title: Dermestes maculatus: an intermediate-germ beetle model system for evo-devo.
abstract: BACKGROUND
  Understanding how genes change during evolution to direct the development of diverse body plans is a major goal of the evo-devo field. Achieving this will require the establishment of new model systems that represent key points in phylogeny. These new model systems must be amenable to laboratory culture, and molecular and functional approaches should be feasible. To date, studies of insects have been best represented by the model system Drosophila melanogaster. Given the enormous diversity represented by insect taxa, comparative studies within this clade will provide a wealth of information about the evolutionary potential and trajectories of alternative developmental strategies.
  
  
  RESULTS
  Here we established the beetle Dermestes maculatus, a member of the speciose clade Coleoptera, as a new insect model system. We have maintained a continuously breeding culture in the lab and documented Dermestes maculatus embryogenesis using nuclear and phalloidin staining. Anterior segments are specified during the blastoderm stage before gastrulation, and posterior segments are added sequentially during germ band elongation. We isolated and studied the expression and function of the pair-rule segmentation gene paired in Dermestes maculatus. In this species, paired is expressed in stripes during both blastoderm and germ band stages: four primary stripes arise prior to gastrulation, confirming an intermediate-germ mode of development for this species. As in other insects, these primary stripes then split into secondary stripes. To study gene function, we established both embryonic and parental RNAi. Knockdown of Dmac-paired with either method resulted in pair-rule-like segmentation defects, including loss of Engrailed expression in alternate stripes.
  
  
  CONCLUSIONS
  These studies establish basic approaches necessary to use Dermestes maculatus as a model system. Methods are now available for use of this intermediate-germ insect for future studies of the evolution of regulatory networks controlling insect segmentation, as well as of other processes in development and homeostasis. Consistent with the role of paired in long-germ Drosophila and shorter-germ Tribolium, paired functions as a pair-rule segmentation gene in Dermestes maculatus. Thus, paired retains pair-rule function in insects with different modes of segment addition.
authors.affiliation: Department of Entomology, University of Maryland, 4112 Plant Sciences Building, College Park, MD 20742 USA ; Program in Molecular and Cell Biology, University of Maryland, 4112 Plant Sciences Building, College Park, MD 20742 USA.
authors.forename: Jie
authors.initials: J
authors.lastname: Xiang
authors.affiliation: Department of Entomology, University of Maryland, 4112 Plant Sciences Building, College Park, MD 20742 USA.
authors.forename: Iain S
authors.initials: IS
authors.lastname: Forrest
authors.affiliation: Department of Entomology, University of Maryland, 4112 Plant Sciences Building, College Park, MD 20742 USA ; Program in Molecular and Cell Biology, University of Maryland, 4112 Plant Sciences Building, College Park, MD 20742 USA.
authors.forename: Leslie
authors.initials: L
authors.lastname: Pick
keywords: Dermestes maculatus; Evo-devo; Pair-rule patterning; RNAi; Segmentation; paired