Left column. Triple FISH of otd, slp1 and gt in wild type (A), run− (B), cic− (C) and run− cic− double mutant (D) embryos. White rectangles indicate the regions used for image quantification. Middle column. Quantification of otd, slp1 and gt in wild type (E), run− (F), cic− (G) and run− cic− double mutant (H) embryos. Y-axis represents normalized intensities. Right column. engrailed (en) mRNA patterns in stage 10–11 embryos. In wild type embryos (I), en is expressed in five head domains that foreshadow the ocular (oc), antennal (an), intercalary (ic), mandibular (md), and maxillary (mx) segments. In run− (J) and cic− (K) mutant embryos, the head region appears enlarged, and all cephalic en domains are still visible. In double mutant embryos (L), only a single cephalic en domain (oc) is observed. Arrow in (L) points to the oc on the opposite side of the embryo.