Russel L. Zimmer

Attachment and metamorphosis of the cheilo-ctenostome bryozoan bugula neritina (linné)†‡

The structure, attachment and subsequent metamorphosis of larvae of the marine bryozoan Bugula neritina were studied by light and electron microscopy.

Fine structure of a potential photoreceptor organ in the larva of Bugula neritina (Bryozoa)

SummaryThe pigmented spots of bryozoan larvae have often been implicated in photoreception due to their preferential occurrence in larvae with positive phototactic behavior. Results of light and electron microscopic studies of Bugula neritina show that the larvae possess two spots, each with a basal sensory cell situated at the base of a pit-like depression. The embedment of the pit and its basal cell in a pad of subepidermal pigment cells allows for directional illumination. The basal sensory cell produces a ball-like mass of non-motile cilia. The configuration of the axoneme is typical of kinocilia and unlike the arrangement previously described for ciliary photoreceptors. Elaboration of receptor organelle membrane surface area is accomplished uniquely by multiple cilia oriented so that large portions of each cilium lie perpendicular to the direction of incident light. The pigmented spot directly contacts the underlying equatorial nerve ring which also connects with the major larval locomotor organ. The pigmented spots of B. neritina are the only potential photoreceptor structures which have been studied by electron microscopy in the three lophophorate phyla. The use of ciliary membranes as the potential photoreceptor organelle allies the bryozoan pigmented spot with the ciliary type photoreceptor which occurs most prevalently in deuterostome animals.

Mycoplasma-Like Organisms: Occurrence with the Larvae and Adults of a Marine Bryozoan

Larvae and adults of the marine bryozoan Watersipora cucullata invariably possess numerous extracellular mycoplasma-like, organisms. Mesodermally encapsulated groups of these atypical bacteria occur in the visceral coeloms of all colony members. In contrast, thousands of the symbionts are externally attached to each larva along a unique superficial groove; the microorganisms are internalized during the complex metamorphosis, thus inoculating the incipient colony. The consequences to the bryozoan of this association are not known.

Larval morphology of the bryozoan Watersipora arcuata (Cheilostomata: Ascophora)

The larva of the ascophoran cheilstome Watersipora arcuata is described on the basis of serial 1‐μ sections, light microscopy of whole mounts, and scanning electron microscopy. Using lightly osmicated specimens, it was possible to map almost every cell on the larval surface. Limited observations on hatching and larval behavior are provided in conjunction with the anatomical description. Tissues of the larva are partitioned between those that function exclusively during the larval period and are degraded at metamorphosis as transitory tissues and those that will have postmetamorphic fates in formation of the ancestrula. Significantly, W. arcuata has two possible anlagen for the ancestrular polypide, the infracoronal cells in the oral hemisphere and the epidermal blastemal cells in the aboral hemisphere, rather than only one or the other of these as reported in other species. Also detailed are the supracoronal flange and groove, which are unique to this genus and are involved in the transmission of mycoplasma‐like organisms between successive generations of adults; two pairs of complex pigment cup ocelli; multiple intercoronal cells that are presumed to have varied sensory and mechanical functions; and the sensory, adhesive, and locomotory components of the pyriform organ. The larval anatomy of W. arcuata is compared with that of the larvae of the ctenostomes Alcyonidium gelatinosum (coronate), Bowerbankia imbricata (coronate), B. gracilis (coronate), and Flustrellidra hispida (shelled lecithotrophic) and of the cheilostomes Bugula neritina (coronate), Electra pilosa (cyphonautes), and Membranipora membranacea (cyphonautes). This study is the first detailed analysis of the larval structure of any ascophoran bryozoan and provides a necessary platform for subsequent analyses of embryology and metamorphosis.

Intercoronal cell complex of larvae of the bryozon Watersipora arcuata (cheilostomata: Ascophora)

The coronate larva of the ascophoran bryozoan Watersipora arcuata has a ring of 32 large, multiciliated coronal cells that are used for swimming. Fourteen pairs of small cells are intercalated between the lateral margins of adjacent coronal cells. These intercoronal cells are arranged in a precise pattern and are polymorphic: seven pairs have multiple cilia and seven pairs are mono‐ or oligociliated. Three pairs of multiciliated intercoronal cells have their cilia arranged as a whorl that is recessed in a pocket formed between the adjacent coronal cells, and they are thought to be photoreceptors that sense general light intensity. Two other pairs of multiciliated cells with cohesive tufts of cilia may be chemo‐ or mechanoreceptors. Roles of the other intercoronal cells in this species are not evident, but it is proposed that the majority, if not all, of them are sensory. The close proximity of all the intercoronal cells to the equatorial nerve ring is compatible with this interpretation. Analyses of the literature on cleavage patterns, pigment cup ocelli, and flagellar tufts that serve as balancers in coronate larvae lead us to propose that (1) an intercoronal cell is the sensory element of most, if not all, pigment cup ocelli of bryozoan larvae; and (2) intercoronal cells are not modified coronal cells but probably are specialized supra‐ and/or infracoronal ones that have migrated to an intercoronal position.