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Fig. 7. Gremlin modulation of mesenchymal-epithelial interaction in the pyloric sphincter is under SOX9 control. (A) Whole-mount in situ hybridization of E7 dissected gut using antisense Gremlin riboprobe. Gremlin expression strongly demarcates the stomach from the duodenum and is present in the mesoderm of the pyloric sphincter (red arrow, A). (B) Histological section of Gremlin-misexpressing E9 gizzard. After mesodermal Gremlin misexpression, the gizzard epithelial cells present bleb like cilia (B). Immunohistochemistry with {alpha}3C2 antibodies show retrovirus infection restricted to the mesoderm of the gizzard (upper inset B). GFP- (C), SOX9- (D) and SOX9{Delta}Cter- (E) misexpressing E7 stomach sections followed by detection of Gremlin expression by in situ hybridization using antisense Gremlin riboprobe. Gremlin is expressed in the mesoderm and adventitia of the pyloric sphincter (C). Ectopic expression of Gremlin in the mesodermal stomach is observed after retroviral SOX9 misexpression in the gizzard (red arrowheads, D). Downregulation of Gremlin expression is correlated with retroviral SOX9{Delta}Cter misexpression in the pyloric sphincter (black arrowheads, E), while Gremlin expression is normal in the adventitia. duo, duodenum; e, endoderm; gizz, gizzard; m, mesoderm; pv, proventriculus; pyl, pyloric structure.