Ultrastructure of gill cilia and ciliary rootlets of Chaetoderma nitidulum Lovén 1844 (Mollusca, Chaetodermomorpha)

Cilia and associated structures on the gill lamellae on the ctenidum of Chaetoderma nitidulum were studied. The gill cilia are very long and have a whip-like narrow portion distally, where only three microtubule doublets continue to the distal tip. In the transition zone between the cilium and the centriolar triplet section of the basal body there is a dense plate, an aggregation of granules and a ciliary necklace with four strands. Further down there is a short cross-striated basal foot and two conical cross-striated ciliary rootlets. The first rootlet is flattened and directed forward. It connects distally with the basal feet of other adjacent cilia. The second rootlet is rounded in cross-section and vertically directed. The epithelial structures of Chaetoderma show similarities with other Mollusca. We found no structural characters that could support the current hypothesis of a close relationship of Xenoturbella to the Mollusca.     

Central nervous system of Chaetoderma japonicum (Caudofoveata, Aplacophora): implications for diversified ganglionic plans in early molluscan evolution

The organization of the central nervous system of an "aplacophoran" mollusc, Chaetoderma japonicum, is described as a means to understand a primitive condition in highly diversified molluscan animals. This histological and immunocytochemical study revealed that C. japonicum still retains a conservative molluscan tetra-neural plan similar to those of neomenioids, polyplacophorans, and tryblidiids. However, the ventral and lateral nerve cords of C. japonicum are obviously ganglionated to various degrees, and the cerebral cord-like ganglia display a lobular structure. The putative chemosensory networks are developed, being composed of sensory cells of the oral shield, eight precerebral ganglia, and eight neuropil compartments that form distinct masses of neurites. In the cerebral cord-like ganglia, three anterior, posterior, and dorsal lobes are distinguished with well-fasciculated tracts in their neuropils. Most neuronal somata are uniform in size, and no small globuli-like cell clusters are found; however, localized serotonin-like immunoreactivity and acetylated tubulin-containing tracts suggest the presence of functional subdivisions. These complicated morphological features may be adaptive structures related to the specialized foraminiferan food in muddy bottoms. Based on a comparative scheme in basal molluscan groups, we characterize an independent evolutionary process for the unique characters of the central nervous systems of chaetoderms.     

Falcidens halanychi, a new species of Chaetodermomorpha (=Caudofoveata) (Mollusca) from the northwest Atlantic Ocean

During a cruise with R/V Oceanus out of Woods Hole organized by the National Science Foundation project WormNet, a new species of chaetodermomorph aplacophoran was found in the northwestern Atlantic on the continental rise, between 1100 and nearly 2900 m. An investigation into the material of older collections revealed that the species had previously been collected on numerous occasions but remained undescribed. We here describe the species under the name Falcidens halanychi sp. nov. The species is compared with F. limifossorides Salvini-Plawen 1992 that has the most similar overall morphology, but occurs in the eastern Pacific at lower slope and abyssal depths between 3700 and 4300 m. We illustrate the radula of limifossorides and the birefringent colours of the sclerites photographically for the first time. We also include two DNA barcodes of F. halanychi (cytochome oxidase I) to aid future molecular identification.     

The cuticle of the Aplacophora and its evolutionary significance in the Mollusca

The cuticle of Proneomenia consists of a mucoid matrix containing calcareous spicules and is secreted by the mantle epithelium at the base of the ventral (pedal) groove and over the general body surface. Histochemical examination shows the matrix to be composed of a glycoprotein complex with high acid mucopolysaccharide and low protein contents in which tanning plays little part in stabilization.
The cuticle of the Aplacophora is tentatively equated with an early mucoid stage in the evolution of the molluscan shell and it is suggested that secretion of additional protein, followed by hardening by quinone-tanning, are necessary further stages before a calcified shell evolves. The aplacophoran cuticle is compared with that of Acanthochitona (Polyplacophora) and, although they are similar in many respects, the latter has in addition a discrete inner cuticular layer whichmay act as a semi-conducting membrane in the deposition of the calcareous plates. The spicules are similar in both groups, each being secreted within a thin cup-like membrane which exhibits somewhat similar properties to the inner cuticle.     

Foraging, feeding and prey remains in middens of juvenile Octopus vulgaris (Mollusca: Cephalopoda)

Juvenile Octopus vulgaris foraged by chemotactile exploration, mainly in crevices and under rocks. They caught small crustaceans and molluscs, including Lima bivalves, and preferred crabs and Lima in a simple laboratory choice test. Octopuses consumed 30% of their prey away from home, and small remains of prey deposited in the midden were selectively removed by waves and currents resulting in an underestimate of the octopus' intake and diet.     

Lethal and sub-lethal effects of UVB on juvenile Biomphalaria glabrata (Mollusca: Pulmonata)

Although Schistosoma mansoni occurs mainly in the tropics, where intense levels of solar radiation are present, the impact of ultraviolet (UV) light on schistosome transmission is not known. The purpose of this study was to investigate potential effects of UVB (290-320nm) on juvenile Biomphalaria glabrata, the snail intermediate host of S. mansoni. Albino and wild-type snails were exposed to doses of UVB from UV-fluorescent lamps, and the following were measured: survival, photoreactivation (light-mediated DNA repair), effects on feeding behavior, and morphological tissue abnormalities. Irradiation with UVB is lethal to B. glabrata in a dose-dependent manner. Exposure to white light subsequent to UVB irradiation enhances survival, probably by photoreactivation. The shell offers some, but not complete, protection. Experiments in which UVB transmittance through the shell was blocked with black nail polish suggest that injury to both exposed (headfoot) and shell-enclosed (mantle and visceral mass) tissues contributes to mortality in lethally irradiated snails. Wild-type (pigmented) snails are less susceptible to lethal effects of UVB than albino snails, and they may be more capable of photoreactivation. UVB exposure inhibits snail feeding behavior, and causes tentacle forks and growths on the headfoot. Thus, UVB may influence the life cycle of S. mansoni by both lethal and sub-lethal damage to the snail intermediate host. However, the ability of snails to photoreactivate may mitigate these effects.     

The surface of Priapulus caudatus (Lamarck, 1816)

A scanning electron microscope study of the surface of Priapulus caudatus is presented. Both adults and larvae have been studied. The pharynx, circumoral region, collar, proboscis, trunk, caudal appendage, and larval lorica are described.     

Ultrastructure of the pericardium in chitons (Mollusca: Polyplacophora), in relation to filtration and contraction mechanisms

Summary The pericardium in Lepidopleurus asellus (Spengler), Tonicella marmorea (Fabricius), T. rubra L., Ischnochiton albus L., and Calleochiton laevis (Montagu), species taxonomically far apart, is described. It consists of a flat, simple epithelium facing the pericardial cavity, a basement membrane, a muscle layer with two types of muscle fibres, nerve processes, glio-interstitial cells, and fibrocytes, embedded in a loose collagen matrix. The epithelium in L. asellus and I. albus have convoluted lateral cell borders, and in L. asellus very long basal cell processes are seen. Type 1 muscle fibres resemble smooth molluscan muscle. Type 2 muscle fibres resemble cardiac muscle fibres in chitons. Nerve processes associated with glio-interstitial cells and cell processes, run free in the matrix. Synapses in type 1 fibres are covered with glio-interstitial cell processes, lacking in type 2 muscle fibres synapses.This work was supported by grants from the Norwegian Research Council for Science and the Humanities     

Dart formation in Helix aspersa (Mollusca,Gastropoda)

Summary Dart formation in Helix aspersa has been investigated by SEM of isolated darts at progressive stages in their development, and by histology of dart sacs at the same times. Dart formation begins at the tip of a tubercle where a small group of epithelial cells secrete an organic material filling a small CaCO₃ cone that is the first mineralized part of the shaft. Subsequent secretory activity by an increasing area of the tubercle epithelium results in an increase in the diameter and anterior lengthening of the shaft. Continued secretion by the tubercle and dart sac epithelium produces the flare and finally the corona. A pattern of deposition is also evident in the fine structure of the mineral. In the shaft and vanes there is an inner layer of spherulitic prismatic structure which is covered by a layer of irregular patches of simple prismatic structure. The outermost layer of the shaft and vanes has a continuous simple prismatic structure. Two layers are present in the flare, an inner granular amorphous layer and an outer spherulitic prismatic layer. The corona consists of a single rarefied prismatic layer. A mechanism of dart formation is suggested that involves two types of organic matrix, calcifying and non-calcifying. Measurements of the calcium content of darts, dart sacs, and collars indicate that the hemolymph is the probable source of calcium for the dart.     

The anatomy of Addisonia (Mollusca,Gastropoda)

Summary The organization of Addisonia lateralis (Requien, 1848) and A. brophyi McLean, 1985 is described. Addisonia species have a thin, asymmetrical, cup-like shell and a very simple shell muscle. Eyes, oral lappets and epipodial tentacles are absent and the right cephalic tentacle is also used as a copulatory organ. Most characteristic is the enormously developed gill which is enlarged into the right subpallial cavity. It is composed of about 30 leaflets with skeletal rods and its epithelia are uniquely arranged. The heart is large and the single auricle is situated anteriorly left. There are two kidneys: the left is small, while the right forms large coelomic cavities and has no connection with the pericardium or the hermaphroditic genital system. Testis and ovary are separate: both have a simple duct proper (vas deferens, oviduct). They are connected to the copulatory organ by an open seminal groove; a small receptaculum is present. The mouth opening is typically triangular, with no jaws or subradular sense organs. Addisonia possesses tuft-like salivary glands, a radula diverticulum and distinct, tubular oesophageal glands. The oesophagus itself is simple. The radula and the posterior alimentary tract are unique; the stomach is completely reduced and the intestine forms a pseudostomach. The streptoneurous, hyoathroid nervous system has pedal cords with three commissures. The visceral loop is also cord-like. A single (left) osphradium is present and the small statocysts have several statocones.The peculiarities and unique combination of primitive and advanced characters in Addisonia reflect a highly enigmatic organization among the Archaeogastropoda. Possible relationships to other archaeogastropod groups are discussed.     

Fertilization in Callochiton castaneus (Mollusca)

A fine-structural study of fertilization in Callochiton castaneus has revealed that the mechanism of sperm penetration into the egg is intermediate between the primitive condition found in members of the order Lepidopleurida and the more derived condition found in the Chitonida. C. castaneus sperm have the long needlelike nuclear filament and reduced acrosome that characterizes all Chitonida, but they have retained several plesiomorphic features such as an unspecialized mid-piece and a lack of flagellar reinforcement. As in some Lepidopleurida but unlike any Chitonida, the egg hull in this species comprises a thick, smooth jelly coat permeated by pores that permit sperm rapid access to the vitelline layer. The jelly coat is delicate and quickly dissolves when a sperm concentrate is used, suggesting that excess acrosomal enzymes may be responsible. Once the sperm have penetrated the vitelline layer, the long nuclear filament bridges the gap to cups in the egg membrane. However, once the fertilization membrane is raised, the perivitelline space exceeds the length of the nuclear filament, preventing other sperm from penetrating the egg. A fertilization cone forms around the nuclear filament of the penetrating sperm, but it does not appear to engulf the body of the sperm. Rather, the nuclear chromatin is injected into the egg as a long thread. The remaining sperm organelles are apparently abandoned on the egg surface. If this is the case, it would be a significant departure from fertilization in other molluscs and many other metazoans, in which sperm organelles, such as centrioles and mitochondria, enter the egg. New sperm and egg characters, as well as significant differences in fertilization, indicate that Callochitonidae are basal to all other members of the order Chitonida and may warrant separation as the sister taxon to the suborders Chitonina and Acanthochitonina.     

Tissue Hemoglobins in Bivalvia (Mollusca)

Distribution and properties of tissue hemoglobins in bivalves are reviewed. The mollusk gills are given particular attention; data on their functions, relative weight, and surface area as well as the content, properties, and functional importance of gill hemoglobin are reviewed.     

Anatomy and affinities of cocculinid limpets (Mollusca, Archaeogastropoda)

The anatomy of seven cocculinid species, representing four genera, is described. Two new genera are erected, Paracocculina and Coccocrater, and a comparative account of all cocculinid species so far anatomically investigated is given. Definitions of six genera, mainly based on anatomical data, are presented. Variation at the species level is also discussed. The closest relatives of the Cocculinidae are found in the Bathysciadiidae, sharing a distinct gill type and similar coelomic conditions. Therefore, both families are united as the Cocculinoidea. The Lepetelloidea (Lepetellidae, Pseudococculinidae, Osteopeltidae, Bathyphytophilidae (?), Cocculinellidae, Addisoniidae and Choristellidae) are more distantly related and are classified, together with the Cocculinoidea, within the suborder Cocculiniformia. The Cocculiniformia are regarded as one of the earliest offshoots at the archaeogastropod level of organization, being secondarily modified in several respects.     

Anatomy and affinities of pseudococculinid limpets (Mollusca, Archaeogastropoda)

The anatomy of 12 pseudococculinid species, representing the two subfamilies and 10 (of 11 known) genera, is described in detail. Three new genera are erected, Yaquinabyssia, Copulabyssia and Amphiplica. Subfamilial and generic systematics are reconsidered and additionally based on anatomical characters. Characters defining subfamilies and genera (Pseudoeoeculininae: Pseudococculina, Notocrater, Tentaoculus, Mcsopelex, Bandabyssia, Kurilabyssia; Caymanabyssiinae: Caymanabyssia, Colotrachelus, Yuquinabyssia, Copulabyssia, Amphiplica) are the modifications in shell and radula, the number and arrangement of gill-leaflets, the size of the pedal gland, the degree of eye-reduction and the modifications of the seminal groove and of the copulatory organ. Separation of the Pseudococculinidae Hiekman, 1983 from the Cocculinidae Dall, 1882 at the superfamilial level is confirmed by great differences in their anatomy. Whereas the Cocculinidae are closely related to the Bathysciadiidae (Cocculinoidea). the Lepctcllidae, Pyropeltidae, Pseudococculinidae,? Bathyphytophilidae, Osteopeltidae, Cocculinellidae, Addisoniidae and Choristellidae form a clearly separated supcrfamily (Lepetelloidea). Both superfamilies compose the Cocculiniformia. a primitive but highly specialized arehaeogastropod suborder.     

Basal chromodorid sperm ultrastructure (Nudibranchia, Gastropoda, Mollusca)

The relationship between three genera considered basal in the Chromodorididae (Cadlina, Tyrinna, Cadlinella) has not yet been resolved by traditional morphological means. Here we examined the sperm ultrastructure of Tyrinna nobilis, Tyrinna evelinae, Cadlina flavomaculata and Cadlina cf. nigrobranchiata, with the expectation of finding phylogenetically informative characters. No Tyrinna or Cadlina species showed sperm similarities to Cadlinella. Both Cadlina species and Tyrinna nobilis (but not T. evelinae) exhibited coarse striations in the acrosomal pedestal. The putative fibers that occurred between the coarse striations of the pedestal are condensed into a layer in Cadlina and Tyrinna, but not in other species that also have coarse striations (Gymnodoris), and may constitute evidence for a close relationship. Tyrinna evelinae possessed fine acrosomal striations, which was shared with other Chromodorididae, Actinocyclidae and the cryptobranchs Rostanga and Aphelodoris. We also examined the sperm ultrastructure of ‘Chromodoris’ ambiguus, an animal which has shown molecular affinities to species of Cadlina, and not Chromodoris. The sperm of ‘C.’ ambiguus did not exhibit the typical Cadlina characteristics, but also showed important differences to other investigated Chromodoris species.     

Origin of the gonad in the juvenile of a solitary ascidian, Ciona intestinalis

In the just-metamorphosed juveniles of Ciona intestinalis, a round mass of tissue debris derived from the resorbed tadpole tail is situated in the broad space enclosed by the peritoneal membrane and the epidermis around the ventral side of the esophagus. In living juveni es, the origin of the gonad rudiment was traced back to the mass of tissue debris. Electron microscopically, the round mass was a clump of irregular-shaped phagocytotic cells engulfing degenerated cell fragments. On the surface of the cell clump, a small number of singly occurring round cells were found and identified as primordial germ cells on the basis of morphological continuity to obvious germ cells in later stages. Presence of nuage around the nucleus characterized the germ cells. In a few days the germ cells assembled to form a solid slender body (gonad rudiment) together with smaller somatic cells. The gonad rudiment left the space around the esophagus, moving into the narrow mesenteric space connecting the stomach and intestine on the fourth day after metamorphosis. It gradually increased in size by proliferation of the germ cells and somatic cells. The solid gonad rudiment changed into an oval vesicle with an eccentrically located cavity on about the seventh day after metamorphosis. The vesicle comprised a thinner wall made of a simple epithelium without germ cells and a thicker wall containing germ cells and somatic cells.     

Ultrastructure of erythrocytes and leucocytes of Priapulus caudatus (de Lamarck) (Priapulida)

The marine priapulid Priapulus caudatus has a voluminous body cavity filled with a blood-like fluid containing erythrocytes and leucocytes (amoebocytes). The hematocrit of animals weighing 0.5–14 gm was 2–10%. The erythrocytes contain a hemerythrin blood pigment. The structure of the coelomocytes was studied by light and electron microscopy. The erythrocytes are nucleated and contain marginal bands, vacuoles and occasionally crystals. The cytoplasm has few organelles. The leucocytes are amoeboid motile cells, the cytoplasm of which contains numerous organelles. The most conspicuous of these are oval particles, probably representing developmental stages of lysosomes. Most of these organelles contain tubules stretching from one pole to another. In the hind part of the animal, certain tissues, primarily the posterior warts contain large numbers of coelomocytes. The histological picture is complicated, showing some resemblance to the lymphoepithelial tissues of vertebrates.