Muscular system of a peculiar parasitic cnidarian Polypodium hydriforme: a phalloidin fluorescence study

Musculature of the free-living stages of Polypodium hydriforme has been studied using phalloidin fluorescence method and confocal microscopy. P. hydriforme is a unique cnidarian possessing only smooth muscle cells situated within the mesoglea, not epithelial muscle cells, like the rest of cnidarians. Phalloidin fluorescence on whole mount preparations demonstrates an extensively developed subepidermal muscle system mostly consisting of long parallel fibers running along the tentacles. For the first time along with contracted muscle fibers we could clearly demonstrate relaxed fibers looking as long spirals. System of thin parallel circular F-actin positive fibers has been discovered outside of longitudinal muscles. The body of the animal and the mouth cone contain weakly developed parallel muscles. No special attachment of the muscle fibers to the tips of the tentacles or to the rim of the mouth has been observed. The results are discussed in connection with the "triploblastic" organization of P. hydriforme and its phylogenetic position.     

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.     

Embryonic muscle development of Convoluta pulchra (Turbellaria-acoelomorpha, platyhelminthes)

We studied the embryonic development of body-wall musculature in the acoel turbellarian Convoluta pulchra by fluorescence microscopy using phalloidin-bound stains for F-actin. During stage 1, which we define as development prior to 50% of the time between egg-laying and hatching, actin was visible only in zonulae adhaerentes of epidermal cells. Subsequent development of muscle occurred in two distinct phases: first, formation of an orthogonal grid of early muscles and, second, differentiation of other myoblasts upon this grid. The first elements of the primary orthogonal muscle grid appeared as short, isolated, circular muscle fibers (stage 2; 50% developmental time), which eventually elongated to completely encircle the embryo (stage 3; at 60% of total developmental time). The first primary longitudinal fibers appeared later, along with some new primary circular fibers, by 60-63% of total developmental time (stage 4). From 65 to 100% of total developmental time (stages 5 to 7), secondary fibers, using primary fibers as templates, arose; the number of circular and longitudinal muscles thus increased, and at the same time parenchymal muscles began appearing. Hatchlings (stage 8) possessed about 25 circular and 30 longitudinal muscles as well as strong parenchymal muscles. The remarkable feature of the body wall of many adult acoel flatworms is that longitudinal muscles bend medially and cross each other behind the level of the mouth. We found that this development starts shortly after the appearance of the ventral mouth opening within the body wall muscle grid. The adult organization of the body-wall musculature consists of a grid of several hundred longitudinal and circular fibers and a few diagonal muscles. Musculature of the reproductive organs developed after hatching. Thus, extensive myogenesis must occur also during postembryonic development. Comparison between the turbellarians and the annelids suggests that formation of a primary orthogonal muscle grid and its subsequent use as a template for myoblast differentiation are the two basic developmental phases in vermiform Spiralia if not in the Bilateria as a whole. Finally, our new data suggest that for the Acoela the orthogonal primary patterning of longitudinal and circular muscles in the body wall is achieved without using originally positional information of the nervous system.     

F-actin framework in Spirorbis cf. spirorbis (Annelida: Serpulidae): phalloidin staining investigated and reconstructed by cLSM

Abstract. Serpulidae encompasses polychaete species whose members have fused anterior ends bearing a tentacular crown, a heteronomous segmented body with a thorax and abdomen, and “chaetal inversion” between the two tagmata. The sessile filter‐feeding organisms live in self‐built, coiled, calcareous tubes on algae. The F‐actin muscular subset of Spirorbis cf. spirorbis was stained with phalloidin and three‐dimensionally reconstructed by means of cLSM, aiming to investigate (1) how the tentacular crown is organized and moved, (2) whether the internal structures, e.g., musculature, follow the thorax–abdomen inversion, and (3) whether circular muscles are present in serpulids. The third aim is by reason of recent investigations suggesting that lack of circular muscle fibers may be a common situation rather than a rare variation in polychaetes. In this manner, this article is part of a comparative evaluation of polychaete muscle systems. We found that longitudinal muscles of the body wall project into the tentacular crown, and that radioli and pinnulae possess three muscle types each, facilitating their great mobility. Operculum, collar, and a pair of unidentified organs possess distinct F‐actin filaments. The trunk is mainly moved by five longitudinal muscle strands, most obvious in the abdomen: two dorsal, two ventral, and an unpaired ventromedian one, out of which the dorsal ones are the strongest. In anterior regions, the two dorsal strands form a single continuous layer; the separated strands lessen posteriorly. Solitary transverse fibers are located ventrally in the middle of each segment, stretching between longitudinal muscles and coelomic lining laterally, where they end. Peripheral and central dorsoventral muscles, two pairs per segment each, are present. Circular fibers as well as bracing muscles were not detected. The results indicate that the musculature does not follow the thorax–abdomen inversion and Serpulidae represents the 15th polychaete taxon in which circular fibers are totally missing.     

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.     

Unique patterns of longitudinal body-wall musculature in the Acoela (Plathelminthes): the ventral musculature of Convolutriloba longifissura

The ventral musculature of Convolutriloba longifissura (Acoela) has been studied using electron microscopy and fluorescently labeled whole mounts to demonstrate filamentous actin. Attention was directed to the reorganization and renewal of musculature during asexual reproduction and the adaptation of muscle sets for special predatory behavior. Three ventral subepidermal muscle layers could be distinguished in adult C. longifissura: (1) outer circular muscles that encircle the body, (2) intermediate modified longitudinal muscles with concentric pattern around the mouth and V-shaped orientation in the posterior part of the animal, and (3) inner special pore muscles with radial alignment fanning out from the mouth. Additionally, a few very fragile muscles were found at the anterior margin of the animal. The anterior ventral muscle system built a funnel with the mouth opening as organizing center. The special radial muscles and the antagonistically concentric muscles are perfectly adapted to catch prey in such a way that the funnel is put over the prey to press it through the mouth into the digestive syncytium. Convolutriloba longifissura shows a unique way of asexual reproduction by a two-step fission which results in three individuals. Immediately after separation from the mother animal, daughter individuals are missing the concentric and the radial muscle sets around the mouth completely, but within 30 h these sets are renewed for the most part. Two to three days after separation, the mouth opening is visible and the animals move for capturing prey. The peculiar course of longitudinal muscles in C. longifissura with concentric rings anteriorly and a V-shape muscle layer posteriorly shows that the pattern of body-wall musculature in such basal Plathelminthes as the Acoela may be highly modified from the original pattern of longitudinal and circular muscles.     

Criteria of the homology and phylogeny of facial muscles in primates including man. I. Prosimia and Platyrrhina]

The facial muscles of primates are derivates of sphincter colli profundus muscle and platysma myoides. A third superficial muscle layer which is present in primitive mammals is found as a rest in Tupaiiformes. The facial muscles of some Lemuriformes must be considered as a model from which originate the facial musculature of other primates. The new formation of muscles takes place at the margine of the original muscle layers; marginal muscle fibers assume another run and get individualized. So it can be seen in facial muscles of prosimians and platyrrhine monkeys that the profound muscles of mouth and nose and some of the rostral margine of the auricle have orginated from sphincter colli muscle, all others from platysma myoides. Primitive and modified muscle forms and intermediate muscle forms can be observed among prosimians as well as among platyrrhine monkeys and by this it is possible to see the homology of the facial muscles.     

The Musculature of <Emphasis Type="Italic">Testudinella patina</Emphasis> (Rotifera: Flosculariacea), Revealed with CLSM

The musculature of Testudinella patina was visualized using phalloidin-linked fluorescent dye by confocal laser scanning microscopy. The conspicuous broad retractors appear to be made up of five separate fibers, of which three anchor in the neck region whereas two extend into the corona. Besides the broad retractors, a total of five paired longitudinal retractors are present and all of them extend into the corona. Incomplete circular muscles are found in groups in the neck region and in the medial and posterior parts of the trunk. The foot musculature comprises eight thin ventral foot muscles and six thicker dorsal foot muscles that all extend from the foot basis to the distal part of the foot. At the basis of the foot, each of the dorsal foot muscles anchors on a smaller, S-shaped subterminal foot muscle. The foot musculature furthermore comprises one pair of paraterminal foot muscles that each anchors basally on a subterminal foot muscle, extends into the most proximal part of the foot and attaches on one of the dorsal foot muscles. The visceral musculature is composed of extremely delicate fibers and is restricted to an area around and posterior to the foot opening. The presence of incomplete circular muscles supports that these muscles are a basal trait for Rotifera, whereas the morphology of the broad retractors and foot muscles is much more specialized and may be autapomorphic for Testudinella or alternatively for this genus and its closest relatives. The present results stress that revealing muscles by staining may produce new information from even well-investigated species, and that this information may contribute to a better understanding of functional as well as phylogenetic aspects of rotifer biology.     

Morphology and ultrastructure of the esophagus during the ontogeny of the spider crab Maja brachydactyla (Decapoda,Brachyura, Majidae)

The esophagus of the eucrustaceans is known as a short tube that connects the mouth with the stomach but has generally received little attention by the carcinologists, especially during the larval stages. By this reason, the present study is focused on the morphology and ultrastructure of the esophagus in the brachyuran Maja brachydactyla during the larval development and adult stage. The esophagus shows internally four longitudinal folds. The simple columnar epithelium is covered by a thick cuticle. The epithelial cells of the adults are intensively interdigitated and show abundant apical mitochondria and bundles of filamentous structures. The cuticle surface has microspines and mutually exclusive pores. Three muscle layers surrounded by the connective tissue are reported: circular muscles forming a broad continuous band, longitudinal muscle bundles adjacent to the circular muscles, and dilator muscles crossing the connective tissue vertically toward the epithelium. The connective tissue has rosette glands. The esophagus of the larvae have epithelial cells with big vesicles but poorly developed interdigitations and filamentous structures, the cuticle is formed by a procuticle without differentiated exocuticle and endocuticle, the connective layer is thin and the rosette glands are absent. The observed features can be explained by his role in the swallowing of the food.     

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.     

Development and embryonic pattern of body wall musculature in the crassiclitellate Eisenia andrei (Annelida,Clitellata)

During early development of Eisenia andrei (Crassiclitellata), a loose arrangement of primary circular and longitudinal muscles encloses the whole embryo. Circular muscles differentiate in an anterior–posterior progression creating a segmental pattern. Primary circular muscles emerge at the segmental borders while later in development the central part of each segment is filled with circular strands. Longitudinal muscles develop in an anterio‐posterior manner as well, but by continuous lengthening. Muscle growth is not restricted by segmental boundaries. The development begins with one pair of prominent longitudinal muscles differentiating ventrally along the right and the left germ band. These first muscles provide a guiding structure for the parallel organization of the afterwards differentiating longitudinal musculature. Additional primary longitudinal muscles emerge and form, together with the initial circular muscles, the primary muscle grid of the embryo. During the following development, secondary longitudinal muscle strands develop and integrate themselves into the primary grid. Meanwhile the primary circular muscles split into thin strands in a ventral to dorsal progression. Thus, a fine structured mesh of circular and longitudinal muscles is generated. Compared to other “Oligochaeta”, embryonic muscle patterns in E. andrei are adapted to the development of a lecithotrophic embryo. Nevertheless, two general characteristics of annelid muscle development become evident. The first is the segmental development of the circular muscles from a set of initial muscles situated at the segment borders. Second, there is a continuous development of primary longitudinal muscles starting at the anterior pole. At least one pair of main primary longitudinal strands is characteristic in Annelida. The space between all primary strands is filled with secondary longitudinal strands during further development. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Three-dimensional reconstruction of the musculature of Cossura pygodactylata Jones, 1956 (Annelida: Cossuridae)

The musculature of adult specimens of Cossura pygodactylata was studied by means of F-actin labelling and confocal laser scanning microscopy (CLSM). Their body wall is comprised of five longitudinal muscle bands: two dorsal, two ventral and one ventromedial. Complete circular fibres are found only in the abdominal region, and they are developed only on the border of the segments. Thoracic and posterior body regions contain only transverse fibres ending near the ventral longitudinal bands. Almost-complete rings of transverse muscles, with gaps on the dorsal and ventral sides, surround the terminal part of the pygidium. Four longitudinal bands go to the middle of the prostomium and 5–14 paired dorso-ventral muscle fibres arise in its distal part. Each buccal tentacle contains one thick and two thin longitudinal muscle filaments; thick muscle fibres from all tentacles merge, forming left and right tentacle protractors rooted in the dorsal longitudinal bands of the body wall. The circumbuccal complex includes well-developed upper and lower lips. These lips contain an outer layer of transverse fibres, and the lower lip also contains inner oblique muscles going to the dorsal longitudinal bands. The branchial filament contains two longitudinal muscle fibres that do not connect with the body musculature. The parapodial complex includes strong intersegmental and segmental oblique muscles in the thoracic region only; chaetal retractors, protractors and muscles of the body wall are present in all body regions. Muscle fibres are developed in the dorsal and ventral mesenteries. One semi-circular fibre is developed on the border of each segment and is most likely embedded in the dissepiment. The intestine has thin circular fibres along its full length. The dorsal blood vessel has strong muscle fibres that cover its anterior part, which is called the heart. It consists of short longitudinal elements forming regular rings and inner partitions. The musculature of C. pygodactylata includes some elements that are homologous with similar muscular components in other polychaetes (i.e., the body wall and most parapodial muscles) and several unique features, mostly at the anterior end.     

Purification and localization of human carbonic anhydrase

Summary Three different isoenzymes of human carbonic anhydrase are now well characterized. Carbonic anhydrase I and II have been known for several years and are located in high amounts in red blood cells as well as in many other tissues.Carbonic anhydrase III, a protein showing CO₂ hydratase and p-nitrophenylphosphatase activity was isolated from skeletal muscle some years ago. Earlier observations based on enzyme activity and radioimmunoassay studies have suggested that this protein is present in greater quantities in red skeletal muscles than in white ones. We have purified CA III from human soleus muscle and using obtained monospecific polyclonal antibody localized this protein in the same muscle fibers which show acid resistant ATPase activity. Using this protein as a marker for type I muscle fibers, fiber classification into type I and II could now be done also from paraffin embedded sections.This study is supported by the Research Council of Physical Education and Sport, Ministry of Education, Finland     

Dopamine produces muscle contractions and modulates motoneuron-induced contractions inAplysia gill

1. Dopamine has been reported to exist in unusually large quantities in Aplysia gill. The physiological role of this neurotransmitter in this organ was examined. 2. The addition of dopamine to a gill perfusate results in the contractions of the lateral and medial external pinnule muscles, the circular and longitudinal muscles of the afferent vessel, and the circular muscles of the efferent vessel. 3. Dopamine-induced contractions persist after chemical synaptic transmission is eliminated in the gill. This suggests that excitatory dopamine receptors are present on gill smooth muscle fibers themselves. 4. Dopamine also potentiates the gill response to action potentials in single identified gill motoneurons. Evidence presented suggests that muscle contractions and modulation of motoneuron contractions are independent phenomena. 5. While modulation may in part be mediated by increases in excitatory junction potential (EJP) amplitude, in many cases large increases in muscle contractions occur while the enhancement of EJPs is disproportionately small. 6. Dopamine's ability to produce muscle contractions suggests that there may be dopaminergic motoneuron innervation of the gill. We suggest that dopamine's modulatory actions may be mediated via modification of excitation-contraction coupling in smooth muscle fibers.     

Arrangement and structure of sinus hair muscles in the big-clawed shrew,Sorex unguiculatus

The arrangement and structure of sinus hair muscles in the snout of the shrew, Sorex unguiculatus, were studied by electron microscopy and serial section light microscopy. Both striated and smooth muscles are directly associated with sinus hair follicles. The striated muscle fibers originate from the base of a follicle and insert onto the superficial portion of adjoining caudally positioned follicles. Some fibers insert into the corium instead of inserting into a follicle. The fibers show a fine structure typical of red fibers. Smooth muscle cells form a network with elastic fibers beneath the corium. Some cells are directly attached to the capsule of the sinus, thus forming a type of M. arrector pili. Striated muscle fibers that appear to end in the corium are connected with the smooth muscle network through the elastic fibers which appear to function as the tendon of these two types of muscle cell.     

Neuromuscular relationships in the abdomen of the Californian shore crab Pachygrapsus crassipes

There are two pairs of muscles in each abdominal segment of the crab; one pair of flexors and one pair of extensors. In the early larval stages the muscles have short sarcomeres--a property of fast fibers--and high thin to thick filament ratios--a property of slow fibers. In the adult the abdominal muscles are intermediate and slow, since they have fibers with intermediate and long sarcomeres, high thin to thick filament ratios, low myofibrillar ATPase activity, and high NADH diaphorase activity. The different fiber types are regionally distributed within the flexor muscle. Microelectrode recordings from single flexor muscle fibers in the adult showed that most fibers are supplied by three excitatory motor axons, although some are supplied by as many as five efferents. One axon supplies all of the flexor muscle fibers in its own hemisegment, and the evoked junctional potentials exhibit depression. This feature together with the innervation patterns of the fibers are similar to those reported for the deep flexor muscles of crayfish and lobsters. Therefore, in the adult crab, the abdominal flexor muscles have some features in common with the slow superficial flexors of crayfish and other features in common with the fast deep flexor muscles.     

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.     

Carbonic anhydrase activity in skeletal muscle fiber types, axons, spindles, and capillaries of rat soleus and extensor digitorum longus muscles

Carbonic anhydrase (CA) activities were studied in soluble extracts and cryostat sections of skeletal muscles from prepubertal and postpubertal rats. Acetazolamide inhibition was utilized to distinguish between activities of the acetazolamide-sensitive (CA I and II) and acetazolamide-resistant (CA III) forms of the enzyme. The inhibition studies indicated that fast-twitch oxidative-glycolytic muscle fibers contained both the sensitive and resistant forms of CA. Acetazolamide-sensitive activity was localized within muscle fibers, axons, myelin, and capillaries. Axoplasmic staining was restricted to subpopulations of myelinated axons in both the dorsal and ventral roots. Soleus muscles exhibited significantly greater activity of CA III than extensor digitorum longus muscles at all ages examined. CA III was richest in slow-twitch oxidative and intrafusal fibers. During puberty, soleus muscle fibers matured and converted from fast-twitch oxidative-glycolytic to slow-twitch oxidative fibers. There was a shift from the sensitive to the resistant form of CA; CA III activity increased about sevenfold. This activity peaked earlier in the muscles of female rats than male rats. These results demonstrated a complex distribution of CA isozymes in the neuromuscular system and pointed out that isozyme content depends on both the type of muscle and the age and sex of the animal.     

Immunohistochemical and ultrastructural analysis of the muscular and nervous systems in the interstitial polychaete Polygordius appendiculatus (Annelida)

The systematic position of Polygordiidae is still under debate. They have been assigned to various positions among the polychaetes. Recent molecular analyses indicate that they might well be part of a basal radiation in Annelida, suggesting that certain morphological characters could represent primitive character traits adopted from the annelid stem species. To test this hypothesis, an investigation of the muscular and nervous systems by means of immunological staining and confocal laser scanning microscopy and transmission electron microscopy was conducted. With the exception of the brain, the nervous system is entirely basiepidermal and consists of the brain, the esophageal connectives, the subesophageal region, the ventral nerve cord and several smaller longitudinal nerves. These are connected by a considerable number of ring nerves in each segment. The ventral nerve cord is made up of closely apposed longitudinal neurite bundles, a median and two larger lateral ones. Since distinct ganglia are lacking, it represents a medullary cord. The muscular system mainly consists of longitudinal fibers, regularly distributed oblique muscles and strong septa. The longitudinal fibers form a right and a left unit separated along the dorsal midline, each divided into a dorsal and ventral part by the oblique muscles. Anteriorly, the longitudinal musculature passes the brain and terminates in the prostomium. There is no musculature in the palps. In contrast to earlier observations, regularly arranged minute circular muscle fibers are present. Very likely, a basiepithelial and non-ganglionic organization of the ventral nerve cord as well as an orthogonal nervous system represent plesiomorphic characters. The same applies for the predominance of longitudinal muscle fibers.     

Walking on inclines: how do desert ants monitor slope and step length

Background

In order to increase the weak database concerning the organogenesis of Acoela – a clade regarded by many as the earliest extant offshoot of Bilateria and thus of particular interest for studies concerning the evolution of animal bodyplans – we analyzed the development of the musculature of Symsagittifera roscoffensis using F-actin labelling, confocal laserscanning microscopy, and 3D reconstruction software.

Results

At 40% of development between egg deposition and hatching short subepidermal fibres form. Muscle fibre development in the anterior body half precedes myogenesis in the posterior half. At 42% of development a grid of outer circular and inner longitudinal muscles is present in the bodywall. New circular muscles either branch off from present fibres or form adjacent to existing ones. The number of circular muscles is higher than that of the longitudinal muscles throughout all life cycle stages. Diagonal, circular and longitudinal muscles are initially rare but their number increases with time. The ventral side bears U-shaped muscles around the mouth, which in addition is surrounded by a sphincter muscle. With the exception of the region of the statocyst, dorsoventral muscles are present along the entire body of juveniles and adults, while adults additionally exhibit radially oriented internal muscles in the anterior tip. Outer diagonal muscles are present at the dorsal anterior tip of the adult. In adult animals, the male gonopore with its associated sexual organs expresses distinct muscles. No specific statocyst muscles were found. The muscle mantles of the needle-shaped sagittocysts are situated along the lateral edges of the animal and in the posterior end close to the male gonopore. In both juveniles and adults, non-muscular filaments, which stain positively for F-actin, are associated with certain sensory cells outside the bodywall musculature.

Conclusion

Compared to the myoanatomy of other acoel taxa, Symsagittifera roscoffensis shows a very complex musculature. Although data on presumably basal acoel clades are still scarce, the information currently available suggests an elaborated musculature with longitudinal, circular and U-shaped muscles as being part of the ancestral acoel bodyplan, thus increasing the possibility that Urbilateria likewise had a relatively complicated muscular ground pattern.