Distinct expression patterns of the two T-box homologues Brachyury and Tbx2/3 in the placozoan Trichoplax adhaerens

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Abstract
Trichoplax adhaerens is the only species known from the phylum Placozoa with one of the simplest metazoan body plans. In the small disc-like organism an upper and a lower epithelium can be distinguished with a less compact third cell layer in between. When Trichoplax was first described in 1883, the relation of these three cell layers with ectoderm, endoderm and mesoderm of higher animals was discussed. Still, little is known about embryonic development of Trichoplax, however, genes thought to be specific for mesoderm in bilaterian animals turned out to be already present in non-bilaterians. Searching for a Brachyury homologue, two members of the T-box gene family were isolated from Trichoplax, Brachyury and a Tbx2/3 homologue. The T-box genes encode a transcription factor family characterized by the DNA-binding T-box domain. T-box genes have been found in all metazoans so far investigated, but in contrast to other transcription factors such as the homeobox family, T-box genes are not present in plants or fungi. The distinct expression patterns of two T-box genes in Trichoplax point to non-redundant functions already present at the beginning of animal evolution. Since the expression patterns derived by in situ hybridization do not overlap with anatomical structures, it can be concluded that this simple animal has more than the four cell types described in the literature. This hidden complexity and the unresolved position in relation to Porifera, Cnidaria, Ctenophora and Bilateria highlight the necessity of the inclusion of Trichoplax in studies of comparative evolutionary and developmental biology.