Mechanisms of rapid morphogenetic movements in the metamorphosis of the bryozoan Bugula neritina (cheilostomata,cellularioidea). I. Attachment to the substratum

The larval morphology, settlement behavior, and the rapid morphogenetic movements that occur during the first 60 sec of metamorphosis of the cellularioid cheilostome bryozoan Bugula neritina have been examined and analyzed by light and electron microscopy. The larva attaches to the substratum at the onset of metamorphosis by the eversion of the internal sac. At the same time, the coronal cilia reverse their direction of beat, spreading an adhesive secreted by the neck region of the everting sac over the metamorphosing larva. During attachment, the larva goes through several configurations that coincide with the sequential contraction and relaxation of certain larval muscles. Histological and ultrastructural evidence indicates that the neck and wall regions of the internal sac are everted by the contraction of the muscles in the equatorial plane of the larva at the same time that the roof region in pulled toward the larval equator by the contraction of the axial muscles. The subsequent relaxation of the axial muscles allows the roof region to be everted by the antagonistic force generated by the sustained contraction of the equatorial musculature. After the roof region attaches to the substratum, the apical disc is temporarily retracted by a second contraction of the axial muscles. The apical disc subsequently reextends as the axial muscles relax just before coronal involution. A comparison of the ontogenetic sequence of rapid morphogenetic movements in the metamorphoses of cheilostome and ctenostome bryozoans indicates that cellularioid cheilostomes have undergone peramorphosis in the aspect of development.     

The larval morphology of the marine bryozoan Bowerbankia gracilis (Ctenostomata: Vesicularioidea)

Summary The larval morphology of the marine bryozoan Bowerbankia gracilis has been investigated by light and electron microscopy. The barrel-shaped larva (200 m long and 150 m in diameter) is light yellow without any apparent eyespots, although it is positively phototactic during its brief free-swimming existence. The primary morphological characteristics of the larva are: (1) a large corona that forms most of the larval surface, (2) a small apical disc without blastemas, (3) a deep pallial sinus lined by an extensive pallial epithelium, (4) an internal sac without regional specializations, and (5) a polypide rudiment in the oral hemisphere. This organization is characteristic of larvae of the ctenostome superfamily Vesicularioidea, and differs radically from the organization of all other bryozoan larvae examined. The major morphological differences occur in the size and organization of the apical disc, the pallial epithelium, and the internal sac. In most bryozoans, these regions of the larval epithelium represent rudiments of the polypide and the body wall epidermis of the ancestrula. The oral polypide rudiment, the extensive pallial epithelium, and the reduced internal sac in vesicularioid larvae indicate that their pattern of metamorphosis also differs radically from the metamorphoses of other bryozoans.Figure Abbreviations AB aboral - acr axial ciliary rootlet - ad apical disc - anc aboral nerve cord - ANT anterior - arm apical retractor muscle - b basal body - bf basal foot process - c corona - cc ciliated cleft - ce centriole - ci cilium - cl cupiform layer of the polypide rudiment - cp ciliary pit - cr ciliary rootlet - enr equatorial neural ring - g glandular cells of the pyriform organ - gl glycocalyx - go Golgi complex - gr granule - hcr horizontal ciliary rootlet - ic intercoronal cell - igf inferior glandular field - ip infrapallial cells - is internal sac - jp juxtapapillary cells - l lipid droplets - L lateral - m mesenchyme - m Type I mesenchyme cell - m Type II mesenchyme cell - m Type III mesenchyme cell - mb median band of the polypide rudiment - mc marginal cells of the apical disc - mi mitochondria - mr microridge - mv microvilli - nn nerve nodule - np neural plate - nu nucleus - O oral - oce oral ciliated epithelium - op opening to the internal sac - ovc oral vesicular collarette - p papilla of the pyriform organ - pa pallial cell - pe pallial epithelium - po pyriform organ - POS posterior - pp parasagittal patches of undifferentiated cells - pr polypide rudiment - rer rough endoplasmic reticulum - sc supracoronal cells - sg secretory granules - sgf superior glandular field - sp suprapallial cells - tc terminal cone - tf transitional filaments - u undifferentiated cells - va vacuole - vc vesicular cell - wc wedge-shaped cells of the apical disc - y yolk granule - za zonula adhaerens Caption Abbreviations Gp Glutaraldehyde-phosphate - Os Osmium     

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. Two points of larval anatomy are of special significance to proper interpretation of the metamorphosis:

• 1 Two cytologically similar blastemal tissues, each laden with free ribosomes, occur as parts of the apical organ complex. The upper blastema directly contacts the larval surface, forming the non-ciliated rows of the apical organ. The lower blastema is internal and is oral to and contiguous with the upper blastema.
• 2 The epidermal tissues of the larva are joined in the following sequence, beginning at the aboral pole: a. apical organ complex; b. apical-connecting cell; c. infolded pallial sinus epithelium; d. vesicular-connecting cell; e. aboral vesicular epithelium; f. corona; g. oral vesicular epithelium; and i., j., and k. internal sac neck, wall and roof regions.

The initial stages of metamorphosis involve a complex sequence of morphogenetic movements, including:

• 1 eversion of the internal sac, permanently attaching the larva to the substrate;
• 2 inrolling of the aboral vesicular epithelium, corona, oral vesicular and ciliated epithelia, and neck region of the internal sac into the larval interior; concomitantly the pallial sinus epithelium evaginates;
• 3 loss of connection between the invaginated tissues and the surface;
• 4 fusion of the pallial sinus epithelium with the wall region of the internal sac, maintaining the integrity of the body surface;
• 5 retraction of the apical organ complex and invagination of the pallial sinus epithelium with the simultaneous elevation of the internal sac wall region to the aboral pole.

At the conclusion of these events the preancestrular surface is covered by the wall and roof regions of the internal sac. Cells of the wall region form the epidermis of the body wall except for the attachment disc and secrete a cuticular exoskeleton that is secondarily calcified; the attachment disc is formed by the roof region of the internal sac. Internally, the ectodermal upper blastema differentiates into the lophophore and digestive tract of the ancestrular polypide, while the lower blastema forms the lining of the lophophoral coelom and the splanchnic (but not the somatic) lining of the visceral coelom. The visceral somatic peritoneum is formed from cells that may originate from the mesodermally derived pigmented cells of the larva to which they are similar in pigmentation and cytology. Such a composite derivation of a coelomic lining has not been described previously.     

Mechanisms of rapid morphogenetic movements in the metamorphosis of the bryozoan Bugula neritina (Cheilostomata,Cellularioidea): II. The role of dynamic assemblages of microfilaments in the pallial epithelium

The rapid morphogenetic movements that internalize the transitory larval epithelium and reorient the presumptive adult epidermis during the metamorphosis of the cellularioid cheilostome bryozoan, Bugula neritina, have been examined by light and electron microscopy and analyzed by experimentation with cytochalasin B (CB) and MgC1₂. The pallial epithelium is gradually drawn out over the aboral hemisphere as the larval ciliated epithelium (the corona and the pyriform organ) involutes. At the end of coronal involution the oral margin of the pallial epithelium constricts and the aboral hemisphere is pulled down against the everted sac. Ultrastructural and experimental evidence indicates that an equatorial contractile ring composed of a temporal alignment of CB-sensitive 5.5 nm microfilaments is responsible for the constriction of the oral margin of the pallial epithelium. This morphogenetic movement, in conjunction with the compression of the aboral hemisphere, juxtaposes the pallial epithelium with the oral epithelium of the everted sac. The pallial epithelium adheres to the neck and wall regions of the everted sac and begins a progressive contraction at its aboral margin, pulling the wall epithelium up over the aboral hemisphere. Ultrastructural examination reveals that the pallial cells contain apical bands of microfilaments and associated vesicles at this stage of metamorphosis. The position and time of appearance of the microfilaments in the pallial epithelium support the hypothesis that they generate the force for wall elevation. Histological and experimental data indicate that the compression of the aboral hemisphere at the umbrella stage and the final retraction of the apical disc are muscle-mediated morphogenetic movements. The constriction of the umbrellar margin and the elevation of the wall epithelium, on the other hand, appear to be caused by two distinct populations of microfilaments that assemble in different regions of the pallial epithelium at specific times during metamorphosis.     

Larval Attachment by Eversion of the Internal Sac in the Marine Bryozoan Bowerbankia gracilis (Ctenostomata: Vesicularioidea): A Muscle-Mediated Morphogenetic Movement

The larval morphology and settlement of the vesicularioid ctenostome bryozoan Bowerbankia gracilis has been investigated by light and electron microscopy in an attempt to elucidate the mechanism of attachment to the substratum at the onset of metamorphosis. The oral epithelium in the free-swimming larva is infolded to form a glandular internal sac at the oral pole. The internal sac is not specialized into distinct regions, but consists of a uniform, simple columnar epithelium filled with secretory granules. The hemispherical internal sac is underlain by a cup-shaped layer of undifferentiated cells that constitutes the polypide rudiment. The cupiform layer of undifferentiated cells is in turn embraced by a network of muscle fibers called the rete muscularis. At the onset of metamorphosis, the larva constricts oro-laterally and the internal sac is everted against the substratum. As the sac everts, the glandular cells secrete an adhesive that is wafted up over the metamorphosing larva by the reversed beating of the coronal cilia. At the same time, the cupiform layer of undifferentiated cells flattens in the plane of the oro-lateral constriction and doubles in thickness. The cells of the cupiform layer undergo a corresponding transformation from short columnar cells to flask-shaped cells that bulge into the glandular cells of the internal sac. The narrow ends of the flask-shaped cells abut the strongly contracted muscle fibers of the rete muscularis. It is hypothesized that the contraction of the muscle fibers of the rete muscularis is responsible for the change in shape of the undifferentiated cells and, consequently, for the eversion of the internal sac. On the basis of this study and a review of the literature, it is concluded that attachment to the substratum at the onset of metamorphosis typically is effected by the eversion of an internal sac in larvae of the ctenostome superfamily Vesicularioidea.     

Invasive cell behavior during Drosophila imaginal disc eversion is mediated by the JNK signaling cascade

Drosophila imaginal discs are monolayered epithelial invaginations that grow during larval stages and evert at metamorphosis to assemble the adult exoskeleton. They consist of columnar cells, forming the imaginal epithelium, as well as squamous cells, which constitute the peripodial epithelium and stalk (PS). Here, we uncover a new morphogenetic/cellular mechanism for disc eversion. We show that imaginal discs evert by apposing their peripodial side to the larval epidermis and through the invasion of the larval epidermis by PS cells, which undergo a pseudo-epithelial-mesenchymal transition (PEMT). As a consequence, the PS/larval bilayer is perforated and the imaginal epithelia protrude, a process reminiscent of other developmental events, such as epithelial perforation in chordates. When eversion is completed, PS cells localize to the leading front, heading disc expansion. We found that the JNK pathway is necessary for PS/larval cells apposition, the PEMT, and the motile activity of leading front cells.     

Fine structure of the doliolaria larva of the feather star Florometra serratissima (Echinodermata: Crinoidea), with special emphasis on the nervous system

The epidermis of the doliolaria larva of the Florometra serratissima is differentiated into distinct structures including an apical organ, adhesive pit, ganglion, ciliary bands, nerve plexus, and vestibular invagination. All these structures possess unique cell-types, suggesting that they are functionally specialized in the larva, except the vestibular invagination that becomes the postmetamorphic stomodeum. The epidermis also contains yellow cells, amoeboid-like cells, and secretory cells. The enteric sac, hydrocoel, axocoel, and somatocoels have differentiated but are probably not functional in the doliolaria stage. Mesenchymal cells, around the enteric sac and coeloms, appear to be actively secreting the endoskeleton and connective tissue fibers. The nervous system is composed of a nerve plexus, ganglion, and sensory receptor cells in the apical organ. The apical organ is a larval specialization of the anterior end; the ganglion is located in the base of the epidermis at the anterior dorsal end of the larva. The nerve plexus underlies most of the epidermis, although it is more prominent in the anterior region. Here, processes from sensory receptor cells of the apical organ, as well as those from nerve cells, contribute to the plexus. These processes contain one or a combination of organelles including vesicles, vacuoles, microtubules, and mitochondria. The configuration of glyoxylic acid-induced fluorescence, revealing catecholamine activity, correlates to the apical organ, nerve cells, and nerve plexus. Morphological evidence suggests that the nervous system may function in initiation and control of settlement, attachment, and metamorphosis. The crinoid larval nervous system is discussed and compared to that found in other larval echinoderms.     

Metamorphosis of cinctoblastula larvae (Homoscleromorpha, porifera)

The metamorphosis of the cinctoblastula of Homoscleromorpha is studied in five species belonging to three genera. The different steps of metamorphosis are similar in all species. The metamorphosis occurs by the invagination and involution of either the anterior epithelium or the posterior epithelium of the larva. During metamorphosis, morphogenetic polymorphism was observed, which has an individual character and does not depend on either external or species specific factors. In the rhagon, the development of the aquiferous system occurs only by epithelial morphogenesis and subsequent differentiation of cells. Mesohylar cells derive from flagellated cells after ingression. The formation of pinacoderm and choanoderm occurs by the differentiation of the larval flagellated epithelium. This is possibly due to the conservation of cell junctions in the external surface of the larval flagellated cells and of the basement membrane in their internal surface. The main difference in homoscleromorph metamorphosis compared with Demospongiae is the persistence of the flagellated epithelium throughout this process and even in the adult since exo- and endopinacoderm remain flagellated. The antero-posterior axis of the larva corresponds to the baso-apical axis of the adult in Homoscleromorpha.     

Changes of the lingual epithelium in Ambystoma mexicanum

Changes in the lingual epithelium during ontogenesis and after induced metamorphosis in Ambystoma mexicanum are described as observed by light microscopy and scanning electron microscopy. The epithelium of the tongue is always multilayered in the larva as well as in the adult. It consists of a stratum germinativum with little differentiated basal cells and a stratum superficiale (superficial layer) with specialized superficial cells and goblet cells. Usually, there are more than two layers because of a stratum intermedium consisting of replacement cells. The apical cell membrane of the superficial cells is perforated by fine pores. Its most typical feature are microridges. Maturing superficial cells possess microvilli. Goblet cells occur in early larvae primarily in the centre of the tongue. They spread throughout the dorsal face of the tongue as their numbers increase during ontogenesis. The small apices of the goblet cells are intercalated in the wedges between the superficial cells. Leydig cells are not found on the larval tongue but on that of adults. Due to metamorphosis, the epithelium of the tongue changes. It is furrowed in its anterior part. The furrows house the openings of the lingual glands. The surface is further modulated by ridges which are densely coated by microvilli and which bear the taste buds. The villi of the tongue which lack extrusion pores show cilia and microvilli but lack microridges. The Leydig cells disappear during metamorphosis. In addition to the two types of goblet cells found in different regions of the glandular tubules, goblet cells occur in the caudal part. They secrete directly into the cavity of the mouth. The posterior part is characterised by a dense coat of cilia.     

Organization of the Nervous System and Sensory Organs in the Larva of the Marine Bryozoan Bowerbankia gracilis (Ctenostomata: Vesiculariidae): Functional Significance of the Apical Disc and Pyriform Organ

The organization of the nervous system and the histology and ultrastructure of the apical disc and the pyriform organ have been investigated by serial sections with light and electron microscopy for the larva of the vesiculariid ctenostome bryozoan Bowerbankia gracilis Leidy 1855. The nervous system consists of four major internal components: (1) a median-anterior nerve nodule; (2) an equatorial, subcoronal nerve ring; (3) paired aboral nerve cords; (4) paired antero-lateral nerve tracts. The nervous system is associated with the ciliated larval surface at the apical disc, the pyriform organ, the corona and the intercoronal cells. The paired aboral nerve cords extend from the apical disc to the nerve nodule, which gives rise to the paired antero-lateral nerve tracts to the pyriform organ and to paired lateral tracts that form the equatorial nerve ring. Ultrastructural evidence is provided for the designation of primary sensory cells in the neural plate of the apical disc and in the juxtapapillary regions of the pyriform organ. Efferent synapses are described between the equatorial nerve ring and the overlying coronal cells, which constitute the primary locomotory organ of the larva. The repertoire of potential functions of the apical disc and pyriform organ are discussed. It is concluded that the apical disc and pyriform organ constitute larval sensory organs involved in orientation and substrate selection, respectively. Their association with the major effector organs of the larva (the corona and the musculature) via the nervous system supports this interpretation.     

Neuromuscular structure of the larva to early ancestrula stages of the cyclostome bryozoan Crisia eburnea

For the first time, the development of a cyclostome bryozoan has been studied with immunochemistry and confocal laser scanning microscopy, with emphasis on nerves and muscles. The larva is covered by multiciliated cells, which are latitudinally strongly elongated and show phalloidin-stained cell junctions. We hypothesize that these cells contract at metamorphosis and squeeze the apical invagination and the adhesive sac out. Ectodermal, longitudinal muscle cells extend from the cells of the inner, conical cuticularized part of the apical invagination to the lower part of the corona, around the adhesive sac pore. These muscles are retained in the ancestrula. Scattered monociliated nerve cells are interspersed between the coronal ciliary cells. An equatorial nerve in the larva disappears at metamorphosis. The central, conical part of the cuticle becomes the terminal membrane of the ancestrula, and the underlying ectodermal and mesodermal cell layers differentiate into the polypide bud, forming a deep narrow invagination, differentiating into vestibule–atrium, mouth ring and pharynx–stomach–rectum. Tentacles develop from the ring of cells around the mouth, and a small ganglion with four nerves innervating each of the tentacles develops at the anal side of the mouth. These new findings yield further support for previous homology statements of bryozoan larvae and development.     

Larval development and metamorphosis in the marine pulmonate Amphibola crenata (Mollusca: Pulmonata)

The development of the free-swimming veliger of Amphibola is followed from hatching to settlement, and the larval structures compared with those of post-metamorphic juveniles and adult snails. Observations of living specimens and light-microscope sections were combined with scanning electron microscopy to build up a composite picture of veliger structure.
Four stages in the development of veligers are recognized, each being characterized by the appearance of organ systems such as the mantle cavity, larval heart, adult heart and kidney, and larval pallial gland. At or after metamorphosis, the larval systems (heart, kidney and pallial gland) disappear, and the developing adult organs move to the positions characteristic of adult snails.
Organogenesis in Amphibola veligers is compared with that of prosobranch and opisthobranch larvae, and with that of pulmonate larvae with direct development. The closest similarity is seen to be with opisthobranch veligers.     

HOMOLOGY OF THE PALLIAL AND PULMONARY CAVITY OF GASTROPODS

The development and morphology of the pallial and pulmonarycavities of various gastropods has been investigated using epoxy-resinserial sections. In the veliger larvae of Cellana sandwicensis(Patellogastro-poda), Gibbula adansonii (Vetigastropoda), Modulustectum (Caenogastropoda) and Ovatella myosotis (Pulmonata) theformation of the pallial cavity is nearly identical. After shellformation a shallow dorsal pallial groove develops beneath themantle edge. During the late veliger stage, the ectoderm formsa deep invagination along the bottom of the pallial groove onthe right side of the larva, giving rise to the pallial cavity.In the ellobiid O. myosotis the pallial cavity becomes the lung(=pulmonary cavity), without any major post-metamorphic modification.Thus, the lung of this species is clearly homologous with thepallial cavity of prosobranchs. The lung of pulmonates withveliger development, as well as of fresh water basommatophoransand stylommatophorans, can be shown to be homologous by comparisonof adult morphology. In contrast to previous views, the pulmonatelung should be regarded as truly homologous with the pallialcavity of prosobranchs and opisthobranchs. In the onchidiidpulmonate Onchidium cf. branchiferum, the larval pallial cavityshifts posteriorly after metamorphosis, where it gives riseto a lung and a cloaca. Contrary to previous interpretations,it can be shown that the onchidiid lung is homologous with atleast part of the pallial cavity. Smeagol climoi has only asmall pallial cavity and no separate lung. The previously described‘lung’ is shown to be a gland. The re-evaluationof the development and morphology of the pulmonate lung hasimportant systematic implications: (1) The pulmonary cavitydoes not represent a synapomorphic character of pulmonates.(2) The gymnomorphs cannot be separated from the remaining pulmonatesbased on lung development. (3) The lack of a lung in the smeagolidsmight give reason to reconsider this group's systematic placementwithin the pulmonates.     

Demonstration of the granular layer and the fate of the hyaline layer during the development of a sea urchin (Lytechinus variegatus)

Summary Employing electron-microscopic methods that help retain polyanionic materials, we describe the extracellular coverings of a sea urchin (Lytechinus variegatus) throughout ontogeny. The surface of the embryo is covered by a two-layered cuticle (commonly called the hyaline layer), which in turn is covered by a granular layer. The granular layer is retained after addition of alcian blue to the fixative solutions, and has not been previously described for any sea urchin. After hatching, the granular layer disappears, but the hyaline layer continues to cover most of the larval surface until settlement and metamorphosis. A few days before metamorphosis, the hyaline layer lining the vestibular invagination of the competent pluteus larva is replaced by a three-layered cuticle resembling that of the adult sea urchin. The hyaline layer covering the rest of the larva is evidently lost at metamorphosis during the involution of the general epidermis.     

Ultrastructural Observations on the Larva of Loxosoma pectinaricola Franzén (Entoprocta,Loxosomatidae)

The larva of Loxosoma pectinaricola Franzén has been studied using scanning and transmission electron microscopy. The embryo develops surrounded by an egg envelope attached to the brood chamber. The newly released larva measures about 100 μm in length and is characterized by a prominent apical organ, stalked vesicles, paired lateral sense organs and a prototroch. The apical organ consists of at least four cell types: (1, 2) two types of ciliated cells, (3) vacuolated cells and (4) myoepithelial cells. The apical organ and frontal ganglion are tightly juxtaposed in the upper tier of the episphere. The stalked vesicles each consisting of two cells are unique evaginations of the epidermis. There are about twenty stalked vesicles with a maximum diameter of about 20.0 μm. The ciliated, knob-shaped, paired lateral sense organs are situated fronto-laterally on the episphere. The prototroch is comprised of a row of contiguous prototroch cells each containing about eighteen long cilia. The apical organ, frontal ganglion and paired lateral sense organs are suggested to be sensory structures that play an important role in active locomotion, settlement site selection and metamorphosis.     

Larval morphology of the Nemertean Carcinonemertes epialti (Nemertea: Hoplonemertea)

The morphology of the newly hatched larva of Carcinonemertes epialti Coe has been examined by light and electron microscopy. The newly hatched larva is covered with cilia and measures about 110 μm in length. Four types of epidermal cells are recognizable: (1) Multiciliated cells, (2) vacuolated cells, (3) mucous cells, and (4) “knob cells”. The knob cells protrude from the posterior end of the larva and contain granules and bundles of microfilaments. The gut is incomplete and is located ventral to the bipartite proboscis. A bilobed brain and two subepidermal ocelli are found in the anterior end of the larva. The anterior and posterior cirri are composed of long, tightly appressed cilia that arise from an invagination of the epidermis at each end of the larva. The anterior cirrus is surrounded by two types of glandular cells. It is proposed that the knob cells have a role in larval attachment, combining the functions of the adhesive cells and anchor cells described in the duo-gland system of turbellarians. The cirri are believed to be larval sensory structures that function in substrate selection. Histological and ultrastructural observations suggest that the larvae of Carcinonemertes are relatively long lived and develop into juveniles without a drastic metamorphosis.     

Ultrastructure of Adult Gonads and Development and Structure of the Larva of Rhabdopleura normani (Hemichordata: Pterobranchia)

Sexually mature adults, embryos and larvae of the pterobranch Rhabdopleura normani from Bermuda were studied with light and electron microscopy. The sexes are separate among the zooids of a colony, but a given colony may contain females and males. In zooids of either sex the single gonad is associated with a large haemal sinus in the trunk sac and is displaced laterally (to the right or to the left). The wall of the gonad is composed of three layers: an outer metasomal peritoneum, an internal lining of germinal epithelium and an intervening genital haemal sinus. The mature gametes lie in the lumen within the gonad. The spermatozoon is characterized by an elongate nucleus, no obvious acrosome, a long mitochondrial filament in a midpiece appendix and a single flagellum with a 9+2 axoneme. Females brood 200 μm eggs and embryos in their distinctive, basally coiled tubes. The yolky eggs undergo radial cleavage and develop into ciliated, lecithotrophic, oblong larvae (400 μm in length) that are characterized by: (1) yellow coloration peppered with black pigment spots; (2) a deep ventral depression; (3) a posterior adhesive organ; (4) an anterior apical sensory organ; (5) an evenly ciliated epitdermis. The ventral depression is not invaginating endoderm, but is instead a glandular epithelium that evidently secretes the larval cocoon and the adult tube. Internally, the peritoneum of the coelomic cavities begins to split from the periphery of a large, central mass of yolky mesenchyme cells. The larva swims using cilia, but also undergoes contractions, evidently powered by the peritoneal cells, which constitute a myoepithelium. The discussion considers pterobranch affinities with other deuterostomes and with lophophorates.