Natl. Acad. Sci. U.S.A. 87:2220-2224 (1990 ).
188.
Weinberg, R.A.: Oncogenes, antioncogenes, and the molecular bases of multistep carcinogenesis. Cancer Res. 49:3713-3721 (1989 ).
189.
Weinreb, M., D. Shinar, andG.A. Rodan: Different pattern of alkaline phosphatase, osteopontin, and osteocalcin expression in developing rat bone visualized by in situ hybridization. J. Bone Min. Res.5:831-842 (1990).
190.
Weiss, M.J., K. Ray, P.S. Henthom, B. Lamb, T. Kadesch, and H. Harris: Structure of the human liver/bone/kidney alkaline phosphatase gene. J. Biol. Chem. 263:12002-12010 (1988).
191.
Whitson, S.W., W. Harrison, M.K. Dunlap, D.E. Bowers, Jr., L.W. Fisher, P.G. Robey, and J.D. Jermine: Fetal bovine cells synthesize bone specific matrix proteins. J. Cell Biol. 88:607-614 (1984).
192.
Wong, G.L. and D.V. Cohn: Target cells in bone for parathormone and calcitonin are different: enrichment for each cell type by sequential digestion of mouse calvaria and selective adhesion to polymeric surfaces. Proc. Natl. Acad. Sci. U.S.A. 72:3167-3171 (1975).
193.
Wong, M.-M., L.G. Rao, H. Ly, L. Hamilton, J. Tong, W. Sturtridge, R. McBroom, J.E. Aubin, and T.M. Murray: Long-term effects of physiologic concentrations of dexamethasone on human bone-derived cells. J. Bone Min. Res. 5:803-813 (1990).
194.
Wozney, J.M., V. Rosen, A.J. Celeste, L.M. Mitsock, M.J. Whitters, R.W. Kriz, R.M. Hewick, and E.A. Wang: Novel regulators of bone formation: molecular clones and activities. Science. 242:1528-1533 (1988).
195.
Wronski, T.J., B.P. Halloran, D.D. Bikle, R.K. Globus, and E.R. Morey-Holton: Chronic administration of 1,25(OH)2D, increased bone but impaired mineralization. Endocrinology. 119:2580-2585 (1986).
196.
Yang-Yen, H.F., J.C. Chambard, Y.L. Sun, T. Smeal, T.J. Schmidt, J. Drouin, and M. Karin: Transcriptional interference between c-jun and the glucocorticoid receptor: mutual inhibition of DNA binding due to direct protein-protein interaction. Cell. 62:1205-1215 (1990 ).
197.
Yoon, K., R. Buenaga, and G.A. Rodan: Tissue specificity and developmental expression of osteopontin. Biochem. Biophys. Res. Comm. 148:1129-1136 (1987 ).
198.
Yoon, K., S.J. Rutledge, R.F. Buenaga, andG.A. Rodan: Characterization of the rat osteocalcin gene: Stimulation of promoter activity by 1,25-dihydroxyvitamin D3, Biochemistry. 27:8581-8526 (1988).
199.
Young, M.F., A.A. Day, P. Dominquez, C.I. McQuillan, and L.W. Fisher: Structure and expression of osteonectin mRNA in human tissue. Connect. Tissue Res. 24:17-28 (1990).
200.
Zambetti, G., J. Stein, and G. Stein: Targeting of a chimeric human histone fusion mRNA to membrane-bound polysomes in HeLa cells. Proc. Natl. Acad. Sci. U.S.A. 84:2683-2687 (1987).
201.
Zhou, H., G.R. Hammonds, Jr., D.M. Findlay, P.J. Fuller, J.T. Martin, and K.W. Ng: Retinoic acid modulation of mRNA levels in malignant, nontransformed, and immortalized osteoblasts. J. Bone Min. Res.6:767-777 (1991).
