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.     

Evolution of body wall musculature

A body wall musculature comprising an outer layer of circularfibers and an inner layer of longitudinal fibers is generallyseen as the basic plan in Annelida. Additional muscles may bepresent such as oblique, parapodial, chaetal, and dorsoventralmuscles. The longitudinal muscle fibers do not form a continuouslayer but are arranged in distinct bands in polychaetes. Mostlythere are four to six bands, usually including prominent ventraland dorsal bands. However, other patterns of muscle band arrangementalso exist. The ventral nerve cord lies between the two ventralbands in certain polychaetes, and is covered by an additionallongitudinal muscle band of comparatively small size. In manypolychaetes with reduced parapodia and in Clitellata a moreor less continuous layer of longitudinal fibers is formed. Clitellatais the only group with a complete layer of longitudinal musculature.Circular fibers are usually less developed than the longitudinalmuscles. However, recent investigations employing phalloidinstaining in combination with confocal laser scanning microscopyrevealed that absence of circular muscles is much more widelydistributed within the polychaetes than was previously known.This necessitates thorough reinvestigations of polychaete musclesystems, and this feature has to be taken into account in furtherdiscussions of the phylogeny and evolution of Annelida.     

Muscular system in polychaetes (Annelida)

The structure of the polychaete muscular system is reviewed. The muscular system comprises the muscles of the body wall, the musculature of the parapodial complex and the muscle system of the dissepiments and mesenteries. Various types of organisation of the longitudinal and circular components of the muscular body wall are distinguished. In Opheliidae, Polygordiidae, Protodrilidae, Spionidae, Oweniidae, Aphroditidae, Acoetidae (=Polyodontidae), Polynoidae, Sigalonidae, Phyllodocidae, Nephtyidae, Pisionidae, and Nerillidae circular muscles are lacking. It is hypothesised that the absence of circular muscles represents the plesiomorphic state in Annelida. This view contradicts the widely accepted idea of an earthworm-like musculature of the body wall comprising an outer layer of circular and an inner layer of longitudinal fibres. A classification of the various types of parapodial muscle construction has been developed. Massive and less manoeuvrable parapodia composed of many components like those of Aphrodita are regarded to represent the plesiomorphic state in recent polychaetes. An analysis of the diversity of the muscular structure supports the hypothesis that the primary mode of life in polychaetes was epibenthic and the parapodial chaetae had a protective function.     

On the absence of circular muscle elements in the body wall of Dysponetus pygmaeus (Chrysopetalidae, 'Polychaeta', Annelida)

The annelid body wall generally comprises an outer layer of circular muscle fibres and an inner layer of longitudinal muscle fibres as well as parapodial and chaetal muscles. An investigation of Dysponetuspygmaeus (Chrysopetalidae) with confocal laser scanning microscopy showed that circular muscles are entirely absent. Further studies indicate that this feature is characteristic for all Chrysopetalidae. A scrutiny of the literature showed a similar situation in many other polychaetes. This lack of circular muscle fibres may either be due to convergence or represent a plesiomorphic character. Since circular muscles are very likely important for burrowing forms but not necessary for animals which proceed by movements of their parapodial appendages or cilia, this problem is also related to the question of whether the ancestral polychaete was epi‐ or endobenthic.     

Functional morphology of musculature in the acoelomate worm, Convoluta pulchra (Plathelminthes)

Convoluta pulchra is a small worm living in the surface sediment of mud flats where it feeds on diatoms. It is roughly teardrop in shape with a ventral groove in which the mouth sits, and it can move in a variety of ways, readily distorting its body in bending, twisting, and turning motions. Fluorescently labeled probes for filamentous actin revealed the musculature in whole mounts of the worm. In the body wall, the musculature consisted of a grid of circular, longitudinal crossover (that is, with a longitudinal orientation in the anterior half of the body but arcing medially to cross over to the contralateral side of the body behind the level of the mouth), and a few diagonal fibers. Inside the body was a strong, irregular brush of muscles originating at the rostral tip of the body and anchoring laterally and posteriorly along the body wall, and strong dorsoventral muscles flanked the ventral groove. Two fans of muscles in the ventral and dorsal body wall reached posteriorly and laterally; that on the dorsal side originated at junctures of the dorsoventral muscles with the body wall and that on the ventral body wall originated from the mouth. By their positions, certain groups of muscles could be correlated with given movements: the crossover muscles with some turning motions and feeding, and the inner muscles with probing and retraction motions of the rostrum and with a tuck-and-turn motion the worm used to turn itself around. Electron microscopy showed numerous maculae adherentes junctions linking all muscle types and special junctions linking the musculature with the epidermis. The latter myoepidermal junctions were of dimensions larger than those of maculae adherentes and contained an interlaminar material which we believe represents islands of basal matrix comparable to basement membrane. Accepted: 12 July 1999     

Pattern of body-wall muscle differentiation during embryonic development of Enchytraeus coronatus (Annelida: Oligochaeta; Enchytraeidae)

The plesiomorphic arrangement of body-wall musculature within the annelids is still under discussion. While polychaete groups show a great variety of patterns in their somatic muscles, the musculature of soil-living oligochaetes was thought to represent the characteristic pattern in annelids. Oligochaete body-wall muscles consist of an outer continuous layer of circular and an inner continuous layer of longitudinal muscles, forming a closed tube. Since designs of adult body musculature are influenced by evolutionary changes, additional patterns found during embryogenesis can give further information about possible plesiomorphic features. In oligochaetes, detailed cell-lineage analyses document the origin of the mesoderm and consequently the muscles, but later processes of muscle formation remain unclear. In the present work, body-wall muscle differentiation was monitored during embryogenesis of thesoil-living oligochaete Enchytraeus coronatus (Annelida) by phalloidin staining. Primary circular muscles form in a discrete anterior-to-posterior segmental pattern, whereas emerging longitudinal muscles are restricted to one ventral and one dorsal pair of primary strands, which continuously elongate towards posterior. These primary muscles establish an initial muscle-template. Secondary circular and longitudinal muscles subsequently differentiate in the previous spaces later in development. The prominent ventral primary longitudinal muscle strands on both sides eventually meet at the ventral midline due to neurulation, which moves the ventral nerve cord into a coelomic position, closing the muscle layers into a complete tube. This early embryonic pattern in E. coronatus resembles the adult body-wall muscle arrangements in several polychaete groups as well as muscle differentiation during embryonic development of the polychaete Capitella sp. I.     

Myoanatomy of Myzostoma cirriferum (Annelida,Myzostomida): Implications for the evolution of the myzostomid body plan

Studies of rare genomic marker systems suggest that Myzostomida are a subgroup of Annelida and phylogenomic analyses indicate an early divergence of this taxon within annelids. However, adult myzostomids show a highly specialized body plan, which lacks typical annelid features, such as external body annulation, coelomic cavities with metanephridia, and segmental ganglia of the nervous system. The putative loss of these features might be due to the parasitic/symbiotic lifestyle of myzostomids associated with echinoderms. In contrast, the larval anatomy and adult locomotory system resemble those of annelids. To clarify whether the myoanatomy of myzostomids reflects their relationship to annelids, we analyzed the distribution of f‐actin, a common component of muscle fibers, in specimens of Myzostoma cirriferum using phalloidin‐rhodamine labeling in conjunction with confocal laser‐scanning microscopy. Our data reveal that the musculature of the myzostomid body comprises an outer circular layer, an inner longitudinal layer, numerous dorsoventral muscles, and prominent muscles of the parapodial complex. These features correspond well with the common organization of the muscular system in Annelida. In contrast to other annelids, however, several elements of the muscular system in M. cirriferum, including the musculature of the body wall, and the parapodial flexor muscles, exhibit radial symmetry overlaying a bilateral body plan. These findings are in line with the annelid affinity of myzostomids and suggest that the apparent partial radial symmetry of M. cirriferum arose secondarily in this species. Based on our data, we provide a scenario on the rearrangements of muscle fibers that might have taken place in the lineage leading to this species. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

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.     

Organization of the dorsoventral musculature in the wings of the pteropod mollusc Clione limacina

The wings of the pteropod mollusc Clione limacina provide forward propulsive force through flapping movements in which the wings bend throughout their length in both dorsal and ventral directions. The musculature of the wings includes oblique, striated muscle bundles that generate the swimming movements of the wings, longitudinal and transverse (smooth) muscle bundles that collapse the wings and pull them into the body during a wing withdrawal response, and dorsoventral muscles that control the thickness of the wings. All muscles act against a hydrostatic skeleton that forms a central hemocoelic space within the wings. Of these muscle types, all have been thoroughly described and studied except the dorsoventral muscles. The fortuitous discovery that the dorsoventral musculature can be intensely labeled with an antibody against the vertebrate hyperpolarization‐activated cation channel (HCN2) provided the opportunity to describe the organization of the dorsoventral musculature in detail. In addition, electrical recordings and microelectrode dye injections supported the immunohistochemical data, and provided preliminary data on the activity of the muscle fibers. The organization and activity of the dorsoventral musculature suggests it may be involved in regulation of wing stiffness during the change from slow to fast swimming.     

Immunocytochemistry of the nervous system and the musculature of the chordoid larva of Symbion pandora (Cycliophora)

To date, the phylum Cycliophora comprises only one described extant species of acoelomate marine invertebrates, Symbion pandora. Adult specimens live commensally on the mouthparts of the Norwegian lobster, Nephrops norvegicus. Its complicated life cycle includes an asexually produced Pandora larva and a sexually produced chordoid larva. Despite detailed TEM investigations and its inclusion in recent molecular phylogenetic analyses, cycliophoran relationships still remain enigmatic. In order to increase the morphological database, I investigated the anatomy of the nervous system and the musculature of the chordoid larva by applying fluorescence-coupled antibodies against the neurotransmitters serotonin and FMRFamide, as well as FITC-coupled phalloidin to label filamentous F-actin, in combination with confocal laser scanning microscopy. The FMRFamidergic nervous system shows a bilobed anterior ganglion and one pair of ventral nerve cords, while serotonin is distributed in a scattered pattern in the anterior ganglion. In addition, there are two pairs of ventral serotonergic nerves, of which the inner pair fuses with the outer nerve cords in the posterior third of the larva. The musculature comprises an outer layer of six units of circular body wall muscles, several helicoid muscle fibers, a set of paired longitudinal muscles that span the entire anterior-posterior axis of the larva, and a few oblique muscle strands. Furthermore, an anterior muscle complex and one pair of posterior muscles are present. The chordoid organ consists of a number of distinct subunits that are each formed by a dense layer of circular muscle fibers.The overall arrangement of the oblique and longitudinal muscles as well as the body wall musculature in the chordoid larva of Symbion pandora exhibits similarities with the condition found in certain rotifers. This is congruent with some recent phylogenies based on 18S rRNA sequences but additional morphological, developmental, and molecular data are needed to clarify the phylogenetic relationships of Cycliophora.     

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.     

Comparative myoanatomy of cycliophoran life cycle stages

The metazoan phylum Cycliophora includes small cryptic epibionts that live attached to the mouthparts of clawed lobsters. The life cycle is complex, with alternating sexual and asexual generations, and involves several sessile and free‐living stages. So far, the morphological and genetic characterization of cycliophorans has been unable to clarify the phylogenetic position of the phylum. In this study, we add new details on the muscular anatomy of the feeding stage, the attached Prometheus larva, the dwarf male, and the female of one of the two hitherto described species, Symbion pandora. The musculature of the feeding stage is composed of myofibers that run longitudinally in the buccal funnel (two fibers) and in the trunk (variable number of fibers). The mouth opening is lined by a myoepithelial ring musculature. A complex myoepithelial sphincter is situated proximal to the anus. In the attached Prometheus larva, three longitudinal sets of myofilaments run dorsally, laterally, and ventrally along the entire anterior‐posterior body axis. The muscular architecture of the dwarf male is complex, especially close to the penis, in the posterior part of the body. An X‐shaped muscle structure is found on the dorsal side, whereas on the ventral side, longitudinal muscles and a V‐shaped muscle structure are present. These muscles are complemented by additional dorsoventral muscles. The mesodermal muscle fibers attach to the cuticle via the epidermis in all life cycle stages studied herein. The musculature of the female is similar to that of the Pandora larva of Symbion americanus and includes dorsoventral muscles and longitudinal muscles that run in the dorsal and ventral body region. Overall, our results reveal striking similarities in the muscular arrangement of the life cycle stages of both Symbion species. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

The organization of the muscle system of the causative agent of dicrocoeliosis,Dicrocoelium dendriticum

The musculature of parasitic flatworms plays a central role in locomotory movement, attachment to the host, and in the function of the digestive, reproductive, and excretory systems. We examine for the first time the muscle system of the flatworm Dicrocoelium dendriticum, a causative agent of the parasitic disease dicrocoeliosis, by use of fluorescently labeled phalloidin and confocal laser scanning microscopy. Somatic musculature of D. dendriticum consists of the circular, longitudinal, and diagonal muscles. The distribution of the muscle fibers in the body wall differed among the anterior, middle, and posterior body regions of the worm. The musculature of the attachment organs, the oral and ventral suckers, includes several types of muscles: the external equatorial and meridional muscles, internal circular and semicircular muscles, and radial muscles. Inside of the ventral sucker the diagonally located muscles were revealed and the supplementary u-shaped muscles were found adjoined to the base of the sucker from outside. The musculature of the internal organs composed of the excretory, reproductive, and digestive systems were characterized. Our results increase our knowledge of the morphology of trematodes and the arrangement of their muscle system.     

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.     

Functional morphology of the muscles in Philodina sp. (Rotifera: Bdelloidea)

Whole-mounts of Philodina sp., a bdelloid rotifer, were stained with fluorescent-labeled phalloidin to visualize the musculature. Several different muscle types were identified including incomplete circular bands, coronal retractors and foot retractors. Based on the position of the larger muscle bands in the body wall, their function during creeping locomotion and tun formation was inferred. Bdelloid creeping begins with the contraction of incomplete circular muscle bands against the hydrostatic pseudocoel, resulting in an anterior elongation of the body. One or more sets of ventral longitudinal muscles then contract bringing the rostrum into contact with the substrate, where it presumably attaches via adhesive glands. Different sets of ventral longitudinal muscles, foot and trunk retractors, function to pull the body forward. These same longitudinal muscle sets are also used in `tun' formation, in which the head and foot are withdrawn into the body. Three sets of longitudinal muscles supply the head region (anterior head segments) and function in withdrawal of the corona and rostrum. Two additional pairs of longitudinal muscles function to retract the anterior trunk segments immediately behind the head, and approximately five sets of longitudinal retractors are involved in the withdrawal of the foot and posterior toes. To achieve a greater understanding of rotifer behavior, it is important to elucidate the structural complexity of body wall muscles in rotifers. The utility of fluorescently-labeled phalloidin for the visualization of these muscles is discussed and placed in the context of rotifer functional morphology.     

Onychophoran‐like myoanatomy of the Cambrian gilled lobopodian Pambdelurion whittingtoni

Arthropods are characterized by a rigid, articulating, exoskeleton operated by a lever‐like system of segmentally arranged, antagonistic muscles. This skeletomuscular system evolved from an unsegmented body wall musculature acting on a hydrostatic skeleton, similar to that of the arthropods’ close relatives, the soft‐bodied onychophorans. Unfortunately, fossil evidence documenting this transition is scarce. Exceptionally‐preserved panarthropods from the Cambrian Lagerstätte of Sirius Passet, Greenland, including the soft‐bodied stem‐arthropod Pambdelurion whittingtoni and the hard‐bodied arthropods Kiisortoqia soperi and Campanamuta mantonae, are unique in preserving extensive musculature. Here we show that Pambdelurion's myoanatomy conforms closely to that of extant onychophorans, with unsegmented dorsal, ventral and longitudinal muscle groups in the trunk, and extrinsic and intrinsic muscles controlling the legs. Pambdelurion also possesses oblique musculature, which has previously been interpreted as an arthropodan characteristic. However, this oblique musculature appears to be confined to the cephalic region and first few body segments, and does not represent a shift towards arthropodan myoanatomy. The Sirius Passet arthropods, Kiisortoqia and Campanamuta, also possess large longitudinal muscles in the trunk, although, unlike Pambdelurion, they are segmentally divided at the tergal boundaries. Thus, the transition towards an arthropodan myoanatomy from a lobopodian ancestor probably involved the division of the peripheral longitudinal muscle into segmented units.     

Evolutionary aspects of pattern formation during clitellate muscle development

As a taxon of the lophotrochozoans, annelids have re-entered scientific investigations focusing on plesiomorphic bilaterian features and the evolutionary changes therein. The view of a clitellate-like plesiomorphic muscle arrangement in annelids has been challenged by recent investigations of polychaete muscle organization. However, there are few investigations of muscle formation in clitellate species that address this problem. Direct comparison of potential homologous muscles between these annelid groups is thus hampered. Somatic muscle formation during embryogenesis of two clitellates-the oligochaete Limnodrilus sp. and the hirudinean Erpobdella octoculata-occurs by distinct processes in each species, even though they share a closed outer layer of circular and an inner layer of longitudinal muscles characteristic of clitellates. In E. octoculata, the first emerging longitudinal muscles are distributed irregularly on the body surface of the embryo whereas the circular muscles appear in an orderly repetitive pattern along the anterioposterior axis. Both primary muscle types consist of fiber-bundles that branch at both their ends. This way the circular muscle bundles divide into a fine muscle-grid. The primary longitudinal muscles are incorporated into a second type of longitudinal muscles, the latter starting to differentiate adjacent to the ventral nerve cord. Those secondary muscles emerge in a ventral to dorsal manner, enclosing the embryo of E. octoculata. In Limnodrilus sp., one dorsal and one ventral bilateral pair of primary longitudinal muscles are established initially, elongating toward posterior. Initial circular muscles are emerging in a segmental pattern. Both muscle layers are completed later in development by the addition of secondary longitudinal and circular muscles. Some features of embryonic longitudinal muscle patterns in Limnodrilus sp. are comparable to structures found in adult polychaete muscle systems. Our findings show that comparative studies of body-wall muscle formation during clitellate embryogenesis are a promising approach to gain further information on annelid muscle arrangements.     

Muscle formation during embryogenesis of the polychaete Ophryotrocha diadema (Dorvilleidae) – new insights into annelid muscle patterns

Background

The standard textbook information that annelid musculature consists of oligochaete-like outer circular and inner longitudinal muscle-layers has recently been called into question by observations of a variety of complex muscle systems in numerous polychaete taxa. To clarify the ancestral muscle arrangement in this taxon, we compared myogenetic patterns during embryogenesis of Ophryotrocha diadema with available data on oligochaete and polychaete myogenesis. This work addresses the conflicting views on the ground pattern of annelids, and adds to our knowledge of the evolution of lophotrochozoan taxa.

Results

Somatic musculature in Ophryotrocha diadema can be classified into the trunk, prostomial/peristomial, and parapodial muscle complexes. The trunk muscles comprise strong bilateral pairs of distinct dorsal and ventral longitudinal strands. The latter are the first to differentiate during myogenesis. They originate within the peristomium and grow posteriorly through the continuous addition of myocytes. Later, the longitudinal muscles also expand anteriorly and form a complex arrangement of prostomial muscles. Four embryonic parapodia differentiate in an anterior-to-posterior progression, significantly contributing to the somatic musculature. Several diagonal and transverse muscles are present dorsally. Some of the latter are situated external to the longitudinal muscles, which implies they are homologous to the circular muscles of oligochaetes. These circular fibers are only weakly developed, and do not appear to form complete muscle circles.

Conclusion

Comparison of embryonic muscle patterns showed distinct similarities between myogenetic processes in Ophryotrocha diadema and those of oligochaete species, which allows us to relate the diverse adult muscle arrangements of these annelid taxa to each other. These findings provide significant clues for the interpretation of evolutionary changes in annelid musculature.     

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

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

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.