title: Differential expression of retinal determination genes in the principal and secondary eyes of Cupiennius salei Keyserling (1877).
abstract: BACKGROUND
  Transcription factors that determine retinal development seem to be conserved in different phyla throughout the animal kingdom. In most representatives, however, only a few of the involved transcription factors have been sampled and many animal groups remain understudied. In order to fill in the gaps for the chelicerate group of arthropods, we tested the expression pattern of the candidate genes involved in the eye development in the embryo of the wandering spider Cupiennius salei. One main objective was to profile the molecular development of the eyes and to search for possible variation among eye subtype differentiation. A second aim was to form a basis for comparative studies in order to elucidate evolutionary pathways in eye development.
  
  
  RESULTS
  We screened the spider embryonic transcriptome for retina determination gene candidates and discovered that all except one of the retinal determination genes have been duplicated. Gene expression analysis shows that the two orthologs of all the genes have different expression patterns. The genes are mainly expressed in the developing optic neuropiles of the eyes (lateral furrow, mushroom body, arcuate body) in earlier stages of development (160 to 220 h after egg laying). Later in development (180 to 280 h after egg laying), there is differential expression of the genes in disparate eye vesicles; for example, Cs-otxa is expressed only in posterior-lateral eye vesicles, Cs-otxb, Cs-six1a, and Cs-six3b in all three secondary eye vesicles, Cs-pax6a only in principal eye vesicles, Cs-six1b in posterior-median, and posterior-lateral eye vesicles, and Cs-six3a in lateral and principal eye vesicles.
  
  
  CONCLUSIONS
  Principle eye development shows pax6a (ey) expression, suggesting pax6 dependence, although secondary eyes develop independently of pax6 genes and show differential expression of several retinal determination genes. Comparing this with the other arthropods suggests that pax6-dependent median eye development is a ground pattern of eye development in this group and that the ocelli of insects, the median eyes of chelicerates, and nauplius eyes can be homologised. The expression pattern of the investigated genes makes it possible to distinguish between secondary eyes and principal eyes. Differences of gene expression among the different lateral eyes indicate disparate function combined with genetic drift.
authors.affiliation: Department of Neurobiology, Centre for Organismal Systems Biology, Faculty of Life Sciences, University of Vienna, Althanstraße 14, 1090 Vienna, Austria.
authors.forename: Leyli
authors.initials: L
authors.lastname: Samadi
authors.affiliation: Department of Neurobiology, Centre for Organismal Systems Biology, Faculty of Life Sciences, University of Vienna, Althanstraße 14, 1090 Vienna, Austria.
authors.forename: Axel
authors.initials: A
authors.lastname: Schmid
authors.affiliation: Department of Neurobiology, Centre for Organismal Systems Biology, Faculty of Life Sciences, University of Vienna, Althanstraße 14, 1090 Vienna, Austria.
authors.forename: Bo Joakim
authors.initials: BJ
authors.lastname: Eriksson
keywords: Eye development; Principal eyes; Retinal determination gene network; Secondary eyes; Spider