title: Comparative gene expression supports the origin of the incisor and molar process from a single endite in the mandible of the red flour beetle Tribolium castaneum.
abstract: BACKGROUND
  The biting edge of the primitive arthropod mandible consists of a biting incisor process and a crushing molar process. These structures are thought to be derived from a structure known as an endite but the precise details of this are not understood. Various hypotheses concerning the number of endites present in the arthropod mandible have been proposed.In the developing embryo, the mandible has an inner and outer lobe that are likely to develop into the incisor and molar processes of the larval mandible; these two lobes are commonly held to be derived from separate endites and to be serially homologous to the galea and lacinia endites of the maxillary appendage respectively (Machida).
  
  
  RESULTS
  We undertook a study of the development of the embryonic mandible of the beetle Tribolium castaneum using the expression of developmental genes as markers of the developing endites in the mandible and maxilla.The Tribolium ortholog of paired (Tc-prd) has expression domains in the developing maxillary and labial endites as well as the inner and outer lobes of the mandible. Following the expression of Tc-prd in the developing mandible through to late stage embryos shows that the molar and incisor process develop from the inner and outer lobes respectively.In addition to Tc-prd, we compared the expression of genes in the endites of the maxilla to the mandible to draw conclusions about the number of endites in the mandible. Homologs of dachshund are typically expressed in the endites of mandibulate gnathal appendages. Comparison of the expression of Tc-prd, Tribolium dachshund (Tc-dac) and Tribolium wingless (Tc-wg) between the endites of the maxilla and the mandible suggest that, while there are two endites in the maxilla only a single endite is present in the mandible.
  
  
  CONCLUSIONS
  Comparative gene expression suggests that the Tribolium mandible has a single endite from which both mandible lobes are derived. Our results do not support Machida's hypothesis homologising the incisor and molar processes of the mandible to the galea and lacinia endites of the maxilla. We propose, instead, that both incisor and molar processes are derived from a single endite serially homologous to the lacinia of the maxilla.
authors.affiliation: Department of Genetics, Environment and Evolution, University College London, Darwin Building, Gower Street, London, WC1E 6BT, UK. m.telford@ucl.ac.uk.
authors.forename: Joshua F
authors.initials: JF
authors.lastname: Coulcher
authors.forename: Maximilian J
authors.initials: MJ
authors.lastname: Telford