Ultrastructure of Presumed Ocelli in Parenterodrilus taenioides (Polychaeta,Protodrilidae) and their Phylogenetic Significance

Abstract Three different types of presumed unpigmented ocelli have been found in the anterior end of Parenterodrilus taenioides, a small gutless interstitial polychaete. The type-1 ocelli are located in the palps and four ocelli have been found along the length of each palp. The type-2 and type-3 ocelli lie close together in the head segment and are located in posterior ganglionic expansions of the brain. There is one pair of the minute type-2 ocelli but at least two pairs of the type-3 organs, which are the largest ocelli. In each ocellus the sensory cells are of the ciliary type and possess two cilia whose plasma membranes branch into numerous microvilli. With the exception of the type-1 ocelli they consist of a sensory cell and a supportive cell. In each ocellus the supportive cell forms a thin cup-shaped envelope around the densely packed ciliary branches. The type-1 ocelli consist of a single cell forming an intracellular vacuole (phaosome) which contains less densely packed microvillus-like structures. In particular, the structure of these ocelli is compared with that in other polychaetes, with special emphasis on the remaining genera of the Protodrilida.     

Ultrastructure of the cephalic sensory organs of adult Pycnophyes dentatus and of the first juvenile stage of P. kielensis (Kinorhyncha, Homalorhagida)

 The ultrastructure of the paired cephalic sensory organs of adult Pycnophyes dentatus and of the first juvenile stage of P. kielensis (Kinorhyncha, Homalorhagida) was investigated by TEM. In both species, each sensory organ is composed of one receptor cell and one enveloping cell which border a common intercellular lumen. A single receptor cilium extends from the receptor cell into this lumen. The cilium expands behind the basal body and branches into numerous processes. A pair of cephalic sensory organs with these characteristics belongs to the ground pattern of, at least, the Pycnophyidae. The sensory organs of these Kinorhyncha correspond closely with the anterior cephalic organs of the Gastrotricha, but differ from the known cephalic receptors of other Nemathelminthes. Currently, it cannot be evaluated conclusively whether the last common ancestor of the Nemathelminthes possessed cephalic sensory organs and, if it did, what these organs looked like. Accepted: 3 December 1996     

Ultrastructural investigations of presumed photoreceptive organs in two Saccocirrus species (polychaeta,saccocirridae)

In addition to the pigmented ocelli, four different types of photoreceptor-like organs without shading pigment have been found in Saccocirrus papillocercus and S. krusadensis. The sensory cells of these presumed ocelli are either ciliary or rhabdomeric with ciliary rudiments. With the exception of the multicellular type-2 ocelli they are bicellular consisting of a sensory cell and a supportive cell. In each ocellus the supportive cell forms a thin cup-shaped envelope around the sensory elements. In the type-2 ocellus, 7 supportive cells form an ovoid cavity leaving openings through which dendritic processes of an equal number of sensory cells enter the cavity. The pigmented ocelli possess an ocellar cavity communicating with the exterior through a pore in the eyecup, ciliary rudiments in both sensory and supportive cell, and additional non-photoreceptive sensory cells in the opening of the eyecup. The sensory organs show characteristic differences between the two species, such as presence or absence of a particular type of ocellus (type 2 is absent in S. krusadensis, type 3 in S. papillocercus), number of cilia in type-4 ocelli, density of microvilli, number of non-photoreceptive sensory cells in the pore of the pigmented ocellus, etc. These differences provide important characters which can be used for discrimination either of species or of subgeneric taxa in Saccocirrus. The phylogenetic significance of the different photoreceptive organs is discussed.     

Lateral organs in sedentary polychaetes (Annelida) – Ultrastructure and phylogenetic significance of an insufficiently known sense organ

Lateral organs are sense organs visible as densely ciliated pits or papillae between the noto‐ and the neuropodia in certain taxa of sedentary polychaetes. Ultrastructural studies in about 10 species of the following taxa Maldanidae, Opheliidae, Orbiniidae, Paraonidae, Magelonidae, Spionidae, Poecilochaetidae and Terebellidae have been designed to evaluate whether these organs are homologous among polychaetes. In spite of great external diversity, the investigations revealed an overall ultrastructural similarity. Differences between species investigated mainly concern the size of the organs as well as the number and arrangement of cells. The organs comprise supportive cells and uniciliated penetrative sensory cells. Their dendrites are closely arranged and thus their cilia may resemble multiciliated cells. There are two types of sensory cells: one type possesses no or mainly thin microvilli of which usually only a few reach the cuticular surface, and in the other type the cilium is consistently surrounded by 10 strong microvilli, which form a pore‐like opening in the cuticle. Further differences occur in the structure of the rootlet system. Basally, a retractor muscle attaches to the organ. The systematic significance of these organs within Annelida is discussed with respect to the conflicting phylogenetic hypotheses explaining the relationships of annelid taxa.     

Central nervous system and sense organs, with special reference to photoreceptor-like sensory elements, in Polygordius appendiculatus (Annelida), an interstitial polychaete with uncertain phylogenetic affinities

Abstract. The phylogenetic position of Polygordius is still pending; relationships with either Opheliidae or with Saccocirrus are the most favored hypotheses. The present study of Polygordius appendiculatus was designed to look for morphological characters supporting either of these two hypotheses. The homology of the anterior appendages, and the structure of the central nervous system and nuchal organ all required clarification; we also examined whether photoreceptor‐like sense organs exist in adults. From their innervation pattern, it is likely that the anterior appendages represent palps. They lack structures typical of palps in Canalipalpata, such as musculature and coelomic cavities, which would be expected in the case of a saccocirrid relationship. Thirteen photoreceptor‐like sense organs were found in front of the brain, the only structures resembling photoreceptors in adults of P. appendiculatus. These multicellular sense organs comprise a supportive cell and several sensory cells enclosing an extracellular cavity. There are three different types of sensory cells: one rhabdomeric and two ciliary. These sensory cells are combined differently into three forms of sense organ: the most frequent uses all three types of sensory cells, the second possesses one rhabdomeric and one ciliary cell type, and the third has two types of ciliary sensory cells. Whereas similar sensory cells are frequently found in various polychaetes, their combination in one sensory organ is unique to Polygordius and is considered to represent an autapomorphy. The nuchal organs exhibit features typical of polychaetes; there are no specific features in common with Saccocirrus. Instead, the covering structures show obvious similarities to Opheliidae, as can also be found in the central nervous system. Altogether, the current observations do not contradict a relationship with opheliids but provide no evidence of a relationship with Saccocirrus as has been found in certain molecular analyses, and thus currently leave the phylogenetic position of Polygordius unresolved.     

The ultrastructure of the eyes in the veliger-larvae of Aporrhais sp. and Bittium reticulatum (Mollusca,Caenogastropoda)

Summary The cerebrally innervated larval eyes of Aporrhais sp. and Bittium reticulatum are investigated by means of transmission electron microscopy. Each organ consists of a pigmented cup containing an acellular lens. The cornea overlaps the anterior portion of the eye. The retina is composed of sensory cells and supportive cells. The sensory cells of Aporrhais sp. bear one cilium and in Bittium reticulatum two cilia, the ciliary membrane being folded into numerous finger-shaped evaginations. The supportive cells contain the pigment granules and most of them bear one or two cilia, the plasmalemma of which is likewise folded. It is supposed that: (a) these cilia have a transportive function for lens material and (b) that the ciliary photoreceptor of Aporrhais sp. and Bittium reticulatum is a functional adaptation to a relatively long larval period.Abbreviations bb basal body - bp basal plate - c cilium - cc corneal cell - cm ciliary membranes - cw ciliary whorl - gd Golgi dictyosomes - gm granular material - l lens - m mitochondrion - mt microtubules - mv microvilli - mvb multivesicular body - n nucleus - pb pigment border - pg pigment granule - rer rough endoplasmic reticulum - sc sensory cell - sj septate junctions - spc supportive cell     

Immunohistochemical and ultrastructural studies on ciliary sense organs of arrow worms (Chaetognatha)

We examined epibenthic and pelagic species of Chaetognatha (Spadellidae and Sagittidae) using immunohistofluorescence and confocal laser scanning microscopy to detect tubulin and cell nuclei in whole-mount preparations and scanning and transmission electron microscopy to visualize the ultrastructural organisation of their ciliary sense organs. All chaetognaths bear three types of ciliary sense organs distributed throughout the body: (1) transversally oriented ciliary fence organs, (2) longitudinally (parallel to the anterior-posterior axis) oriented ciliary tuft organs, and (3) a ciliary loop, the corona ciliata. This study targets the ciliary fence as well as the ciliary tuft organs. Two types of primary receptor cells constitute the ciliary fence and ciliary tuft organs. The first type of receptor cells forms a single cell line along the midline axis of the organs, whereas the second type of receptor cells forms multiple lines of cells at either side of type 1 cells. Each receptor cell extends a single, non-locomotory cilium from its narrow apex collared by slender, non-reinforced microvilli; however, both types of sensory cells considerably differ on ultrastructural level. Type 1 sensory cells have thicker cilia than those protruded by the type 2 sensory cells which are characterized by rootlets consisting of an elongated, amorphous distal as well as a cross-striated proximal portion. These results likely reveal that both types of sensory cells have distinct functions.     

The ciliary photoreceptor in the teleplanic veliger larvae of Smaragdia sp. and Strombus sp. (Mollusca,Gastropoda)

The cerebrally innervated eyes of the veliger larvae of Smaragdia sp. and Strombus sp. are composed of a lens, a cornea, and an everse retina. The retina contains two different types of cells, ciliary sensory cells and supportive cells which bear one or two cilia. It is suggested that: (a) the ciliary photoreceptors of these teleplanic veliger larvae are correlated with a long pelagic life in the ocean, which can last up to twelve months, and (b) that structural details of the photoreceptors can change during ontogenesis (ciliary vs rhabdomeric). Furthermore, the cilia of the supportive cells apparently tranport lens material and thus play an important role in lens formation. A decomposition mechanism of pigment granules is examined.Abbreviations bb basal body - bp basal plate - c cilium - cc corneal cell - cm ciliary membranes - cw ciliary whorl - ecm extracellular matrix - gr electron-dense granules - l lens - lb lamellar body - mp membranous pieces - mt microtubules - mv microvilli - n nucleus - oc optic cavity - on optic nerve - pg pigment granule - sc sensory cell - sj septate junction - spc supportive cell - v vesicles     

Ultrastructure of nuchal organs in some marine polychaetes

Polychaetes normally possess one pair of nuchal organs at the posterior edge of the prostomium or peristomium. They have been regarded as chemosensory organs. The nuchal organs of four marine polychaete species with different habits were investigated by electron microscopy. Although the shapes of nuchal organs can vary greatly from simple ciliary bands (Scolelepis squamata, Spionidae) to retractile tongue-like, piston- or finger-shaped forms (Eteone longa, Anaitides mucosa, Phyllodocidae; Heteromastus filiformis, Capitellidae), the structural components, including the ciliated supporting cells, sensory cells, and nuchal epidermal cells, are essentially similar. The differences basically concern 1) the position of the sensory cells with relation to the ciliated supporting cells, 2) the location and structure of the nuchal nerve, and 3) the structure of the nuchal cuticle. The diverging nature of this modified cuticle is described and discussed in detail. Comparisons are made with the fine structure of nuchal organs of other polychaete species. Similarities of cellular components of nuchal organs are found not only in the four species studied here but also in all nuchal organs investigated so far. This is hypothesized to be due to the fact that the polychaete stem species already possessed nuchal organs with the respective cell types. Differences in the number and distribution of cellular components and in the overall shape of nuchal organs are thought to have evolved in correlation with the equipment of other cephalic appendages and with different habits and modes of nutrition.     

Composition,Seasonal Dynamics,and Long-Term Fluctuations in the Density of Pelagic Polychaetes in Amurskii Bay,Sea of Japan

This is the first survey of the composition and seasonal density dynamics of the pelagic polychaete larvae in Amurskii Bay. This is a summary of the results of two years of observations (1996–1998). The larvae of 14 polychaete families have been recorded. The greatest number of species have been found in the family Spionidae. The most common were the larvae of Harmothoe imbricata (family Polynoidae) and Paraprionospio sp. (family Spionidae). The larvae of polychaetes have been found in plankton throughout the year and have two density peaks. The first peak was due to the spawning of H. imbricata; the second one was provided by representatives of several species of the family Spionidae, mostly Paraprionospio sp., Polydora ciliata, and P. caeca.     

Head sensory organs of Dactylopodola baltica (Macrodasyida, Gastrotricha): a combination of transmission electron microscopical and immunocytochemical techniques

The anterior and posterior head sensory organs of Dactylopodola baltica (Macrodasyida, Gastrotricha) were investigated by transmission electron microscopy (TEM). In addition, whole individuals were labeled with phalloidin to mark F-actin and with anti-alpha-tubulin antibodies to mark microtubuli and studied with confocal laser scanning microscopy. Immunocytochemistry reveals that the large number of ciliary processes in the anterior head sensory organ contain F-actin; no signal could be detected for alpha-tubulin. Labeling with anti-alpha-tubulin antibodies revealed that the anterior and posterior head sensory organs are innervated by a common stem of nerves from the lateral nerve cords just anterior of the dorsal brain commissure. TEM studies showed that the anterior head sensory organ is composed of one sheath cell and one sensory cell with a single branching cilium that possesses a basal inflated part and regularly arranged ciliary processes. Each ciliary process contains one central microtubule. The posterior head sensory organ consists of at least one pigmented sheath cell and several probably monociliary sensory cells. Each cilium branches into irregularly arranged ciliary processes. These characters are assumed to belong to the ground pattern of the Gastrotricha.     

Fine structure of the sensilla of Peripatopsis moseleyi (Onychophora)

Summary Three types of sensilla occurring on the lips and on the antennae of Peripatopsis moseleyi have been investigated by scanning and transmission electron microscopy. On the lips sensory spines can be found which contain numerous cilia originating from bipolar receptor cells. They reach the tip of the spine where the cuticle is modified. The perikarya of the sensory cells, a large supporting cell with a complicated surface and a second type of receptor, form a bud-like structure and are surrounded by a layer of collagen fibrils. The second receptor cell bears apical stereocilia as well as a kinocilium which are directed towards the centre of the animal — thus the cell appears to be turned upside down. The sensilla of the antennae are 1) sensory bristles containing two or three kinds of receptor cells, one of which bears an apical cilium and one kind of supportive cell and 2) sensory bulbs located within furrows consisting of receptor cells with branched cilia and two kinds of supportive cells which are covered by a modified thin cuticle. According to the electron microscopical findings the sensory spines on the lips are presumably chemoreceptors. The sensory bristles on the antennae can be regarded as mechanoreceptors and the sensory bulbs as chemoreceptors.Supported by the Deutsche Forschungsgemeinschaft (Sto 75/3)     

Structure of the auditory system of the weta Hemideina crassidens (Blanchard, 1851) (Orthoptera,Ensifera, Gryllacridoidea,Stenopelmatidae)

Summary This study of the ultrastructure of the auditory sensilla of the New Zealand weta, Hemideina crassidens, is the first such study on a member of the orthopteran Superfamily Gryllacridoidea. Ultrastructure of the auditory sensilla is similar in all of the tibial mechanosensory organs, here called subgenual organ, intermediate organ and crista acoustica by analogy with comparable structures in Tettigoniidae.Distal to each sensory soma is a dendrite containing multiple ciliary rootlets that fuse into a single ciliary root. This splits into nine root processes that pass around the outside of the proximal basal body and then rejoin at the level of the distal basal body, distal to which the dendrite has a modified ciliary structure with a circlet of nine peripheral paired tubes and rods as it passes through the proximal extracellular space. It is then enclosed by a zone of scolopale cell cytoplasm before expanding into a dilatation within the distal extracellular space. In some sensilla this space is partially occluded by electron dense material which is part of the scolopale cell. Distal to the dilatation the cilium shrinks and ends surrounded by the scolopale cap.Accessory cells consist of glia enwrapping the sensory neuron in the region of its soma, the scolopale cell surrounding the ciliary portion of the dendrite, and the attachment cell surrounding the scolopale cell and scolopale cap and connected to them by desmosomes. The attachment cells are filled with microtubules in differing densities and orientations. Lamellae are present in the acellular matrix surrounding the attachment cells. Banded fibres, presumably of collagen, are also present in the matrix.     

Ultrastructure of the “statocysts” inProtodrilus species (Polychaeta): Reconstruction of the cellular organization with morphometric data from receptor cells

Summary The statocysts inProtodrilus ciliatus, P. oculifer, P. haurakiensis andP. helgolandicus are situated in the prostomium anterior to the palps and have been investigated by electron microscopy. The sensory organs were reconstructed from serial sections, volumes were calculated from areas of consecutive section profiles, and additional data on surface area of distal receptor elements have been determined. In spite of variations in size (diameter 8–20 m) their structure is nearly identical. The organs consist of one cup-shaped supportive cell, one large bi- or multiciliated sensory cell and two small uni- or biciliated sensory cells forming an extracellular cavity. This cavity is completely filled with microvillus-like or paracrystalline structures and there are no signs of statoliths composed of extracellular material. The most striking feature is the occurrence of paracrystals made up of undulating ciliary membranes extending from the large sensory cell and occupying 75–90% of the cavity inP. ciliatus, P. oculifer andP. haurakiensis. The remaining space is filled with microvilli or dendritic processes of the sensory cells. InP. helgolandicus the ciliary paracrystals are almost completely replaced by microvillus-like branches of cilia of the corresponding sensory cell. Paracrystals fill less than 10% of the cavity and are formed of flattened membranes. These sensory organs enclose large surface areas of membranes (15,000–38,000 m²). The surface areas of the paracrystals composed of undulating membranes is almost identical to that of densely arranged arrays of microvilli (about 25 m² per m³). These sensory organs are so different from all known statocysts that it is likely that they have another function. Their greatest structural correspondence is to light-receptive organs, especially in the structure and arrangement of microvilli. The role the paracrystals play is discussed: they might bear photopigments or simply represent a lens — a transparent, refractile and crystalline structure. These sensory organs are completely different from pigmented ocelli and phaosomes occurring in some protodrilids and represent a type of sensory organ thus far undescribed in polychaetes.     

Problems in polychaete systematics

Some of the intriguing issues in current polychaete systematics are reviewed. (1) The root of the `polychaete' tree. Currently there are two major hypotheses concerning the root position among polychaetes. One is based on rooting cladograms with outgroups such as Mollusca and result in simple-bodied taxa such as Opheliidae and Questidae forming a basal annelid grade along with Clitellata. Other hypotheses do not use outgroup rooting but involve scenarios on the evolution of the group and would place taxa in Aciculata as basal annelids, thus making Aciculata and Phyllodocida paraphyletic. Molecular sequence data has been of little help in resolving this issue thus far, largely due to limited taxon sampling. (2) Paraphyly. Owing, in part, to a tradition involving the emphasis on differences among taxa, and the application of Linnean ranks (e.g., family), paraphyly is undoubtedly a widespread phenomenon in polychaete systematics. An example of this has been proposed already for Spionidae. If the tree topology and rooting used by Blake & Arnofsky (1999) is correct, Spionidae is made paraphyletic by the recognition of the following four family-ranked taxa; Trochochaetidae, Poecilochaetidae, Longosomatidae and Uncispionidae. Another possible example is seen with Cirratulidae. A preliminary cladistic analysis shows that it is entirely possible that seven other taxa recognised as families may be nested within Cirratulidae. These include Acrocirridae, Ctenodrilidae, Fauveliopsidae, Flabelligeridae, Flotidae, Poeobiidae and Sternaspidae. (3) Problematic taxa. Apart from the problems exposed by the analysis of Cirratuliformia, the position of some of these groups, such as Aberranta, Alciopidae, Hesionides,Lopadorhynchidae, Microphthalmus,Nerillidae, Spinther,Tomopteridae and Sabellariidae, is discussed.     

TEM studies on the lattice organs of cirripede cypris larvae (Crustacea, Thecostraca, Cirripedia)

 Lattice organs consist of five pairs of sensory organs situated on the dorsal carapace in cypris larvae of the Crustacea Cirripedia. The lattice organs in cypris larvae of Trypetesa lampas (Acrothoracica) and Peltogaster paguri (Rhizocephala) represent the two main types found in cirripedes, but only minor differences exist at the TEM level. Each lattice organ is innervated by two bipolar, primary receptor cells. The inner dendritic segment of each receptor cell carries two outer dendritic segments. The outer dendritic segments contain modified cilia with a short ciliary segment (9×2+0 structure). Two sheath cells envelop the dendrite except for the distal ends of the outer dendritic segments. This distal end enters a cavity in the carapace cuticle and reaches a terminal pore situated at the far end of the cavity. The cuticle above the cavity is modified. In both species the epicuticle is partly perforated by numerous small pores and the underlying exocuticle is much thinner and less electron dense than the regular exocuticle. Lattice organs very probably have a chemosensory function and are homologous with the sensory dorsal organ of other crustacean taxa. Accepted: 18 August 1998     

Ultrastructural investigation of the nuchal organs of Pygospio elegans (Polychaeta)

Summary The differentiation of the dorsal organs as well as the structure of the nuchal organs and their relation to the central nervous system in adult Pygospio elegans were studied by electron microscopy and compared to the nuchal organs of the larvae. The nuchal organs are represented by paired ciliary bands on the dorsal side of the first setiger, delimiting a median caruncle that is completely filled with epidermal and nervous tissue. They are composed of ciliated supporting cells and bipolar primary sensory cells constituting the nuchal ganglia, which are integrated into the brain. Microvillus-like processes of the ciliated cells give rise to a secondary covering layer over the sensory epithelium. The size of the nuchal organs is a sexually dimorphic feature.Dorsal organ formation is concomitant with the onset of sexual maturation in the male sex only. They appear as metameric ciliary bands on the dorsal side of the anterior body region and consist of ciliated cells accompanied by lateral accumulations of tubular gland cells. In the gametogenic segments they are structurally associated with the male genital pores and may be involved in reproduction. The results refute previous theories that dorsal organs are sensory and have a common origin to nuchal organs.Abbreviations ac anterior commissure of the brain - ace anterior circumesophageal connective - bb basal body - bl basal lamina - c cuticle - ca caruncle - cc ciliated cell - ci sensory cilium - co microvillar cover - d septate desmosome - db dorsal blood vessel - dn dorsal nerve cord - ea efferent axons - ec epidermal cell - eg elementary granules - g Golgi complex - i filamentous inclusion - lm longitudinal muscles - ly lysosome - mc motile cilia - mv microvillus - n neuron - ng nuchal ganglion - nn nuchal nerve - nu nucleus - oc olfactory chamber - pa palp - pc posterior commissure of the brain - pce posterior circumesophageal connective - rer rough endoplasmic reticulum - sI setiger I - sb sensory bulb - sc sensory cell - sd sensory dendrite - ser smooth endoplasmic reticulum - tf tonofilament bundle - v clear vesicles - za zonula adherens     

Sensory endings of the ascidian static organ (Chordata,Ascidiacea)

Summary The ultrastructure of the static organ is examined in larvae of Diplosoma macdonaldi, a colonial ascidian, and Styela plicata, a solitary ascidian; the results are similar. As previous workers found, the cell body of a unicellular statocyte lies in the lumen of the sensory vesicle and contains the statolith. A narrow neck connects the cell body to an anchoring foot in the floor of the sensory vesicle. Two previously undescribed sensory endings project into the lumen just to the left of the statocyte, one anterior and one posterior to the neck. A network of fine processes from each ending contacts the statocyte body. It is proposed that movements of the statocyte cell body are detected by these endings. They arise from neurons in the ventral wall of the sensory vesicle that project axons to the visceral ganglion. The placement of the sensory endings may allow discrimination of the directon of statocyte deflection.Abbreviations ax axons - bb ciliary basal body - bl basal lamina - c cilium - cr striated ciliary rootlet - ec ependymal cells - en endoderm - h hemocoel - ly lysosome - mv microvilli - n neuron - nf neurofilaments - ns neck of the statocyte - sb statocyte cell body - sd sensory dendrite - sn sensory neuron - sp sensory processes - stf statocyte foot - svl sensory vesicle lumen - zo zonula occludens     

Elektronenmikroskopische Untersuchungen zum Problem der Sekretion der bindegewebigen Interzellularsubstanz

Zusammenfassung Die Sinnesorgane des Vorderendes von Linens ruber werden elektronen-mikroskopisch vor allem im Hinblick auf die rezeptorischen Strukturen untersucht. Am Kopfvorderrand liegen zwei Rezeptortypen: Typ 1 ist mit 20–40, Typ 2 nur mit einer Zilie besetzt. Die Seitenwurzeln der Zilien von Typ 1 sind parallel angeordnet. Die Perikarya beider Rezeptoren liegen unterhalb des Epithelverbandes, dessen Großteil von Typ 1 eingenommen wird. In den Kopfspalten finden sich primäre Sinneszellen mit eingesenkten Zilien, deren Membran regelmäßig aufgeschwollen ist. Im Cerebralorgan dominiert ebenfalls eine primäre Sinneszelle, die reichlich efferent innerviert wird. In den Lichtrezeptoren findet sich eine Zilie unter dem distalen Mikrovillisaum.Alle beschriebenen Zilien sind deutlich voneinander und von den Wimpern der Epidermis unterschieden. Sie werden mit den Zilien der Rezeptoren anderer Tiergruppen verglichen.

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The fine structure of the sensory organs of Lineus ruber O. F. Müller (Nemertini, Heteronemertini)

Summary The sensory organs located in the head reagion of the ribbon worm Lineus ruber have been investigated with the electron microscope. Attention has been focussed on the apical cell-structures, which are assumed to be related to sensory functions. At the anterior edge of the head two receptor-types occur: type 1 bears 20–40 apical cilia, type 2 only 1 cilium per cell. The lateral ciliary rootlets of type 1 are oriented strictly parallel. The pericarya of both receptor types of which type 1 is particularly dominant, are situated below the basement lamina of the epithelium. In the cephalic clefts primary sensory cells occur, the cilia of which are invaginated into the apical cytoplasm. Their ciliary membrane exhibits regular inflations. In the cerebral organ also primary sensory cells dominate which receive an abundant efferent nerve supply. In the light receptors only one cilium is to be found below the distal villi. The cilia of the sensory cells and also of the ordinary epidermis cells show individual characteristics not shared by the other types. Their structural peculiarities are compared with those of sensory organs of other groups of animals.

Herrn Prof. Dr. Drs. h. c. W. Bargmann zum 65. Geburtstag gewidmet.     

Sense Organs of Two Marine Arthrotardigrades (Heterotardigrada,Tardigrada)

The sense organs of the marine arthrotardigrades Halechiniscus greveni Renaud-Mornant & Deroux and Batillipes noerrevangi Kristensen were investigated with the aid of interference phase-contrast and transmission electron microscopy. The sense organs of the two species are quite different in outer cuticular morphology, but the inner cellular parts are constructed after the same model: the arthropod sensory setae. All investigated sensilla contained ciliary structures. Regeneration of the cirri during moult is very similar to the regeneration process in insects and spiders, but in the two arthrotardigrades the trichogen cell is not retracted from the cuticular part of the setae after moult. The clava is presumed to be olfactory; the cephalic cirri and the leg spines are contact chemoreceptors with one mechanoreceptive cilium terminating at the sensillum shaft. Cirrus E is compared with the trichobothrium of arthropods and the phylogenetic implications of the investigated structures are discussed.