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.     

Evolution de la musculature desNéréidiens (Annélides polychètes) au cours de l'Epitoquie

Résumé Au cours de l'épitoquie des Nereidiens, les fibres musculaires longitudinales ne sont pas forméesde novo, à partir de cellules indifferenciées ou myoblastes, mais proviennent des fibres anciennes atoques. Celles-ci subissent une véritable dédifférenciation plus ou moins synchrone d'une redifférenciation. Les deux processus ne sont pas successifs mais simultanés, et une dédifférenciation complète est absente.Les premières cellules en évolution appartiennent à la couche musculaire externe; ensuite, les fibres des assises plus profondes se transforment à leur tour.Les transformations consistent en: 1) La dédifférenciation du bord interne ou coelomique de la fibre. Les structures contractiles disparaissent dans cette zone et de nombreuses particules de glycogène se différencient sans relation avec le reticulum endoplasmique ou les ribosomes. Aucun lysosome ou signe précurseur ne peuvent être observés avant la disparition des filaments contractiles et des éléments Z. 2) Le bord coelomique s'hypertrophie. Dans la région axiale de la fibre, de nombreuses mitochondries et particules et de glycogène remplacent le matériel contractile. Corrélativement, l'épaisseur des bandes A et I diminue. 3) La fibre hétéronéreidienne ou épitoque est constituée et présente deux parties: un cortex myoplasmique et une médulla sarcoplasmique, remplie de mitochondries et de glycogène. Le noyau renfermant un nucléole volumineux est situé dans une hernie sarcoplasmique latérale.

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Evolution of muscles inNereidae (Annelida polychaeta) during Epitoky. III. Dedifferentiation of the longitudinal fibres

Summary During epitoky inNereidae, the longitudinal muscle fibres are not formedde novo from undifferentiated cells or myoblasts, but arise from the old atokous fibres. These undergo a true dedifferentiation more or less synchronously with a redifferentiation. The two processes are not successive but simultaneous and there is no complete dedifferentiation.The first cells that develop are in the outside muscle layer; then the fibres of the inside layers are transformed in their turn.The transformations consist of: 1) Dedifferentiation of the edge of the inner or coelomic fibre. The contractile structures disappear in this part and numerous glycogen particles differentiate, unrelated to endoplasmic reticulum or ribosomes. No lysosomes or precursory markings are observed before the disappearance of contractile filaments and Z rods. 2) The coelomic edge becomes enlarged. In the axial region of the fibre, numerous mitochondria and and glycogen particles take the place of the contractile material. Consequently, the thickness of A and I bands decreases. 3) The heteronereid or epitokous fibre is formed and shows two parts: a myoplasmic cortex and a sarcoplasmic medulla, filled with mitochondria and glycogen. The nucleus with a voluminous nucleolus settles inside a lateral sarcoplasmic swelling.

     

Ultrastructure of neuromuscular junctions in Drosophilal: Comparison of wild type and mutants with increased excitability

The ventral longitudinal muscles of the Drosphila larval body wall are innervated by at least four types of synaptic terminals that can be distinguished on morphological grounds at the light microscopical level. The innervation of these muscles has been previously shown to be regulated by neuronal activity. In this report we investigate the ultrastructural basis for synaptic bouton differences by using serial sections, and examine the structure of synaptic terminals in mutants with increased excitability. We report that individual identifiable muscle fibers are innervated by terminals containing two to three types of synaptic boutons that can be distinguished in terms of synaptic vesicle population, presynaptic and postsynaptic specialization, and general shape. We propose a model to account for the bouton types observed at the light microscopical level. We find that in the hyperexcitable mutant eag Sh, there are dramatic ultrastructural alterations at synaptic boutons. These alterations include a partial depletion of two types of synaptic vesicles and a change in appearance of a third type, changes in number and appearance of synaptic densities, and the presence of multivesicular bodies. Our results show that an increase in neuronal excitability produces profound effects in synaptic terminal structure. © 1993 John Wiley & Sons, Inc.     

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.     

Evolution of interstitial Polychaeta (Annelida)

An update of the systematics is given for the eight most important interstitial polychaete families: Diurodrilidae, Nerillidae, Protodrilidae, Protodriloididae, Saccocirridae, Parergodrilidae, Polygordidae and Psammodrilidae. Additional information and new observations are presented for the Diurodrilidae, Nerillidae and Psammodrilidae. Three new supplementary evolutionary hypotheses for these families are here suggested: (I) basal position of Diurodrilidae in Polychaeta, (2) evolution of Nerillidae in mud, and (3) evolution from meio- to macrofaunal forms of Psammodrilidae.     

Phylogenetic inference regarding Parergodrilidae and Hrabeiella periglandulata ('Polychaeta', Annelida) based on 18S rDNA, 28S rDNA and COI sequences

Parergodrilidae and Hrabeiella periglandulata are Annelida showing different combinations of clitellate-like and aclitellate characters. Similarities between both of these taxa and Clitellata have widely been regarded as the result of convergent evolution due to similar selection pressures. The position of the three taxa in the phylogenetic system of Annelida is still in debate. However, in analyses based on 18S rDNA sequences a close relationship of Parergodrilidae with Orbiniidae and Questidae was suggested. To infer their phylogeny the sequences of the 28S rDNA and of the cytochrome oxidase I (COI) gene of Stygocapitella subterranea , Parergodrilus heideri and H. periglandulata were determined. The data were extended by sequences of various species including species from Clitellata and Orbiniidae. Prior to tree reconstruction the dataset was analysed in detail for phylogenetic content by applying a sliding window analysis, a likelihood mapping and Modeltest V.3.04. Subsequently, generalized parsimony and maximum likelihood methods were employed. Clade robustness was estimated by bootstrapping. In addition, combined analyses of the sequences of 18S rDNA and 28S rDNA as well as of 18S rDNA, 28S rDNA and COI were performed. The combination of the data of the two structure genes and a mitochondrial gene improved the resolution obtained with the single datasets slightly. These analyses support a close relationship of Parergodrilidae and Orbiniidae but cannot resolve the position of H. periglandulata . In every analysis Clitellata cluster within 'Polychaeta', confirming previous investigations.     

Number and arrangement of extraocular muscles in primitive gnathostomes: evidence from extinct placoderm fishes

Exceptional braincase preservation in some Devonian placoderm fishes permits interpretation of muscles and cranial nerves controlling eye movement. Placoderms are the only jawed vertebrates with anterior/posterior obliques as in the jawless lamprey, but with the same function as the superior/inferior obliques of other gnathostomes. Evidence of up to seven extraocular muscles suggests that this may be the primitive number for jawed vertebrates. Two muscles innervated by cranial nerve 6 suggest homologies with lampreys and tetrapods. If the extra muscle acquired by gnathostomes was the internal rectus, Devonian fossils show that it had a similar insertion above and behind the eyestalk in both placoderms and basal osteichthyans.     

The loss of the lower temporal arcade in diapsid reptiles

The fossil record of diapsid reptiles does not provide conclusive evidence that the loss of the lower temporal arcade is correlated with the development of streptostyly. Analysis of the structure of the external jaw adductor in extant lizards results in the formulation of an alternative hypothesis to explain the loss of the lower temporal arcade in lepidosaurs. Extant lizards show the development of a posteroventral portion of the superficialis layer (lb) of the external adductor, which invades the lateral surface of the lower jaw, thus escaping the original confines of the external adductor. The development and expansion of a posteroventral lb muscle would have to be correlated with the loss of at least the posterior portion of the lower temporal arcade. Fossil lepidosaurs such as Clevosaurus, Prolacerta and Macrocnemus show the reduction of the lower temporal arcade from back to front, which is consistent with the hypothesis of the development of a posteroventral lb muscle.     

JUNCTIONAL PORES AND CELL WALL DEPRESSIONS OF HORMOSCILLA PRINGSHEIMII (CYANOPHYCEAE)1

Sassoon's isolate identified as Borzia trilocularis (but recently renamed Hormoscilla pringsheimii Anagnostidis and Komárek (1988) was studied with transmission electron microscopy because of its unusual combination of longitudinal wall features, described here in development for the first time. Junctional pores (linear rows of circumferential L-II layer pores near crosswalls) developed into multiple, parallel series, unlike the pores in many other species, which form only single rows. In dividing cells, two single pore rows appear opposite crosswalls after crosswall initiation, but an additional parallel row is usually added to each row by completion of fission. Elongating cells reveal 3–6 parallel and uniformly spaced pore rows developing on each side of crosswall pairs; these rows may end up toward the center of the cell wall after cell elongation. Pores are 18–24 nm wide with a center-to-center and row-to-row distance of 24–36 nm and occur in an especially thick L-II area. The second group of pit-like pores of longitudinal cell walls are 50–135 nm-wide depressions and have a center-to-center distance of 100–1000 nm. These depressions arise when the L-II layer fails to form and appear next to a row-pair of junctional pores soon after fission. Most depressions form single rows, but when they form several rows they may cover much of the surface of the cell. The L-III and L-IV wall layers line these L-II layer cavities; the outer surface of the L-IV layer around and within the depressions is covered with fibrous mucilage. Given their diversity, pores and depressions of longitudinal walls deserve further attention from functional and taxonomic points of view.     

Do life history traits account for diversity of polychaete annelids?

Abstract. Within many phylogenetic assemblages, a pattern of domination has been observed: one or a few clades have had many more speciation events or fewer extinctions than other clades in a particular assemblage. We investigated this phenomenon in the polychaete annelids. Polychaetes comprise ∼9000 described species classified in over 70 families and exhibit a great variety of life history strategies. Our goal was to test whether diverse polychaete families are characterized by species with short generation times, high reproductive output, small body size, or with planktotrophic larval development. Each of these factors has been advanced as cause for high diversity in other taxonomic assemblages. Here, we establish that the diversification pattern of polychaete families is non-random, but the data collected show no significant correlations between familial diversity and several life history traits including age at first reproduction, life span, body size, fecundity, and egg size. Pairwise comparisons of sister families do not reveal any trends between familial diversity and any of the life history traits. The great variability of life history traits within polychaete families may explain the lack of significant results; perhaps no trends are seen because polychaete life history traits cannot be generalized at the family level.     

The musculature of three species of gastrotrichs surveyed with confocal laser scanning microscopy (CLSM)

The muscular system of gastrotrichs consists of circular, longitudinal and helicoidal bands that when analysed with confocal laser scanning microscopy, provide new insights into their functional organization and phylogenetic importance. We therefore undertook a comparative study of the muscle organization in three species of Gastrotricha from the orders Macrodasyida (Paradasys sp., Lepidodasyidae; Turbanella sp., Turbanellidae) and Chaetonotida (Polymerurus nodicaudus, Chaetonotidae). The general muscle organization of the marine interstitial macrodasyidans, Paradasys and Turbanella, not only confirms earlier observation on other species but also adds new details concerning the organization and number of helicoidal, longitudinal and other muscle bands (e.g. semicircular band). The freshwater, epibenthic–epiphytic chaetonotid, Polymerurus nodicaudus, has a similar muscular organization to other species of Chaetonotidae, especially species of Chaetonotus, Halichaetonotus and Lepidodermella. Perhaps unique to Polymerurus is the combined presence of an unbranched Rückenhautmuskel (also in Halichaetonotus and Lepidodermella) and a specialized dorsoventral caudal muscle, which flank the splanchnic component of the longitudinal muscles (only in Chaetonotus and Lepidodermella). This combination, together with the presence of splanchnic dorsoventral muscles, known only in Xenotrichulidae, implies a unique phylogenetic position for Polymerurus, and indicates a potential basal position of this taxon among the Chaetonotidae studied so far (i.e. Aspidiophorus, Chaetonotus, Halichaetonotus and Lepidodermella).     

Muscular and osseous anatomy of the primate anterior temporal fossa and the functions of the postorbital septum

Many adaptive explanations for anthropoid origins incorporate hypotheses regarding the function of the postorbital septum. Two hypotheses are evaluated here: Cachel's ([1979b] Am. J. Phys. Anthropol. 50:1–18) hypothesis that the anthropoid postorbital septum evolved to augment muscle attachment area in the anterior temporal fossa and Cartmill's ([1980] in RL Ciochon and AB Chiarelli (eds.): Evolutionary Biology of the New World Monkeys and Continental Drift. New York: Plenum, pp. 243–274.) hypothesis that the septum evolved to insulate the foveate eye of haplorhines from movements in the temporal fossa during mastication. Dissections of the masticatory muscles of 55 species of primates, with emphasis on the anatomy of the anterior temporal fossa, reveal that in all anthropoids the temporal muscles take origin from the portion of the septum formed by the frontal bone. In some platyrrhines this muscle is anterior temporalis, and in others it is zygomatico-mandibularis. In tarsiers and most platyrrhines, muscle attachment to the zygomatic portion of the postorbital septum is very restricted (and of possibly varying homologies), whereas in catarrhines the zygomatico-mandibularis arises from the postorbital ridge on the zygomatic portion of the septum. This suggests that, contrary to Cachel's hypothesis, the earliest anthropoids did not have extensive areas of muscle attachment on the postorbital septum, a suggestion supported by the bony morphology of Catopithecus browni. Dissections also indicate that in all haplorhines the anteriormost temporal fibers curve around the postorbital septum between origin and insertion, implying that, were the septum not present, the anterior temporal muscles would disturb the orbital contents when contracting. This suggests that insulation may have been the septum's original function, even in the absence of a retinal fovea. In anthropoids, the rostral migration of the line of action of the anterior temporal muscles relative to the eye is attributed to their possession of extreme degrees of both orbital frontation and convergence; in tarsiers it is attributed to their possession of both massively hypertrophied eyes and moderately convergent and frontated orbits. It is argued that the postorbital septum is most likely to have evolved in a morphological context similar to that exhibited by omomyids. © 1995 Wiley-Liss, Inc.     

Electromyography of the gluteal muscles in Hylobates,Pongo, and pan: Implications for the evolution of hominid bipedality

The gluteal musculature of primates has been a focus of great research interest among those who study human evolution. Most current theorists agree that gluteus superficialis (= maximus) need not have changed its action in the step from pongid to hominid, but dispute has arisen over a purported change in action and role of the gluteus medius. To clarify the functions of gluteus medius, gluteus superficialis, and tensor fasciae femoris during ape locomotion, we conducted a telemetered electromyographic study of these muscles in two gibbons, one orangutan, and four chimpanzees as they walked bipedally on the ground and on a horizontal tree trunk, walked quadrupedally on the same substrates, and climbed a vertical tree trunk. The results indicate that the gluteus medius of apes is not, as has been previously suggested, primarily an extensor of the thigh; its action is chiefly that of medial rotation. The role of the gluteus medius during bipedality is the same in apes and humans–to provide side-to-side balance of the trunk at the hip. The change in the hominid lateral balance mechanism can be viewed as primarily osteological, allowing preservation of the same muscle function with an extended thigh. As a result, the stride length is increased and there occurs a diminution of the demands placed on other muscles to maintain anteroposterior balance at the hip and knee. Our data also support the view that vertical climbing may be specifically preadaptive to bipedalism. One may picture the earliest hominid as part biped, when on the ground traveling between scattered food trees, and part climber, when moving from the ground to food.