The reproductive system, embryonic development, larval development and metamorphosis of the sea urchin Heliocidaris erythrogramma (Val.) (Echinoidea : Echinometridae)

Abstract

H. erythrogramma has an annual reproductive cycle, spawning asynchronously. The female cycle has five phases: resting, regenerating, mature, spent and regressing; the male cycle omits the regression phase. Regression and resting occur only during April-August; regeneration begins in September; during December-March individual urchins pass through the mature, spent and regenerating phases two or three times. The egg averages 400 �m diameter and floats at the water surface. At 24-25�C, equal, total, radial cleavage yields a wrinkled coeloblastula by 6-7 hours; this undergoes egression, becoming a spherical stereoblastula by 13-14 hours; during this process an acellular central yolk mass is segregated from a peripheral columnar blastoderm. The ciliated stereoblastula gastrulates by a typical diphasic invagination, beginning at 15 hours, and the gastrulating embryo hatches as a ciliated, planktonic yolk-larva at 15-16 hours. Lecithotrophic development of the yolk-larva to a uniformly ciliated, planktonic vitellaria is complete by 1.5 days, and the larva metamorphoses gradually to a lecithotrophic juvenile over the next 3.5 days. It remains planktonic during the first half of this period (to 3.75 days), but settles during the second half (to 5 days). The juvenile begins to feed >3 weeks after settling. Vegetally proliferating mesenchyme invades the yolk mass during gastrulation. Invagination of the archenteron is followed by evagination of a primary pair of coelomic pouches from its wall; the left enterocoel forms the hydrocoel. Other coelomic cavities are re-formed by schizocoely. The larval gut rudiment temporarily loses connection with the exterior after the blastopore closes. The proctodaeum and stomodaeum of the juvenile form secondarily during and after metamorphosis. Ectodermal development of the echinus rudiment, though gradual, is typical of echinoids. The blastulation of H. evythrogramma is unique among large-egged echinoderms in combining egression of a wrinkled coeloblastula with segregation of a blastoderm around a central yolk mass. The development of the species supports the view that the vitellaria larva is secondarily evolved within the Echinoidea.