Ultrastructural Investigation of the Integument of Tubiluchus philippinensis (Priapulida, Tubiluchidae)

The integument of Tubiluchus philippinensis van der Land, 1984 has been investigated by means of scanning and transmission electron microscopy. The ultrastructure of the cuticle corresponds principally to what has been found in Priapulidae. The tumuli are mere cuticular thickenings. Setae, tubuli, flosculi and scalids are receptor organs. Tubuli additionally serve a second function: they produce a secretion. The male genital area is equipped with various receptor organs, the internal morphology of which has been described. All receptor cells are characterized by apical cilia, which may be surrounded by a circlet of microvilli. They sometimes bear a rather complicated rootlet apparatus.     

Scanning and Transmission Electron Microscopical Observations on the Larvae of Priapulus caudatus (Priapulida)

Abstract Scanning electron microscopical studies revealed four distinct morphological larva-types (not instars) of Priapulus caudatus whose lorica-length measured 82–860 μm. The smallest of the larvae are round to oval, have 20 longitudinal ridges, a series of transverse ridges, and have two pairs of laterally situated tubuli near the posterior limits. The second larva-type is dorsoventrally compressed, has a single dorsal and single unsculptured ventral plate each with a prominent midridge near the anterior limits, three infolded lateral plates with a pair of tubuli near the posterior limits. The third and fourth larva-types remain dorsoventrally flattened; the third larva-type has less pronounced sculpturing than the fourth larva-type and is smaller; two pairs of tubuli are situated as in the previous stage. The number and arrangement of scalids on the introvert and pharyngeal teeth differ according to each of the four larva-types and are described. The ultrastructure of all organ systems is described. All scalids are sensilla equipped with ciliated receptor cells. The cuticle is similar to the cuticle in the adult and differs fundamentally from the cuticle of larval Halicryptus. The trunk is richly supplied with sensory structures. Trunk tubuli contain secretory cells. Posteriorly, a gland complex was found which presumably is the precursor of the equivalent gland in postlarvae and adults. The implications of the first larva-type's structural similarity to the larva of Tubiluchus are discussed.     

Microscopic anatomy and ultrastructure of the introvert of Priapulus caudatus and P. tuberculatospinosus (Priapulida)

Introverts of Priapulus caudatus and P. tuberculatospinosus bear 25 rows of scalids, as well as 8 spines and scattered papillae in the region the circumoral lip. These, as well as the first ring of pharyngeal teeth in P. tuberculatospinosus, are sensory organs. Although superficially they differ between species, they are all characterized by apical and/or subapical openings which are located on tiny cuticular tubules. All sensory organs contain cilia bearing bipolar receptor cells. The 8 sensory spines situated between the circumoral area and the beginning of the scalids are ultrastructurally similar to the scalids. The introvert and pharyngeal body walls, and associated muscles are described. © 1994 Wiley-Liss, Inc.     

Morphology and ultrastructure of the sensory appendages of a kinorhynch introvert

The structure and arrangement of appendages (scalids) on the head of the homalorhagid kinorhynch Kinorhynchus phyllotropis Brown & Higgins, 1983 are named, described and illustrated. In adults of this species, seven rings of external scalids are separated by segment boundary structures from the oral styles and three rings of internal scalids. All of these appendages contain ciliated receptor cells which pass to pores at the scalid tips, and all but the two anterior rings are jointed. All of these appendages are radially arranged in multiples of five, and closely associated with the ten lobed circumoral nerve ring. The most posterior scalid ring consists of fourteen trichoscalids, of which six are longer than the other eight. The six longer trichoscalids are arranged in bilateral symmetry, two to each of the three facets of the body. Similarities between scalid arrangement in Kinorhyncha and Loricifera are discussed.     

Hatching and earliest larval stages of the priapulid worm Priapulus caudatus

Abstract. Here we describe the hatching and morphology of the earliest larval stages of the priapulid worm Priapulus caudatus for the first time. The hatching larva differs considerably from previously described larvae not only in its general body shape but also in its lack of a proper lorica including the typical lorica tubuli. Furthermore, no mouth opening or pharyngeal teeth have formed as yet, and the number and arrangement of scalids differ from that of later larvae. The hatching larva molts and emerges as the first lorica larva. This larva partially resembles earlier described lorica larvae, but there are a number of important differences; the first lorica larva is smaller, and the mouth opening as well as pharyngeal teeth are still yet to form. The second lorica larva is equipped with four rings of pharyngeal teeth; it shows striking similarity to the previously described larva of P. caudatus , i.e., the larva-type 2 , only differing in the scalid pattern. We conclude that the first two larval stages of P. caudatus have not been described previously. We suggest that discrepancies between the earliest lorica larvae described here and in earlier publications might depend on sub-speciation or ecophenotypic modification of larvae collected from different localities. Our findings highlight the importance of studying the development of non-model organisms such as priapulids under controlled laboratory conditions.     

Ultrastructure of introvert and pharynx of Halicryptus spinulosus (priapulida)

The introvert of Halicryptus spinulosus bears three kinds of sensilla: buccal papillae, ordinary scalids, and dentoscalids. They are all characterized by bipolar monociliary receptor cells. The former two have apical openings at which the sensory cilia are in close contact with the ambient sea water. The pharyngeal teeth are composed of slender epithelial cells the tips of which are devoid of organelles and a thick cuticle. The anatomy of the muscle arrangement of the pharynx is described. Glands occur at the junction of the pharynx and midgut.     

Rugiloricus bacatus sp. nov. (Loricifera ‐Pliciloricidae) and a ghost‐larva with paedogenetic reproduction

Abstract

A new species of Loricifera, Rugiloricus bacatus sp. nov. is described together with the diagnoses of two other Rugiloricus species, Rugiloricus sp. nov. A and B, from the Faroe Bank (North Atlantic). Characteristic for all three species is the presence of a new type of reduced larva, a ghost‐larva. This type of reduced larva was discovered in 1986 by Jeanne Renaud‐Mornant, but it was with the Faroe Bank material that it was first discovered that the ghost‐larvae belonged to the phylum Loricifera. The ghost‐larvae are eitherfound inside late instar Higgins‐larvae, called penultimate Higgins‐larvae, or in the sediment. The three types of Higgins‐larvae from the Faroe Bank can be distinguished by characters such as anterior setae, posterior setae and toes. The adults of Rugiloricus bacatus sp. nov. are characterised by a prominent ruff resembling a pearl necklace; two of the eight clavoscalids are modified in the 1st row; the 2nd row of leg‐shaped scalids are very large and robust, and the 9th row with 30 beak‐like scalids alternating with 30 alternating plates. The postlarvae are free‐living and their scalids on the introvert are reduced to protoscalids. Postlarvae and adult stages have not been found for Rugiloricus sp. nov. A and B and therefore only diagnoses of these two species are presented here.     

Ultrastructure of multiciliated collar cells in the pilidium larva of Lineus bilineatus (Nemertini)

Summary The ultrastructure of the apical plate of the free-swimming pilidium larva of Lineus bilineatus (Renier 1804) is described with particular reference to the multiciliated collar cells. In the multiciliary collar cells there are several, up to 12, cilia surrounded by a collar of about 20 microvilli extending from the cells' apical surface. The cilia have the typical 9+2 axoneme arrangement and are equipped with striated caudal rootlets extending from the basal bodies. No accessary centriole or rostral rootlet were observed. Microvilli surrounding the cilia are joined in a cylindrical manner by a mucus-like substance to form a collar. In comparison with many sensory receptor cells built on a collar cell plan the multiciliary collar cells of the pilidium larva apical plate are rather simple and unspecialized. In other pilidium larvae monociliated collar cells are found in the apical plate. The possible function and phylogenetic implications of multiciliated collar cells in Nemertini are briefly discussed.List of Abbreviations a axoneme - b basal body - c cilia or flagella - d desmosome - G Golgi apparatus - m mitochondria - mf microfilaments - mu mucus - mv microvilli - n nucleus - nt neurotubules - pm plasma membrane - r rootlet - ri ribosomes - v secretory vesicles     

Ultrastructural observations of the larva ofTubiluchus corallicola (Priapulida)

Larvae ofTubiluchus corallicola van der Land 1968 were investigated by scanning and transmission electron microscopy. The scalids are sensory organs, each has a bipolar receptor cell with a single apical cilium similar to the scalid in the adult. Muscle cells of the larva are more differentiated than previously reported for other Priapulida; the larval arrangement of circular and longitudinal musculature differs from that of the adult, and a diaphragm is reported for the first time in Priapulida. The diaphragm may function in hydrostatic control of eversion and inversion of the introvert and mouth cone. The functional morphology of these two structures is discussed and contrasted with the Kinorhyncha.     

Symmetry of priapulids (Priapulida). 1. Symmetry of adults

Priapulids possess a radial symmetry that is remarkably reflected in both external morphology and internal anatomy. It results in the appearance of 25-radial (a number divisible by five) symmetry summarized as a combination of nonaradial, octaradial, and octaradial (9+8+8) symmetries of scalids. The radial symmetry is a secondary appearance considered as an evolutionary adaptation to a lifestyle within the three-dimensional environment of bottom sediment. The eight anteriormost, or primary, scalids retain their particular position because of their innervation directly from the circumpharyngeal brain. As a result of a combination of the octaradial symmetry of primary scalids, pentaradial symmetry of teeth, and the 25-radial symmetry of scalids, the initial bilateral symmetry remains characterized by the single sagittal plane.     

The development of the stomach in Clarias lazera and the intestinal absorption of protein macromolecules

Summary The development of the stomach of the teleost, Clarias lazera, during the early posthatching period, is described, and the developing stomach is compared with that of adult Clarias.The stomach develops in two distinct parts: the corpus, which differentiates first, and the pylorus. The corpus contains a mucous surface epithelium, arranged in folds, and a tubular gland system containing only one type of gland cell, to which the secretion of pepsinogen and HCl is attributed. The pyloric region does not contain tubular glands.From the ultrastructure of the gland cells, the ³H-thymidine labeling index, and the onset of acid production (as determined with pH indicators) it is concluded that a functional stomach is present in juveniles with a standard length of ± 11 mm (approximately 12 days after fertilization at 23–24° C).The ultrastructure of the intestinal epithelium has also been studied. The intestine consists of three segments, similar to those described for stomachless teleosts and a number of fish larvae. In larvae as well as in juveniles, the enterocytes of the second segment show pinocytosis of horseradish peroxidase, although in the juveniles the stomach has already developed. This second segment has the same relative length in all studied larvae and juveniles and is also present in adult Clarias.It is therefore concluded that the capacity to absorb protein macromolecules is not specifically related to the absence of a functional stomach in this teleost species.     

Introvert,mouth cone,and nervous system of Echinoderes capitatus (Kinorhyncha,Cyclorhagida) and implications for the phylogenetic relationships of Kinorhyncha

Summary The introvert, mouth cone, and nervous system of Echinoderes capitatus were examined by transmission and scanning electron microscopy. The introvert bears seven rings of primarily quincunxial sensory scalids, including type 1 and 2 spinoscalids as well as trichoscalids; the latter two types are additionally provided with glandular cells. The mouth cone bears one ring of decamerous sensory oral styles and three rings of quincunxial sensory pharyngeal styles. The intra- to basiepithelial, bilateral nervous system consists of a circumentric nerve ring in the introvert, a terminal and proximal nerve ring in the mouth cone, a ventral chain of ganglia, one in each trunk zonite, and a caudal ganglion. The introvert, the neck, and the trunk zonites are innervated from the forebrain; the mouth cone and the pharyngeal bulb are innervated from the hindbrain. The monophyly of the Kinorhyncha is based upon the following autapomorphic characters: (1) a mouth cone, (2) a neck with 16 placids, (3) a trunk with 11 zonites, (4) scalids of three types: type 1 and type 2 spinoscalids, and trichoscalids, (5) an anteriormost ring of ten type 1 spinoscalids (sensory organs divided into a basal and a terminal part), (6) a posteriormost ring of 14 trichoscalids (glandular sensory organs which are undivided), (7) rings in between the anteriormost and posteriormost are type 2 spinoscalids (glandular sensory organs divided into a basal and a terminal part), (8) a mouth cone with a terminal and a proximal nerve ring, (9) nine sensory oral styles with decamerous symmetry (the dorsal style is missing) and (10) three rings of sensory pharyngeal styles with, from anterior to posterior, ten, five, and five styles with quincunxial arrangement. The following characters are assumed to be autapomorphic for the taxon Nematoda+Gastrotricha+Kinorhyncha+Loricifera+Priapulida: (1) a basiepithelial circumentric brain and (2) a neuropileous nerve ring in a subterminal position. The following characters are assumed to be autapomorphic for the taxon Kinorhyncha+Loricifera+Priapulida: (1) a neuropileous nerve ring in a terminal position, (2) an introvert with scalids, (3) an eversible foregut and (4) tanycytes.The unpublished doctoral thesis of B. Neuhaus [1991 Zur Ultrastruktur, Postembryonalentwicklung und phylogenetischen Verwandtschaft der Kinorhyncha. PhD thesis. University of Götingen, Germany] was finished simultaneously with the completion of this study.     

The presence of two populations of sensory-type cells in the pineal organ of the five-bearded rockling,Ciliata mustela L. (Teleostei)

The pineal organ of the five-bearded rockling, Ciliata mustela L., was examined by means of electron microscopy. Two categories of sensory cells are described: 1) Sensory cells 1 (or photoreceptor cells sensu stricto) showing the characteristic ultrastructure of photoreceptor cells with a well-developed receptor pole (outer segment) and a transmitter pole (ribbon-type synapse in the basal pedicle contacting dendritic processes), and a segmental organization of organelles. 2) Sensory cells 2 (or photoneuroendocrine cells) displaying no particular segmentation. The ultrastructure of the receptor pole (outer segment) is variable in shape (with either long or short disks) and in the number of disks; some outer segments are simple cilia of the 9 + 0 type. This second cell category is rich in smooth endoplasmic reticulum, beta-particles of glycogen, dense inclusions of variable size and content, and dense-core vesicles 130 nm in diameter. These cells have an extended contact area with the perivascular space. The functional significance of both cell categories is discussed in terms of the known physiological responses of the pineal organ. A possible confusion in identification of interstitial cells and neuroendocrine cells in some teleost species is discussed.     

Spinitectus acipenseri n. sp. (Nematoda: Cystidicolidae) from the lake sturgeonAcipenser fulvescens (Rafinesque) in Canada

Spinitectus acipenseri is described as a new species from the muscular stomach of the lake sturgeonAcipenser fulvescens from Manitoba and Saskatchewan. This new species closely resemblesS. gracilis Ward & Magath, 1917, but the major differences are the arrangement and larger size of spines (circlets and semicirclets of spines reaching to the anus in females ofS. acipenseri). Other differences include total body dimensions of adults (length and width relationships) and a 1:4–1:5 ratio of oesophagus to body length.     

Oogenesis in the viviparous phoronid,Phoronis embryolabi

The study of gametogenesis is useful for phylogenetic analysis and can also provide insight into the physiology and biology of species. This report describes oogenesis in the Phoronis embryolabi, a newly described species, which has an unusual type of development, that is, a viviparity of larvae. Phoronid oogonia are described here for the first time. Yolk formation is autoheterosynthetic. Heterosynthesis occurs in the peripheral cytoplasm via fusion of endocytosic vesicles. Simultaneously, the yolk is formed autosynthetically by rough endoplasmic reticulum in the central cytoplasm. Each developing oocyte is surrounded by the follicle of vasoperitoneal cells, whose cytoplasm is filled with glycogen particles and various inclusions. Cytoplasmic bridges connect developing oocytes and vasoperitoneal cells. These bridges and the presence of the numerous glycogen particles in the vasoperitoneal cells suggest that nutrients are transported from the follicle to oocytes. Phoronis embryolabi is just the second phoronid species in which the ultrastructure of oogenesis has been studied, and I discuss the data obtained comparing them with those in Phoronopsis harmeri. Finally, I discuss the distribution of reproductive patterns across both, molecular and morphological phylogenetic trees in Phoronida proving that parental care has evolved independently several times in this phylum.     

Occurrence and Ultrastructure of Collar Cells in the Stomach Gastrodermis of Polypodium hydriforme Ussov (Cnidaria)

The existence of collar cells lining the stomach gastrodermis in free-living Polypodium hydriforme and their ultrastructure are described. The collar cells are provided with a collar consisting of 9–10 microvilli which encircles a central flagellum and forms a flagellar pit. At the bottom of the pit around the basal part of the flagellum there is fine crystalline material which extends also in the spaces between the microvilli and keeps them straight. The flagellum has a typical axoneme (9+2), its basal body is located below the apical surface of the collar cell and continues into a striated rootlet. An accessory centriole is situated close to the upper part of the rootlet. The cell nucleus is located in the basal part of the cell. Prominent mitochondria with tubular cristae, Golgi cisternae and fragments of rough endoplasmic reticulum are situated mostly in the basal part of the cytoplasm. Discoidal vesicles are abundant in the apical cytoplasm. The collar cells are connected to each other by septate junctions and interdigitations. The ultrastructure of collar cells described here is discussed in comparison to that of other Cnidarians and in connection with the problem of Polypodium's systematic position.     

Ultrastructure of the spermatozoon of the digenean Plagiorchis elegans (Rudolphi, 1802) (Plagiorchioidea,Plagiorchiidae)

The ultrastructure of the mature spermatozoon of the type genus of the Plagiorchiidae Plagiorchis elegans (Rudolphi, 1802), a parasite of the Golden hamster, Mesocricetus auratus is described. This study is the first ultrastructural study of the spermatozoon of a Plagiorchis, the second of a plagiorchiid species and only the third in the Plagiorchioidea. Previously data on spermatozoon ultrastructure existed only for the plagiorchiid Enodiotrema reductum and the omphalometrid Rubenstrema exasperatum. The mature spermatozoon of P. elegans exhibited the general pattern described in most digenean species, namely two axonemes of the 9 + “1” Trepaxonemata pattern, nucleus, mitochondria, external ornamentation of the plasma membrane, spine‐like bodies, and glycogen granules. However, the rather typical expansion of the plasma membrane is not found in P. elegans. Another peculiarity of the spermatozoon of P. elegans is the presence of a structure called thin cytoplasm termination. Spermatozoon ultrastructure of P. elegans is compared with that of E. reductum and R. exasperatum. Spermatozoon of P. elegans conforms to the general pattern described in E. reductum. Thus, this study further expands our knowledge on the spermatozoon ultrastructure among the members of the Plagiorchioidea, one of the most phylogenetically derived groups of the digenea. J. Morphol. 274:965–972, 2013. © 2013 Wiley Periodicals, Inc.     

Ultrastructure of the lateral-line sense organs of the ratfish,Chimaera monstrosa

Summary The ultrastructure of the lateral-line neuromasts in the ratfish, Chimaera monstrosa is described. The neuromasts rest at the bottom of open grooves and consist of sensory, supporting, basal and mantle cells. Each sensory cell is equipped with sensory hairs consisting of a single kinocilium and several stereocilia. There are several types of sensory hair arrangement, and cells with a particular arrangement form patches within the neuromast. There are two types of afferent synapse. The most common afferent synapse has a presynaptic body and is typically associated with an extensive system of anastomosing tubules on the presynaptic side. When the tubules are absent, vesicles surround the presynaptic body. These synapses are often associated into synaptic fields, containing up to 35 synaptic sites. The second type of afferent synapse does not have a presynaptic body and is not associated with the tubular system. The afferent synapses of the second type do not form synaptic fields and are uncommon. The efferent synapses are either associated with a postsynaptic sac or more commonly with a strongly osmiophilic postsynaptic membrane. The accessory cells are similar to those in the acoustico-lateralis organs of other aquatic vertebrates. A possibility of movement of the presynaptic bodies and of involvement of the tubular system in the turnover of the transmitter is discussed. A comparison of the hair tuft types in the neuromasts of Ch. monstrosa with those in the labyrinth of the goldfish and of the frog is attempted.