Orthodenticle and Empty Spiracles Genes Are Expressed in a Segmental Pattern in Chelicerates

Members of the orthodenticle (otd/Otx) and empty spiracles (ems/Emx) gene families are head gap genes that encode homeodomain-containing DNA-binding proteins. Although numerous studies show their central role in developmental processes in brain specification, a surprisingly high number of other developmental processes have been shown to involve their expression. In this paper, we report the identification and expression of ems and otd in two chelicerate species: a scorpion, Euscorpius flavicaudis (Chactidae, Scorpiona, Arachnida, Euchelicerata) and a spider, Tegenaria saeva (Aranea, Arachnida, Euchelicerata). We show that both ems and otd are expressed not only in an anterior head domain but also along the entire anterior-posterior axis during embryonic development. The expression patterns for both genes are typically segmental and concern neurectodermal territories. During patterning of the opisthosoma, ems and otd are expressed in the lateral ectoderm just anterior to the limb bud primordia giving rise to respiratory organs and spinnerets (spider). This common pattern found in two divergent species thus appears to be a conserved character of chelicerates. These results are discussed in terms of evolutionary origin of respiratory organs and/or functional pathway recruitment.