Evolutionary modifications of morphogenetic mechanisms and alternate life history strategies in ascidians

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Abstract
Comparative embryological studies using anural and urodele ascidians provide an experimental system to study interactions between developmental and evolutionary mechanisms that produce alternate life history strategies. In this paper, cellular features of anural morphogenesis in Molgula pacifica are compared to morphogenesis in species that develop tailed (urodele) larvae and other anural molgulid species. The results of these studies are discussed with regard to possible mechanisms responsible for the evolution of anural morphogenesis and the ecological consequences of anural development. Early developmental processes including ooplasmic segregation, cleavage patterns, and the site and timing of gastrulation were similar in M. pacifica compared to urodele embryos and embryos produced by other anural species. The limited extent of invagination caused by large, yolky cells that restricted vegetal pole cell movements in M. pacifica gastrulae contrasted with the extensive movements of vegetal pole cells that accompanied invagination in M. provisionalis embryos and the embryos of four urodele species. The modified mode of gastrulation exhibited by M. pacifica embryos is likely due to the relatively high yolk content of their eggs. The developmental fates of muscle and epidermal progenitor cells in M. pacifica embryos were altered compared to urodele embryos. Ultrastructural studies and acetylcholinesterase histochemistry experiments indicate that muscle progenitor cells have lost their potential to develop muscle cell features. This loss in myogenic potential suggests that muscle progenitor cells were re-programmed to die. However, this possibility was not supported by the results of ultrastructural studies. A second possibility is discussed in that muscle progenitor cells may have been re-specified to differentiate into adult cells after metamorphosis. Evidence is presented suggesting that the timing mechanism responsible for controlling the onset of metamorphosis, first evident by the outgrowth of epidermal ampullae, was modified in M. pacifica. This paper concludes with a discussion of how anural morphogenesis altered the ancestoral urodele life cycle and the possible ecological benefits of these evolutionary alterations.