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A three-step model for BMP signaling in the Helobdella O–P equivalence group (orange): 1) short-range signaling from q bandlet to p bandlet mediated by BMP5-8; 2) BMP5-8 signaling specifies the P fate (red), including up-regulation of gremlin; 3) gremlin signaling specifies O fate (yellow) in the neighboring bandlet by inhibiting broadly occurring BMP2/4 signaling (see text for further details).
