title: Multifaceted signaling regulators of chondrogenesis: Implications in cartilage regeneration and tissue engineering.
abstract: Defects of articular cartilage present a unique clinical challenge due to its poor self-healing capacity and avascular nature. Current surgical treatment options do not ensure consistent regeneration of hyaline cartilage in favor of fibrous tissue. Here, we review the current understanding of the most important biological regulators of chondrogenesis and their interactions, to provide insight into potential applications for cartilage tissue engineering. These include various signaling pathways, including: fibroblast growth factors (FGFs), transforming growth factor β (TGF-β)/bone morphogenic proteins (BMPs), Wnt/β-catenin, Hedgehog, Notch, hypoxia, and angiogenic signaling pathways. Transcriptional and epigenetic regulation of chondrogenesis will also be discussed. Advances in our understanding of these signaling pathways have led to promising advances in cartilage regeneration and tissue engineering.
authors.affiliation: The University of Chicago Pritzker School of Medicine, Chicago, IL 60637, USA; Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Jordan D
authors.initials: JD
authors.lastname: Green
authors.affiliation: The University of Chicago Pritzker School of Medicine, Chicago, IL 60637, USA; Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Viktor
authors.initials: V
authors.lastname: Tollemar
authors.affiliation: The University of Chicago Pritzker School of Medicine, Chicago, IL 60637, USA; Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Mark
authors.initials: M
authors.lastname: Dougherty
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA; Department of Orthopaedic Surgery, The Second Affiliated Hospital of Chongqing Medical University, Chongqing 400016, China.
authors.forename: Zhengjian
authors.initials: Z
authors.lastname: Yan
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA; Department of Orthopaedic Surgery, The Second Affiliated Hospital of Chongqing Medical University, Chongqing 400016, China.
authors.forename: Liangjun
authors.initials: L
authors.lastname: Yin
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA; School of Bioengineering, Chongqing University, Chongqing, China.
authors.forename: Jixing
authors.initials: J
authors.lastname: Ye
authors.affiliation: The University of Chicago Pritzker School of Medicine, Chicago, IL 60637, USA; Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Zachary
authors.initials: Z
authors.lastname: Collier
authors.affiliation: The University of Chicago Pritzker School of Medicine, Chicago, IL 60637, USA; Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Maryam K
authors.initials: MK
authors.lastname: Mohammed
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Rex C
authors.initials: RC
authors.lastname: Haydon
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Hue H
authors.initials: HH
authors.lastname: Luu
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Richard
authors.initials: R
authors.lastname: Kang
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Michael J
authors.initials: MJ
authors.lastname: Lee
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Sherwin H
authors.initials: SH
authors.lastname: Ho
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Tong-Chuan
authors.initials: TC
authors.lastname: He
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Lewis L
authors.initials: LL
authors.lastname: Shi
authors.affiliation: Department of Orthopaedic Surgery and Rehabilitation Medicine, The University of Chicago Medical Center, Chicago, IL 60637, USA.
authors.forename: Aravind
authors.initials: A
authors.lastname: Athiviraham
keywords: BMPs; Cartilage; Cell signaling; Chondrogenesis; FGF; Hedgehog; Regenerative medicine; Sox9; TGFβ; Wnt
references.citation: PLoS Genet. 2011 Nov;7(11):e1002356
references.pmid: 22072985
references.citation: PLoS One. 2013;8(2):e55296
references.pmid: 23383321
references.citation: Exp Cell Res. 1998 Jan 10;238(1):265-72
references.pmid: 9457080
references.citation: Am J Sports Med. 2012 Oct;40(10):2273-80
references.pmid: 22922521
references.citation: Osteoarthritis Cartilage. 2001;9 Suppl A:S69-75
references.pmid: 11680692
references.citation: Exp Cell Res. 1986 Jul;165(1):127-38
references.pmid: 3519255
references.citation: Science. 2005 Jul 8;309(5732):310-1
references.pmid: 15919954
references.citation: Int J Mol Med. 2010 Mar;25(3):377-84
references.pmid: 20127042
references.citation: Biomaterials. 2005 May;26(15):2479-89
references.pmid: 15585250
references.citation: J Cell Physiol. 2005 May;203(2):398-409
references.pmid: 15521064
references.citation: BMB Rep. 2008 Jul 31;41(7):485-94
references.pmid: 18682032
references.citation: J Cell Biol. 1988 Nov;107(5):1969-75
references.pmid: 2460474
references.citation: FASEB J. 2001 Jun;15(8):1436-8
references.pmid: 11387249
references.citation: Dev Cell. 2001 Aug;1(2):277-90
references.pmid: 11702786
references.citation: Cell Death Differ. 2012 Mar;19(3):461-9
references.pmid: 21869831
references.citation: J Cell Sci. 2010 Jun 15;123(Pt 12):2068-76
references.pmid: 20501701
references.citation: J Bone Miner Res. 1999 May;14(5):757-63
references.pmid: 10320524
references.citation: Cell Tissue Res. 1999 Apr;296(1):111-9
references.pmid: 10199971
references.citation: Proc Natl Acad Sci U S A. 2014 May 13;111(19):6940-5
references.pmid: 24778247
references.citation: Sci Rep. 2013 Oct 23;3:3022
references.pmid: 24149372
references.citation: Biomaterials. 2005 Feb;26(6):599-609
references.pmid: 15282138
references.citation: Annu Rev Cell Dev Biol. 2005;21:659-93
references.pmid: 16212511
references.citation: Nat Rev Genet. 2004 Sep;5(9):691-701
references.pmid: 15372092
references.citation: Exp Mol Med. 2010 Mar 31;42(3):166-74
references.pmid: 20164682
references.citation: Genes Dev. 2005 Dec 1;19(23):2783-810
references.pmid: 16322555
references.citation: Dev Cell. 2002 Sep;3(3):439-49
references.pmid: 12361605
references.citation: Dev Cell. 2001 Aug;1(2):251-64
references.pmid: 11702784
references.citation: Dev Biol. 2007 Dec 15;312(2):572-81
references.pmid: 17997399
references.citation: Cytotherapy. 2009;11(3):261-7
references.pmid: 19308769
references.citation: Proc Natl Acad Sci U S A. 1985 Apr;82(8):2267-71
references.pmid: 3857579
references.citation: Osteoarthritis Cartilage. 2009 Aug;17(8):1065-75
references.pmid: 19254740
references.citation: J Cell Physiol. 2008 Sep;216(3):708-15
references.pmid: 18366089
references.citation: Bone. 2008 Jul;43(1):64-71
references.pmid: 18456591
references.citation: PLoS One. 2007 May 16;2(5):e450
references.pmid: 17505543
references.citation: J Orthop Sci. 2006 Oct;11(5):491-6
references.pmid: 17013738
references.citation: Dev Biol. 1997 May 1;185(1):104-18
references.pmid: 9169054
references.citation: Genes Cells. 2008 Apr;13(4):355-63
references.pmid: 18363966
references.citation: J Biol Chem. 2002 Dec 27;277(52):50668-75
references.pmid: 12381733
references.citation: Development. 1997 Aug;124(15):2945-60
references.pmid: 9247337
references.citation: Genes Dev. 1998 Jun 1;12(11):1571-86
references.pmid: 9620845
references.citation: Development. 1999 Dec;126(23):5515-22
references.pmid: 10556075
references.citation: J Cell Sci. 2002 Jul 15;115(Pt 14):3015-25
references.pmid: 12082161
references.citation: Mech Dev. 2011 Jan-Feb;128(1-2):104-15
references.pmid: 21093584
references.citation: Osteoarthritis Cartilage. 2008 Mar;16(3):279-86
references.pmid: 17945514
references.citation: J Biomech. 2012 Jan 3;45(1):17-26
references.pmid: 21899847
references.citation: Mod Rheumatol. 2008;18(3):213-9
references.pmid: 18351289
references.citation: Artif Organs. 2013 Jul;37(7):648-55
references.pmid: 23495957
references.citation: J Cell Sci. 2002 Dec 15;115(Pt 24):4809-18
references.pmid: 12432069
references.citation: J Cell Biol. 2006 Nov 20;175(4):555-62
references.pmid: 17101698
references.citation: Differentiation. 2009 Dec;78(5):260-8
references.pmid: 19716643
references.citation: Development. 2013 Jan 1;140(1):76-81
references.pmid: 23154410
references.citation: Biomaterials. 2011 Sep;32(27):6425-34
references.pmid: 21652067
references.citation: Annu Rev Biomed Eng. 2012;14:431-55
references.pmid: 22655599
references.citation: Biochem Biophys Res Commun. 2002 Mar 8;291(4):775-9
references.pmid: 11866432
references.citation: Cell. 1993 Nov 5;75(3):579-87
references.pmid: 8221896
references.citation: Proc Natl Acad Sci U S A. 2001 Mar 27;98(7):3802-7
references.pmid: 11274398
references.citation: Acta Med Okayama. 2010 Dec;64(6):351-7
references.pmid: 21173804
references.citation: Dev Biol. 2009 Apr 1;328(1):40-53
references.pmid: 19389374
references.citation: Development. 1999 Feb;126(5):987-98
references.pmid: 9927599
references.citation: Clin Orthop Relat Res. 2000 May;(374):212-34
references.pmid: 10818982
references.citation: J Bone Miner Res. 2004 Mar;19(3):463-70
references.pmid: 15040835
references.citation: Arthritis Res Ther. 2007;9(3):R55
references.pmid: 17537234
references.citation: Osteoarthritis Cartilage. 2012 Jun;20(6):593-602
references.pmid: 22425884
references.citation: Dev Biol. 2000 Mar 15;219(2):237-49
references.pmid: 10694419
references.citation: PLoS One. 2014 Feb 13;9(2):e89025
references.pmid: 24551211
references.citation: Genes Dev. 2004 May 1;18(9):1072-87
references.pmid: 15132997
references.citation: J Biomech Eng. 1991 May;113(2):143-51
references.pmid: 1652042
references.citation: Biomech Model Mechanobiol. 2005 Mar;3(3):141-6
references.pmid: 15668766
references.citation: Mol Endocrinol. 2010 Aug;24(8):1581-93
references.pmid: 20573686
references.citation: Cell Metab. 2009 Mar;9(3):287-302
references.pmid: 19254573
references.citation: Crit Rev Eukaryot Gene Expr. 2005;15(3):243-54
references.pmid: 16390320
references.citation: Cell. 2001 Feb 9;104(3):341-51
references.pmid: 11239392
references.citation: Semin Cell Dev Biol. 2005 Aug-Oct;16(4-5):539-46
references.pmid: 16144691
references.citation: Dev Cell. 2007 Jun;12(6):931-41
references.pmid: 17543865
references.citation: Tissue Eng. 2007 Apr;13(4):737-46
references.pmid: 17371156
references.citation: BMB Rep. 2011 Jan;44(1):28-33
references.pmid: 21266103
references.citation: J Biol Chem. 2009 Apr 24;284(17):11326-35
references.pmid: 19251704
references.citation: J Cell Sci. 1995 Apr;108 ( Pt 4):1497-508
references.pmid: 7615670
references.citation: Orthod Craniofac Res. 2005 Aug;8(3):209-18
references.pmid: 16022723
references.citation: J Clin Invest. 2001 Feb;107(3):295-304
references.pmid: 11160153
references.citation: Stem Cell Rev Rep. 2014 Dec;10(6):841-55
references.pmid: 24923882
references.citation: J Orthop Res. 2009 Jun;27(6):793-9
references.pmid: 19051248
references.citation: J Biol Chem. 2003 Oct 17;278(42):41227-36
references.pmid: 12893825
references.citation: Genes Dev. 2002 Nov 1;16(21):2813-28
references.pmid: 12414734
references.citation: Osteoarthritis Cartilage. 2012 Dec;20(12):1638-46
references.pmid: 22947280
references.citation: Int J Mol Sci. 2013 Mar 14;14(3):5966-77
references.pmid: 23493060
references.citation: J Cell Biochem. 2012 Aug;113(8):2687-95
references.pmid: 22441842
references.citation: J Biol Chem. 2005 May 27;280(21):20509-15
references.pmid: 15781473
references.citation: Osteoarthritis Cartilage. 2015 Mar;23(3):443-53
references.pmid: 25464167
references.citation: Clin Cancer Res. 2007 Jan 1;13(1):275-82
references.pmid: 17200366
references.citation: EMBO J. 2006 Apr 19;25(8):1646-58
references.pmid: 16601693
references.citation: J Orthop Res. 2010 Jan;28(1):89-95
references.pmid: 19623616
references.citation: Orthod Craniofac Res. 2005 Aug;8(3):200-8
references.pmid: 16022722
references.citation: Biochim Biophys Acta. 2000 Feb 2;1495(2):168-82
references.pmid: 10656974
references.citation: Biomaterials. 2006 Jan;27(1):91-9
references.pmid: 16023192
references.citation: Biotechnol Bioeng. 1999 Jan 20;62(2):166-74
references.pmid: 10099526
references.citation: J Cell Biochem. 2006 Jan 1;97(1):33-44
references.pmid: 16215986
references.citation: Dev Genet. 1996;19(1):43-50
references.pmid: 8792608
references.citation: Development. 2015 Mar 1;142(5):817-31
references.pmid: 25715393
references.citation: J Cell Biochem. 2015 Jul;116(7):1391-400
references.pmid: 25708223
references.citation: J Bone Miner Res. 2010 Dec;25(12):2680-9
references.pmid: 20593410
references.citation: Stem Cells Int. 2013;2013:232896
references.pmid: 24082888
references.citation: J Biol Chem. 2010 Aug 6;285(32):24381-7
references.pmid: 20529846
references.citation: J Biol Chem. 2002 Sep 6;277(36):33398-410
references.pmid: 12052822
references.citation: Genes Dev. 2002 Apr 1;16(7):859-69
references.pmid: 11937493
references.citation: J Cell Biol. 2003 Sep 1;162(5):833-42
references.pmid: 12952936
references.citation: Cold Spring Harb Perspect Biol. 2013 Jan 01;5(1):a008334
references.pmid: 23284041
references.citation: Dev Biol. 2008 Feb 15;314(2):341-50
references.pmid: 18191119
references.citation: J Biol Chem. 2012 Jun 22;287(26):21926-35
references.pmid: 22544738
references.citation: Proc Natl Acad Sci U S A. 2005 Feb 15;102(7):2414-9
references.pmid: 15699338
references.citation: J Orthop Res. 2004 Jul;22(4):742-50
references.pmid: 15183429
references.citation: Poult Sci. 2008 Aug;87(8):1556-64
references.pmid: 18648049
references.citation: PLoS One. 2013 Oct 03;8(10):e73673
references.pmid: 24098328
references.citation: J Cell Sci. 2000 Jan;113 ( Pt 1):59-69
references.pmid: 10591625
references.citation: Tissue Eng Part A. 2010 Feb;16(2):545-56
references.pmid: 19728793
references.citation: Dev Biol. 2007 Jul 15;307(2):300-13
references.pmid: 17544391
references.citation: J Biol Chem. 2008 Oct 3;283(40):27154-64
references.pmid: 18644788
references.citation: Nature. 2003 May 15;423(6937):332-6
references.pmid: 12748651
references.citation: Osteoarthritis Cartilage. 2005 Jul;13(7):623-31
references.pmid: 15896984
references.citation: Tissue Eng. 2004 May-Jun;10(5-6):762-70
references.pmid: 15265293
references.citation: Tissue Eng. 2007 Jun;13(6):1159-71
references.pmid: 17518738
references.citation: Science. 2001 Oct 26;294(5543):853-8
references.pmid: 11679670
references.citation: Biomaterials. 2002 Feb;23(4):1187-95
references.pmid: 11791922
references.citation: Arthritis Rheum. 2002 Nov;46(11):2867-77
references.pmid: 12428226
references.citation: Development. 2014 Oct;141(20):3978-87
references.pmid: 25294942
references.citation: Dev Biol. 1999 May 1;209(1):11-27
references.pmid: 10208739
references.citation: J Cell Biol. 1990 Jun;110(6):2195-207
references.pmid: 2351696
references.citation: Cell. 1993 Dec 3;75(5):843-54
references.pmid: 8252621
references.citation: Biomaterials. 2005 Dec;26(34):7082-94
references.pmid: 15985292
references.citation: J Biol Chem. 2014 May 16;289(20):13810-20
references.pmid: 24711445
references.citation: Biorheology. 2004;41(3-4):445-58
references.pmid: 15299276
references.citation: Int J Biochem Cell Biol. 2003 Sep;35(9):1323-40
references.pmid: 12798347
references.citation: Mol Cell Biol. 2009 Aug;29(15):4262-73
references.pmid: 19470758
references.citation: Osteoarthritis Cartilage. 2008 Dec;16(12):1450-6
references.pmid: 18541445
references.citation: Ann N Y Acad Sci. 1996 Jun 8;785:254-5
references.pmid: 8702145
references.citation: Development. 2000 Apr;127(7):1445-53
references.pmid: 10704390
references.citation: Biomaterials. 2004 Aug;25(18):4163-73
references.pmid: 15046906
references.citation: J Biol Chem. 2012 Jun 22;287(26):22206-15
references.pmid: 22547066
references.citation: Endocr Relat Cancer. 2007 Sep;14(3):875-86
references.pmid: 17914116
references.citation: Osteoarthritis Cartilage. 2004 Aug;12(8):599-613
references.pmid: 15262240
references.citation: Clin Orthop Relat Res. 1971 Oct;80:167-73
references.pmid: 5133323
references.citation: Tissue Eng Part B Rev. 2009 Mar;15(1):29-41
references.pmid: 19063663
references.citation: Dev Biol. 2002 Nov 1;251(1):142-56
references.pmid: 12413904
references.citation: J Bone Joint Surg Am. 1972 Mar;54(2):323-32
references.pmid: 4651264
references.citation: Development. 2009 Apr;136(7):1093-104
references.pmid: 19224984
references.citation: J Clin Invest. 1988 Feb;81(2):277-81
references.pmid: 3276726
references.citation: Tissue Cell. 2004 Dec;36(6):431-8
references.pmid: 15533458
references.citation: Arthritis Rheum. 2012 Oct;64(10):3302-12
references.pmid: 22576962
references.citation: EMBO J. 2010 Apr 7;29(7):1192-204
references.pmid: 20224554
references.citation: PLoS One. 2011;6(7):e22887
references.pmid: 21818404
references.citation: PLoS One. 2013 Apr 23;8(4):e62582
references.pmid: 23626837
references.citation: Dev Dyn. 1997 Aug;209(4):377-86
references.pmid: 9264261
references.citation: Methods. 2015 Aug;84:109-14
references.pmid: 25828645
references.citation: Genes Dev. 2001 Nov 1;15(21):2865-76
references.pmid: 11691837
references.citation: Stem Cell Res Ther. 2012 Mar 02;3(2):9
references.pmid: 22385573
references.citation: Stem Cells. 2007 Nov;25(11):2786-96
references.pmid: 17656642
references.citation: Tissue Eng Part A. 2010 Jun;16(6):1845-55
references.pmid: 20067399
references.citation: J Cell Physiol. 2001 Dec;189(3):275-84
references.pmid: 11748585
references.citation: Stem Cells Dev. 2015 Apr 15;24(8):995-1007
references.pmid: 25519748
references.citation: Ann N Y Acad Sci. 2002 Jun;961:134-8
references.pmid: 12081882
references.citation: Int Orthop. 2015 May;39(5):995-1003
references.pmid: 25512139
references.citation: Annu Rev Cell Dev Biol. 1998;14:59-88
references.pmid: 9891778
references.citation: Arthritis Rheum. 2009 Jan;60(1):155-65
references.pmid: 19116905
references.citation: Plast Reconstr Surg. 1991 Nov;88(5):753-9
references.pmid: 1924560
references.citation: Dev Growth Differ. 1999 Feb;41(1):29-40
references.pmid: 10445500
references.citation: Cell Physiol Biochem. 2015;35(3):841-57
references.pmid: 25632940
references.citation: BMC Dev Biol. 2010 Jun 21;10:69
references.pmid: 20565961
references.citation: Dev Biol. 1991 May;145(1):99-109
references.pmid: 2019328
references.citation: Tissue Eng. 1999 Dec;5(6):545-54
references.pmid: 10611546
references.citation: Dev Biol. 1991 Aug;146(2):406-15
references.pmid: 1713866
references.citation: Dev Biol. 1988 Nov;130(1):324-34
references.pmid: 3053296
references.citation: Arthritis Rheum. 2009 Sep;60(9):2723-30
references.pmid: 19714579
references.citation: Biotechnol Bioeng. 1999 Nov 5;65(3):274-81
references.pmid: 10486125
references.citation: Med J Malaysia. 2004 May;59 Suppl B:7-8
references.pmid: 15468792
references.citation: Proc Natl Acad Sci U S A. 2006 Dec 12;103(50):19004-9
references.pmid: 17142326
references.citation: Osteoarthritis Cartilage. 2004 Jun;12(6):476-84
references.pmid: 15135144
references.citation: Biomaterials. 2004 Jul;25(16):3211-22
references.pmid: 14980416
references.citation: Development. 2006 Feb;133(4):651-62
references.pmid: 16421188
references.citation: Hum Gene Ther. 2003 Mar 1;14(4):393-402
references.pmid: 12659680
references.citation: J Cell Biol. 2007 Jun 18;177(6):1105-17
references.pmid: 17576802
references.citation: J Orthop Res. 2009 Apr;27(4):529-35
references.pmid: 18973239
references.citation: Blood. 2008 Jul 15;112(2):295-307
references.pmid: 18332228
references.citation: J Biol Chem. 1998 Feb 27;273(9):5026-32
references.pmid: 9478951
references.citation: Hum Mol Genet. 1998 May;7(5):813-22
references.pmid: 9536085
references.citation: Dev Biol. 2004 Sep 1;273(1):134-48
references.pmid: 15302603
references.citation: J Tissue Eng Regen Med. 2015 Apr;9(4):332-42
references.pmid: 23576364
references.citation: J Neurosurg Spine. 2005 Apr;2(4):457-65
references.pmid: 15871486
references.citation: Nat Commun. 2013;4:2850
references.pmid: 24276541
references.citation: Dev Dyn. 2001 Nov;222(3):522-33
references.pmid: 11747085
references.citation: J Med Dent Sci. 2005 Sep;52(3):153-62
references.pmid: 16350840
references.citation: J Clin Invest. 2008 Sep;118(9):3098-108
references.pmid: 18677406
references.citation: FASEB J. 2009 Oct;23(10):3539-52
references.pmid: 19561124
references.citation: Trends Cell Biol. 2006 Mar;16(3):151-8
references.pmid: 16466918
references.citation: J Biol Chem. 2005 Apr 22;280(16):15872-9
references.pmid: 15703179
references.citation: Dev Biol. 2008 Nov 15;323(2):152-65
references.pmid: 18796301
references.citation: Curr Opin Rheumatol. 2006 Jan;18(1):64-73
references.pmid: 16344621
references.citation: Biomaterials. 2005 Jan;26(1):63-72
references.pmid: 15193881
references.citation: Mol Cell Biol. 2001 Aug;21(15):5132-41
references.pmid: 11438668
references.citation: J Biol Chem. 2004 Nov 5;279(45):47081-91
references.pmid: 15337766
references.citation: Development. 2000 Jul;127(14):3141-59
references.pmid: 10862751
references.citation: PLoS One. 2013 Dec 12;8(12):e82166
references.pmid: 24349210
references.citation: Clin Orthop Relat Res. 2004 Oct;(427 Suppl):S89-95
references.pmid: 15480081
references.citation: J Biol Chem. 2012 Apr 27;287(18):14760-71
references.pmid: 22351759
references.citation: Arthritis Rheum. 2008 Sep;58(9):2754-63
references.pmid: 18759300
references.citation: BMC Dev Biol. 2006 Feb 20;6:7
references.pmid: 16504022
references.citation: Tissue Eng Part A. 2011 Apr;17(7-8):1157-67
references.pmid: 21142619
references.citation: Dev Biol. 2002 Aug 15;248(2):240-50
references.pmid: 12167401
references.citation: Science. 2002 Aug 2;297(5582):848-51
references.pmid: 12161657
references.citation: Dev Cell. 2001 Sep;1(3):411-22
references.pmid: 11702952
references.citation: J Orthop Res. 2004 Sep;22(5):1114-9
references.pmid: 15304287
references.citation: J Biol Chem. 1987 Feb 25;262(6):2869-74
references.pmid: 3469200
references.citation: J Bone Joint Surg Am. 2007 Apr;89(4):780-5
references.pmid: 17403800
references.citation: Osteoarthritis Cartilage. 2012 Oct;20(10):1170-8
references.pmid: 22796508
references.citation: Stem Cells. 2005 Aug;23(7):958-64
references.pmid: 16043460
references.citation: Development. 2006 Aug;133(15):3039-49
references.pmid: 16818445
references.citation: Osteoarthritis Cartilage. 2005 Sep;13(9):828-36
references.pmid: 16006153
references.citation: J Cell Sci. 2004 Jan 26;117(Pt 3):417-25
references.pmid: 14702386
references.citation: Adv Exp Med Biol. 2006;589:120-33
references.pmid: 17076278
references.citation: Biomaterials. 2012 Nov;33(31):7699-711
references.pmid: 22841919
references.citation: Cell Signal. 2014 Mar;26(3):468-82
references.pmid: 24333667
references.citation: Tissue Eng Part B Rev. 2014 Apr;20(2):106-25
references.pmid: 23815376
references.citation: J Cell Biochem. 2011 Dec;112(12):3582-93
references.pmid: 21793044
references.citation: Int J Mol Med. 2015 Mar;35(3):579-86
references.pmid: 25543998
references.citation: Cell Tissue Res. 2012 Jan;347(1):11-20
references.pmid: 21701805
references.citation: Tissue Eng. 2006 Apr;12(4):969-79
references.pmid: 16674308
references.citation: Mol Endocrinol. 2006 Oct;20(10):2432-43
references.pmid: 16728531
references.citation: Curr Top Dev Biol. 2009;89:163-92
references.pmid: 19737646
references.citation: Tissue Eng. 2004 Mar-Apr;10(3-4):575-84
references.pmid: 15165474
references.citation: Genes Dev. 1999 Jan 15;13(2):225-37
references.pmid: 9925646
references.citation: Genes Dev. 2003 Feb 1;17(3):394-409
references.pmid: 12569130
references.citation: J Artif Organs. 2006;9(3):179-84
references.pmid: 16998703
references.citation: Biomaterials. 2012 Sep;33(26):6042-51
references.pmid: 22677190
references.citation: J Biol Chem. 2005 Mar 4;280(9):8343-50
references.pmid: 15623506
references.citation: Cell Res. 2009 Jan;19(1):128-39
references.pmid: 19114990
references.citation: Development. 2007 Apr;134(7):1243-51
references.pmid: 17329370
references.citation: J Orthop Res. 2001 Jul;19(4):738-49
references.pmid: 11518286
references.citation: Osteoarthritis Cartilage. 2005 Jun;13(6):527-36
references.pmid: 15922187
references.citation: Biochem Biophys Res Commun. 2007 Jul 20;359(1):108-14
references.pmid: 17532297
references.citation: Cells Tissues Organs. 2008;187(4):275-85
references.pmid: 18187938
references.citation: Mech Dev. 2014 Feb;131:24-34
references.pmid: 24269712
references.citation: J Cell Physiol. 2011 Jun;226(6):1683-93
references.pmid: 21413026
references.citation: Dev Biol. 2007 Oct 15;310(2):304-16
references.pmid: 17822689
references.citation: Genes Dev. 1999 Aug 15;13(16):2072-86
references.pmid: 10465785
references.citation: Development. 2005 Sep;132(17):3977-88
references.pmid: 16049113
references.citation: Tissue Eng Part A. 2013 Sep;19(17-18):1960-71
references.pmid: 23611597
references.citation: J Musculoskelet Neuronal Interact. 2008 Oct-Dec;8(4):316-7
references.pmid: 19147955
references.citation: Biorheology. 2006;43(3,4):355-70
references.pmid: 16912408
references.citation: Arthritis Res Ther. 2012 Jul 20;14(4):R168
references.pmid: 22817660
references.citation: Tissue Eng. 2004 May-Jun;10(5-6):736-46
references.pmid: 15265290
references.citation: J Orthop Surg Res. 2014 Oct 28;9:102
references.pmid: 25346272
references.citation: Proc Natl Acad Sci U S A. 2003 Aug 5;100(16):9360-5
references.pmid: 12878728
references.citation: Tissue Eng. 1999 Feb;5(1):1-11
references.pmid: 10207185
references.citation: Biotechnol Bioeng. 2005 Aug 5;91(3):338-55
references.pmid: 15959891
references.citation: Curr Top Dev Biol. 2004;60:91-126
references.pmid: 15094297
references.citation: Dev Biol. 2004 Apr 15;268(2):441-7
references.pmid: 15063179
references.citation: Development. 1997 Jun;124(11):2235-44
references.pmid: 9187149
references.citation: Gene Regul Syst Bio. 2008 Apr 22;2:177-91
references.pmid: 19787082
references.citation: PLoS Genet. 2012;8(11):e1003002
references.pmid: 23133399
references.citation: J Cell Biol. 1987 Dec;105(6 Pt 2):2861-76
references.pmid: 3320058
references.citation: Front Biosci (Landmark Ed). 2013 Jan 01;18(2):493-503
references.pmid: 23276938
references.citation: Circ Res. 2007 Sep 14;101(6):581-9
references.pmid: 17673669
references.citation: J Cell Biol. 1987 Jul;105(1):457-63
references.pmid: 3475276
references.citation: Nucleic Acids Res. 2008 May;36(9):3011-24
references.pmid: 18390577
references.citation: Biochem Biophys Res Commun. 2011 Nov 4;414(4):653-9
references.pmid: 21986536
references.citation: Int J Exp Pathol. 2009 Feb;90(1):1-15
references.pmid: 19200246
references.citation: J Cell Physiol. 2008 Oct;217(1):228-41
references.pmid: 18506848
references.citation: J Anat. 2003 Jun;202(6):495-502
references.pmid: 12846471
references.citation: J Cell Biol. 2009 Nov 30;187(5):685-99
references.pmid: 19948490
references.citation: Dev Cell. 2003 Sep;5(3):367-77
references.pmid: 12967557
references.citation: Cell Physiol Biochem. 2013;32(2):486-98
references.pmid: 23988723
references.citation: Arthritis Rheum. 2012 Jun;64(6):1909-19
references.pmid: 22143896
references.citation: Genes Cells. 1996 Jan;1(1):1-9
references.pmid: 9078362
references.citation: J Cell Biol. 2002 Jul 8;158(1):39-51
references.pmid: 12105181
references.citation: Tissue Eng. 1998 Winter;4(4):415-28
references.pmid: 9916173
references.citation: Nat Genet. 2003 Nov;35(3):215-7
references.pmid: 14528307
references.citation: Science. 2004 Sep 3;305(5689):1437-41
references.pmid: 15284456
references.citation: Stem Cell Reports. 2015 Mar 10;4(3):459-72
references.pmid: 25733021
references.citation: Biomaterials. 2012 Jun;33(18):4500-7
references.pmid: 22449550
references.citation: J Bone Miner Res. 2000 Sep;15(9):1659-68
references.pmid: 10976986
references.citation: Sci Rep. 2013;3:2683
references.pmid: 24042188
references.citation: Cell Physiol Biochem. 2015;36(1):44-60
references.pmid: 25924688
references.citation: Cell. 1987 Nov 20;51(4):589-99
references.pmid: 3677168
references.citation: Proc Natl Acad Sci U S A. 2004 Dec 28;101(52):18087-92
references.pmid: 15601778
references.citation: Dev Biol. 1998 Aug 1;200(1):35-45
references.pmid: 9698454
references.citation: PLoS One. 2011;6(7):e21679
references.pmid: 21799743
references.citation: Dev Biol. 2013 Sep 15;381(2):423-33
references.pmid: 23806211
references.citation: Dev Dyn. 1997 Apr;208(4):516-25
references.pmid: 9097023
references.citation: Dev Cell. 2002 Jun;2(6):695-706
references.pmid: 12062082
references.citation: Development. 2007 Sep;134(18):3283-95
references.pmid: 17699607
references.citation: Stem Cells. 2005;23(1):113-23
references.pmid: 15625128
references.citation: J Biol Chem. 2001 Sep 21;276(38):35290-6
references.pmid: 11466306
references.citation: Proc Natl Acad Sci U S A. 2009 Aug 25;106(34):14420-5
references.pmid: 19590010
references.citation: Dev Dyn. 2009 May;238(5):1150-65
references.pmid: 19334275
references.citation: J Cell Sci. 1998 Jul 30;111 ( Pt 14):2067-76
references.pmid: 9645953
references.citation: J Bone Miner Metab. 2003;21(6):344-52
references.pmid: 14586790
references.citation: J Cell Biol. 2004 Jul 5;166(1):85-95
references.pmid: 15226309
references.citation: Cell Tissue Res. 2007 Mar;327(3):539-51
references.pmid: 17093926
references.citation: Arthritis Rheum. 2007 Oct;56(10):3297-306
references.pmid: 17907154
references.citation: J Biol Chem. 2000 Feb 25;275(8):5613-9
references.pmid: 10681543
references.citation: Dev Dyn. 2003 Oct;228(2):208-16
references.pmid: 14517992
references.citation: Osteoarthritis Cartilage. 2004 Jul;12(7):559-67
references.pmid: 15219571
references.citation: Int J Dev Biol. 1997 Feb;41(1):91-102
references.pmid: 9074941
references.citation: J Clin Invest. 2008 Feb;118(2):429-38
references.pmid: 18246193
references.citation: Proc Natl Acad Sci U S A. 2005 Apr 5;102(14):5062-7
references.pmid: 15781876
references.citation: Arthritis Rheum. 2009 Oct;60(10):3038-48
references.pmid: 19790048
references.citation: Mol Cell Biol. 2015 Jun;35(12):2119-30
references.pmid: 25870110
references.citation: Arthritis Rheum. 2004 Nov;50(11):3561-73
references.pmid: 15529345
references.citation: Biochem Biophys Res Commun. 2007 Nov 23;363(3):513-8
references.pmid: 17889828
references.citation: Arthritis Res Ther. 2005;7(4):R904-14
references.pmid: 15987493
references.citation: Genes Dev. 2002 Aug 1;16(15):1990-2005
references.pmid: 12154128
references.citation: Biochem Biophys Res Commun. 2004 Jul 30;320(3):914-9
references.pmid: 15240135
references.citation: Nat Med. 1999 Jun;5(6):623-8
references.pmid: 10371499
references.citation: Cytotechnology. 2008 Jan;56(1):1-7
references.pmid: 19002835
references.citation: Genes Dev. 2006 Nov 1;20(21):2937-42
references.pmid: 17050674
references.citation: Front Physiol. 2010 Oct 14;1:135
references.pmid: 21423376
references.citation: Biochem Biophys Res Commun. 2005 Aug 12;333(4):1300-8
references.pmid: 15979579
references.citation: Ann Biomed Eng. 2005 Jul;33(7):943-50
references.pmid: 16060535
references.citation: Mol Pharmacol. 2006 Nov;70(5):1469-80
references.pmid: 16887934
references.citation: Genes Dev. 2004 Apr 15;18(8):952-63
references.pmid: 15107406
references.citation: Exp Cell Res. 2002 Feb 15;273(2):197-203
references.pmid: 11822875
references.citation: Am J Vet Res. 2005 Apr;66(4):599-605
references.pmid: 15900939
references.citation: J Biol Chem. 2002 Aug 23;277(34):30870-8
references.pmid: 12077113
references.citation: J Bone Miner Res. 2007 Mar;22(3):366-74
references.pmid: 17181398
references.citation: Braz J Med Biol Res. 2014 Apr;47(4):279-86
references.pmid: 24652327