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Bone morphogenetic protein 2 induces noncanonical Wnt signaling in osteoblast precursors through the receptor ROR2. Bone morphogenetic proteins (BMPs) are members of the transforming growth factor-β superfamily and are key factors in skeletal development. A subset of BMPs exert their activity through noncanonical Wnt signaling, and a role for noncanonical Wnt signaling in bone morphogenesis is suggested by the phenotype of mice lacking the receptor Frizzled-4 (Fzd4). However, the effect of BMP signaling on the noncanonical Wnt pathway is largely unexplored, and the identity of the noncanonical receptor complex has remained controversial. Here, we show that BMP2 induces noncanonical Wnt signaling in osteoblast precursors and that the receptor ROR2 is the core component of the noncanonical signaling complex. We also show that BMP2-mediated noncanonical Wnt signaling is dispensable for osteoblast differentiation. However, ectopic expression of ROR2 in calvarial osteoblasts results in a robust increase in both endochondral bone formation and periosteal bone formation, with loss of the periosteal envelope, indicating that ROR2 is a key regulator of periosteal bone formation. Taken together, our data suggest that BMP2 signaling to ROR2 represents a noncanonical Wnt signaling pathway critical for regulating periosteal bone formation.The present invention relates to a new and distinct cultivar of Illicium plant, botanically known as Illicium verum ‘Cameo’ and will be referred to hereafter by its cultivar name, ‘Cameo’. ‘Cameo’ represents a new evergreen shrub grown for landscape use. ‘Cameo’ derived from a naturally occurring mutation of Illicium verum ‘Cameo’ (not patented). ‘Cameo’ was discovered by the Inventor in October of 2012 in a containerized planting of 2,000 plants of ‘Cameo’ in a research nursery in Boskoop, The Netherlands. Asexual propagation of the new cultivar was first accomplished by stem cuttings in February of 2013 in Boskoop, The Netherlands. Asexual propagation by stem cuttings has determined that the characteristics of the new cultivar are stable and are reproduced

 

 

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