The functional anatomy and histology of the alimentary canal of the maldanid polychaetes Clymenella torquata and Euclymene oerstedi

The alimentary canal of the maldanid polychaetes Clymenella torquata (Leidy), and Euclymene oerstedi (Claparède (= Caesicirrus neglectus Arwidsson, 1911) resembles, in many ways, that of the arenicolids. It is divided into buccal mass, pharynx, oesophagus, stomach and intestine, the three latter regions showing further subdivision. The buccal mass and anterior pharynx together form an eversible proboscis. The pharynx, oesophagus, and greater part of the intestine are ciliated. Simple feeding experiments, and histochemical tests, suggest that the stomach is concerned with the digestion and absorption of proteins, fats and carbohydrates, that the anterior intestine is a digestive and major absorptive region, and that the posterior intestine is a storage region. Waste materials are stored mainly in the wall of the oesophagus. A certain amount of intracellular digestion is carried out in the intestine of Euclymene but not in Clymenella. The difference is attributed to the richer, diatomaceous diet of Clymenella. British individuals of this species, being apparently selective feeders, differ not only from Euclymene but also from American ones, both of which ingest the substratum non-selectively.
The pharynx, oesophagus and rectum are surrounded by plexuses of blood capillaries, while the remaining regions are associated with a blood sinus system which varies in position and form in the different regions, lying deepest in the absorptive intestine. The gut muscle seems to be more concerned with moving the blood forward through the sinus system and into the anterior plexus than with moving the food backward. One region of the stomach musculature is especially concerned with this circulation. Rectal respiration probably occurs.     

Feeding, growth, and fecundity of Abarenicola pacifica in relation to sediment organic concentration

For marine deposit-feeding invertebrates, the distribution of species with different life history strategies has long been known to be correlated with sediment organic concentration. Large populations of opportunistic species are found in sediments with enriched organic concentration, while equilibrium species populate low organic concentration sediments. Differences in both behavioral (e.g. feeding rate) and physiological (e.g. growth rate, reproductive output) adaptations determine the ability of species to establish populations in different environments. By systematically documenting differences in the way these factors vary as sediment organic concentration varies for both opportunistic and equilibrium species, we can better understand the mechanisms underlying this correlation between sediment organic concentration and species distributions. Here, we present the results of experiments examining the interactions among food concentration, feeding rate, growth rate, and reproductive output (measured as egg number and size) for the equilibrium species Abarenicola pacifica. A. pacifica is a large, long-lived, iteroparous, sub-surface deposit-feeding polychaete. Individual worms were reared throughout most of one generation in sediments differing only in the concentration of organic matter. Juveniles (<20 mg AFDW) had higher feeding rates and growth rates in sediments of higher organic concentration throughout the range tested. These results are consistent with the predictions from optimal foraging theory. As worms grew, however, these patterns changed. Once worms reached a mean body size of approximately 50 mg AFDW, feeding rate was greater on sediments of lower organic concentration (although it took worms in the sediments with lower organic concentration longer to reach this size). Differences in growth rates among treatments decreased as worms grew. For worms >100 mg AFDW, growth rates were uniformly low ( approximately 1%/day) on all sediments, but the early advantage obtained by worms in the high organic treatments resulted in much greater body sizes after 200 days. Worms had higher tissue triacylglyceride concentrations and produced more eggs (independent of worm size) as sediment organic concentration increased. We conclude that A. pacifica alters its feeding rate in response to variations in food resources in such a way as to maximize its energy intake and thereby maximize fitness. Future studies should investigate whether opportunistic species (as well as other equilibrium species) also have this ability.     

Neuromuscular control systems in the proboscis of Paranemertes peregrina

Neural systems underlying muscular control in the proboscis of Paranemertes peregrina were investigated through the effects of various neuropharmacological agents on electrical and contractile activity.

Peristalsis in the circular muscle was elicited by catecholaminergic drugs. Each wave was accompanied by a spindle‐shaped burst of electrical activity. The occurrence and rate of peristalsis could be manipulated separately, suggesting the presence of a polyneuronal pathway. Acetylcholine elicited contraction of the longitudinal musculature, with little or no obvious accompanying electrical activity. The circular and longitudinal control systems exhibited reciprocal inhibition, which appeared to occur prior to the postsynaptic neuromuscular junction. Additionally, both systems were excited by serotonin.

Histochemical localisations revealed cholinesterases in the longitudinal musculature. Norepinephrine in the proboscideal nerve cords and serotin in the proboscideal nerve cords and plexus were demonstrated through fluorescence spectrophotometry.     

Homology conundrum among foreguts of caenogastropod molluscs: A view from comparative patterns of development

Evolution of two novel feeding strategies among caenogastropod molluscs, suspension feeding in calyptraeids such as Crepidula fornicata and predatory feeding with a pleurembolic proboscis among neogastropods, may have both involved elongation of the anterior esophagus. Emergence of predatory feeding with a proboscis is particularly significant because it correlates with the rapid adaptive radiation of buccinoidean and muricoidean neogastropods during the Cretaceous. However, the notion that this important evolutionary transition involved elongation of the anterior esophagus to extend down a long proboscis has been disputed by evidence that it may have been the wall of the buccal cavity that elongated. We undertook a comparative study on foregut morphogenesis during larval and metamorphic development in C. fornicata and in three species of neogastropods with a pleurembolic proboscis to examine the hypothesis that the same region of foregut has elongated in all. We approached this by identifying a conserved marker for the boundary between buccal cavity and anterior esophagus, which was recognizable before the developing foregut showed regional differences in length. A survey of four species of littorinimorph caenogastropods suggested that the site of neurogenic placodes for the buccal ganglia could serve as this marker. Results showed that foregut lengthening in C. fornicata involved elongation posterior to neurogenic placodes for buccal ganglia, an area that corresponded to the anterior esophagus in the other littorinimorphs. However, foregut elongation occurred anterior to neurogenic placodes for buccal ganglia in two buccinoidean and one muricoidean neogastropod. The elongated foregut within the pleurembolic proboscis of these neogastropods qualifies as anterior esophagus only if the definition of the anterior esophagus is expanded to include the dorsal folds that run down the roof of the buccal cavity. Regardless of how the anterior esophagus is defined, comparative developmental data do not support the hypothesis of homology between the elongated adult foregut regions in C. fornicata and in neogastropods with a pleurembolic proboscis.     

Feeding strategy of the megamouth shark Megachasma pelagios (Lamniformes: Megachasmidae)

The feeding biology of the planktivorous megamouth shark Megachasma pelagios was investigated. Morphological examination disclosed that the megamouth has a suite of unique characteristics among sharks, such as large mouth, large bucco-pharyngeal cavity, elongate jaw cartilages, long palatoquadrate levator and preorbital muscles, long ethmopalatine ligament and elastic skin around the pharynx. The combination of these characters suggests that the megamouth shark performs engulfment feeding that is typically seen in the rorqual and humpback whales. Engulfment is a new feeding method for sharks, and the detailed mechanism of the engulfment feeding is discussed.     

THE TOXOGLOSSAN PROBOSCIS: STRUCTURE AND FUNCTION

The structure of the proboscis and anterior alimentary systemis described and reviewed in the three families (Conidae, Terebridae,Turridae) of the Conoidea (=Toxoglossa). A number of proboscisforms are shown, characterized by the presence of a permanentrhynchodeum, modified alimentary system and a venom apparatus.Despite the diversity of prey types in the Conidae (polychaetes,molluscs and fish) they have a uniform proboscis structure witha long buccal tube and functional buccal and venom apparati.This homogeneity is not present in the Turridae and the Terebridaewhich show a variety of proboscis structures. Terebrids feedonly on polychaetes or hemichordates but appear to have threeprimary feeding modes based on significant difference in thestructure and function of the buccal tube and associated buccalorgans. The greatest variety of proboscis type is found in thediverse Turridae which feed mainly on polychaetes, but includesipunculans, gastropods and nemerteans in their diet.     

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.     

Laboratory observations on the feeding behaviour, reproduction and morphology of Galeodea echinophora (Gastropoda: Gassidae)

Galeodea echinophora fed on Echinocardium cordatum in an aquarium. Every few days, each G. echinophora emerged from the sand and foraged for 1–3 h. On detecting an Echinocardium , the Galeodea stopped locomotion and attacked the buried prey from the surface of the sand. The proboscis was extended down to the prey and a small area of test was cleared of spines. A disc was cut out of the softened test, leaving a hole of about 2 mm in diameter. All flesh except the gut was removed from the prey, the entire procedure taking 50–180 min.
One Galeodea echinophora laid 120 egg capsules on the side of the aquarium. These were kept at 13^oC and hatched after 112–159 days. Within the capsule, veligers fed on the yolk cells of abortive embryos. Juveniles hatched at the crawling stage, and at first secreted strings of mucus acting as drogues, but after 1 day the juveniles crawled over the substratum. They readily attached themselves to adult Echinocardium cordatum , apparently feeding on the epithelium between the spines or tube feet. Dissection revealed entry of the acinar glands into the proboscis gland ducts, a widening of these ducts as they empty into the buccal cavity and a dorsal fold in the anterior oesophagus. Radular teeth and jaws of Galeodea differ little among species; a taxonomic review of the genus is needed.     

Nemertean predation on the tropical fiddler crab Uca musica

We saw 79 predatory interactions between a new speciesof monostiliferous, suctorial hoplonemertean and thefiddler crabs Uca musica (77 cases) and U.stenodactylus (2 cases). At an intertidal sand barin the Pacific entrance of the Panama Canal, worms ateabout 0.1% of the adult crab population per day. Themode of attack and the spatial and temporaldistributions of interactions suggest the worm is anambush predator. When struck by a worms sticky,mucous-covered proboscis, crabs produced copious foamfrom their buccal area. Mucous-laden crabs thatescaped, again foamed indicating that the foam maycounteract the mucus. If the attack led to a kill,the struggling crab soon became quiescent, as istypical in other nemertean-prey interactions. Theworm inverted its proboscis, found ingress to thecrabs body and fed. Crabs escaped by autotomizingappendages entwined by the proboscis, by forcefullypulling away and by remaining quiescent, then movingaway when the worm inverted its proboscis and beforeit entered the crab. Immobility, a response to visualpredators, may falsely indicate paralysis to the wormand cause it to invert its proboscis, therebyproviding the crab with an opportunity to escape. This predator-prey interaction seems to incorporategeneralized predator tactics and fortuitous preydefenses that give worms and crabs about an evenchance of success.     

The feeding ecology of Axiothella rubrocincta (Johnson) (Polychaeta: Maldanidae)

The feeding ecology of Axiothella rubrocincta (Johnson) from Tomales Bay, California, is described. This worm inhabits a U-shaped tube of agglutinated sand grains and mucus. Morphological adaptations such as nuchal and anal plaques prevent the tube from becoming clogged. Foreign debris entering the tube is either consumed or incorporated into the tube wall. The proboscis and notosetae are probably used to clean the tube wall. A. rubrocincta combines feeding and burrowing activities to form the funnel and complete the tube: it does not ingest sediment while burrowing. Organic matter deposited into the funnel is buried there by sand slides originating at the rim of the funnel. The concentration of organic matter within the funnel is significantly higher than for non-funnel sediments. A. rubrocincta consumes food from the upper 2 cm of the substratum and is 4.6% efficient as a depositfeeder. It also feeds within the funnel and can ingest large quantities of food. This feeding process is described. A. rubrocincta irrigates its tube at a rate of 5.1 ml sea water/g/h while feeding, and briefly reverses this current to a rate of 0.1 ml/g/h when defaecating. The rhythmic activity patterns are integrated: the mean defaecation and inverse pumping time is 14.4 min at 15 ± 1 C.     

Observations on the feeding mechanism, diet and digestive physiology of Histriobdella homari Van Beneden 1858: an aberrant polychaete symbiotic with North American and European lobsters.

1. The aberrant annelid Histriobdella homari (Polychaeta:Eunicida) lives in the branchial chambers of the marine lobsters Homarus americanus and H. vulgaris where it feeds on the rich microflora of bacteria, blue-green algae and related organisms which grow on the inner surface of the branchial chamber, the setae fringing the edges of the carapace, the gill filaments and, especially, the surfaces and setae of the epipodite plates between the gills. H. homari, therefore, is to be regarded as an epizoic microphagous cleaning symbiote of the lobsters. 2. The alimentary canal consists of mouth, buccal cavity, oesophagus, proventriculus, stomach, intestine and anus. The much-modified proboscis lies ventrally below the oesophagus and proventriculus, with its anterior portions protruding into the rear of the buccal cavity. 3. The proboscis consists of two fixed parallel mandibles, a transverse carrier which slides upon the mandibles and to which is attached, posteriorly, a median flexible dorsal rod and, anteriorly, four pairs of movable articulated maxillae, paired external and internal retractor muscles and various tensor, flexor and extensor muscles. 4. Contraction of the retractor muscles withdraws the carrier and maxillae posteriorly, causing bowing of the dorsal rod which is fixed at its posterior end. Relaxation of the muscles allows the rod to straighten and, thus, causes protraction of the carrier and protraction and lateral expansion of the maxillae. Contraction and relaxation of the relaxation of the retractor muscles are supplemented by appropriate changes in the other muscular components of the proboscis. 5. During feeding the serrated anterior ends of the mandibles are applied to the food, the maxillae are fully expanded and then dawn ventro-posteriorly toward the mid-line by contraction of the retractor muscles in the effective movement of the feeding mechanism. This draws the food organisms across the anterior ends of the mandibles, detaching them from the substratum and allowing their ingestion by ciliary action. The first pair of maxillae are also capable of independent action and can be used while the remainder of the proboscis apparatus is held in the protracted position. 6. Detached microorganisms are entangled in a sticky mucous secretion from the salivary glands; other salivary secretions provide a transport medium for the clumped particles and a third set contain C-esterases which initiate digestion. 7. Ingested food is held briefly in the proventriculus, then passed to the stomach where gland cells secrete A- and C-esterases which continue and extend the digestion initiated by the salivary C-esterases. 8. Some soluble products of gastric digestion are taken up by absorptive cells in the stomach wall and their digestion is completed intracellularly by enzymes which include beta-glucuronidase. Others pass into the intestine for absorption and completion of digestion by cells similar to the gastric absorptive cells but which lack beta-glucuronidase...     

The structure of the buccal cavity and pharynx in relation to the method of feeding of Reighardia sternae Diesing 1864 (Pentastomida)

The buccal complex of the haematophagous pentastomid Reighardia sternae has been described. It consists of a buccal capsule containing a prominent oral papilla communicating with a heavily chitinized pharynx which is crescentic in cross section. The chitinous skeleton of the buccal apparatus provides a framework for the attachment of various muscle systems, the contractions of which have been related to the mode of operation of the oral papilla and pharnyx during the feeding process. Particular emphasis was placed on the possible function of the oral papilla in the initial rupture of a blood capillary, and on the pumping action of the pharynx during the ingestion of the liberated blood.     

Ultrastructure of introvert and pharynx of Halicryptus spinulosus (priapulida)

The introvert of Halicryptus spinulosus bears three kinds of sensilla: buccal papillae, ordinary scalids, and dentoscalids. They are all characterized by bipolar monociliary receptor cells. The former two have apical openings at which the sensory cilia are in close contact with the ambient sea water. The pharyngeal teeth are composed of slender epithelial cells the tips of which are devoid of organelles and a thick cuticle. The anatomy of the muscle arrangement of the pharynx is described. Glands occur at the junction of the pharynx and midgut.     

Food and feeding behavior of the hoplonemertean Oerstedia dorsalis

The monostiliferous nemertean Oerstedia dorsalis was collected from eelgrass (Zostera marina) beds located along the coast of New Jersey, and feeding responses to amphipods and isopods were observed in the laboratory. Tests with 46 worms showed that they fed suctorially on Ampelisca vadorum, Ampithoe longimana, Corophium acherusicum and C. tuberculata. Corophiids were preferred. Upon contact with an amphipod, the proboscis is everted and strikes the prey on the ventral side, immobilizing it in a few minutes. The worm probes the sternal region with its head and everts its proboscis one or more times during the process. The exoskeleton is eventually penetrated by the head, and the stomach is everted into the hemocoel as a flattened funnel-like structure. Peristaltic undulations of the body signify the suctorial action that removes the living contents from the exoskeleton. The actual feeding process (from head penetration to removal of the head) takes about 7 min. O. dorsalis is only the third species within the Prosorhochmidae for which the feeding behavior has been documented. The other two are terrestrial species, and are also suctorial.     

The Stylet Apparatus of Monostiliferous Hoplonemerteans

This paper compares the ultrastructure, development, and functionalmorphology of the stylet apparatus in monostiliferous hoplonemerteans(Phylum Nemertea: Class Enopla). The apparatus occurs in themiddle region of the worm's eversible proboscis and consistsof two main components: 1) a central stylet that is attachedto an anchoring device called the basis; and 2) reserve styletsacs that contain nail-like reserve stylets. The knob-shapedproximal piece of the central stylet is embedded in the anteriorend of the basis, and the tapered shaft of the stylet is directedanteriorly. When the proboscis is fully everted, the centralstylet is used to stab such prey as polychaete worms and smallcrustaceans. Wounds inflicted by the stylet allow the introductionof paralytic neurotoxins that are produced by glandular cellsin the epithelial lining of the proboscis. Reservestylets formintracellularly within the reserve stylet sacs and serve toreplace the central stylet when it becomes lost or damaged.Fully developed stylets typically measure 50 to 250 µmin length and comprise an organic core surrounded by an inorganiccortex that contains calcium phosphate. In juvenile worms thatare four days to several weeks old, reserve stylets are assembledbefore other components of the stylet apparatus are fully differentiated.Subsequently, a reserve stylet is transferred to the centralregion of the proboscis and placed on the basis. Stages in theprocess of stylet attachment are described, and the probablesequence by which the stylet apparatus co-evolved with toxin-producingcells is discussed.     

Morphology and evolution of the nervous system in Gnathostomulida (Gnathifera,Spiralia)

Within Spiralia, Gnathifera may represent the deepest branching lineage comprising the jaw worms Gnathostomulida and their sister group Micrognathozoa + Syndermata. Yet, very few nervous system studies have been conducted on this lineage of microscopic, jaw-bearing worms, limiting our understanding of the evolution of this organ system in Spiralia. The nervous system of representatives from all major groups of Gnathostomulida was here mapped using confocal laser scanning microscopy and immunohistochemistry. Their intra-epidermal, unsegmented nervous systems comprise an anterior brain and three to five ventral and two to four dorsal longitudinal nerves, connected by few transverse commissures. Neurites of the stomatogastric nervous system were found lining the pharynx and connecting to a prominent buccal ganglion. Supposedly, sensory ciliated cells in the pharynx and the gut were documented for the first time. Based on these morphological results, primary homologies of neural structures in Gnathostomulida and other Gnathifera were hypothesized and thereafter tested using parsimony. This first neurophylogeny of Gnathostomulida resulted in a topology congruent with molecular data, supporting the monophyly of Bursovaginoidea, Conophoralia, and Scleroperalia. From this topology, the evolution of the gnathostomulid nervous system was reconstructed. It suggests a specialization and diversification of cords and serotonin-like immunoreactive cell patterns from a plesiomorphic neuroarchitecture of three unsegmented nerve cords and a compact anterior brain and buccal ganglion. These plesiomorphic states resemble the nervous system of Micrognathozoa, and possibly the ancestral states of Spiralia.     

Mechanical properties of the peristomial membrane of the cidaroid sea-urchin Stylocidaris affinis

The peristomial membrane (PM) is the area of flexible body wall which surrounds the mouth of regular sea-urchins. In cidaroid sea-urchins, like Stylocidaris. affnis (Phil.), it contains an extensive endoskeleton of overlapping plates. This paper describes the basic mechanical properties of the PM of S. affinis and compares them with those of the PM of the previously studied echinacean euechinoid Paracentrotus lividus. When subjected to vertical deformation, the PM of S. affinis generates J-shaped tension-deformation curves with an inflexion point at a deformation corresponding to about 3% of the perignathic girdle diameter. It is considerably stiffer than the P. lividus PM and, unlike the latter, it shows asymmetrical properties, i.e. greater resistance to aboral flexion (retraction) than to oral flexion (protraction). In interambulacral regions the junction between the PM endoskeleton and test of S. affinis takes the form of a wide ligament-filled gap which acts as a compliant hinge. In marked contrast to the echinacean PM, that of S. affinis provides little evidence for the presence of mutable collagenous tissue (MCT). The cidaroid PM is interpreted as a design which maximizes mechanical defence whilst retaining the capacity for some flexibility. A possible correlation between the lantern systems and feeding habits of sea-urchins is discussed.