Muscles advance the teeth in sand dollars and other sea urchins

We demonstrate the action of the dental promoter muscles in advancing the continuously growing teeth of sand dollars and sea urchins. Teeth wear at the occlusal end, while new calcite is added to the opposite end. Dental ligaments rigidly hold teeth during chewing, but soften and reform during advancement. The source of forces that advance the teeth was unknown until our discovery of the dental promoter muscles. The muscles, which underly the tooth, attach centrally to the stereom of the pyramid of the Aristotle''s lantern (jaw) and peripherally to a membrane that covers the distal end of the tooth. The muscles shorten along an axis nearly parallel to the long axis of the tooth. We stimulated contraction by addition of acetylcholine, with increasing concentrations of acetylcholine generating higher forces. Forces exerted by this muscle are appropriate for its size and are 1000 times lower than forces exerted by interpyramidal muscles that generate chewing forces. In sand dollars, a single muscle contraction of the dental promoter muscle can account for half the mean daily advancement of the teeth.     

Ultrastructural Observations on the Dwarf Male of Bonellia viridis (Echiura)

The organization of the dwarf male of Bonellia viridis was studied by electron microscopy. The epidermis is formed by two types of epithelial cells: the majority are multiciliated cells; highly vacuolated, non-ciliated cells are less abundant. The body wall musculature consists of an outer circular, a diagonal, and a longitudinal layer. As a unique feature in coelomate spiralians it was found that the perikarya of all muscle cells are located internal to the entire contractile muscular layer. The muscles are solitary myocytes embedded in extracellular matrix. Masses of secretory and indifferent cells occur inside the muscles. Two types of secretory cells were distinguished. Both of them apparently undergo holocrine secretion. A complete lining of thin peritoneal cells delimits the body cavity. Also, the gut and sperm sac have a complete peritoneal lining. The coelomic lining of the gut is a single-layered myoepithelium, that of the sperm sac a pseudo-stratified myoepithelium. The vas deferens was seen to be ciliated. The entrance of the sperm sac is formed by a ciliated funnel that leads into the reservoir by means of a thin, ciliated canal. The existence of repeated transverse nerves and of four longitudinal nerve cords is described for the first time.     

The lantern of Aristotle: organization of its coelom and origin of its muscles (Echinodermata,Echinoida)

Summary Comparative ultrastructural analyses of the muscles that work the lantern of Aristotle support the opinion that the muscles in question are myoepithelially organized or derivatives of myoepithelia. They are part of the epithelium of the peripharyngeal cavity (=lantern coelom). The coelom epithelium may become multiplelayered in certain regions and is composed of (1) a layer of muscle cells that vary in number and size, (2) nerve cells and their processes that are interspersed between the muscle layer and (3) monociliated adluminal cells that build a continuous cell lining and completely cover the muscle layer. According to their complexity, the lantern muscles exhibit consecutive stages of myoepithelial variations and may finally simulate subepithelial musculature. The results of this study support the hypothesis of a histological development of subepithelial musculature from simple myoepithelia, although both epithelial and mesenchymal musculature may occur in the Echinodermata. Detailed knowledge of the organization of the lantern's coelom space was a prerequisite for the present study. In contrast to previous examinations the lantern coelom is not a continuous space, but is subdivided into several cavities that are partially completely separated from each other. On the one hand, this subdivision is probably caused by the sophisticated arrangement of the lantern's ossicles and on the other by the septa that give rise to muscles that fulfill different functions. lanter's ossicles and on the other by the septa that give     

The compass depressors ofParacentrotus lividus (Echinodermata,Echinoida): ultrastructural and mechanical aspects of their variable tensility and contractility

Summary The compass depressors are bands of soft tissue which connect the compass ossicles of the echinoid lantern to the inner edge of the test. They are essentially ligaments with on one side a thin layer of muscle cells. The ligamentous component consists mainly of a parallel array of collagen fibrils with interspersed 12 nm microfibrils. The most notable cellular constituents are granule-containing cell bodies and their processes which resemble the juxtaligamental cells that have been found in all echinoderm mutable collagenous tissues and which may control the tensility of these tissues. The muscle cells occupy about 8% of the total cross-sectional area of the compass depressor and are located in a richly innervated pseudostratified myoepithelium. When subjected to constant low loads in creep tests the compass depressor stretches to a fixed length beyond which there is no further extension. The length at this creep limit coincides with the maximum length to which the compass depressor is stretched by natural movements of the intact lantern. Stress-strain tests show that treatment with 1 mM acetylcholine or 100 mM K⁺ ions can increase reversibly the stiffness of the compass depressor to an extent that cannot be due to contraction of the myoepithelium, suggesting that the mechanical properties of the ligament are under physiological control. Tension-length data on the myoepithelium suggest that it generates a maximum active tension when the compass depressor is stretched to the creep limit. The implications of these results for the function of the compass depressors are discussed.     

Ultrastructural study of the proboscis endothelium of Riseriellus occultus (Nemertea, Heteronemertea)

We have examined with transmission electron microscopythe epithelial layer exposed to the rhynchocoel fluidof the proboscis in the heteronemertine Riseriellus occultus. This epithelium is organized asa monociliated, pseudostratified myoepitheliumconsisting of two cell types: apically situatedmonociliated supportive cells and subapical myocyteslacking cilia. The low supportive cells form acontinuous adluminal sheet and reach with numerouscytoplasmic processes into the extracellular matrix;these cells are characterized by numerous, irregularlyshaped, apical folds projecting into the rhynchocoelfluid, delimiting broad extracellular spaces. Theauthors suppose that both apical and basal folds couldaccommodate stretching of the endothelium when theproboscis is everted. The apical folds of thesupportive cells increase the interface of these withthe rhynchocoel fluid; this feature, together with thepresence of pinocytotic vesicles in such cells,suggest that they could be involved in the exchange ofsubstances between the rhynchocoel fluid and theproboscis. The myocytes are scattered singly withinthe monociliated pseudostratified myoepithelium. Theyare situated between the supportive cells and thesubjacent extracellular matrix. Basement membraneseparating both cells types is lacking. Myofibrillarparts protrude basally from the myocyte somata. Themyofibrillar parts lie in direct apposition to theextracellular matrix, and are oriented circular to thelongitudinal axis of the proboscis. We consider themyocytes to be intra-epithelial, myoepithelial cells.     

Mechanical properties of sea-urchin lantern muscles: a comparative investigation of intact muscle groups in Paracentrotus lividus (Lam.) and Stylocidaris affinis (Phil.) (Echinodermata, Echinoidea)

The jaw apparatus, or lantern, of sea-urchins contains five pairs of retractor and protractor muscles which are responsible for lantern displacement. Using intact retractor or protractor groups, the force-length relations of these muscles were compared in two taxonomically distant species, Paracentrotus lividus and Stylocidaris affinis. The total contractile forces generated by the muscles can be resolved into vertical and horizontal components. It was found that the vertical component of the retractors is maximal at a lantern position which is significantly lower (i.e. more protruded) in Paracentrotus than in Stylocidaris. Total forces generated by the retractors were in both species maximal at or above the lantern `resting positions'. In Paracentrotus alone, the total force-displacement curves tended to be bimodal. It is hypothesized that the retractors of Paracentrotus contain two populations of muscle fibres, one adapted for jaw opening and one for lantern retraction. No significant differences in the properties of the protractors of the two species could be identified. The lantern of Paracentrotus is more mobile than that of Stylocidaris and is able to exploit a wider range of food sources. This investigation has shown that the force-length relations of the lantern muscles match their differing working conditions. Accepted: 3 November 1997     

Organization and mechanical behaviour of myocyte–ligament composites in a sea-urchin lantern: the compass depressors of Stylocidaris affinis (Echinodermata, Echinoida)

 Certain components of the jaw apparatus, or lantern, of regular sea-urchins form a ’compass system’ the function of which has still to be established. This system includes ten compass depressors (CDs) which connect the compass ossicles to the inner edge of the test. Previous studies focused on the CDs of euechinoid sea-urchins. This paper provides the first detailed account of the organization and mechanical behaviour of the CDs of a cidaroid sea-urchin, Stylocidaris affinis. The outermost layer of its CDs is a coelothelium comprising apical peritoneocytes, sparse subapical myocytes and two types of granule-containing cells. The central core of the CD is dominated by longitudinally orientated collagen fibrils arranged in bundles surrounded by networks of beaded microfibrils. Myocytes are scattered throughout the core and are always surrounded by a basal lamina which usually encloses the same two types of granule-containing cells observed in the coelothelium. Each CD is attached to the outer coelomic septum by a mesentery, both the connective tissue and coelothelia of which lack myocytes and granule-containing cells. From data on the relationship between the vertical position of the lantern and the passive and active forces developed by intact CD sets, it appears that the CDs resist elevation of the lantern above its ’resting position’ and that they develop a maximal contractile force when their length corresponds to that at the resting position. The connective tissue of the CDs has the capacity to undergo nervously mediated changes in stretch resistance, although this is expressed more weakly than in the euechinoid CDs. These results are related to current views on the physiological role of the compass system. Accepted: 15 March 1998     

A spicule-reinforced contractile mesentery: organisation and mechanical behaviour of the exterior coelomic septum of Stylocidaris affinis (Echinodermata, Echinoida)

The exterior coelomic septum (ECS) is a mesentery-like structure that encloses the lantern of regular sea-urchins and connects it to the inner surface of the test. This paper describes the ultrastructure and microarchitecture of the ECS in Stylocidaris affinis (Cidaridae, Echinoida) and provides information on its contractile and passive mechanical properties. The ECS forms five interambulacral pouches each of which has adthecal (test-facing) and adambulacral (ambulacrum-facing) walls. The ECS wall comprises two coelothelia separated by a layer of connective tissue. The outer coelothelium is a single layer of monociliated cuboidal peritoneocytes and basally located axon-like processes. The inner coelothelium is a single layer of squamous peritoneocytes overlying axon-like processes and, in the adthecal regions only, parallel arrays of elongated myocytes orientated obliquely or horizontally. The intraseptal connective tissue consists mainly of collagen fibrils with sparsely distributed spherule cells and cells containing heterogeneous vesicles. In the adambulacral regions of the ECS hollow beaded microfibrils 20–23 nm in diameter form fibre-like aggregations. This layer also contains calcite spicules of variable size, shape, abundance and orientation. Isolated preparations of the ECS show concentration-dependent contractile responses to K⁺ ions and acetylcholine. The magnitude of the contractile force varies with the vertical position of the lantern (which determines the starting length of the ECS) in an unusual pattern. Cyclical loading-unloading tests indicate that, as the lantern is raised, the ECS shows low stiffness until the lantern reaches its normal resting position. It is concluded that the adthecal regions of the ECS help to set a limit to lantern retraction and that their contractility assists the protractor muscles in exerting a downward pull on the lantern. Accepted: 24 August 2000     

The structure of the proboscis musculature in <Emphasis Type="Italic">Baseodiscus delineatus</Emphasis> (Delle Chiaje, 1825) (Heteronemertea) and the comparative analysis of the proboscis musculature in heteronemerteans

The proboscis musculature was studied in the nemertean Baseodiscus delineatus using confocal laser scanning and electron transmission microscopy. Three muscle layers were differentiated in the proboscis wall: the outer-longitudinal, the diagonal, and the inner-circular layer. The endothelium consists of two cell types: apical supportive cells with rudimentary cilia and subapical myocytes making up the inner-circular musculature of the proboscis. The supportive cells have thin processes attached to the basal extracellular matrix and their perikarya are spread over the apical surfaces of myocytes. The endothelium of B. delineatus is characterized by a folded basal layer of the extracellular matrix and by different heights of myocyte processes, giving an impression that the inner-circular musculature is multilayered. Comparative analysis shows that the diagonal musculature of Baseodiscus is not homologous to that of other heteronemerteans. An assumption is made that the inner-circular muscles have endothelial origin in all heteronemerteans.     

Functional anatomy of the valves in the ambulacral system of sea urchins (Echinodermata,Echinoida)

Summary The water vascular system of sea urchins is examined with special reference to the valves positioned between the radial vessel and the ampullae of the tube feet. The lips of the valve protrude into the ampulla. Thus the valve functions mainly like a check valve that allows the unidirectional flow of fluid towards the ampulla. Each ampulla-tube foot compartment acts as a semi-autonomous hydraulic system. The lumina of the ampulla and the tube foot are lined with myoepithelia except for the interconnecting channels that pierce the ambulacral plate. The contraction of the ampulla results in an increasing hydraulic pressure that protrudes the tube foot, provided that the valve is closed. The retraction of the tube foot results in a backflow of fluid independent of the condition of the valve. The lips of the valve are folds of the hydrocoel epithelium. The pore slit lies in the midline. The perradial faces of the lips are covered with the squamous epithelium of the lateral water vessel. The ampullar faces are specialized parts of the ampulla myoepithelium. Turgescent cells which form incompressible cushions take the place of the support cells. The valve myocytes run parallel to the pore slit and form processes that run along the base of the ampulla and the perradial channel up to the podial retractor muscle. The findings lead to the hypothesis of multiple control of the ampulla-tube foot system: (1) The mutual activity of the ampulla and the tube foot is indirectly controlled by the lateral and podial nerves which release transmitter substances that diffuse through the connective tissue up to the muscle layers. (2) A muscle-to-muscle conduction causes the simultaneous contraction of the ampulla or the podial retractor muscles. (3) The valve muscles are directly controlled by the processes of the valve myocytes which make contact with the podial retractor. In extreme conditions a backflow of hydrocoel fluid towards the radial water vessel occurs.     

Ultrastructure of the subepidermal musculature of Xenoturbella bocki, the adelphotaxon of the Bilateria

 Xenoturbella bocki is the only species of the high-ranked taxon Xenoturbellida. The species lives on marine mud bottoms at a depth of 20–120 m and moves extremely slowly by ciliary gliding. Nevertheless it possesses a well-developed body wall musculature with outer circular muscles, a prominent layer of inner longitudinal muscles and radial muscles that extend from the outer circular myocytes to the musculature surrounding the gastrodermis. The longitudinal myocytes are not compact cells, but form fascicles of fibrils running parallel to each other. Fine cytoplasmic cords connect the fibres of a cell to each other and with its nuclear region. The muscles are embedded within a sometimes expansive extracellular matrix (ECM) that lacks any fibrillar components. All muscle cells display conspicuous and numerous cytoplasmic extensions that are intermingled with each other. Tight coupling between adjacent cell membranes is not found, but zonula adhaerens-like junctions exist. Fibrils belonging to different myocytes, but also fibrils of the same cell, are coupled by such cytoplasmic extensions. Circular, radial and at least the peripheral longitudinal myocytes display cell-matrix connections with the internal lamina, a component of the subepidermal ECM. This internal lamina projects down into the centres of the fascicles with longitudinal muscle fibrils and forms extensive attachment zones with the muscle cells, reminiscent of focal contacts. For the ingestion of food, X. bocki opens the simple mouth pore and protrudes the aciliated gastrodermis. The body wall musculature is responsible for this protrusion and also for the withdrawal of the gastrodermis. In the past, possible phylogenetic kinships with the Acoelomorpha (Plathelminthes) or the Enteropneusta and Holothuroidea were discussed, but, on the basis of all information available, X. bocki is hypothesized to be the sister taxon of the Bilateria. Accepted: 2 April 1997     

Basal lamina directs acetylcholinesterase accumulation at synaptic sites in regenerating muscle

In skeletal muscles that have been damaged in ways which spare the basal lamina sheaths of the muscle fibers, new myofibers develop within the sheaths and neuromuscular junctions form at the original synaptic sites on them. At the regenerated neuromuscular junctions, as at the original ones, the muscle fibers are characterized by junctional folds and accumulations of acetylcholine receptors and acetylcholinesterase (AChE). The formation of junctional folds and the accumulation of acetylcholine receptors is known to be directed by components of the synaptic portion of the myofiber basal lamina. The aim of this study was to determine whether or not the synaptic basal lamina contains molecules that direct the accumulation of AChE. We crushed frog muscles in a way that caused disintegration and phagocytosis of all cells at the neuromuscular junction, and at the same time, we irreversibly blocked AChE activity. New muscle fibers were allowed to regenerate within the basal lamina sheaths of the original muscle fibers but reinnervation of the muscles was deliberately prevented. We then stained for AChE activity and searched the surface of the new muscle fibers for deposits of enzyme they had produced. Despite the absence of innervation, AChE preferentially accumulated at points where the plasma membrane of the new muscle fibers was apposed to the regions of the basal lamina that had occupied the synaptic cleft at the neuromuscular junctions. We therefore conclude that molecules stably attached to the synaptic portion of myofiber basal lamina direct the accumulation of AChE at the original synaptic sites in regenerating muscle. Additional studies revealed that the AChE was solubilized by collagenase and that it remained adherent to basal lamina sheaths after degeneration of the new myofibers, indicating that it had become incorporated into the basal lamina, as at normal neuromuscular junctions.     

Evolution of a novel muscle design in sea urchins (Echinodermata: Echinoidea)

The sea urchin (Echinodermata: Echinoidea) masticatory apparatus, or Aristotle's lantern, is a complex structure composed of numerous hard and soft components. The lantern is powered by various paired and unpaired muscle groups. We describe how one set of these muscles, the lantern protractor muscles, has evolved a specialized morphology. This morphology is characterized by the formation of adaxially-facing lobes perpendicular to the main orientation of the muscle, giving the protractor a frilled aspect in horizontal section. Histological and ultrastructural analyses show that the microstructure of frilled muscles is largely identical to that of conventional, flat muscles. Measurements of muscle dimensions in equally-sized specimens demonstrate that the frilled muscle design, in comparison to that of the flat muscle type, considerably increases muscle volume as well as the muscle's surface directed towards the interradial cavity, a compartment of the peripharyngeal coelom. Scanning electron microscopical observations reveal that the insertions of frilled and flat protractor muscles result in characteristic muscle scars on the stereom, reflecting the shapes of individual muscles. Our comparative study of 49 derived "regular" echinoid species using magnetic resonance imaging (MRI) shows that frilled protractor muscles are found only in taxa belonging to the families Toxopneustidae, Echinometridae, and Strongylocentrotidae. The onset of lobe formation during ontogenesis varies between species of these three families. Because frilled protractor muscles are best observed in situ, the application of a non-invasive imaging technique was crucial for the unequivocal identification of this morphological character on a large scale. Although it is currently possible only to speculate on the functional advantages which the frilled muscle morphology might confer, our study forms the anatomical and evolutionary framework for future analyses of this unusual muscle design among sea urchins.     

Androgen receptor expression in satellite cells of the neonatal levator ani of the rat

Androgens are thought to mediate sexual differentiation of spinal nucleus of the bulbocavernosus (SNB) motoneurons via actions on androgen receptors (ARs) within their target muscles bulbocavernosus and levator ani (LA). However, the cells within these muscles which mediate masculinization of the SNB remain undefined. Until recently, myocytes were thought to be the most likely candidate cell type. However, genetic tests of AR function in myocytes have failed to support a sufficient role for these cells in producing masculine SNB morphology, suggesting the involvement of other cell types. To identify other candidate cell types in the LA, we evaluated whether satellite cells or fibroblasts express AR. Fluorescent immunohistochemistry and confocal microscopy were used to evaluate whether satellite cells and fibroblasts express AR in neonatal male and female rats in the LA and an adjacent sexually monomorphic control muscle (CM). We found that a small proportion of satellite cells in the LA express AR and that this proportion is significantly greater in the LA compared to the CM. No sex differences were found between the proportions of satellite cells expressing AR in either muscle. Less colocalization of satellite cells and AR was seen in postnatal day 3 muscle than in postnatal day 1 muscle. In contrast, only negligible amounts of fibroblasts labeled with S100A4 express AR in either the LA or the CM. Together, findings support satellite cells, but not fibroblasts, as a candidate cell type involved in the sexual differentiation of the SNB neuromuscular system. © 2012 Wiley Periodicals, Inc. Develop Neurobiol 73: 448–454, 2013.     

Mammary ductal elongation: differentiation of myoepithelium and basal lamina during branching morphogenesis

Elongation of mammary ducts in the immature mouse takes place as a result of rapid growth in end buds. These structures proliferate at the apex of elongating ducts and are responsible for penetration of the surrounding adipose stroma; by turning and branching, end buds give rise to the characteristic open pattern of the mammary ductal tree. We have used a variety of techniques to determine the cellular and structural basis for certain of these end bud activities, and now report the following. (1) The end bud tip is covered with a monolayer of epithelium, the "cap cells," which are characterized by a relative lack of intercellular junctions and other specialized features. (2) The cap cell layer extends along the end bud flank and neck regions where it is continuous with the myoepithelium which surrounds the subtending mature duct. A linear sequence of differentiative changes occur in the cap cells in this region as they progressively alter in shape and accumulate the cytological features of mature myoepithelium. Cap cells may therefore be defined as a stem cell population providing new myoepithelial cells for ductal morphogenesis and elongation. (3) Differentiation of cap cells into myoepithelium is associated with conspicuous changes in the basal lamina. At the tip, cap cells form a 104-nm lamina similar to that described in expanding mammary alveoli and in embryonic tissues. Along the end bud flanks the basal lamina is raised from the cell surface and extensively folded, resulting in a greatly thickened lamina, measuring as much as 1.4 microns. At the surface of the subtending ducts the lamina becomes structurally simplified and resembles that at the tip, but has a significantly greater thickness, averaging 130 nm. (4) The codifferentiation of myoepithelium and its basement membrane is associated with changes in the surrounding stroma. Undifferentiated mesenchymal-like cells attach to the surface of the basal lamina in the midportion of the end buds and become increasingly numerous in the neck region, forming a monolayer over the myoepithelial basal lamina. These stromal cells progressively differentiated into fibrocytes which participate in collagen fibrillogenesis and give rise to the fibrous components of the stroma surrounding the mature duct.     

Morphology of connective tissue in skeletal muscle

The arrangement and distribution of connective tissue in six different skeletal muscles and smooth muscle was examined by scanning electron microscopy. The endomysial arrangement of collagen was similar in all types of muscle and consisted of three components: (1) myocyte-myocyte connectives; (2) myocyte-capillary connectives; and (3) a weave network of collagen intimately associated with the basal laminae of the myocytes. The perimysium of the different muscles was qualitatively similar but quantitatively dissimilar. The perimysium consisted of large tendon-like bundles of interwoven collagen which connected with the dense weave collagen that surrounded groups of muscles. The arrangement of the collagen in the perimysium and endomysium would explain differences in the mechanical properties of the different muscle. The contribution of the connective tissue to mechanical properties of muscle is discussed.     

Arrangement of smooth muscle cells and intramuscular septa in the taenia coli

Summary Bands of electron-dense material beneath the cell membrane of smooth muscle cells of the guinea-pig taenia coli provide attachment to thin myofilaments and to intermediate (10 nm) filaments; about 50% of the cell membrane is occupied by dense bands in muscle cells transversely sectioned at the level of their nucleus, and between 50 and 100% in smaller cell profiles nearer the cell's ends. In addition to the known cell-to-cell junctions (intermediate contacts), more complex apparatuses anchor muscle cells together, either end-to-end or end-to-side or side-to-side. They consist of elaborate folds, invaginations and protrusions accompanied by large amounts of basal lamina material. In the end-to-end anchoring apparatuses numerous finger-like and laminar processes from the two cells interdigitate. Other muscle cells have a star-shaped profile in the last few microns of their length, or show longitudinal invaginations occupied by a thickened basal lamina and occasionally by collagen fibrils. The septa of connective tissue extend only for a few hundred microns along the length of the taenia. In taeniae fixed in condition of mild stretch the muscle cells form an angle of about 5° with the septa. In muscles fixed during isotonic contraction the angle increases to about 20–22°, and in longitudinal sections the muscle cells appear arranged in a herring-bone pattern. The collagen concentration in the taenia coli is 4–6 times greater that in skeletal and cardiac muscles. These various structures are discussed in terms of their possible role in the mechanism of force transmission.I thank Mr. S.J. Sarsfield and Miss E.M. Franke for expert technical assistance, and Dr. Adam Yamey for much help in the experiments on collagen content. This work is supported by grants from the Medical Research Council     

The significance of muscle cells for the origin of mesoderm in bilateria

Muscle tissue may have played a central role in the early evolutionof mesoderm. The first function of myocytes could have beento control swimming and gliding motion in ciliated vermiformorganisms, as it still is in such present-day basal Bilateriaas the Nemertodermatida. The only mesodermal cells between epidermisand gastrodermis in Nemertodermatida are myocytes, and conceivablythe myocyte was, in fact, the original mesodermal cell type.In Nemertodermatida as well as the Acoela, myocytes are subepithelialfiber-type muscle cells and appear to originate from the gastrodermalepithelium by emigration of single cells. Other mesodermal cellsin the acoels are the peripheral parenchyma (connective tissue)and tunica cells of the gonads, and these also arise from thegastrodermis. Musculature in many of the coelomate protostomesand deuterostomes, on the other hand, is in the form of epitheliomuscular(myoepithelial) cells, and this cell type may also have beenan early form of the mesodermal myocyte. The mesodermal bandsin the small annelid Polygordius and in juvenile enteropneustshave cells intermediate between mesenchymal and epithelial intheir histological organization as they develop into myoepithelia.If acoelomates were derived from coelomates by progenesis, thenthe fiber-type muscles of acoelomates could be products of foreshorteneddifferentiation of such tissue. The precise serial patterningof circular muscle cells along the anterior-posterior axis duringembryonic development in the acoel Convoluta pulchra providesa model for early steps in the gradual evolution of segmentationfrom iterated organ systems.     

Structure of the digestive tract of the pluteus larva of Dendraster excentricus (Echinodermata: Echinoida)

Summary Histological and ultrastructural observations of the digestive tract of eight-armed plutei of Dendraster excentricus are reported. The esophagus is divided into two regions. The uppermost is a narrow tube comprised of ciliated cells that assist in transporting food to the more bulbous lower esophagus where food particles are formed into a bolus prior to entering the stomach. The esophagus is surrounded by a network of smooth muscle fibers that are predominantly oriented circumferentially in the upper esophagus, and longitudinally in the lower esophagus. The musculature of the upper esophagus produces peristaltic contractions, whereas contractions of the muscle of the lower esophagus open the cardiac sphincter and force food from the lower esophagus into the stomach. Axons are associated with the ciliated cells and the muscles of the upper esophagus. The cardiac sphincter consists of a ring of myoepithelium, with cross-striated myofibrils oriented around the bases of the cells. The gastric epithelium is comprised of two cell types. Type I cells, which predominate, absorb and store nutrients, and may be the source of secreted digestive enzymes. Type II cells apparently phagocytize and intracellularly digest whole algal cells. The intestine is comprised of relatively unspecialized cells and probably functions primarily as a conductive tube for the elimination of undigested materials.