Organization of the coelomic lining and a juxtaposed nerve plexus in the suckered tube feet of Parastichopus californicus (Echinodermata: Holothuroida)

The coelomic lining of the water-vascular canal in a suckered tube foot from the sea cucumber, Parastichopus californicus, is a pseudostratified myoepithelium consisting of flagellated adluminal cells and myofilament-bearing retractor cells. The bodies of adluminal cells flank the water-vascular canal and send basal processes between the underlying retractor cells to confront the podial connective tissue. Retractor cells have a contractile apparatus of unregistered thick and thin myofilaments. The contractile apparatus is confined to the medullary sarcoplasm and oriented parallel to the primary axis of a tube foot. The bodies and processes of retractor cells intermingle with the basal processes of adluminal cells at the basal lamina of the coelomic lining. A ganglionated nerve plexus in the podial connective tissue approximates the basal lamina. Neuronal connectives link the ganglia to one another and to the nerve plexus in deep sectors of the podial epidermis. External laminae enveloping the ganglia and connectives in the podial connective tissue are continuous with the basal lamina of the epidermis. The adventitial nerve plexus, since it merges with the epidermal nerve plexus, is a component of the ectoneural division of the echinoderm nervous system.     

Functional morphology and evolutionary aspects of unusual antennal circulatory organs in Labidura riparia Pallas (Labiduridae), Forficula auricularia L. and Chelidurella acanthopygia géné (Forficulidae) (Insecta : Dermaptera)

The antennal circulatory organs in the earwigs Labidura riparia Pallas (Labiduridae), Forficula auricularia L. and Chelidurella acanthopygia Géné (Forficulidae) (Dermaptera) represent a functional type that has not been found in other insects. An independent organ exists for each antenna, consisting of a pulsatile ampulla connected to an antennal blood vessel. The ampulla is attached to the frontal cuticle medial to the antenna base and forms a thin-walled sac with a valved ostium on its ventral side. The ampulla wall epithelium is not muscular, but consists of elastic connective tissue. The pumping movements are affected by a precerebral frontopharyngeal muscle, which causes systolic compression of the ampulla upon contraction. The elasticity of both the ampulla and a band of connective tissue, which suspends it in the head capsule, passively effect diastole. The antennal vessel is quite voluminous in the head capsule, but narrows remarkably upon entrance into the antenna. It extends with a constant diameter to the apex, where it opens with a terminal pore. At the base of the vessel, near the ampulla, is a very delicate valve flap which prevents hemolymph backflow during diastole. A comparison of the antennal heart types in insects revealed fundamental differences in the attachments and functions of the associated muscles. In the Dermaptera, the involvement of a precerebral frontopharyngeal muscle suggests an ancestral condition.     

Specializations for excitation-contraction coupling in the podial retractor cells of the starfish Stylasterias forreri

Summary Ultrastructural examination of the podium of the asteroid echinoderm Stylasterias forreri has revealed that cells of the coelomic epithelium and cells of the retractor muscle should be considered as components of a single epithelium. The podial retractor cells are, therefore, myoepithelial in nature. This report concentrates on those ultrastructural features of the retractor cells that are most likely involved with excitation-contraction coupling. The spatial arrangement of the sarcoplasmic reticulum, the couplings between the sarcoplasmic reticulum and sarcolemma, and an intramembranous specialization of the sarcolemma are documented and discussed. Current concepts regarding the innervation of the retractor cells of the podium and the protractor cells of the ampulla are reviewed, and specific proposals for further investigation of podial innervation are outlined.This investigation was supported by Research Operating Grant A0484 from the Natural Sciences and Engineering Research Council of Canada (M.J.C.) and by general research funds from the Department of Anatomy of the University of Southern California (R.L.W.). Mr. Steve Osborne and Ms. Aileen Kuda provided technical assistance. A portion of this study was conducted at the Friday Harbor Laboratories of the University of Washington, and the authors gratefully acknowledge the cooperation and hospitality of the Director, Dr. A.O. Dennis Willows     

Ontogenetic torsion in two basal gastropods occurs without shell attachments for larval retractor muscles

Results of this study on two species of vetigastropods contradict the long-standing hypothesis, originally proposed by Garstang (1929), that the larval retractor muscles power the morphogenetic movement of ontogenetic torsion in all basal gastropods. In the trochid Calliostoma ligatum and the keyhole limpet Diodora aspera, the main and accessory larval retractor muscles failed to establish attachments onto the protoconch (larval shell) when the antibiotics streptomycin sulfate and penicillin G were added to cultures soon after fertilization. Defects in protoconch mineralization were also observed. Despite these abnormalities, developing larvae of these species accomplished complete or almost complete ontogenetic torsion, a process in which the head and foot rotate by 180 degrees relative to the protoconch and visceral mass. Analysis by using phalloidin-fluorophore conjugate and transmission electron microscopy showed that myofilaments differentiated within myocytes of the larval retractor muscles and adherens-like junctions formed between muscle and mantle epithelial cells in both normal and abnormal larvae. However, in abnormal larvae, apical microvilli of mantle cells that were connected to the base of the larval retractor muscles failed to associate with an extracellular matrix that normally anchors the microvilli to the mineralized protoconch. If morphogenesis among extant, basal gastropods preserves the original developmental alteration that created gastropod torsion, as proposed by Garstang (1929), then the alteration involved something other than the larval retractor muscles. Alternatively, the developmental process of torsion has evolved subsequent to its origin in at least some basal gastropod clades so that the original alteration is no longer preserved in these clades.     

Ultrastructure of coelomic lining in echinoderm podia: significance for concepts in the evolution of muscle and peritoneal cells

Summary Ultrastructural data are presented on the histological organization of coelomic lining in the podia of ten species of the five major groups of extant echinoderms. Further evidence of the incorporation of podial retractor muscle cells (myocytes) into a monociliated myoepithelial coelomic lining is provided. In the podia of the crinoid Nemaster rubinginosa and the ophiuroid Ophiophragmus wurdemani as well as in the feeding tentacles of the holothurian Leptosynapta tenuis, coelomic linings are organized as simple myoepithelia consisting of non-contractile peritoneal cells (peritoneocytes) and myocytes. Coelomic linings in the holothurian Thyonella gemmata, the echinoids Eucidaris cf. tribuloides and Lytechinus variegatus, and the asteroids Asterias forbesi and Astropecten sp. are pseudostratified or bipartite pseudostratified myoepithelia consisting of subapical myocytes and apically situated peritoneocytes. The ophiuroid podia of Ophioderma brevispinum and Ophiothrix angulata exhibit transitions from simple myoepithelia to partially pseudostratified epithelia. Intermediate forms between the extremes in myoepithelial organization also occur in the podial lining of single species (e.g. Eucidaris cf. tribuloides). These data supplement recent ultrastructural studies on the podial lining of echinoderms and, in conjunction with published ultrastructural data on the myoepithelial organization of other coelomic linings in echinoderms and in other coelomates, suggest myoepithelial organization of the coelomic lining is a plesiomorph feature in Bilateria.     

THE ORGANISATION OF THE PEDAL MUSCULATURE AND ITS CONNECTION TO THE DORSOVENTRAL MUSCULATURE IN SCAPHOPODA

Scaphopoda possess one or two pairs of dorsoventral muscles.At the level of the diaphragm between the intestinal and theperianal sinus these muscles divide into a latero-dorsal portionand a medio-ventral portion. The entire medio-ventral portionand the ventral parts of the latero-dorsal portion form thepedal longitudinal musculature. This is the general patternin both orders, Dentaliida and Gadilida. In Gadilida exceptthe family Entalinidae, these muscle portions are reduced comparedto an additional pair of central pedal retractor muscles. Themusculature of the pedal wall is four-layered: outer circularmuscles, two layers of helical muscles, and inner longitudinalmuscles. Because of differences in the organization of the musculature,the foot consists of three functional units: a pedal base, amiddle piece, and an anchoring organ. In the dentalud foot,all of the longitudinal muscles are in contact with the pedalwall. In the Entalinidae, three or four pairs of central retractormuscles become free of the pedal wall at the base of the middlepiece. In all other Gadilida the two central retractors arecontinuous from the dorsoventral muscles into the anchoringorgan. The elongation of the foot is purely hydraulic in Gadilida.In Dentaliida, however, the principles of hydraulic and muscular-hydrostatsare combined. (Received 17 July 1991; accepted 28 October 1991)     

Evidence that gastropod torsion is driven by asymmetric cell proliferation activated by TGF-beta signalling

Gastropods are characterized by their asymmetric bodyplan, which develops through a unique ontogenetic process called 'torsion'. Despite several intensive studies, the driving force of torsion remains to be determined. Although torsion was traditionally believed to be driven by contraction of the retractor muscle connecting the foot and the shell, some recent reports cast doubt on that idea. Here, we report that torsion is accompanied by left-right asymmetric cell proliferation in the mantle epithelium in the limpet Nipponacmea fuscoviridis. Furthermore, we found that pharmacological inhibition of the transforming growth factor-β (TGF-β) signalling pathway, including that of Nodal, blocked torsion. We confirmed that the blocking was brought about through failure of the activation of cell proliferation in the right-hand side of the mantle epithelium, while the retractor muscle apparently developed normally. These results suggest that limpet torsion is driven by left-right asymmetric cell proliferation in the mantle epithelium, induced by the TGF-β pathway.     

A numerical analysis of the backflow between the leaflets of a St Jude Medical cardiac valve prosthesis

Clinically significant, unexplained hemolysis has been reported to occur in St Jude Medical (SJM) cardiac valve prostheses. The leakage phase of backflow is identified as having the most hemolytic potential, when compared with the other phases of the cardiac cycle. A two-dimensional, laminar, constant fluid property finite element analysis (FEA) is used to calculate the peak shear stress obtained in the flow through the narrow slit formed by the closed leaflets of an idealized size 29 mm (tissue annulus diameter) SJM cardiac valve prosthesis, during the leakage phase of backflow. The flow geometry was such that the simple laminar flow theory (SLFT) could be used to model the flow through the region of peak shear stresses with reasonable accuracy. The SLFT predicts that the maximum shear stress depends upon the thickness of the clearance space and the average velocity through that space. These results suggest that the magnitude of the peak shear stress is of the order of 700-1000 Pa for a duration of the order of 0.5-0.4 ms for pressure drops across the valve of 150-300 torr. This suggests that hemolysis is possible for certain unfortunate combinations of clearances and pressure conditions. However, further research is needed before this flow phenomenon can be associated with the reported clinical hemolysis.     

Influence of tongue muscle contraction and dynamic airway pressure on velopharyngeal volume in the rat.

The mammalian pharynx is a collapsible tube that narrows during inspiration as transmural pressure becomes negative. The velopharynx (VP), which lies posterior to the soft palate, is considered to be one of the most collapsible pharyngeal regions. I tested the hypothesis that negative transmural pressure would narrow the VP, and that electrical stimulation of extrinsic tongue muscles would reverse this effect. Pressure (-6, -3, 3, and 6 cmH2O) was applied to the isolated pharyngeal airway of anesthetized rats that were positioned in a 4.7-T MRI scanner. The volume of eight axial slices encompassing the length of the VP was computed at each level of pressure, with and without bilateral hypoglossal nerve stimulation (0.1-ms pulse, one-third maximum force, 80 Hz). Negative pressure narrowed the VP, and either whole hypoglossal nerve stimulation (coactivation of protrudor and retractor muscles) or medial nerve branch stimulation (independent activation of tongue protrudor muscles) reversed this effect, with the greatest impact in the caudal one-third of the VP. The dilating effects of medial branch stimulation were slightly larger than whole nerve stimulation. Positive pressure dilated the VP, but tongue muscle contraction did not cause further dilation under these conditions. I conclude that the narrowest and most collapsible segment of the rat pharynx is in the caudal VP, posterior to the tip of the soft palate. Either coactivation of protrudor and retractor muscles or independent contraction of protrudor muscles caused dilation of this region, but the latter was slightly more effective.     

Determinants of valve gating in collecting lymphatic vessels from rat mesentery

Secondary lymphatic valves are essential for minimizing backflow of lymph and are presumed to gate passively according to the instantaneous trans-valve pressure gradient. We hypothesized that valve gating is also modulated by vessel distention, which could alter leaflet stiffness and coaptation. To test this hypothesis, we devised protocols to measure the small pressure gradients required to open or close lymphatic valves and determine if the gradients varied as a function of vessel diameter. Lymphatic vessels were isolated from rat mesentery, cannulated, and pressurized using a servo-control system. Detection of valve leaflet position simultaneously with diameter and intraluminal pressure changes in two-valve segments revealed the detailed temporal relationships between these parameters during the lymphatic contraction cycle. The timing of valve movements was similar to that of cardiac valves, but only when lymphatic vessel afterload was elevated. The pressure gradients required to open or close a valve were determined in one-valve segments during slow, ramp-wise pressure elevation, either from the input or output side of the valve. Tests were conducted over a wide range of baseline pressures (and thus diameters) in passive vessels as well as in vessels with two levels of imposed tone. Surprisingly, the pressure gradient required for valve closure varied >20-fold (0.1-2.2 cmH(2)O) as a passive vessel progressively distended. Similarly, the pressure gradient required for valve opening varied sixfold with vessel distention. Finally, our functional evidence supports the concept that lymphatic muscle tone exerts an indirect effect on valve gating.     

Major muscle systems in the larval caenogastropod,Ilyanassa obsoleta,display different patterns of development

This study describes the anatomical and developmental aspects of muscular development from the early embryo to competent larval stage in the gastropod Ilyanassa obsoleta. Staining of F‐actin revealed differential spatial and temporal patterns of several muscles. In particular, two major muscles, the larval retractor and pedal retractor muscles originate independently and display distinct developmental patterns similar to observations in other gastropod species. Additionally, together with the larval retractor muscle, the accessory larval muscle developed in the embryo at the trochophore stage. Therefore, both these muscles develop prior to ontogenetic torsion. The pedal retractor muscle marked the most abundant growth in the mid veliger stage. Also during the middle stage, the metapodial retractor muscle and opercular retractor muscle grew concurrently with development of the foot. We show evidence that juvenile muscles, such as the buccal mass muscle and siphon muscle develop initially during the late veliger stage. Collectively, these findings substantiate that larval myogenesis involves a complex sequence of events that appear evolutionary conserved within the gastropods, and set the stage for future studies using this model species to address issues concerning the evolution and eventual fates of larval musculature in molluscs. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

Further pharmacological study on the cholinergic and glutaminergic properties of the radula muscles of a mollusc,Rapana thomasiana

1. In the radula protractor of the prosobranch mollusc Rapana thomasiana, both twitch contractions and acetylcholine contractions were markedly depressed or blocked by propantheline (10⁻⁵ M) and strychnine (10⁻⁵ M), but in the radula retractor, only acetylcholine contraction was markedly affected by the antagonists,2. Glutamate contractions of both of the muscles were little or slightly affected by the drugs.3. Twitch contraction of the protractor was slowly depressed when the muscle was immersed in concanavalin A (0.3 mg/ml), while that of the retractor was first potentiated and then slowly depressed in it.4. In both of the muscles, glutamate contractions were markedly enhanced by the lectin, but acetylcholine contractions were not affected.5. These results support the notion that the principal excitatory neurotransmitter in the protractor is acetylcholine, whereas that in the retractor is glutamate.     

The properties of the extraocular muscles of the frog. III. Morphological, mechanical and pharmacological properties of the isolated retractor bulbi muscle.

Some morphological, physiological, and pharmacological properties of the retractor bulbi muscle of the frog were tested. The enzyme-histochemical investigation shows that the retractor bulbi muscle contains twitch muscle fibres only. Two types of twitch muscle fibres, which are especially different in their diameter and in the content of mitochondria, build the muscle in an irregular arrangement; tonic muscle fibres were not observed. On the average, the isolated retractor bulbi muscle has at room temperature a contraction time of 26 ms, a half-relaxation time of 28 ms, a fusion frequency of 75 stimuli/s, and a twitch-tetanus ratio of 0.28. The fatigability of this muscle is higher than in oculorotatory eye muscles but lower than in skeletal muscles of the frog. An increase of the extracellular K+-concentration elicits in retractor bulbi muscles a quickly transient contracture; the mechanical threshold of the muscle fibres is found in a range between 20 and 25 mM K+ in Ringer solution. Similar short-lasting contractures, which are probably caused by twitch fibres, rich in mitochondria, are also evoked by application of depolarizing drugs like acetylcholine. The properties of the retractor bulbi muscle are compared with those of the sartorius muscle of the frog, which likewise contains twitch muscle fibres only.     

Biology and functional morphology of Kendrickiana gen. nov. veitchi (Bivalvia: Anomalodesmata: Clavagelloidea) from southern Australia

Abstract. The largest extant species of the adventitious tube-building Clavagelloidea has hitherto been placed in the genus Foegia (itself formerly a subgenus of Brechites ), the type species of which is the Western Australian F. novaezelandiae (B ruguiére 1792). Following examinations of and comparison with F. novaezelandiae and the southern Australian F. veitchi , the latter is herein placed in its own new genus Kendrickiana .
Individuals of both F. novaezelandiae and K. veitchi are essentially amyarian in terms of adductor and pedal retractor muscles, but in the latter species the connection to the adventitious tube is located dorsally by an unique horseshoe-shaped array of muscular papillae, which are inserted into holes in the tube. K. veitchi is different from other clavagelloids too in that the siphons are capable of only limited retraction into the tube. Their extension in K. veitchi is almost exclusively by hydraulic means because the complex internal siphonal muscles seen in other species of the Clavagelloidea, and which act antagonistically with extensive blood-filled haemocoels, are vestigial. Kendrickiana can also be separated from Foegia in other anatomical respects. For example, members of the former have paired anterior suspensory muscles, and vestigial posterior pedal retractor muscles with pericardial proprioreceptors associated with them (as in Humphreyia and Dianadema ), whereas the latter does not. Similarly, in Foegia there is a muscularized pedal disc, not found in Kendrickiana . Members of F. novaezelandiae are inhabitants of intertidal hypoxic muds, whereas those of K. veitchi apparently live exclusively in subtidal sea grass beds.     

Paramyosin in invertebrate muscles. I. Identification and localization

By sodium dodecyl sulfate-polyacrylamide gel electrophoresis and immunodiffusion, we identified paramyosin in two smooth invertebrate "catch" muscles (Mytilus anterior byssus retractor and Mercenaria opaque adductor) and five invertebrate striated muscles (Limulus telson levator, Homarus claw muscle, Balanus scutal depressor, Lethocerus air tube retractor, and Aequipecten striated adductor). We show that (a) the paramyosins in all of these muscles have the same chain weights and (b) they are immunologically similar. We stained all of these muscles with specific antibody to Limulus paramyosin using the indirect fluorescent antibody technique. Paramyosin was localized to the A bands of the glycerinated striated muscles, and diffus fluorescence was seen throughout the glycerinated fibers of the smooth catch muscles. The presence of paramyosin in Homarus claw muscle, Balanus scutal depressor, and Lethocerus air tube retractor is shown here for the first time. Of the muscles in this study, Limulus telson levator is the only one for which the antiparamyosin staining pattern has been previously reported.     

Torsion in Patella caerulea (Mollusca, Patellogastropoda): ontogenetic process, timing, and mechanisms

Abstract. Torsion is a process in gastropod ontogenesis where the visceral body portion rotates 180° relative to the head/foot region. We investigated this process in the limpet Patella caerulea by using light microscopy of living larvae, as well as scanning electron microscopy (SEM) of larvae fixed during the torsion process. The completion of the 180° twist takes considerably less time in larvae of Patella caerulea than previously described for other basal gastropod species. At a rearing temperature of 20–22°C, individuals complete ontogenetic torsion in ?2 h. Furthermore, the whole process is monophasic, i.e., carried out at a constant speed, without any evidence of distinct ‘fast” or ‘slow” phases. Both larval shell muscles—the main and the accessory larval retractor—are already fully contractile before the onset of torsion. During the torsion process both retractors perform cramp‐like contractions at ～30 s intervals, which are followed by hydraulic movements of the foot. However, retraction into the embryonic shell occurs only after torsion is completed. The formation of the larval operculum is entirely in‐dependent from ontogenetic torsion and starts before the onset of rotation, as does the mineralization of the embryonic shell. The reported variability regarding the timing (mono‐ versus biphasic; duration) of torsion in basal gastropod species precludes any attempt to interpret these data phylogenetically. The present findings indicate that the torsion process in Patella caerulea, and probably generally in basal gastropods, is primarily caused by contraction of the larval shell muscles in combination with hydraulic activities. In contrast, the adult shell musculature, which is independently formed after torsion is completed, does not contribute to ontogenetic torsion in any way. Thus, fossil data relying on muscle scars of adult shell muscles alone appear inappropriate to prove torted or untorted conditions in early Paleozoic univalved molluses. Therefore, we argue that paleontological studies dealing with gastropod phylogeny require data other than those based on fossilized attachment sites of adult shell muscles.     

Evolution of locomotion in arachnida: The hydraulic pressure pump of the giant whipscorpion,Mastigoproctus Giganteus (Uropygi)

Many arachnids lack extensor muscles at the femoropatellar (knee) joint of their legs and extend this joint with hydraulic pressure during locomotion. Pressure is generated through compression of the prosoma, but there is disagreement about which muscles are involved in this process. Many arachhnologists consider contraction of the musculi laterales, a group of modified extrinsic leg muscles, as the cause of high prosomal pressure and regard hydraulic extension as a derived feature. However, integration of results from phylogenetic and comparative anatomical studies supports the view that hydraulic extension is primitive in Arachnida and that fluid pressure is generated by contraction of endosternal suspensor muscles. The functional predictions of the musculi laterales and endosternite hypotheses were tested by measuring muscle activity and prosomal pressure during unrestrained locomotion in a primitively “extensorless” arachnid, the giant whipscorpion. The results corroborate the endosternite model and refute the musculi laterales model. Changes in the prosomal pressure baseline were correlated with changes in endosternal muscle activity, while the musculi laterales fired in a step-coupled pattern of discrete bursts that appeared to be incapable of generating the pressure observed during locomotion. Step-coupled fluctuations in prosomal pressure were observed but were apparently caused by rapid flexing of the femoropatellar joints of the fourth leg pair rather than contraction of the musculi laterales.     

Fine structure of the axial complex of Sphaerechinus granularis (Lam.) (Echinodermata: Echinoidea)

Summary Three regions of the axial complex in Sphaerechinus granularis can be distinguished: 1) The axial organ which protrudes from one side of the axial sinus; the sinus septum which separates the sinus from the body cavity and encloses the stone canal; the pulsating vessel which runs along the inside of the axial organ. 2) The blindly-ending terminal sinus in which the pulsating vessel broadens out to the contractile terminal process. 3) The ampulla of the stone canal which connects the axocoel and water vascular system and which opens out through the madreporite.A single-layered, monociliated coelomic epithelium surrounds all regions of the axial complex. This epithelium contains smooth muscle cells at the contractile areas. Canaliculi, surrounded by basal lamina, are formed through infolding of epithelia; they end blindly in the fluid and connective tissue -matrix of the inner structures.The lacunae of the dorso-ventral mesentery connect the periesophageal and the perianal haemal ring with the axial organ. The axial organ contains many coelomocytes rich in pigment and granules. These coelomocytes are separated into compartments by elastic fibres. Phagocytosis of whole cells and transformational stages of coelomocytes suggest storage and degradation functions. An excretory function via the water vascular system is also suggested.     

Action and immunoreactivity of neuropeptides in the buccal neuromuscular system of a prosobranch mollusc,Rapana thomasiana

Summary In a prosobranch mollusc, Rapana thomasiana, the catch-relaxing peptide H-Ala-Met-Pro-Met-Leu-Arg-Leu-NH₂ (CARP) was found to depress the contraction of the radula protractor and retractor elicited by electrical stimulations. The action of CARP was in contrast to that of other neuropeptides, H-Phe-Met-Arg-Phe-NH₂ (FMRFamide) and H-Phe-Leu-Arg-Phe-NH₂ (FLRFamide), which enhanced the contraction of the radula protractor and retractor, respectively. By immunohistochemical examinations, FMRFamide-like immunoreactive neurons were found on the rostral side of the right buccal ganglion and the caudal side of the left ganglion, where some CARP-like immunoreactive neurons were also distributed, indicating a possible coexistence of FMRFamide and CARP. FMRFamide- and CARP-like immunoreactivities were also detected in the neuropile of buccal ganglia, radula nerves arising from the ganglia, and nerve fibers in the radula muscles. The present results suggest that FMRFamide- and CARP-like peptides are involved in the regulation of the contraction of the radula muscles.