title: A detailed staging scheme for late larval development in Strongylocentrotus purpuratus focused on readily-visible juvenile structures within the rudiment.
abstract: BACKGROUND
  The purple sea urchin, Strongylocentrotus purpuratus, has long been the focus of developmental and ecological studies, and its recently-sequenced genome has spawned a diversity of functional genomics approaches. S. purpuratus has an indirect developmental mode with a pluteus larva that transforms after 1-3 months in the plankton into a juvenile urchin. Compared to insects and frogs, mechanisms underlying the correspondingly dramatic metamorphosis in sea urchins remain poorly understood. In order to take advantage of modern techniques to further our understanding of juvenile morphogenesis, organ formation, metamorphosis and the evolution of the pentameral sea urchin body plan, it is critical to assess developmental progression and rate during the late larval phase. This requires a staging scheme that describes developmental landmarks that can quickly and consistently be used to identify the stage of individual living larvae, and can be tracked during the final two weeks of larval development, as the juvenile is forming.
  
  
  RESULTS
  Notable structures that are easily observable in developing urchin larvae are the developing spines, test and tube feet within the juvenile rudiment that constitute much of the oral portion of the adult body plan. Here we present a detailed staging scheme of rudiment development in the purple urchin using soft structures of the rudiment and the primordia of these juvenile skeletal elements. We provide evidence that this scheme is robust and applicable across a range of temperature and feeding regimes.
  
  
  CONCLUSIONS
  Our proposed staging scheme provides both a useful method to study late larval development in the purple urchin, and a framework for developing similar staging schemes across echinoderms. Such efforts will have a high impact on evolutionary developmental studies and larval ecology, and facilitate research on this important deuterostome group.
authors.forename: Andreas
authors.initials: A
authors.lastname: Heyland
authors.affiliation: Hopkins Marine Station of Stanford University, Pacific Grove, CA 93950, USA. seastar@stanford.edu.
authors.forename: Jason
authors.initials: J
authors.lastname: Hodin
mesh_terms: D000818:Animals; D015509:Developmental Biology / Q000379:methods; D005260:Female; D007814:Larva / Q000254:growth & development; D008018:Life Cycle Stages; D008297:Male; D008675:Metamorphosis, Biological; D047270:Strongylocentrotus purpuratus / Q000196:embryology / Q000254:growth & development*; D013997:Time Factors