Bone Morphogenetic Proteins, Their Antagonists, and the Skeleton
Ernesto Canalis,
Aris N. Economides and
Elisabetta Gazzerro
Department of Research (E.C., E.G.), Saint Francis Hospital and Medical Center, Hartford, Connecticut 06105; University of Connecticut School of Medicine (E.C., E.G.), Farmington, Connecticut 06030; and Regeneron Pharmaceuticals, Inc. (A.N.E.), Tarrytown, New York 10591
Correspondence: Address all correspondence and requests for reprints to: Ernesto Canalis, M.D., Department of Research, Saint Francis Hospital and Medical Center, 114 Woodland Street, Hartford, Connecticut 06105-1299. E-mail: ecanalis{at}stfranciscare.org
Skeletal homeostasis is determined by systemic hormones andlocal factors. Bone morphogenetic proteins (BMP) are uniquebecause they induce the differentiation of mesenchymal cellstoward cells of the osteoblastic lineage and also enhance thedifferentiated function of the osteoblast. However, the activityof BMPs needs to be tempered by intracellular and extracellularantagonists. BMPs bind to specific receptors and signal by phosphorylatingthe cytoplasmic proteins mothers against decapentaplegic (Smad)1 and 5, which form heterodimers with Smad 4, and after nucleartranslocation regulate transcription. BMP antagonists can becategorized as pseudoreceptors that compete with signaling receptors,inhibitory Smads that block signaling, intracellular bindingproteins that bind Smad 1 and 5, and factors that induce ubiquitinationand proteolysis of signaling Smads. In addition, a large numberof extracellular proteins that bind BMPs and prevent their bindingto signaling receptors have emerged. They are the componentsof the Spemann organizer, noggin, chordin, and follistatin,members of the Dan/Cerberus family, and twisted gastrulation.The antagonists tend to be specific for BMPs and are regulatedby BMPs, indicating the existence and need of local feedbackmechanisms to temper BMP cellular activities.
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