Intracapsular Feeding by Embryos of the Gastropod Genus Littorina

Many gastropod species develop within egg capsules within which larvae are provided with extraembryonic nutrients. Species with encapsulated development frequently have transitory embryonic organs, such as "larval kidneys," that may represent specializations for consumption of intracapsular nutrition. Larvae of Littorina species with nonplanktonic, encapsulated development consume intracapsular albumen, but they lack obvious morphological modifications for albumen consumption. To determine the mechanism and location of protein uptake, larvae of seven species of Littorina (L. keenae, L. littorea, L. plena, L. saxatilis, L. scutulata, L. sitkana, L. subrotundata) were exposed to solutions of either fluorescently labeled protein (FITC-bovine serum albumen) or ferritin. Under fluorescence microscopy, larvae of all species with encapsulated, nonplanktonic development displayed strong regional affinity for FITC in the ciliated cells of the velum, whereas hatched larvae of planktotrophic Littorina species did not. Transmission electron microscopy of epithelial cells of nonplanktotrophic veligers exposed to ferritin supported the interpretation that localized affinity for labeled protein indicated endocytotic protein uptake. Planktotrophic Littorina and Littorina with encapsulated, nonplanktonic development were shown to share equivalent velar width/larval length ratios during early embryonic development, whereas a literature search suggested that in other nonplanktotrophic prosobranchs the velum is relatively smaller than in planktotrophs. Retention of a large velum in Littorina that develop entirely within egg capsules may facilitate feeding on intracapsular protein, in the absence of specialized assimilative organs found in other species with encapsulated development.