Die Ultrastruktur der Photorezeptoren von Lithobius forficatus L. (Chilopoda: Lithobiidae)

Zusammenfassung Die Untersuchung der feinstrukturellen Organisation der Ocellen von Lithobius forficatus L. ergab, daß der dioptrische Apparat aus einer lamellenartig gebauten, ungleichseitig bikonvexen Cornealinse besteht. Ein Glaskörper und spezielle Pigmentzellen fehlen. Der Augenbecher wird von 35–110 Sinneszellen gebildet, unter denen 2 morphologisch distinkte Typen unterschieden werden können. Die großen Sehzellen des distalen Bereiches besitzen einseitig inserierende Rhabdomere, die in radiärsymmetrischer Anordnung ein umfangreiches geschlossenes Rhabdom bilden. Der proximale Teil des Augenbechers wird von kleineren, konischen Basalzellen in Form einer undeutlich abgesetzten Retinula eingenommen. Durch enge Verzahnung ihrer zirkumapikal oder zweiseitig angeordneten Mikrovilli entstehen stelzenförmige Doppel- und Mehrfachrhabdomere, die mit dem zentralen Rhabdom in Verbindung stehen. Alle Sehzellen sind durch eine Gliederung in verschiedene Zonen gekennzeichnet. Sie sind bei distalen Rezeptoren senkrecht zur optischen Achse, bei Basalzellen transversal zur Längsachse der Zelle angeordnet. Auf die rhabdomerischen Mikrovilli des Augenzentrums folgt nach außen eine Schaltzone aus Elementen des ER und anderen vesilukären Bildungen. Diese Schaltzone stellt wahrscheinlich eine mit dem Adaptationszustand des Auges korrelierte Funktionsstruktur dar. In der cytoplasmatischen Zone fällt die Zahl verschiedenartiger multivesikulärer Korpuskel neben wenigen großen multilamellären Körpern auf. Die funktionelle Bedeutung des Ocellusaufbaus bei Lithobius wird diskutiert.

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Ultrastructure of the Photoreceptors of Lithobius forficatus L. (Chilopoda: Lithobiidae)

Summary The ultrastructure of the ocelli of Lithobius forficatus L. was investigated by means of conventional electron microscopy. The dioptric apparatus consists of an unequal biconvex corneal lens which has a lamella-like fine structure. Crystalline cones and special pigment cells are lacking. The eye cup is composed of 35 to 110 sense cells of two different morphological types. The large visual cells of the distal region are characterized by unilaterally inserted rhabdomeres which form in a radial symmetrical arrangement the extended closed rhabdome. The proximal part of the eye cup is occupied by somewhat smaller basal cells of conical shape, showing an indistinct retinula. These cells have numerous microvilli either in the apex region or laterally which interdigitate to form stiltlike double or multiple rhabdomeres in close communication to the central rhabdome. In both types of sense cells particular zones appear because of the characteristic distribution of certain cell elements. They are arranged perpendicularly to the optical axis in the large receptors and vertical in the main axis of the basal cells. Elements of ER and other vesicles constitute a Schaltzone which borders the microvilli of the rhabdomer. This Schaltzone probably is a functional structure correlated to the adaptional state of the eye. The cytoplasmatic zone contains numerous different multivesicular and few large multilamellar bodies. The functional meaning of the organization of the Lithobius ocellus is discussed.

Mit dankenswerter Unterstützung durch die Deutsche Forschungsgemeinschaft.     

The water relations of Lithobius forficatus and the role of the coxal organs (Myriapoda: Chilopoda)

The water relations of the common brown centipede, Lithobius forficatus (L.), are examined with regard to the possible involvement of the coxal organs in water uptake. Centipedes with blocked and open coxal pores are subjected to a variety of dehydration and rehydration regimes and weight change over time measured. Weight loss and regain is related to body surface area and is not influenced by the coxal organs.
The conclusion of the paper is that the coxal organs of L. forficatus do not contribute significantly to the ability of these animals to recover water lost through cuticular transpiration.     

Immunohistochemical evidence for ecdysteroid-like material in the putative molting glands of Lithobius forficatus (Chilopoda)

Summary Ecdysteroid-like material was demonstrated by means of immunhistochemistry in the anterior body region of Lithobius forficatus with the use of an antiserum against an ecdysone-methoxim-BSA-conjugate in conjunction with a modified PAP-method (Sternberger and Joseph 1979). This material is restricted to a tissue formed by podocytes loosely surrounding the salivary glands. Earlier ultrastructural, experimental and biochemical in vitro investigations indicated that this tissue represents the ecdysial glands; this interpretation is now strengthened by immunohistochemical evidence. Reactivity within the cells occurs predominantly in cytosomes.The authors dedicate this paper to Professor Günther Cleffmann, Institut für Tierphysiologie, Justus-Liebig-Universität Giessen, on the occasion of his 60^th birthday, January 27, 1988     

Chromosome study of Lithobius forficatus (Lithobiomorpha, Chilopoda)

The diploid number 2n = 46 and the chromosome arm number NF = 74 are described in Lithobius forficatus from Olsztyn (Poland). Analyses of silver and CMA3-stained mitotic chromosomes suggest that a single chromosome pair has active NORs which correspond to G-C-rich (CMA3-positive) chromatin. Heteromorphism of the largest metacentric chromosome pair was observed. The sex chromosomes were not identified. Size polymorphism of the first chromosome pair was found.     

Influence of the ventral nerve cord on the spermatogenetic cycle in Lithobius forficatus L. (Myriapoda,Chilopoda)

The role of the ventral nerve cord has been investigated in Lithobius forficatus L., using ganglionic destructions, ganglionic implantations and ganglionic electrostimulations. It seems that a stimulating principle is present in the sub-oesophageal ganglion. Its action is not so important as that of the pars intercerebralis.     

The chemosensory behaviour of Lithobius forficatus. 1. Evidence for a pheromone released by the coxal organs (Myriapoda: Chilopoda)

The behaviour of the Common brown centipede, Lithobius forficatus (.L.) is investigated with reference to the function of the coxal organs which are a peculiar and characteristic feature of Chilopoda. Previous studies of these organs in centipedes have been largely morphological and have relied on ultrastructural comparisons with other arthropod epithelia of disparate functions. Evidence is presented which suggests that the coxal pores release a sex-specific pheromone. The nature of intraspecific interactions in L. forficatus is discussed.     

The peristomatic structures of Lithobiomorpha (Myriapoda, Chilopoda): comparative morphology and phylogenetic significance

A comparative survey of the epipharynx and hypopharynx of lithobiomorph centipedes by light and scanning electron microscopy examines 18 species that sample the major groups of both families, the Lithobiidae and Henicopidae. Cladistic analysis of 11 characters of the peristomatic structures together with 29 additional morphological characters serves as a basis for interpreting the evolution of the lithobiomorph peristomatic structures. Scutigeromorpha is used for outgroup comparison in the framework of a homology scheme for the basic components of the epi- and hypopharynx. Compared to other chilopods, the monophyly of Lithobiomorpha is supported by a row of distinctive bottle-shaped gland openings at the border between the labral and clypeal parts of the epipharynx, as well as by a distinctive shape of the hypopharynx. Paired rows of elongate spines on the clypeal part of the epipharynx are an apomorphic character of Lithobiidae. The transformation of these spine rows into a few groups of branching spines is characteristic for the Monotarsobius group sensu Verhoeff. Similar groups of branching clypeal spines characterize the Anopsobiinae within Henicopidae, whereas Henicopinae possess a dense cluster of short, simple spines instead. The recently described genus Dzhungaria is resolved closer to Henicopinae than to Anopsobiinae, a hypothesis supported by a field of grooves on the medial labral part of the epipharynx. Monophyly of Henicopidae does not receive unique support from the peristomatic structures although two homoplastic characters contribute to this node; among these, the reduction of a median spine field between clypeal and labral parts of the epipharynx to a narrow transverse band also supports a close relationship between the Ezembius group and Hessebius within Lithobiidae. An Ezembius+Hessebius clade is additionally supported by the absence of a transverse bulge between the clypeal and labral parts of the epipharynx, a character otherwise present in all lithobiomorph species studied so far. Lithobius is resolved as polyphyletic, with different species being most closely related to such genera as Australobius, Hessebius and Pleurolithobius.     

Sinnesorgane auf den Antennen vonLithobius forficatus L. (Myriapoda,Chilopoda)

Zusammenfassung Sensilla trichodea auf den Antennen vonLithobius forficatus L. wurden licht-, raster- und durchstrahlungselektronenmikroskopisch untersucht. Jede Antenne tr?gt beim ausgewachsenen Tier etwa 2000 dieser Organe, von denen jedes über 18 Sinneszeüen und mindestens 3 Hüllzellen verfügt. Der Haarschaft ist ungef?hr 100–150Μm lang, sehr elastisch und beweglich aufgeh?ngt. Er verfügt über eine Pore in der Spitze, bis zu der 17 Dendriten ziehen. Ein Dendrit endet mit einem Tubulark?rper in einer komplizierten Cuticuladifferenzierung an der Haarbasis. Die innerste Hüllzelle bildet den Sensillenliquorraum I, in den die Dendriten einmünden und Cilienstruktur annehmen. Sie verfugt über ein reich ausgebildetes granul?res ER und bildet eine innere Dendritenscheide, die die Dendriten bis ins Haar einhüllt. Die zweite Hüllzelle ist die trichogene Zelle. Sie enth?lt zahlreiche Microtubuli und entsendet einen schmalen Ausl?ufer in das Haarlumen, der durch die innere Scheide von den Dendriten getrennt wird. Die dritte Hüllzelle ist die tormogene Zelle, sie bildet den Sensillenliquorraum II. Eine zweite cuticul?re Scheide liegt zwischen trichogener und tormogener Zelle; sie ist mit der K?rperwand verbunden. In der frisch abgestreiften Exuvie des Sensillums findet man keine Reste mehr von Dendriten und anderen Zellausl?ufern; diese müssen vor der Ecdysis zurückgezogen worden sein. Unmittelbar nach der H?utung verfügt das noch im Aufbau befindliche Organ weder über die komplizierte, für die Mechanoreception wichtige Cuticuladifferenzierung an der Haarbasis noch über den Sensillenliquorraum II. Das Sensillum wird als Kombination von Kontaktchemo- und Mechanoreceptor angesehen. Seine m?gliche Bedeutung für die Orientierung vonLithobius wird diskutiert.

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Sense organs on the antennae ofLithobius forficatus L. (Myriapoda, Chilopoda)

Summary Sensilla trichodea on the antennae ofLithobius forficatus L. were investigated by means of light-, scanning- and transmission-electron-microscopy. Each antenna of the adult animal carries about 2000 of these organs, each of which consists of 18 sensory cells and at least 3 enveloping cells. The elastic shaft is about 100–150Μm long and easily movable. It has a terminal pore, up to which extend 17 dendrites. One dendrite, containing a tubular body, ends in a complicated cuticular structure at the hair base. The innermost enveloping cell forms the “Sensillenliquorraum I”, into which enter the dendrites and assume ciliary structure. This cell has a rich granular endoplasmic reticulum and forms an inner dendritic sheath which envelops the dendrites up into the hair lumen. The second enveloping cell is the trichogen cell. It contains numerous microtubules and sends off a narrow process into the hair lumen, which is separated from the dendrites by the inner cuticular sheath. The third enveloping cell is the tormogen cell, it forms the “Sensillenliquorraum II”. A second cuticular sheath is lying between the trichogen and the tormogen cell, it is connected with the body wall. In the freshly cast exuvia of the sensillum no remains of the dendrites or other cells can be found, they must have been retracted before moulting. Immediately after the ecdysis, the new-developing organ does neither possess the complicated cuticular structure at the base, which is important for the mechanoreception, nor the “Sensillenliquorraum II”. The sensillum is supposed to be a combined contactchemo- and mechanoreceptor. Its possible signification for the orientation ofLithobius is discussed.

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1. Teil einer Dissertation am Fachbereich 23 (Biologie) der Freien Universit?t Berlin (D 188). Herrn Prof. Dr. J. Brandenburg danke ich für sein stetiges Interesse am Verlauf dieser Arbeit. Frau C.St. Friedemann danke ich für die Anfertigung der Zeichnungen, Herrn Rau von der Zentraleinrichtung für Elektronenmikroskopie der Technischen Universit?t Berlin (Leiter: Prof. Helmcke) für die Bedienung des Rasterelektronenmikroskopes und Herrn Prof. Dr. U. Thurm für die kritische Durchsicht des Manuskripts.     

Ultrastructure and regeneration of midgut epithelial cells in Lithobius forficatus (Chilopoda,Lithobiidae)

Lithobius forficatus (Myriapoda, Chilopoda, Lithobiidae) is a widespread species of centipede that is common across Europe. Its midgut epithelial cells are an important line of defense against toxic substances that originate in food, such as pathogens and metals. Despite this important role, the biology of the midgut epithelium is not well known. Here we describe the ultrastructure of the midgut epithelium, as well as the replacement of degenerated midgut epithelial cells. The midgut epithelium of L. forficatus is composed of digestive, secretory, and regenerative cells. The cytoplasm of digestive cells shows regionalization in organelle distribution, which is consistent with the role of these cells in secretion of enzymes, absorption of nutrients, and accumulation of lipids and glycogen. Secretory cells, which do not reach the luminal surface of the midgut epithelium, possess numerous electron‐dense and electron‐lucent granules and may have an endocrine function. Hemidesmosomes anchor secretory cells to the basal lamina. Regenerative cells play the role of midgut stem cells, as they are able to proliferate and differentiate. Their proliferation occurs in a continuous manner, and their progeny differentiate only into digestive cells. The regeneration of secretory cells was not observed. Mitotic divisions of regenerative cells were confirmed using immunolabeling against BrdU and phosphohistone H3. Hemocytes associate with the midgut epithelium, accumulating between the visceral muscles and beneath the basal lamina of the midgut epithelium. Hemocytes also occur among the digestive cells of the midgut epithelium in animals infected with Rickettsia‐like microorganisms. These hemocytes presumably have an immunoprotective function in the midgut.     

The chemosensory behaviour of Lithobius forficatus (Myriapoda: Chilopoda). 2. Bioassay and chemistry of the coxal pheromone

The common brown centipede, Lithobius jorJcatus (L.), releases a sex-specific pheromone which emanates from distinctive cuticular openings on the coxae of the hind legs-the coxal pores (Littlewood & Blower, 1987). The pheromone can be isolated from tissue extracts of the coxal organs (surrounding the coxal pores) and from substrata which have been 'marked by male or female L. forjcatus and presented to centipedes in the form of a behavioural bio-assay.
In this paper, further evidence is presented for a centipede pheromone, the chemical components of male and female secretions are separated using Gas Layer Chromatography (GLC), a qualitative chemical analysis is described and the nature of the pheromone is discussed.     

Etude ultrastructurale des spermatogonies et de la croissance spermatocytaire chez Lithobius forficatus L. (Myriapode Chilopode)

Résumé L'étude ultrastructurale des spermatogonies et des spermatocytes a été envisagée chez Lithobius forficatus L. (Myriapode Chilopode). Les spermatogonies présentent un noyau à chromatine dispersée dont le nucléole est condensé. Leur cytoplasme renferme des ribosomes, libres ou associés en polysomes; les mitochondries et les dictyosomes sont peu abondants.Au cours de la croissance spermatocytaire, le noyau et le cytoplasme augmentent considérablement de volume. La dispersion de la chromatine est plus importante et le nucléole présente de nombreuses figures de bourgeonnement. Les ribosomes, les mitochondries, les dictyosomes et le reticulum endoplasmique sont très abondants. De nouvelles formations sont observables: 1) des amas de saccules réticulaires concentriques, délimitant une zone cytoplasmique; 2) des empilements lamellaires ayant leur origine soit dans le reticulum banal, soit dans les amas concentriques.L'activité synthétique se manifeste lors de la croissance spermatocytaire par: 1) d'abondants échanges nucléo-cytoplasmiques; 2) une augmentation du nombre de mitochondries et des ribosomes; 3) une intense activité golgienne. Elle est vraisemblablement en rapport avec le gigantisme spermatocytaire observé chez les Myriapodes Chilopodes.

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Ultrastructural study of spermatogonia and spermatocyte growth of Lithobius forficatus L. (Myriapoda, Chilopoda)

Summary The ultrastructure of spermatogonia and spermatocytes of Lithobius was studied. The spermatogonia show a nucleus with dispersed chromatin and a compact nucleolus. Their cytoplasm contains free ribosomes and polysomes; mitochondria and dictyosomes are not abundant.During spermatocyte growth, nucleus and cytoplasm increase markedly in volume. The dispersion of the chromatin is more pronounced; the nucleolus shows frequent budding. Ribosomes, mitochondria, dictyosomes and endoplasmic reticulum are very numerous. New structures are (1) accumulations of concentric reticular saccules with a cytoplasmic center, (2) piles of narrow cisternae originating from the regular endoplasmic reticulum or from concentric saccules.The synthetic activity during spermatocyte growth manifests itself by (1) numerous signs of nucleo-cytoplasmic exchanges, (2) an increase in the number of mitochondria and ribosomes, (3) a high activity of the Golgi apparatus. This is probably related to the gigantism of spermatocytes in Chilopoda.

     

Nature et origine des réserves vitellines dans l'ovocyte de Lithobius forficatus L. (Myriapode Chilopode)

Résumé Dans l'ovocyte de Lithobius forficatus L., les trois types classiques de réserves vitellines apparaissent successivement: glycogène, globules lipidiques et vitellus protéique. La synthèse du glycogène semble effectuée au contact des membranes ergastoplasmiques. Les globules lipidiques paraissent élaborés à partir d'un matériel qui transite par le reticulum puis l'appareil de Golgi. Le vitellus protéique est d'origine exogène et pénètre dans l'ovocyte par pinocytose.L'ovocyte mûr est très riche en réserves vitellines et ne renferme qu'une mince couche cytoplasmique périphérique, pauvre en organites.

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Nature and origin of the vitelline reserves in the oocyte of lithobius forficatus L. (myriapoda chilopoda)

Summary In the oocyte of Lithobius forficatus L., the three classical types of vitelline reserves appear successively: glycogen, lipid droplets, and protein yolk. Glycogen synthesis seems to occur in contact with the rough endoplasmic reticulum. Lipid droplets appear to be elaborated from a material which passes through the endoplasmic reticulum and the Golgi apparatus. Protein yolk originates elsewhere and enters the oocyte by pinocytosis.The mature oocyte is almost completely filled with yolk. There remains only a thin outer coat of cytoplasm with very few organelles.

     

Barcoding Fauna Bavarica: Myriapoda - a contribution to DNA sequence-based identifications of centipedes and millipedes (Chilopoda, Diplopoda)

We give a first account of our ongoing barcoding activities on Bavarian myriapods in the framework of the Barcoding Fauna Bavarica project and IBOL, the International Barcode of Life. Having analyzed 126 taxa (including 122 species) belonging to all major German chilopod and diplopod lineages, often using four or more specimens each, at the moment our species stock includes 82% of the diplopods and 65% of the chilopods found in Bavaria, southern Germany. The partial COI sequences allow correct identification of more than 95% of the current set of Bavarian species. Moreover, most of the myriapod orders and families appear as distinct clades in neighbour-joining trees, although the phylogenetic relationships between them are not always depicted correctly. We give examples of (1) high interspecific sequence variability among closely related species; (2) low interspecific variability in some chordeumatidan genera, indicating that recent speciations cannot be resolved with certainty using COI DNA barcodes; (3) high intraspecific variation in some genera, suggesting the existence of cryptic lineages; and (4) the possible polyphyly of some taxa, i.e. the chordeumatidan genus Ochogona. This shows that, in addition to species identification, our data may be useful in various ways in the context of species delimitations, taxonomic revisions and analyses of ongoing speciation processes.     

The hemocytes of the centipedeLithobius forficatus (Chilopoda,Lithobiomorpha)

Summary Five different types of hemocytes are found within the hemolymph ofLithobius forficatus: (1) small prohemocytes, (2) very actively spreading plasmatocytes, (3) granulocytes which have a lower spreading ability but tend to agglutinate, (4) spherulocytes which are filled with spherules, and (5) presumably, a coagulocyte, characterized by instant disintegration. Cystocytes as described forL. forficatus in the literature are preparation artifacts. Cell types are characterized by electron microscopy and in vitro and vital staining techniques at the light microscopic level. Results are discussed with reference to different nomenclatures and functions of hemocytes in other arthropods.     

Fine structure and function of the coxal glands of lithobiomorph centipedes: Lithobius forficatus and L. crassipes (Chilopoda,Lithobiidae)

The ultrastructure of the coxal glands and associated tissues in the centipedes Lithobius forficatus and Lithobius crassipes has been examined in the light of two contrasting functional hypotheses postulated by different authors. Lithobiomorph chilopods possess eight sets of pores on the posterioventral border of the coxal podomeres of leg pairs 12–15 in adult (maturus) and subadult (pseudomaturus) stadia. A modified cuticular hypodermis, known as the coxal gland, surrounds the distal portion of each blindended pore. Each gland is made up of cells which contain large numbers of hypertrophied mitochondria and a highly folded apical and basal plasma membrane. The similarity of the coxal gland to so called “transporting epithelia” is discussed and further comparisons are made between these and secretory glands in arthropods. A careful consideration of both functional hypotheses (osmoregulation or pheromone release) has revealed the possibility that the coxal gland may encompass both functions.     

A common terminology for the external anatomy of centipedes (Chilopoda)

A common terminology for the external morphological characters of centipedes (Chilopoda) is proposed. Terms are selected from the alternatives used in the English literature, preferring those most frequently used or those that have been introduced explicitly. A total of 330 terms are defined and illustrated, and another ca. 500 alternatives are listed.     

Coxal organs in geophilomorpha (Chilopoda). Organization and fine structure of the transporting epithelium

Summary The coxal organs of different Geophilomorpha were studied by scanning and by transmission electron microscopy.1) The coxae of the last trunk-segment contain pores in different arrangements and numbers. They are the openings of the coxal organs.2) The coxal organs are formed by four different cell types: the main epithelium consists of radially arranged transporting cells, surrounded by junctional cells, gland cells, and the cells of the pore channel.3) The cells of the transporting epithelium show an enlargement of the apical and basal surface. Deep and narrow extracellular channels of the apical infoldings are closely associated by mitochondria (plasmalemma-mitochondrial complexes). The epithelium is covered by a prominent cuticle with a spacious subcuticle.4) A distinct mucous layer covers the cuticle of the transporting epithelia, and is secreted by the gland cells.5) A small cellular sheath separates the epithelium of the coxal organ against the haemolymph.6) The possible function of the coxal organs in ion and fluid transport is discussed.     

A redescription of the large geophilid centipede Chilenophilus corralinus (Attems, 1903) (Myriapoda: Chilopoda: Geophilomorpha), with new distribution records from southern Chile

The little known geophilomorph centipede Chilenophilus corralinus (Attems, 1903) (Myriapoda: Chilopoda, Geophilidae), a large geophilid species from South America is herein redescribed and illustrated based on new specimens collected in the Andes of Southern Chile. New morphological features of specific value are also given for the taxon. Chilenophilus corralinus is reported for the first time from the following Chilean localities: Region XIV (de Los Ríos region): Valdivia province: Pirihueico; camping “La Herradura”; Mafil. Region X (de Los Lagos region): Chiloé province: Chiloé Island. Llanquihue province: Parque Nacional Alerce Andino. Palena province: Hualaihué; Chaitén; Palena; Futaleufú.