An ultrastructural approach to feeding behaviour in Philodina roseola and Brachionus calyciflorus (rotifers) II. The oesophagus

The cuticular oesophagus is a simple expansion of the dorsal pharyngeal wall of the mastax. The ciliary oesophagus is the cellular anterior wall of the stomach lumen, but seems to have the same embryological origin as the pharynx.In Brachionus calyciflorus, its cilia are surrounded by cuticular velums which have the same myelin-like structure and the same function as the buccal velum of Philodina roseola. In all cases, the oesophagus prevents the return of food particles from the stomach to the mastax lumen.     

Sensory receptors involved in the feeding behaviour of the rotifer Asplanchna brightwelli

A study of the anterior sensory receptors of male and female Asplanchna brightwelli by scanning electron microscopy reveals some important differences in the region surrounding the mouth. In the male, the ventrolateral sensory bristles, the pseudotrochus, the inner and the outer buccal tufts and the mastax receptors are absent. The oral receptors are reduced. Transmission electron microscopy of these receptors shows that they consist of ciliated sensory cells surrounded by epithelial supporting cells. The distal ends of the cilia of the mastax receptors are modified; the cilia of the other receptors differ only in their length and rootlet structure from the locomotor cilia of the cingulum. A consideration of the feeding behaviour of Asplanchna leads us to suppose that these sensory cilia function in mechanoreception and in chemoreception.     

On the Fine Structure of Rastrognathia macrostoma gen. et sp.n. Placed in Rastrognathiidae fam.n. (Gnathostomulida)

Rastrognathia macrostoma gen. et sp.n. is described as belonging to Rastrognathiidae fam.n. The ultrastructure is described, especially the nervous and digestive systems and the epidermal specializations. In the epidermis the single cilia belong to the choanocyte-like cell type. An epidermal sensory cell with several cilia is described. The intraepithelial nervous system shows an orthogon pattern. Two ventral nerve trunks unite into a buccal ganglion. The jaws are secreted by the buccal epidermis. The endoderm consists of a single cell type. The food consists of bacteria. The mesodermal elements consist of six main longitudinal muscles.     

Combining confocal laser scanning and transmission electron microscopy for revealing the mastax musculature in Bryceella stylata (Milne, 1886) (Rotifera: Monogononta)

The rotiferan jaw apparatus (mastax) is characterized by enormous plasticity and according to morphology and feeding strategy, different mastax types can be distinguished. The cuticular hard parts (trophi) of the mastax are often highly specialized and have both a major taxonomic and phylogenetic relevance. Owing to numerous light and scanning electron microscopic studies, the morphology of the trophi is well known but only few attempts have been made to analyze the morphology and functionality of the mastax as a whole. Particularly, the complex muscular system connecting the individual trophi elements and moving them against each other was disregarded in the past. Therefore, the subject of the present study is a detailed analysis of the mastax musculature of the proalid rotifer Bryceella stylata using a combination of transmission electron and confocal laser scanning microscopic techniques, previously applied for revealing the somatic musculature in rotifers exclusively. Based on ultrathin serial sections and phalloidin-dyed specimens, a total number of six paired and two unpaired individual mastax muscles have been identified for the modified malleate trophi system of B. stylata. Possibly homologous muscles in other, so far investigated rotifer species are discussed as well as functional considerations of the individual mastax muscles and their interaction when moving the trophi elements are suggested.     

Combining confocal laser scanning and transmission electron microscopy for revealing the mastax musculature in Bryceella stylata (Milne, 1886) (Rotifera: Monogononta)

The rotiferan jaw apparatus (mastax) is characterized by enormous plasticity and according to morphology and feeding strategy, different mastax types can be distinguished. The cuticular hard parts (trophi) of the mastax are often highly specialized and have both a major taxonomic and phylogenetic relevance. Owing to numerous light and scanning electron microscopic studies, the morphology of the trophi is well known but only few attempts have been made to analyze the morphology and functionality of the mastax as a whole. Particularly, the complex muscular system connecting the individual trophi elements and moving them against each other was disregarded in the past. Therefore, the subject of the present study is a detailed analysis of the mastax musculature of the proalid rotifer Bryceella stylata using a combination of transmission electron and confocal laser scanning microscopic techniques, previously applied for revealing the somatic musculature in rotifers exclusively. Based on ultrathin serial sections and phalloidin-dyed specimens, a total number of six paired and two unpaired individual mastax muscles have been identified for the modified malleate trophi system of B. stylata. Possibly homologous muscles in other, so far investigated rotifer species are discussed as well as functional considerations of the individual mastax muscles and their interaction when moving the trophi elements are suggested.     

Stomatogenesis during sexual and asexual reproduction in an amicronucleate strain of Paramecium caudatum.

Amicronucleate cells of Paramecium caudatum, whose micronuclei have been artifically removed by micropipetting, are characterized by the appearance of a deciliated area at the posterior part of the buccal opening. These cells form food vacuoles at a slightly lower rate than micronucleate cells. Their mean interfission time is longer than that in micronucleates. The exconjugants of amicronucleate cells can not form food vacuoles and eventually die witout fission, though conjugation proceeds normally in them as well as in their micronucleate mate. The oral apparatus of amicronucleate exconjugants seems to be shallower than that of micronucleates. The membranellar cilia, therefore, can be seen through the buccal overture by scanning electron microscope. The results obtained from the cross of micronucleate and amicronucleate strains and from the induction of autogamy in amicronucleate strains suggest that the micronucleus has a primary role in developing the normal oral apparatus after nuclear reorganization.     

Ultrastructure and function of the mastax in Dicranophorus forcipatus (Rotifera: Monogononta)

Rotifers are characterized by a complex set of cuticularized jaw elements in the pharynx. The fine structure of the jaw elements has been the subject of SEM studies for some time, but only very limited information exists on the ultrastructure of the jaw elements and their function beyond taxonomic considerations. Drawing on SEM and TEM techniques, the present study presents a detailed analysis of the mastax in Dicranophorus forcipatus, a carnivorous monogonont rotifer species from freshwater habitats characterized by an extrusible, grasping jaw apparatus. Based on ultrathin serial sections, the jaw elements are reconstructed and, in total, nine paired and two unpaired muscles identified. Possibly homologous muscles in other rotifer species are discussed and functional considerations of the forcipate mastax are suggested.     

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...     

Mechanics of the feeding action of a cyprinid fish

The pressures in the buccal cavities of Golden orfe (Idus idus) have been recorded as the fish took food from a tube connected to a pressure transducer. Pressures 50–105 cm water below the pressure of the surrounding water were recorded as the buccal cavity expanded to suck the food in. Calculations based on the dimensions of the muscles lead to the conclusion that the levatores hyoidei, sternohyoideus, rectus abdominis, epaxial trunk muscles and obliquus internus probably all had to exert almost their maximum isometric tensions, to produce the largest reductions of pressure. These muscles seem nicely balanced in strength for their functions in feeding.     

Study of architectonics of rotifer musculature by the method of fluorescence with use of confocal microscopy

Musculature of two species of rotifers Testudinella patina (Testudinellidae) and Platyias patulus (Brachiomidae) was studied in confocal laser scanning microscope (CLSM) using fluorescent-labeled phalloidin. It includes cutaneous, visceral, and cutaneus-visceral musculature. The common pattern of structure of the cutaneous musculature is represented by postcoronal circular or transverse muscles and connected with them 2–3 pairs of retractors of the trunk, dorsolateral muscles (17-4), two pairs or bundles of lateral retractors of the corona, circular muscles of the foot, and 10-2 retractors of the foot. Visceral musculature includes muscles of the mastax of both kinds. Spiral-like muscle of cloaca of the T. patina and associated with it V-shaped one as well as strong dorsolateral retractors consisting of 6 longitudinal muscle bundles are typical of Testudinellidae only. Three pairs of cutaneus-visceral muscles bind the musculature of mastax with the body surface in T. patina. Differences in localization and thickness of some elements of musculature of these species are determined by morphological peculiarities of structure of the corona, mastax, and foot, as well as by the rotifer body shape.     

The musculature of Squatinella rostrum (Milne, 1886) (Rotifera: Lepadellidae) as revealed by confocal laser scanning microscopy with additional new data on its trophi and overall morphology

Wilts, E.F., Wulfken, D., Ahlrichs, W.H. and Martínez Arbizu, P. 2012. The musculature of Squatinella rostrum (Milne, 1886) (Rotifera: Lepadellidae) as revealed by confocal laser scanning microscopy with additional new data on its trophi and overall morphology.—Acta Zoologica (Stockholm) 93 : 14–27. The monogonont rotifer Squatinella rostrum was investigated with light, scanning electron and confocal laser scanning microscopy to reveal new morphological data on its inner and outer anatomy. In total, the visualized somatic musculature displays five paired longitudinal muscles (musculi longitudinales I–V) and nine circular muscles (musculi circulares I–IX). Compared to other species, S. rostrum is characterized by the absence of several longitudinal and circular muscles (e.g. musculus longitudinalis capitis, corona sphincter and pars coronalis). A reconstruction of the mastax musculature revealed a total number of seven paired and two unpaired mastax muscles. Possibly homologous somatic and mastax muscles in other, thus far investigated rotifers are discussed. Moreover, we provide a phylogenetic evaluation of the revealed morphological characters and suggest possible autapomorphic characters supporting Squatinella and Lepadellidae. Finally, we refer to some striking similarities in the morphology, ecology and way of movement of Squatinella and Bryceella that may indicate a closer relationship of both taxa.     

Rhinoglena frontalis (Rotifera,Monogononta): a scanning electron microscopic study

Females and males of Rhinoglena frontalis (Monogononta, Epiphanidae) are observed by SEM and their external morphologies are compared. The two sexes differ in size and shape of the body. The female body is fusiform with a short, conical foot, while the male body is more slender and has a rather long foot. The rotatory apparatus (or corona) of both sexes is similar with only minor differences and consists of rows and tufts of cilia arranged around the mouth opening. The corona is made of two paired lobes lateral to the mouth and of a third prominent dorsal lobe, usually called proboscis. The three lobes are lined externally by dense rows of cilia, which constitute the cingulum, used for swimming. The central surface of the proboscis is covered with numerous longitudinal rows of cilia bent towards the mouth. The lateral lobes show, on their central surfaces, two concentric arcs of cirri (made of tightly packed cilia) bent towards the mouth. The similar organization of the rotatory apparatus of both sexes is related to the fact that the male, in this species, is able to feed and has a developed mastax and digestive system. The trophi of both sexes are illustrated and compared.     

Rotatory apparatus in Bdelloids

All Bdelloid Rotifers have the same body plan: elongated body, ciliated apical region (rotatory apparatus or corona), telescopically retractable foot and head with pseudosegments, paired gonads, single dorsal antenna, apical rostrum, ramate mastax. Bdelloids use the rotatory apparatus for both locomotion and collecting food and therefore the shape of the corona and arrangement of the cilia, both related to the animal's life style, probably are important for the fitness of the rotifers. We have analyzed the fine morphology of the corona and the rostrum from species belonging to the four families, Habrotrochidae, Philodinidae, Adinetidae and Philodinavidae, each with its own form of feeding and locomotion. In the rostrum one can distinguish a sensorial area and a ciliated area. The former is common to all bdelloids, while the latter is lacking in the Adinetidae. Three models of corona can be recognized: 1) a simple ciliated field of undifferentiated cilia (Adinetidae), 2) a well developed ciliated field with specialized cilia forming the paired trochi on the disks and the cingulum (Habrotrochidae and Philodinidae), and 3) a ciliated field with a single trochus encompassing rudimentary pedicels and cingulum (Abrochtha). We propose (1) to no longer use Digononta as a taxon, (2) to assign the class rank to Bdelloidea, (3) to distinguish three orders, grouping Philodinidae and Habrotrochidae under a single order and (4) to retain the current families.     

Mechanics of the feeding action of various teleost fishes

Pressure changes have been recorded from the buccal cavities of nine species of teleost as they sucked food into their mouths. Some of the pressures recorded are discussed in relation to the sizes of the muscles responsible. Cinematograph films have been used to estimate how much the volume of the orobranchial chamber increases as food is sucked in. Some peculiarities in the feeding movements of Taurulus, Ictalurus and Anguilla are described and discussed.     

Functional adaptations of the digestive system of the carnivorous mollusc Pleurobranchaea californica MacFarland, 1966

The opisthobranch mollusc Pleurobranchaea californica feeds on whole organisms and the functional morphology of the digestive system reflects this behavior. By a rhythmic behavior involving well-developed extrinsic buccal muscles and hemocoelic fluid, the buccal mass is protracted to the tip of the everted oral tube. Here a series of repeated protractions and retractions of the intrinsic buccal muscles associated with the flat radular ribbon and jaws draws the prey into the buccal cavity and conveys it to the dorsal esophagus, where by peristaltic action it is passed to the expansible crop for storage. Prey entering the buccal cavity is mixed with acid from a large single gland and secretion from the paired salivary glands. Prey is retained in the crop over long periods of time while it is slowly broken down and passed via the stomach into the digestive glands. Special modifications that allow flexibility of the digestive organs include elongated salivary gland ducts with propulsive bulbs, long flexible nerve cords connecting the ganglia, a long, large muscular duct for storage of the acid secretion, large jaws for muscle attachment and grasping the prey, and a broad radular ribbon with many teeth that acts as a conveyor belt to move food. Additional modifications for handling whole prey include a buccal membrane that aids in maintaining hemocoelic fluid pressure, the extensive acid gland for immobilization of prey, and the expansible crop for storage of food.     

CILIARY MEMBRANE DIFFERENTIATIONS IN TETRAHYMENA PYRIFORMIS : Tetrahymena Has Four Types of Cilia

We have examined thin sections and replicas of freeze-fractured cilia of Tetrahymena pyriformis. The ciliary necklace located at the base of all freeze-fractured oral and somatic cilia has been studied in thin sections. Since electron-dense linkers have been found to connect both microtubule doublets and triplets to the ciliary membrane at the level of the necklace, the linkers and the associated necklace seem to be related to the transition region between the doublets and triplets of a cilium. Plaque structures, consisting of small rectangular patches of particles located distal to the ciliary necklace, are found in strain GL, but are absent in other strains examined in this study. In freeze-cleaved material, additional structural differentiations are observed in the distal region of the ciliary membranes of somatic and oral cilia. Somatic cilia contain many randomly distributed particles within their membrane. Oral cilia can be divided into three categories on the basis of the morphology of their freeze-fractured membranes: (a) undifferentiated cilia with very few randomly distributed particles: (b) cilia with particles arranged in parallel longitudinal rows spaced at intervals of 810–1080 Å that are located on one side of the cilium; and (c) cilia with patches of particles arranged in short rows oriented obliquely to the main axis of the cilium. The latter particles, found on one side of the cilium, seem to serve as attachment sites for bristles 375–750 Å long and 100 Å wide which extend into the surrounding medium. The particles with bristles are located at the tips of cilia in the outermost membranelle and may be used to detect food particles and/or to modify currents in the oral region so that food particles are propelled more efficiently into the buccal cavity. Examination of thin-sectioned material indicates that the particles in oral cilia which form the longitudinal rows could be linked to microtubule doublets. Linkage between microtubule doublets and adjacent membrane areas on one side of the cilium could modify the form of ciliary beat by restricting the sliding of the microtubules. It is suggested that membrane-microtubule interactions may form the basis for the various forms of ciliary beat observed in different organisms.     

A comparison of the surface features of Philophthalmus megalurus and Philophthalmus gralli rediae by scanning electron microscopy.

General features observed on the surface of Philophthalmus megalurus and Philophthalmus gralli rediae include 2 rounded ambulatory buds, a tapered tail, a slitlike birth pore, and an oral opening surrounded by uniciliate sensory receptors. The tegument is folded in a ribbed pattern and is covered with small microvilli. The sensory receptors are concentrated on the lip of the oral opening but are less dense inside the buccal cavity. Both the birth pore and the buccal cavity are lined with a honeycomblike surface that may support the structural integrity of these highly extensible orifices. Major differences between the 2 species were noted in the structure of the sensory receptors. In P. megalurus they have flexible cilia that originate from a bulbous tegumental base. Receptors of P. gralli rediae are less dense, appear to have rigid cilia, and lack a bulbous base. In general P. megalurus rediae resemble more closely rediae of the philophthalmid Parorchis acanthus than they do those of P. gralli.     

The role of the frontal ganglion in foregut movements of the moth,Manduca sexta

Two types of rhythmic foregut movements are described in fifth instar larvae of the moth, Manduca sexta. These consist of posteriorly-directed waves of peristalsis which move food toward the midgut, and synchronous constrictions of the esophageal region, which appear to retain food within the crop. We describe these movements and the muscles of the foregut that generate them.The firing patterns of a subset of these muscles, including a constrictor and dilator pair from both the esophageal and buccal regions of the foregut, are described for both types of foregut movement.The motor patterns for the foregut muscles require innervation by the frontal ganglion (FG), which lies anterior to the brain and contains about 35 neurons. Eliminating the ventral nerve cord, leaving the brain and FG intact, did not affect the muscle firing patterns in most cases. Eliminating both the brain and the ventral nerve cord, leaving only the FG to innervate the foregut, generally resulted in an increased period for both gut movements and muscle bursts. This manipulation also produced increases in burst durations for most muscles, and had variable effects on the phasing of muscle activity. Despite these changes, the foregut muscles still maintained a rhythmic firing pattern when innervated by the FG alone.Two nerves exit the FG to innervate the foregut musculature: the anteriorly-projecting frontal nerve, and the posteriorly-directed recurrent nerve. Cutting the frontal nerve immediately and irreversibly stopped all muscle activity in the buccal region, while cutting the recurrent nerve immediately stopped all muscle activity in the pharyngeal and esophageal regions. Recordings from the cut nerves leaving the FG showed that the ganglion was spontaneously active, with rhythmic activity continuing within the nerves. These observations indicate that all of the foregut muscle motoneurons are located within the FG, and the FG in isolation produces a rhythmic firing pattern in the motoneurons. We have identified several motoneurons within the FG, by cobalt backfills and/or simultaneous intracellular recordings and fills from putative motoneurons and their muscles.Abbreviations BC Buccal Constrictor - BC1 buccal constrictor motoneuron 1 - BC2 buccal constrictor motoneuron 2 - BD Buccal Dilator - BD1 buccal dilator motoneuron 1 - EC Esophageal Dilator - EC1 esophageal dilator motoneuron 1 - EC2 esophageal dilator motoneuron 2 - EC3 esophageal dilator motoneuron 3 - ejp excitatory junction potential - FG frontal ganglion - psp postsynaptic potential     

The ultrastructure of the mastax of Filinia longiseta (Flosculariaceae, Rotifera): Informational value of the trophi structure and mastax musculature

The study contributes to the discussion of mastax evolution within Rotifera by giving an insight into the ultrastructure of the mastax in the rotifer species Filinia longiseta (Flosculariacea) and additionally into the bdelloid rotifer species Adineta vaga and Zelinkiella synaptae. The existence of cuticularized jaw elements (trophi) in the mastax, a muscular pharynx, is one of the defining rotiferan characters and the basis on which the monophyletic taxon Gnathifera Ahlrichs 1995a, comprising Rotifera, Gnathostomulida, Micrognathozoa and Acanthocephala, was erected. By means of SEM observations of the trophi and ultrathin serial sections (TEM) of the mastax, the internal and external organization of the jaw elements of F. longiseta is reconstructed. TEM sections of the incus of Filinia demonstrate that the fulcrum and the rami are built up by multitudes of tiny cuticular tubes. While tubular substructures in the rotiferan fulcrum have been described previously, distinct cuticular tubes as a substructure of the ramus have only been described for species belonging to the taxa Seisonidea and Bdelloidea so far ( [Koehler and Hayes, 1969] and [Ahlrichs, 1995b]). By comparing the appearance and arrangement of the cuticular tubes in the rami of F. longiseta to those found in species of Seisonidea and Bdelloidea, a higher degree of resemblance between the structures in F. longiseta and Bdelloidea can be reported. The occurrence of the ramus substructures in species of Seisonidea (Paraseison annulatus and Seison nebaliae) is given consideration to represent an intermediate between the ramus substructure of Bdelloidea/Flosculariacea and Ploima. Additionally, the mastax musculature of F. longiseta, being associated with the trophi, is described: A total of seven muscles are found that directly insert the jaw elements or are indirectly associated with them via muscle-to-muscle connections.     

On the structure and function of the mastax ofBrachionus plicatilis (Rotifera), a scanning electron microscope analysis

Sectioned specimens ofBrachionus plicatilis were examined by scanning electron microscopy (SEM). This technique was used to characterize the out-side appearance as well as the position and arrangement of the hard parts (trophi) in the interior of the mastax. Two strips of muscle stretch dorsally across the width of the mastax. Below these muscle strips, all the other components of the mastax form a continuous structure which is smoothly locked to the outside. Some nerve structures and glands of the mastax are found on the dorsal side, outside these muscle strips. By comparing the shape of parts of the trophi, exposed by sectioning, with the views of preparations of isolated trophi, it was possible to estimate how the trophi are positioned in relation to other components of the mastax and in relation to the animal as a whole. The results are used to complement and modify statements and conclusion from a previous study.