Fine Structure of Tentacles,Arms and Associated Coelomic Structures of Cephalodiscus gracilis (Pterobranchia,Hemichordata)

The tentacles of the pterobranch Cephalodiscus, a hemisessile ciliary feeder, originate from the lateral aspects of the arms and are covered by an innervated epithelium, the majority of its cells bearing microvilli. Each side of a tentacle has two rows of ciliated cells and additional glandular cells. The coelomic spaces in the tentacles are lined by cross-striated myoepithelial cells, allowing rapid movements of the tentacles. One, possibly two, blood vessels accompany the coelomic canal. On their outer sides the arms are covered by a simple ciliated epithelium with intra-epithelial nerve fibres; the inner side is covered by vacuolar cells. On both sides different types of exocrine cells occur. The collar canals of the mesocoel are of complicated structure. Ventrally their epithelium is pseudostratified and ciliated; dorsally it is lower and forms a fold with specialized cross-striated myoepithelial cells of the coelomic lining. Arms, tentacles, associated coelomic spaces and the collar canal of the mesocoel are considered to be functionally interrelated. It is assumed that rapid regulation of the pore width is possible and even necessary when the tentacular apparatus is retracted, which presumably leads to an increase of hydrostatic pressure in the coelom.     

Fine Structure of Heart,Pericardium and Glomerular Vessel in Cephalodiscus gracilis M'Intosh, 1882 (Pterobranchia,Hemichordata)

Heart, pericardium and glomerular vessel of Cephalodiscus gracilis have been studied with the electron microscope. The lumen of the heart is lined by a basal lamina and an associated epithelium, composed of myoepithelial cells with well developed thin and thick myofilaments. The heart is located in the pericardial cavity, which is deliminated by the pericardium. The latter is composed of two flat layers of myoepithelia with fused basal laminae. The outer layer of the pericardium is the protocoelomic lining, and the inner layer is the ‘parietal’ pericardial epithelium. The myoepithelium forming the heart wall can be considered to represent the ‘visceral’ pericardial epithelium. The spacious glomerular vessel is lined by a basal lamina, on which typical podocytes rest. These cells indicate that ultrafiltration takes place through the wall of the glomerular vessel. The lumen of the vessel contains fine granular material (presumably precipitated blood proteins), fibrils with a faint cross striation, suggesting that they represent collagen, and stellate cells, which in part line the vessel. Since ultrafiltration requires hydrostatic pressure, it is inferred that the blood flow is from the dorsal region then through the heart and into the glomerular vessel.     

Cephalodiscus reproductive biology (Pterobranchia,Hemichordata)

Sexually mature adults and embryos and larvae of Cephalodiscus nigrescens and C. gracilis were studied by light and electron microscopy. Contrary to claims in the literature, individual coenecial cavities are inhabited by colonies of up to 15 joined zooids and not by single individuals, which is important for the interpretation of the mode of life of the related fossil group the graptolites. Some aspects of the reproductive apparatus and reproduction in Cephalodiscus are reported. The ultrastructure of the spermatozoon is described for the first time. Coelom formation is by schizocoely. The structure of the larva at several developmental stages is illustrated. Not all fertilised eggs are destined to become motile larvae and some develop into zooids omitting the motile stage. The lumen of the oviduct is much larger than previously supposed. Spermatozoa are shed into the cavity of the coenecium. It is proposed that fertilisation takes place within the coenecium. The ultrastructure of the enigmatic black ‘Comma Body’ is described and a reproductive function is proposed. Budding takes place from a base common to several zooids. This base probably also serves as an attachment foot. Large masses of yolk have been discovered within the coelom of some zooids and muscle stalks. It is inconceivable that a colony of Cephalodiscus nigrescens could survive unless it spent most of its life outside the coenecium.     

The habitat and behaviour of Cephalodiscus gracilis (Pterobranchia, Hemichordata) from Bermuda

Cephalodiscus gracilis lives in shallow water around Bermuda. The zooids secrete a transparent coenecium. Several zooids can be attached to a common point. The zooids may be of differing maturity, having from none to five pairs of arms. The mature zooids feed by extending their arms like meridians around a globe with the tentacles of adjacent arms interdigitating to make a spherical filter net. Feeding currents are induced by cilia. The mucus flows along the external surfaces of the arms, around the collar and into the mouth. The rejection current runs on the inside surface of the arms. The rejected material is stored in pellets near the arm tips. It is'flicked'away at intervals.
The larvae are found in densely pigmented stalks attached to the common sucker. The zooids also reproduce by budding.     

Feeding and gut contents in Cephalodiscus nigrescens (Hemichordata, Pterobranchia) from the Weddell Sea

The gut of the pterobranch hemichordate Cephalodiscus nigrescens contains plankton of sizes from less than 1 μm to over 100 μm in diameter. Some of the smaller plankton are clumped together in spherical bolus that is mucus-bound. Most plankton types known from the habitat are respresnted amongst the gut contents.     

An intertidal Rhabdopleura (Hemichordata, Pterobranchia) from Fiji

Rhabdopleura has been discovered living in coral rubble on reefs in Fiji. The habitat is unusual, being the underside of coral boulders in the intertidal zone. Some features of the environment and the fauna associated with the Rhabdopleura are briefly described. The Rhabdopleura zooids exhibit other modes of tube building besides the regular cylindrical horizontal and erect tubes. The colony ramifies through interstices in the dead coral and the zooids can line larger cavities within the coral with coenecial tissue. It is probably these cavities, together with the overall porosity of coral, that retain enough water to keep the zooids alive while the intertidal reef flat is exposed to the air. Within the depths of the coral the black stolons are frequently naked. The species is probably R. normani Allman.     

The prosicular stage of Rhabdopleura (Pterobranchia: Hemichordata)

After settling, the larva of Rhabdopleura surrounds itself with a collagenous dome. Later, the zooid breaks through the wall of the dome and builds the horizontal tube part of the coenecium on to the dome.
The dome is a layered structure, unknown in other parts of the coenecium. whereas the horizontal tube is made up of rings in the classical manner of the adult coenecium. The construction of these two parts is different. The techniques used to reinforce the horizontal tube show a marked similarity to the cortical bandages recently described in the fossil graptolites, and give support to the claim that they are ancestral to Rhabdopleura. There are two sorts of early horizontal tube, one is a straight tube, and the other is longer and coiled. The hole in the dome through which the zooid emerges to build the horizontal tube is probably produced by a chemical boring of the zooid, and supports the hypothesis that the zooids can bore holes in shells and corals.     

Graptolite (Hemichordata,Pterobranchia) preservation and identification in the Cambrian Series 3

Due to inadequate preservation, pterobranchs are often difficult to identify in the fossil record, and a better understanding of preservational modes and diagenetic and metamorphic effects is needed for their recognition. Pterobranch hemichordates are common in Cambrian Stage 5 and younger sedimentary rocks, but are frequently overlooked. Often, pterobranch hemichordate colonies have been considered to be algal remains or hydroids. Re‐examination of Cambrian Burgess Shale algae reveals that the genera Yuknessia and Dalyia can be recognized as putative early representatives of pterobranch hemichordates. Distinct fusellar construction of the individual zooidal tubes and branching of the creeping proximal part of the colonies are found in the morphologically similar rhabdopleurid pterobranch genus Sphenoecium. The erect tubes of Sphenoecium do not branch and can reach a length of several centimetres. The development of the fusellar construction in this taxon shows a highly irregular development of the suture patterns, but a fairly consistent height of the individual fuselli. The taxon is widely distributed in the Cambrian Series 3, but has regularly been identified as a hydroid or an alga. Sphenoecium wheelerensis from the Cambrian Wheeler Shale of Utah is described as new.     

Fine Structure of the Hepatic Sacculations of Glossobalanus minutus (Enteropneusta,Hemichordata)

Abstract The hepatic region of Glossobalanus minutus is characterized by deep foldings of the dorsal side of the gut epithelium which affect the neighbouring tissues and structures: coelomic spaces, musculature and epidermis. The following cell types of the gut epithelium are described: vacuolated cells, undifferentiated cells, two types of mucous cells and two types of granular secretory cells. The nature and function of the different cell types are discussed. Data on the general ciliation and subepithelial nerve plexus of the gut epithelium are also given, with special mention of a possible neuroendocrine secretion towards the subjacent blood spaces. A well-developed blood sinus (gut sinus) lies between the gut and the visceral peritoneum. The ultrastructural features of the gut epithelium and its close association with the blood sinus point to an absorptive function. The coelomic cavity is reduced to a narrow space limited by two peritoneal sheets (visceral and parietal) of myoepithelial nature. Amoebocyte-like cells (coelomocytes) occur free in the coelomic fluid, and muscular, unicellular bridges are attached to both peritoneal walls across the coelomic space. The dorsal epidermis follows the gut foldings and is formed by flat, overlapping cells. The present observations are compared with previous histological, histochemical and ultrastructural data.     

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.     

Erect tube growth in Rhabdopleura compacta (Hemichordata: Pterobranchia) from off Start Point, Devon

Patterns of tube construction in the upright tubes of Rhabdopleura compacta are described. Tube building is seen to be a highly regular process, with growth extending over more than one season. Participation in the building of any one tube can involve multiple generations of zooids. Spatial awareness in a zooid adding new material to a pre-existing tube can be demonstrated. This shows that the construction of the tube is strictly determined, either by environmental or genetic mechanisms, rather than being a function of developing zooid morphology, as previously suggested, or random processes.     

Fine structure of eyespots in tornarian larvae (Phylum: Hemichordata)

Summary The eyespots of tornariae of enteropneusts (Ptychodera flava from Hawaiian waters and an unknown species from southern California) were studied by electron microscopy. An ocellus is composed of two types of cells: sensory and supportive. The former is characterized by a bulbous cilium (with 9+2 axoneme) at its distal end, one or sometimes two arrays of microvilli from its sides below the cilium, and a basal axon. The latter features large, clear vesicles which presumably contained the reddish-orange pigment seen in the ocellus of a living larva. Five-day old tornariae of P. flava are positively phototactic. Both cilium and microvilli may function as photoreceptors. The tornarian ocellus studied is compared with eyespots of other invertebrates, and the evolutionary significance of its putative photoreceptors is discussed.We acknowledge the kind assistance of Drs. Michael G. Hadfield, University of Hawaii, and Russel L. Zimmer, Santa Catalina Marine Biological Laboratory, and the support of grant 10292 from the USPHS.     

Structure and Ultrastructure of the Endodermal Region of the Alimentary Tract in the Freshwater Shrimp Neocaridina heteropoda (Crustacea,Malacostraca)

The freshwater shrimp Neocaridina heteropoda (Crustacea, Malacostraca, Decapoda) originates from Asia and is one of the species that is widely available all over the world because it is the most popular shrimp that is bred in aquaria. The structure and the ultrastructure of the midgut have been described using X-ray microtomography, transmission electron microscopy, light and fluorescence microscopes. The endodermal region of the alimentary system in N. heteropoda consists of an intestine and a hepatopancreas. No differences were observed in the structure and ultrastructure of males and females of the shrimp that were examined. The intestine is a tube-shaped organ and the hepatopancreas is composed of two large diverticles that are divided into the blind-end tubules. Hepatopancreatic tubules have three distinct zones – proximal, medial and distal. Among the epithelial cells of the intestine, two types of cells were distinguished – D and E-cells, while three types of cells were observed in the epithelium of the hepatopancreas – F, B and E-cells. Our studies showed that the regionalization in the activity of cells occurs along the length of the hepatopancreatic tubules. The role and ultrastructure of all types of epithelial cells are discussed, with the special emphasis on the function of the E-cells, which are the midgut regenerative cells. Additionally, we present the first report on the existence of an intercellular junction that is connected with the E-cells of Crustacea.     

Structure of the digestive tract of tornaria larva inEnteropneusta (Hemichordata)

The ultrastructure of the digestive tract of tornaria larva of enteropneusts was investigated. It showed that the digestive tract consists of three parts: esophagus, stomach, and intestine. The esophagus epithelium consists of two types of multiciliated epithelial cells and solitary muscle cells. Axonal tracts and neurons were found in the ventral wall of the esophagus. The cardiac sphincter contains an anterior band of strongly ciliated cells and a posterior band of cells with long vacuolized processes which partition the sphincter lumen. The stomach consists of three cell types: (1) cells with electron-opaque cytoplasm, bearing a fringed border on their apical sides; (2, 3) sparse cells with electron-light cytoplasm and different patterns of apical microvilli. Cells of the pyloric sphincter bear numerous cilia and almost no microvilli. The intestine consists of three parts. The anterior part is formed of multiciliated cells which bear the fringed border. The middle part consists of flattened cells bearing rare cilia and vast numbers of mace-like microvilli. The posterior part of the intestine is formed of cells bearing numerous cilia and few microvilli. Muscle cells were not found in either stomach or intestine epithelium. One noticed that the structure of the digestive tract of enteropneust tornaria larva differs from that of echinoid pluteus larva.     

The central and peripheral nervous system of <Emphasis Type="Italic">Cephalodiscus gracilis</Emphasis> (Pterobranchia,Deuterostomia)

Nervous systems are important in assessing interphyletic phylogenies because they are conservative and complex. Regarding nervous system evolution within deuterostomes, two contrasting hypotheses are currently discussed. One that argues in favor of a concentrated, structured, central nervous system in the last common ancestor of deuterostomes (LCAD); the other reconstructing a decentralized nerve net as the nervous system of the LCAD. Here, we present a morphological analysis of the nervous system of the pterobranch deuterostome Cephalodiscus gracilis Harmer, 1905 based on transmission electron microscopy, confocal laser scanning microscopy, immunohistochemistry, and computer-assisted 3D reconstructions based on complete serial histological sections. The entire nervous system constitutes a basiepidermal plexus. The prominent dorsal brain at the base of the mesosomal tentacles contains an anterior concentration of serotonergic neurons and a posterior net of neurites. Predominant neurite directions differ between brain regions and synapses are present, indicating that the brain constitutes a centralized portion of the nervous system. Main structures of the peripheral nervous system are the paired branchial nerves, tentacle nerves, and the ventral stalk nerve. Serotonergic neurites are scattered throughout the epidermis and are present as concentrations along the anterior border of the branchial nerves. Serotonergic neurons line each tentacle and project into the brain. We argue that the presence of a centralized brain in C. gracilis supports the hypothesis that a nerve center was present in the LCAD. Moreover, based on positional and structural similarity, we suggest that the branchial nerves in C. gracilis could be homologous to branchial nerves in craniates, a hypothesis that should be further investigated.     

Fine Structure of the Optic Tract in the Atlantic Hagfish,Myxine glutinosa

The fine structure of the the optic tract in Myxine glutinosa was studied in three animals. The nerve fibers are unmyelinated and occur either singly or in groups surrounded by glial cytoplasm. 810 and 1467 nerve fibers were counted in two tracts. In the zone where the optic tract leaves the eye cup, a cell type is described and suggested to correspond to ganglion cells. Presynaptic terminals containing agranular vesicles were seen adjacent to this cell type and to other processes in the area. The results are discussed in view of previous reports.     

On the Fine Structure of the Nervous System of Tubiluchus philippinensis (Tubiluchidae,Priapulida)

At the mouth tube/introvert border a circumenteric intraepithelial nerve ring occupies a circular ridge protruding into the body cavity. The ring has a centrally located neuropile nearly free of perikarya and two zones of different perikarya above and below the neuropile. Presumably non-neuronal perikarya have an oval nucleus, large heterochromatin clumps and marked filament bundles. Such elements resemble tanycytic glial cells. Two types of presumably neuronal perikarya contain small cytoplasmic granules, similar to those in nerve fibre profiles. One of these neurons has a pale nucleus with a prominent nucleolus, the other a rather inconspicuous nucleus similar to that of the tanycytic cells. The neuronal processes of the fibre ring differ in diameter and contain clear and dense core vesicles, small granules (high or medium electron density) or granules with a dense periphery and a light centre. Sometimes neighbouring processes seem interconnected by electrical synapses. Images suggesting chemical synapses are rare. A large intraepithelial nerve lies in the wall of the introvert and ventral body wall close to the musculature, possibly innervated by this nerve. Frontal of the anus lies an intraepithelial ganglion demonstrating again a central neuropile. two neuronal types and tanycytic elements with filament bundles. Comparative aspects of the characters of the Tubiluchus nervous system are also discussed.