Evolution of cell fate specification modes in spiral cleavage

Spiral cleavage is a highly stereotypical early embryonic program, and the ancestral, defining feature to Spiralia, a major phylogenetic clade including almost half of the animal phyla. Remarkably, spiral-cleaving embryos specify homologous cell fates (e.g. the progenitor cell of posterodorsal structures) conditionally –via cell interactions– or autonomously –via segregation of maternal inputs. This variation occurs naturally, even between closely related species, and has been related to the precocious formation of adult characters (adultation) in larvae of autonomous spiral-cleaving species. How spiralian lineages repeatedly shifted between these two cell fate specification modes is largely unexplored, because the mechanisms controlling spiral cleavage are still poorly characterized.This project tests the hypothesis that maternal chromatin and transcriptional regulators differentially incorporated in oocytes with autonomous spiral cleavage explain the evolution of this mode of cell fate specification. Through a comparative and phylogenetic-guided approach, we will combine bioinformatics, live imaging, and molecular and experimental techniques to: (i) Comprehensively identify differentially supplied maternal factors among spiral cleaving oocytes with distinct cell fate specification modes using comparative RNA-seq and proteomics; (ii) Uncover the developmental mechanisms driving conditional spiral cleavage, which is the ancestral embryonic mode; and (iii) Investigate how maternal chromatin and transcriptional regulators define early cell fates, and whether these factors account for the repeated evolution of autonomous specification modes.Our results will fill a large gap of knowledge in our understanding of spiral cleavage and its evolution. In a broader context, this project will deliver fundamental insights into two core questions in evolutionary developmental biology: how early embryonic programs evolve, and how they contribute to phenotypic change.