Gizzard formation and the role of Bapx1.

The anterior-posterior gut pattern is formed from three broad domains: fore-, mid-, and hindgut that have distinct functional, morphological, and molecular boundaries. The stomach demarcates the posterior boundary of the foregut. Avian stomachs are composed of two chambers: the anterior chamber (proventriculus) and the thick muscular posterior chamber (gizzard). Expression of candidate pattern formation control factors are restricted in the chick stomach regions such that Bmp4 and Wnt5a are not expressed in the gizzard. We previously implicated Bmp4 as controlling growth and differentiation of the gut musculature. Bmp4 is not expressed in the developing gizzard but is expressed in the rest of the gut including the adjacent proventriculus and midgut. Bapx1 (Nkx3.2) is expressed in the gizzard musculature but not in the proventriculus or midgut. We show ectopic expression of Bapx1 in the proventriculus results in a gizzard-like morphology and inhibits the normal proventricular expression of Bmp4 and Wnt5a. Overexpression of a reverse-function Bapx1 construct can result in a small stomach and ectopic extension of Bmp4 and Wnt5a expression into the gizzard. We suggest the role of Bapx1 is to regulate the expression of Bmp4 and Wnt5a to pattern the avian stomach.