Alexander B. Tzetlin

Pharynx and intestine

The alimentary canal of polychaetes consists of a foregut, midgut, and hindgut. The alimentary canal shows different specializations even in homonomously segmented polychaetes. The foregut gives rise to the buccal cavity, pharnyx and oesophagus, the midgut may be divided into a stomach and the intestine proper. Since polychaetes use a wide spectrum of food sources, structures involved in feeding vary as well and show numerous specializations. In the foregut these specializations may be classified as one of the following types: dorsolateral folds, ventral pharynx, axial muscular pharynx, axial non-muscular proboscis and dorsal pharynx. The latter, typical of oligochaetous Clitellata, occurs rarely in polychaetes. The structure, evolution and phylogenetic importance of these different types are described and discussed. Axial muscular and ventral pharynges may be armed with jaws, sclerotized parts of the pharyngeal cuticle. Terminology, structure, occurrence and development of the jaws are briefly reviewed. Special attention has been paid to the jaws of Eunicida including extinct and extant forms. Conflicting theories about the evolution of the jaws in Eunicida are discussed. The epithelia of the intestine may form a pseudostratified epithelium composed of glandular cells, absorptive cells and ciliated cells or only one cell type having similar functions. A conspicuous feature in the intestine of certain polychaetes is the occurrence of unicellular tubular structures, called enteronephridia. So far these enteronephridia are only known in a few meiofauna species.

Muscular system in polychaetes (Annelida)

The structure of the polychaete muscular system is reviewed. The muscular system comprises the muscles of the body wall, the musculature of the parapodial complex and the muscle system of the dissepiments and mesenteries. Various types of organisation of the longitudinal and circular components of the muscular body wall are distinguished. In Opheliidae, Polygordiidae, Protodrilidae, Spionidae, Oweniidae, Aphroditidae, Acoetidae (=Polyodontidae), Polynoidae, Sigalonidae, Phyllodocidae, Nephtyidae, Pisionidae, and Nerillidae circular muscles are lacking. It is hypothesised that the absence of circular muscles represents the plesiomorphic state in Annelida. This view contradicts the widely accepted idea of an earthworm-like musculature of the body wall comprising an outer layer of circular and an inner layer of longitudinal fibres. A classification of the various types of parapodial muscle construction has been developed. Massive and less manoeuvrable parapodia composed of many components like those of Aphrodita are regarded to represent the plesiomorphic state in recent polychaetes. An analysis of the diversity of the muscular structure supports the hypothesis that the primary mode of life in polychaetes was epibenthic and the parapodial chaetae had a protective function.

Reconstruction of the musculature of Magelona cf. mirabilis (Magelonidae) and Prionospio cirrifera (Spionidae) (Polychaeta, Annelida) by phalloidin labeling and cLSM

Recent investigations have suggested that a lack of circular muscle fibers may be a common situation rather than a rare exception in polychaetes. As part of a comparative survey of polychaete muscle systems, the F-actin musculature subset of Magelona cf. mirabilis and Prionospio cirrifera were labeled with phalloidin and three-dimensionally analyzed and reconstructed by means of cLSM. Obvious similarities are sublongitudinal lateral, circumbuccal, palp retractor, dominating dorsal longitudinal, perpendicular lateral and ventral transverse muscles. Differences between M. cf. mirabilis and P. cirrifera are: (1) two types of prostomial muscles (transversal and longitudinal) in M. cf. mirabilis versus one type (diagonal) in P. cirrifera; (2) one type of palp muscles (longitudinal) in M. cf. mirabilis versus three types (longitudinal, diagonal, circular) in P. cirrifera; (3) five ventral longitudinal muscles (ventromedian, paramedian, ventral) in M. cf. mirabilis versus four (two paramedian, two ventral) in P. cirrifera. Ventral and lateral transverse fibers are present in the thorax, but absent in the abdomen of M. cf. mirabilis. The triangular lumen of the pharynx in M. cf. mirabilis is surrounded by radial muscle fibers; three sets of pharynx diductors attach to its dorsal side. The unique features of P. cirrifera are one pair of brain muscles and segmentally arranged dorsal transverse muscles, the latter located outside the longitudinal muscles. The transverse lateral muscles are restricted to the sides and lie beneath the longitudinal muscles, a pattern described here for the first time. A true, outer layer of circular fibers is absent in both species of Spionida that were investigated.

Ultrastructure of the Body Wall, Body Cavity, Nephridia and Spermatozoa in Four Species of the Chrysopetalidae (Annelida, “Polychaeta”)

Abstract The Chrysopetalidae are a small taxon of polychaetous Annelida with a controversial position in the phylogenetic system. Although they most likely belong to the Phyllodocida, their position is difficult to infer because they share some characters with most taxa of this group. Little information is available about the internal anatomy of the Chrysopetalidae. In order to provide additional data for clarifying the systematic position of the group, we here describe the structure of the body wall, coelomic organization, nephridia, oogenesis, spermatozoa and copulatory organs in four species. The Chrysopetalidae comprise a group of relatively large, epibenthic shallow-water species and a group of small endobenthic forms; two representatives of each group have been chosen, two Chrysopetalum spp. and two Dysponetus spp. All species have comparatively short segments, no circular body wall muscles, a highly developed system of transverse, oblique, dorsoventral and parapodial muscles, metanephridia and a closed circulatory system without a heart. Spermatozoa of the primitive type and gonochorism were observed in both Chrysopetalum species, whereas Dysponetus pygmaeus is a protandric hermaphrodite possessing filiform spermatozoa showing mitochondrial interpolation and male copulatory organs. These features of the reproductive system and a relatively simple organization of the other structures investigated are regarded as apomorphies and adaptations typical of meiobenthic polychaetes. The results of the present investigation are indicative of a fairly basal position of the Chrysopetalidae within the Phyllodocida.

Fine structure of the pharyngeal apparatus of the pelagosphera larva in Phascolosoma agassizii (Sipuncula) and its phylogenetic significance

Sipuncula is a small taxon of worm-like marine organisms of still uncertain phylogenetic position. Sipunculans are characterized by an unsegmented body composed of a trunk into which the anterior part, the introvert, can be withdrawn. The group has been placed at various positions within Metazoa; currently, it is either seen as sister group of a clade comprising Mollusca and Annelida or as sister to each of these. An in-group position in either Mollusca or Annelida has usually been precluded till now due to the lack of so-called annelid or molluscan “key-characters” such as segmentation and chaetae or the radula. In the development of certain taxa the trochophore stage is followed by a planktonic larva, the pelagosphera, which might exhibit phylogenetically important structures. Among these is the buccal organ, which has been considered homologous either to the ventral pharyngeal organ present in many sedentary polychaetes or to the radular apparatus of molluscs. In the present paper, the ventral pharynx of the pelagosphera larva of Phascolosoma agassizii is investigated by transmission electron microscopy. The pharynx comprises dorsolateral ciliary folds, a muscle bulb formed by transverse muscle fibres with large intercellular spaces, and an investing muscle. A tongue-like organ is lacking. These results show great structural correspondences to the ventral pharynx of polychaetes, especially to that of the flabelligerid Diplocirrus longisetosus. In contrast, there are no signs of structural similarities to the corresponding structures of molluscs. Thus evidence increases that Sipuncula are closely related to annelids; moreover, an in-group position of Sipuncula within Annelida, as suggested by recent molecular studies, is not precluded by the present data. Instead these studies find additional support. Hence the lack of segmentation and chitinous chaetae in Sipuncula would be a secondary rather than a primary situation, as has recently been shown for Echiura and Pogonophora.

Fauna associated with detached kelp in different types of subtidal habitats of the White Sea

The fauna, associated with Laminaria and other largebrown macroalgae was studied by using SCUBA anddredging in two different types of underwater habitatsof the White Sea.In shallow water fjords and bays, with a depth of nomore than 30–40 m, detached kelp (mainly Laminaria saccharina, L. digitata and Alaria esculenta) formed large accumulations. One ofthese benthic accumulations, which has existed morethan 20 years, was studied. It covers about2000 m2, and is about 2 m thick. The upper layerof the accumulation of fronds is characterized by highturbulence and is well aerated. The lower layer ischaracterized by anoxic conditions. Mats of sulphurbacteria were not observed, although fronds in themiddle layer were covered by layers of cyanobacteria.About 50 species of macroinvertebrates were found,mainly species that are normally associated withliving kelp, such as the detritivorous species Ophiura robusta and Gammarus oceanicus, and fewspecies that are specific inhabitants of organic-richbiotopes in the White Sea such as Capitellacapitata, Ophryotrocha irinae and Nebaliabipes. It was remarkable that in the shallow waterbasins of the White Sea, the process of decompositionof brown algae in the sublittoral takes place withoutsea urchins, and no other macrofaunal form plays anecological role in the mechanical breakdown of theplant substratum, even not in the large accumulationsof detached kelp.Along the open rocky shoreline, communities associatedwith dead detached kelp were situated at a depth of60–90 m, 40–50 m below the belt of living kelp. Inthis deep zone, no macroinvertebrates typical of thekelp community in the photic zone were found. Duringthe passage from the shoreline to the deeper benthiccommunity, where sea urchins were dominant, all plantdebris became fragmented. These deeper benthiccommunities appeared to be the zone for decompositionof the detached kelp.

Ultrastructure of the body wall and gametogenesis in Cossura cf. longocirrata (Cossuridae Polychaeta)

Summary There is little information available about the internal anatomy of the polychaete family Cossuridae Day, 1963. In the interests of clarifying the systematic position of this group, we describe the structure of the abdominal body wall, coelomic organization, spermatogenesis and oogenesis of Cossura cf. longocirrata Webster and Benedict, 1887. There is a spacious coelomic cavity in the abdominal region unlike the thoracic region of the body, though as in the thorax there is no outer peritoneal lining. Muscle layers are virtually absent in certain areas of the abdomen, explaining the tendency for cossurids to disintegrate very easily. Oogenesis in this species is extraovarian but each oocyte is ensheathed by a few follicle cells. The follicle cells appear to be of the “inactive” type since they show no microvilli of the outer surfaces, and no rER or Golgi bodies. Spermiogenesis occurs with synchronously developing clusters of cells attached to a central cytophore. Mature sperm have a spherical head,...

A New Species of the Spionidae (Polychaeta) with Asexual Reproduction Associated with Sponges

Polydorella smurovi sp.n. is described from an underwater coral bank near the Dachlak Archipelago (Red Sea). This new species has alternating sexual and asexual (paratomy) reproduction, short caruncle, specialised broom‐like setae on segment 5 and capillary setae in the neuropodia of the same segment. Worms inhabited the surface of a red sponge covering a block of a dead coral. The genus Polydorella Augener is re‐established and its diagnosis altered.

Morphology and ultrastructure of the anterior end of Diplocirrus longisetosus Marenzeller, 1890 (Flabelligeridae, Polychaeta, Annelida)

The morphology and ultrastructure of the sedentary polychaete Diplocirrus longisetosus Marenzeller, 1890, collected from the White Sea, were studied using dissection, histological methods, light microscopy, and both scanning and transmission electron microscopy. The prostomium and peristomium carry a pair of palps, eight branchiae, a pair of nuchal organs and two nephridiopores, ciliated folds and the mouth. The prostomium, peristomium and the first chaetigerous segment with all appendages comprise the so-called siphon complex. The mouth leads to a pharyngeal organ that is closed ventrally and composed of a ventral muscle bulb adjoined dorsally by two folds projecting into the pharyngeal lumen. These parts are connected and enveloped by the longitudinal investing muscle. No tongue-like organ is present. The nervous system of the siphonal part comprises the brain, the circum-oesophageal connectives and the ganglia of the peristomium and first chaetigerous segment.

Morphology of the jaw apparatus in 8 species of Patellogastropoda (Mollusca, Gastropoda) with special reference to Testudinalia tesulata (Lottiidae)

The fine structure of the jaw apparatus was studied by scanning electron microscopy in eight species of Patellogastropoda. The jaw apparatus is an unpaired two-layered dorsolateral structure with anterior and posterior wings attached to the odontophore by muscles. The jaw of Testudinalia tesulata (O.F. Müller, 1776) is a derivative of the cuticle typical for the foregut. The tissue forming the jaw is a specialized foregut epithelium (gnathoepithelium), consisting of a special type of cells called gnathoblasts. The jaw grows in areas of the epithelium characterized by high concentration of electron-dense vesicles, ER and long microvilli that penetrate deep into the jaw plate. This indicates that the gnathoblasts take an active part in jaw growth. In most cases, these areas of the gnathoepithelium are highly folded. The main differences between the species studied are form and thickness of the frontal edge of the jaw. These differences do not correlate with the systematic position of the species studied but likely depend more on the feeding mode. The transmission electron microscopy studies yielded new morphological criteria for comparison between various gastropod species and other members of Trochozoa, in particular, Annelida. The jaws of Annelida are cuticular structures formed on the surface of specialized epithelial cells, often also called gnathoblasts. The jaw of Patellogastropoda can be attributed to the first type of annelid jaw formation characterized by an epithelium with long microvilli and continuous growth.