Ultrastructure of the pharynx of Prorhynchus (Platyhelminthes, Lecithoepitheliata)

The pharynx variabilis of Prorhynchus is strongly muscular, with a small pharyngeal fold and a thin surrounding sheath. There is one row of inner longitudinal musclcs, up to six rows of inner circular muscles, many radial muscles, one row of outer circular and one row of outer longitudinal muscles, with no sphincter muscle groups. Three kinds of secretion, produced in a cluster of gland cell bodies posterior to the pharynx, enter the pharynx wall. They travel anteriorly in ducts and two kinds unite in a common duct just prior to discharging into the anterior region of the pharynx lumen. The perikarya of lumen epithelial cells lie within the pharynx musculature and, at the anterior and posterior margins of the pharynx, external to the pharynx. Bundles of ciliated receptors are numerous at the anterior and posterior constrictions. Similarities in the ultrastructure of flame bulbs of Rhabdocoela and Lecithoepitheliata suggest a relationship between these groups. However, the usefulness of pharynx ultrastructure for platyhelminth phylogeny cannot be assessed until complete ultrastructural studies of various groups of Rhabdocoela have been made.     

Musculature of Notholca acuminata (Rotifera: Ploima: Brachionidae) revealed by confocal scanning laser microscopy

Abstract. The body-wall and visceral musculature of Notholca acuminata was visualized using phalloidin-linked fluorescent dye under confocal laser scanning microscopy. The body-wall musculature includes dorsal, lateral, and ventral pairs of longitudinally oriented body retractor muscles, two pairs of head retractors, three pairs of incomplete circular muscles, which are modified into dorso-ventral muscles, and a single pair of dorsolateral muscles. The visceral musculature consists of a complex of thick muscles associated with the mastax, as well as several sets of delicate fibers associated with the corona, stomach, gut, and cloaca, including thin longitudinal gut fibers and viscero-cloacal fibers, never before reported in other species of rotifers. The dorsal, lateral, and ventral retractor muscles and the incomplete circular muscles associated with the body wall appear to be apomorphies for the Rotifera. Muscle-revealing staining shows promise for providing additional information on previously unrecognized complexity in rotifer musculature that will be useful in functional morphology and phylogenetic analyses.     

The formation of syncytia within the visceral musculature of the Drosophila midgut is dependent on duf, sns and mbc.

The visceral musculature of the Drosophila midgut consists of an inner layer of circular and an outer layer of longitudinal muscles. Here, we show that the circular muscles are organised as binucleate syncytia that persist through metamorphosis. At stage 11, prior to the onset of the fusion processes, we detected two classes of myoblasts within the visceral trunk mesoderm. One class expresses the founder-cell marker rP298-LacZ in a one- to two-cells-wide strip along the ventralmost part of the visceral mesoderm, whereas the adjacent two to three cell rows are characterised by the expression of Sticks-and-stones (SNS). During the process of cell fusion at stage 12 SNS expression decreases within the newly formed syncytia that spread out dorsally over the midgut. At both margins of the visceral band several cells remain unfused and continue to express SNS. Additional rP298-LacZ-expressing cells arise from the posterior tip of the mesoderm, migrate anteriorly and eventually fuse with the remaining SNS-expressing cells, generating the longitudinal muscles. Thus, although previous studies proposed a separate primordium for the longitudinal musculature located at the posteriormost part of the mesoderm anlage, our cell lineage analyses as well as our morphological observations reveal that a second population of cells originates from the trunk mesoderm. Mutations of genes that are involved in somatic myoblast fusion, such as sns, dumbfounded (duf) or myoblast city (mbc), also cause severe defects within the visceral musculature. The circular muscles are highly unorganised while the longitudinal muscles are almost absent. Thus the fusion process seems to be essential for a proper visceral myogenesis. Our results provide strong evidence that the founder-cell hypothesis also applies to visceral myogenesis, employing the same genetic components as are used in the somatic myoblast fusion processes.     

Some structural and functional properties of nerves and muscles in the oviduct of the pink bollworm moth,Pectinophora gossypiella (Saund.)

The oviduct of the pink bollworm is innervated by an intrinsic neural network arising from 4 nerves from the terminal ganglion. The nerve tracts in this network often contained elliptical swellings, each with a central nucleus.

A distinct surface topography was evident in the muscular sheath of the common and lateral oviduct. A very thin muscular envelope consisting of an inner band of circular fibers and an outer layer of longitudinal fibers was also found in the ovarioles. Although conventional A and I bands were recognized, the z-disk was composed of an irregular and loose meshwork, suggesting that the visceral muscles of the reproductive tract possess super-contracting properties. Even when the oviduct and the ovarioles were isolated from the central nervous system, an endogenous rhythmic activity was evident.

Two types of spontaneous excitatory postsynaptic potentials were detected in the oviduct. The type most frequently observed had a complex of multiple spikes with a duration of 18–32 msec. The other type had a saw-tooth shape and a duration of 80–160 msec. Spontaneous action potentials with a plateau-type configuration and a duration of 280–320 msec were also observed. After the removal of the terminal ganglion, endogenous electrical activity distinct from the events just described was found in the midand upper common oviduct. Such discharges seem to originate from the intrinsic neural network and had durations similar to those found for neurosecretory cells.     

Anatomy of the planarian Dugesia japonica I. The muscular system revealed by antisera against myosin heavy chains

The planarian Dugesia japonica has two genes encoding myosin heavy chain, DjMHC-A and B (Kobayashi et al., 1998). We produced antibodies specifically recognizing each myosin heavy chain protein using their carboxyl terminal regions expressed in E. coli as antigens. Immunohistochemical analyses of sections and whole-mount specimens revealed the detailed structure and distribution of each type of muscle fiber in the planarian. In general, the MHC-A muscle fibers were distributed beneath the epithelial layers, namely, they were observable in the pharynx, the mouth, the intestine, the eyes and the body wall. In the pharynx, only MHC-A muscle fibers were present. In contrast, the MHC-B muscle fibers were distributed in the mesenchyme as dorso-ventral and transverse muscles, and in the body wall. The body-wall muscles were composed of an outer layer of circular MHC-A muscles and inner longitudinal and intermediate diagonal MHC-B muscle layers. Thus, two types of muscle fibers were distinguished by their distribution in the planarian.     

The adult musculature of two pseudostomid species reveals unique patterns for flatworms (Platyhelminthes,Prolecithophora)

We analyzed the adult musculature of two prolecithophoran species, Cylindrostoma monotrochum (von Graff, 1882) and Monoophorum striatum (von Graff, 1878) using a phalloidin-rhodamine technique. As in all rhabdithophoran flatworms, the body-wall musculature consisted of three muscle layers: on the outer side was a layer of circular muscle fibers and on the inner side was a layer of longitudinal muscle fibers; between them were two different types of diagonally orientated fibers, which is unusual for flatworms. The musculature of the pharynx consisted of a basket-shaped grid of thin longitudinal and circular fibers. Thick anchoring muscle fibers forming a petal-like shape connected the proximal parts of the pharynx with the body-wall musculature. Male genital organs consisted of paired seminal vesicles, a granular vesicle, and an invaginated penis. Peculiar ring-shaped muscles were only found in M. striatum, predominantly in the anterior body part. In the same species, seminal vesicles and penis only had circular musculature, while in C. monotrochum also longitudinal musculature was found in these organs. Female genital organs were only present in M. striatum, where we characterized a vagina interna, and a bursa seminalis. Transverse, crossover, and dorsoventral muscle fibers were lacking in the middle of the body and greatly varied in number and position in both species.     

Correlative morphometric and electrochemical measurements of serotonin content in earthworm muscles

Distribution of serotonin (5-HT) content of nervous fibers in both the somatic and the visceral muscle of Eisenia fetida have been investigated using immunocytochemical staining and voltammetric measurements. The somatic muscles in the body wall are richer innervated with serotoninergic fibers than the visceral ones in the pharynx and gizzard. The relative density of immunopositive fibers in the circular muscle layer of the body wall was found to be 2.73% while in the prostomium it was 1.02%. In the case of the muscle in pharynx 1.12% and in gizzard 1.28% density values were found. Differential Pulse Voltammetric (DPV) measurements with carbon fiber electrodes in the above mentioned muscle layers gave 272.5 nA, 135.0 nA, 122.5 nA, 137.5 nA peak heights, respectively. In the statistical analysis T-test was used at a confidence level of 95% (p<0.05). DPV current peak (i(p)) values reflect clearly the 5-HT concentration differences. Significant correlation was found between the innervation density and the i(p) values recorded in different areas. The i(p) values recorded at different times in different locations are determined by instantaneous serotonin concentration of the living tissue. As far as we know this is the first report using in vivo voltammetry investigating serotonin content in earthworm, E. fetida.     

Phalloidin-rhodamine preparations of Macrostomum hystricinum marinum (Plathelminthes): morphology and postembryonic development of the musculature

Summary A whole-mount fluorescence technique using rhodamine-labeled phalloidin was used to demonstrate for the first time the whole muscle system of a free-living plathelminth, Macrostomum hystricinum marinum. As expected, the body-wall musculature consisted of circular, longitudinal, and diagonal fibers over the trunk. Also distinct were the musculature of the gut and of the mouth and pharynx (circular, longitudinal, and radial). Dorsoventral fibers where restricted in this species to the head and tail regions. Circular muscle fibers in the body wall were often grouped into bands of up to four parallel strands. Surprisingly, diagonal fibers formed two distinct sets, one dorsal and one ventral. Certain diagonal muscle fibers entered the wall of the mouth and were continuous with some longitudinal muscles of the pharynx. Dorsoventral fibers in the rostrum occurred partly in regularly spaced pairs, a fact not known for free-living Plathelminthes. All muscle fibers appeared to be mononucleated. During postembryonic development, the number of circular muscle fibers can be estimated to increase by a factor of 3.5 and that of longitudinal muscles by a factor of 2. Apparently as many as 700–800 circular muscle cells must be added in the region of the gut alone during postembryonic development. Stem cells (neoblasts), identified by TEM in the caudalmost region of the gut, lie along the lateral nerve cords. In the same body region most perikarya of circular muscle cells occurred in a similar position. This suggests that the nucleus-containing part of the cell remains in the position where differentiation starts.     

The stomatogastric nervous system of the house cricket Acheta domesticus L. I. The anatomy of the system and the innervation of the gut

The distribution of the ganglia and nerves of the stomatogastric nervous system and the innervation of the extrinsic and intrinsic muscles are described. Median unpaired frontal and hypocerebral ganglia and paired ingluvial ganglia are present. The anterior pharynx is innervated by branches of the frontal nerve and by the anterior and posterior pharyngeal nerves, originating from the frontal ganglion. The posterior pharyngeal nerves are linked to nerves innervating the posterior part of the pharynx which have their origin in the hypocerebral ganglion, the anterior portion of which has previously been regarded as part of the recurrent nerve. Paired esophageal nerves run the length of the esophagus and crop between the hypocerebral and and ingluvial ganglia, innervating the muscularis by serial side branches. From each ingluvial ganglion runs an ingluvial nerve which innervates the gizzard and a cecal nerve which innervates the midgut and its ceca. At the posterior end of the midgut there is a poorly developed nerve ring. Nerves running posteriorly from this nerve ring link the stomatogastric nervous system with the proctodeal innervation from the terminal abdominal ganglion. Multipolar peripheral neurons are present on the muscularis of the whole of the foregut, rather randomly distributed on the crop and gizzard but forming fairly definite groupings at some points on the pharynx. Though of varied appearance, these cells could not be divided into discrete morphological categories. Peripheral neurons on the midgut are of different and characteristic morphology, though a few cells of the same appearance as those of the foregut occur at the midgut-hindgut boundary. Nerve fibers on the gut almost invariably terminate on the fibers of the muscularis.     

Reconstruction of the musculature of <Emphasis Type="Italic">Magelona</Emphasis> cf. <Emphasis Type="Italic">mirabilis</Emphasis> (Magelonidae) and <Emphasis Type="Italic">Prionospio cirrifera</Emphasis> (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.     

Blown fuse regulates stretching and outgrowth but not myoblast fusion of the circular visceral muscles in Drosophila

Circular visceral muscles of Drosophila are binuclear syncytia arising from fusion of two different kinds of myoblasts: a circular visceral founder cell and one visceral fusion-competent myoblast. In contrast to fusion leading to the somatic body-wall musculature, myoblast fusion for the circular visceral muscles does not result in massive syncytia but instead in syncytia interconnected with multiple cytoplasmic bridges, which differentiate into large web-shaped muscles. Here, we show that these syncytial circular visceral muscles build a gut-enclosing network with the interwoven longitudinal visceral muscles. At the ultrastructural level, during circular visceral myoblast fusion and the first step of somatic myoblast fusion prefusion complexes and electron-dense plaques were not detectable which was surprising as these structures are characteristic for the second step of somatic myoblast fusion. Moreover, we demonstrate that Blown fuse (Blow), a cytoplasmic protein essential for the second step of somatic myoblast fusion, plays a different role in circular visceral myogenesis. Blow is known to be essential for progression beyond the prefusion complex in the somatic mesoderm; however, analysis of blow mutants established that it has a restricted role in stretching and outgrowth of the syncytia in the circular visceral muscles. Furthermore, we also found that in the visceral mesoderm, Blow is expressed in both the fusion-competent myoblasts and circular visceral founders, while expression in the somatic mesoderm is initially restricted to fusion-competent myoblasts. We also demonstrate that different enhancer elements in the first intron of blow are responsible for this distinct expression pattern. Thus, we propose a model for Blow in which this protein is involved in at least two clearly differing processes during Drosophila muscle formation, namely somatic myoblast fusion on the one hand and stretching and outgrowth of circular visceral muscles on the other.     

The musculature of Draculiciteria tessalata (Chaetonotida, Paucitubulatina): implications for the evolution of dorsoventral muscles in Gastrotricha

The muscular system of the marine interstitial gastrotrich Draculiciteria tessalata (Chaetonotida, Paucitubulatina) was analyzed with fluorescent phalloidin. Muscles in circular, longitudinal, helicoidal and dorsoventral orientations were found. Circular muscles were present as discreet rings on the pharynx only. Five pairs of longitudinal muscles were found in dorsal, lateral and ventral positions. One of the two pairs of lateral muscles is newly described for the species. Helicoidal muscles, external to the circular muscles and some longitudinal bands, spiraled around the pharynx and anterior portion of the intestine. Two pairs of segmentally-arranged dorsoventral muscles were also present. Lateral dorsoventral muscles extended from the base of the pharynx to the anterior part of the caudal furca. Medial dorsoventral muscles extended from the pharyngeal-intestinal junction into each ramus of the caudal furca. A hypothesis on the evolution of dorsoventral muscles in D. tessalata is proposed which includes a splitting of circular muscles into separate somatic and splanchnic components with a further displacement of both muscle sets into a dorsoventral orientation.     

Distribution of three types of muscle fiber in the skeletal muscles of Rana temporaria frogs (according to histochemical findings)]

On 15 skeletal muscles with different functional profile (flexors and extensors, trunk muscles and muscles of the head) and different embryonic origin (somatic and visceral), it has been shown that muscles of R. temporaria are mixed. Muscle fibers are divided into three groups which differ in the content of lipids, the activity of succinic dehydrogenase and localization of myofibrils on a cross section. On the basis of the data obtained and those in literature, these fiber groups are qualified as phasic, tonic and transitional (slow phasic) ones. Both phasic and tonic fiber groups are uniform, whereas transitional fibers form a broad specturm. Most of the muscles studied contain all the three types of fibers. In muscles of the extremities (especially in flexors), tonic fibers together with transitional ones usually form more or less compact bundles. The highest content of tonic and transitional fibers was found in muscles which do not directly participate in locomotion. If muscles contain only two types of fibers (phasic and transitional ones), these fibers form a mosaic structure.     

Myoneural junctions and neurosecretory endings on spermathecal muscle fibers of two weevils,Sitophilus granarius and Hypera postica

The fine structure of muscle fibers connecting the two arms of the spermatheca and their innervating axons was studied with the electron microscope. The muscle fibers appear to be a sub-set of skeletal and not visceral muscles. Neurosecretory axons with electron dense granules are adjacent to the muscle fibers in young females O-day post-eclosion but not in the ovipositing adult. The typical nerves form synaptic junctions with muscle fibers at all ages but the nerves are divided into two types based on the morphology of the synaptic vesicles they contain, either spherical or flattened.     

The ovaries and changes in their structural components at the end of vitellogenesis and during vitelline membrane formation in the butterfly, Calpodes

The eight ovarioles of Calpodes ethlius are meroistic and polytrophic with seven nurse cells per follicle. The follicles consisting of oocyte, nurse cells and surrounding follicular cells are connected by interfollicular bridges, whose cells are characterized by bundles of microtubules which appear, with some fine filaments, to terminate on or near to the plasma membrane at hemidesmosomes. The ovariolar sheath consists of tightly knit circular and longitudinal muscles which are heavily tracheated. The ovariolar duct consists of more loosely arranged circular and longitudinal muscles and an inner epithelial layer, also tracheated. The vitelline membrane appears to be secreted largely by the oocyte first as an electron-lucent layer which becomes gradually more electron-dense, probably as a result of addition of material from the follicular cells. Overlapping plate-like structures on the outer surface of the fully formed vitelline membrane may provide waterproofing.     

Functions for Drosophila brachyenteron and forkhead in mesoderm specification and cell signalling.

The visceral musculature of the larval midgut of Drosophila has a lattice-type structure and consists of an inner stratum of circular fibers and an outer stratum of longitudinal fibers. The longitudinal fibers originate from the posterior tip of the mesoderm anlage, which has been termed the caudal visceral mesoderm (CVM). In this study, we investigate the specification of the CVM and particularly the role of the Drosophila Brachyury-homologue brachyenteron. Supported by fork head, brachyenteron mediates the early specification of the CVM along with zinc-finger homeodomain protein-1. This is the first function described for brachyenteron or fork head in the mesoderm of Drosophila. The mode of cooperation resembles the interaction of the Xenopus homologues Xbra and Pintallavis. Another function of brachyenteron is to establish the surface properties of the CVM cells, which are essential for their orderly migration along the trunk-derived visceral mesoderm. During this movement, the CVM cells, under the control of brachyenteron, induce the formation of one muscle/pericardial precursor cell in each parasegment. We propose that the functions of brachyenteron in mesodermal development of Drosophila are comparable to the roles of the vertebrate Brachyury genes during gastrulation.     

Ultrastructural Studies of the Visceral Muscles of Chaetognaths

The visceral musculature of Chaetognaths was studied with special attention given to the digestive apparatus muscle. In the head the digestive apparatus muscle is relatively thick; individual muscles are difficult to distinguish at the anatomical level; in the anterior part of the oesophagus discontinuous bundles and layers of cross-striated fibers are found. As a group however, the oesophageal musculature completely covers the oesophageal epithelium. The prominent muscle layers around the oesophagus probably help to force food into the intestine against the turgor pressure of the trunk cavity which tends to collapse the intestine. Around the intestine the musculature is largely circular and smooth. The intestinal epithelium is ciliated despite its muscular covering. Muscle fibers are not individually innervated. They form myoepithelial structures with various intercellular junction types. In the intestinal muscle the fibers show myoendothelial-like junctions. Sphincters composed of myoepidermal cells surround the anus and the female gonopores. The somatic side of the general cavity is lined with a polymorphic squamous epithelium. Sometimes myoepithelial cells are found, with the occasional presence of extracellular matrix basal to the layer of the squamous epithelium. The ontogenetic relations between the polymorphic epithelium and the composite ‘mesenteries’ remain to be established. We have now some idea about the architecture of the body of Chaetognaths in relation to contractile structures.     

THE ORGANIZATION OF FLIGHT MUSCLE IN AN APHID, MEGOURA VICIAE (HOMOPTERA) : With a Discussion on the Structure of Synchronous and Asynchronous Striated Muscle Fibers

The organization of the indirect flight muscle of an aphid (Hemiptera-Homoptera) is described. The fibers of this muscle contain an extensive though irregularly disposed complement of T system tubules, derived as open invaginations from the cell surface and from the plasma membrane sheaths accompanying the tracheoles within the fiber. The sarcoplasmic reticulum is reduced to small vesicles applied to the T system surfaces, the intermembrane gap being traversed by blocks of electron-opaque material resembling that of septate desmosomes. The form and distribution of the T system and sarcoplasmic reticulum membranes in flight muscles of representatives of the major insect orders is described, and the extreme reduction of the reticulum cisternae in all asynchronous fibers (to which group the aphid flight muscle probably belongs), and the high degree of their development in synchronous fibers is documented and discussed in terms of the contraction physiology of these muscle cells.     

Correlative morphometric and electrochemical measurements of serotonin content in earthworm muscles

Distribution of serotonin (5-HT) content of nervous fibers in both the somatic and the visceral muscle of Eisenia fetida have been investigated using immunocytochemical staining and voltammetric measurements. The somatic muscles in the body wall are richer innervated with serotoninergic fibers than the visceral ones in the pharynx and gizzard. The relative density of immunopositive fibers in the circular muscle layer of the body wall was found to be 2.73% while in the prostomium it was 1.02%. In the case of the muscle in pharynx 1.12% and in gizzard 1.28% density values were found. Differential Pulse Voltammetric (DPV) measurements with carbon fiber electrodes in the above mentioned muscle layers gave 272.5 nA, 135.0 nA, 122.5 nA, 137.5 nA peak heights, respectively. In the statistical analysis T-test was used at a confidence level of 95% (p < 0.05). DPV current peak (i_p) values reflect clearly the 5-HT concentration differences.Significant correlation was found between the innervation density and the i_p values recorded in different areas. The i_p values recorded at different times in different locations are determined by instantaneous serotonin concentration of the living tissue. As far as we know this is the first report using in vivo voltammetry investigating serotonin content in earthworm, E. fetida.     

金丝猴食管和胃连接部的组织学研究

本文研究了金丝猴食管和胃连接处的组织结构。金丝猴的食管粘膜为典型的复民鳞状上皮,食管末端含有粘膜腺,粘膜表面有轻微的角质化。管壁外纵肌层有少量的横纹肌。与胃粘膜的连接均位于胃的贲门部以内,两种上皮的连接是突然的,不存在过渡。 贲门腺为少量的分枝管状腺,短而直,由粘膜细胞构成,对PAS染色呈阳性反应。