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2B). Taken together, the Pv-twi sequence clearly belongs to the twist class of bHLH genes. Expression of Pv-twi during mesoderm development The invariant cleavage pattern of spiralian embryos such as that of Patella allows the tracing of blastomere fates during development. It has thus been established that spiralian embryos have two sources of mesoderm: ectomesoderm and endomesoderm (Verdonk and van den Biggelaar 1983; Boyer and Henry 1998; Damen and Dictus 2001). The ectomesoderm in Patella is derived from derivatives of the so-called 2b, 3a and 3b cells and the endomesoderm from the progeny of the 4d cell, or mesentoblast (van den Biggelaar 1977; Dictus and Damen 40 twist Figure 3.

49 Chapter 4 Introduction Comparative analysis of genes involved in development of multicellular organisms has revealed conserved sets of transcription factors that are deployed by numerous species. One of these is the class of homeodomain transcription factors, characterized by a 60 amino acid DNA binding domain. Many members of this class can be found throughout the metazoans. The best-known example is the Hom/Hox cluster that is involved in anterior/posterior patterning in various species (Ferrier and Holland 2001).

However, without knowing the role of twist genes in mesoderm development in lophotrochozoans, we cannot conclude that this common feature is the ancestral role of twist genes in triploblasts. In order to study whether twist is involved in mesoderm formation in lophotrochozoans, we set out to clone a putative twist ortholog of the gastropod mollusc Patella vulgata. We present here the cloning, characterization and the early spatio-temporal expression pattern of the twist ortholog of Patella vulgata, Pv-twi.

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