The midgut of Tomocerus minor lubbock (Insecta,Collembola): Ultrastructure,cytochemistry, ageing and renewal during a moulting cycle

The midgut cells of Tomocerus minor (Insecta, Collembola) were examined with the electron microscope and cytochemically. The midgut epithelium consists of a series of cells characterised by numerous mineral concretions scattered throughout the cytoplasm. Mitochondria are abundant; microvilli are well developed at the apical surface of the cell. A zonula continua (continuous junction) characterises the apical contact region of these cells. Polysaccharides, glycoproteins and carbohydrate components have been demonstrated on the surface of microvilli. Peritrophic membranes surround the food bolus and preserve midgut cells from mechanical abrasion. Lysosomes are present during the alimentary period and show strong acid phosphatase activity. During an intermoulting cycle, two stages can be observed: (1) the postexuvial feeding period during which cytoplasmic extrusions appear at the apical part of the cell: lysosomes increase in number and autophagic vacuoles appear. (2) The preexuvial fasting period; a new epithelium grows beneath the old one and pushes it into the lumen. Degeneration processes can be observed in the old epithelium. This excretory reactivity of the midgut epithelium has been compared to the cycle of the cuticle.     

Embryogenesis of a springtail,Tomocerus ishibashii (Collembola,tomoceridae): External morphology

The external features of the developing embryos of the springtail, Tomocerus ishibashii, are described. The clypeolabral anlage arises as a single, unpaired swelling. The entognathy is completed by the ventral growth of the tergal anlagen of mandibular, maxillary, and labial segments. These anlagen also form the posterior part of the cranium. The palpi of maxilla and labium are homologous with the telopodites, and proximal parts of these head appendages are homologous with the coxopodites. The sternal element of the labial segment does not participate in the postmentum formation. The anlagen of abdominal appendages appear in the first to the fourth abdominal segments. The first, third, and fourth appendage anlagen form the ventral tube, tenaculum, and furcula, respectively. The fused proximal parts of the first, third, and fourth appendage anlagen are homologous with the coxopodites, and the distal parts which do not fuse are homologous with the telopodites. The anlagen of the second abdominal appendages become flattened and spread over the ventral side of this segment. The ventral structures of the first four abdominal segments are appendicular in origin.     

Das Doppelauge vonEntomobrya muscorum Nicolet (Insecta,Collembola)

Zusammenfassung Der bodenlebende CollembolaEntomobrya muscorum hat an jeder Kopfseite ein loses Aggregat von acht Ommatidien, von denen zwei einen deutlich kleineren Linsendurchmesser aufweisen. Die sechs großen Ommatidien werden als Haupt-, die zwei kleinen als Nebenaugen bezeichnet. Beide Ommatidientypen haben verschieden gebaute Rhabdome, so daß der Komplex ein Doppelauge darstellt. Es wird der Bau der Hauptaugen geschildert. Ihr sensibler Teil setzt sich aus acht Retinulazellen zusammen, die ein offenes Rhabdom in zwei Lagen bilden. Eine zentrale sechste Zelle in der distalen Schicht weicht in ihrer Feinstruktur deutlich von den übrigen ab. Die Rhabdome beider Ommatidientypen werden miteinander verglichen, und ihre funktionelle Bedeutung vor allem in Hinblick auf eine mögliche Arbeitsteilung für eine Polarisationswahrnehmung wird erörtert. Es wird kurz die phylogenetische Bedeutung dieser Augen für die Insekten und die Collembolen selbst angeschnitten.

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The double eye ofEntomobrya muscorum nicolet (Insecta, Collembola)

Summary The ground living CollembolaEntomobrya muscorum has on each side of the head eight circular ommatidia in an arrangement characteristic of the species. Of these two have a distinctly smaller lens. The six big ommatidia are called primary eyes, the two small ones accessory eyes. The two types of ommatidia have different rhabdom structures, so justifying the term double eye. The structure of the primary eyes is described. The photosensitive part is constructed of eight retinula cells which form an open rhabdom in two layers. The central sixth cell in the distal layer is quite different from the others. The rhabdoms of both types are compared. The possible function of the accessory eyes in perception of the e-vector is discussed. The phylogenetic significance of these eyes for the Insecta and the Collembola is considered.

     

Polymorphism of polytene chromosomes in Bilobella aurantiaca (Insecta: Collembola)

Polytene chromosomes in the salivary glands of Bilobella aurantiaca are analysed in a population from Sierra de Gredos (Central Spain). They display a high degree of heterozygosity. Only a short part of the total length of the seven pairs of homologous chromosomes, examined in a single favourable specimen, was homozygous. Concerning the short arm of chromosomes III and IV, the respective proportion of heterozygosity for the whole population was 38.6% and 92.9%. Some hypotheses are formulated to account for these results: differential funtioning of homologous chromosomes, generalisation of minute duplications and deficiencies, complex inversions (which cannot be specified owing to the asynapsis of homologous chromosomes), and possibly local large deletions.     

Studies on populations of Folsomia candida (Insecta: Collembola)

Summary Experiments are described that were designed to investigate the effects of food availability and rate of exploitation on the growth and production of populations of Folsomia candida (Willem). In an initial experiment in which there was excess food it was found that increasing the rate of exploitation resulted in increases in both the numerical and biomass productivity of the populations. In a second experiment it was shown that, when there is severe competition for food, the rate of exploitation does not affect either the biomass or the numerical production. It is concluded that the effect of overcrowding, in the form of competition for space, does contribute to the growth in numbers of populations, but that the supply of food plays a more important role in regulating the population. Anomalous results, showing that exploitation has a lesser effect when there is severe competition for food, are discussed.     

Cytochemistry and X-ray microprobe analysis of the midgut of Tomocerus minor lubbock (Insecta,Collembola) with special reference to the physiological significance of the mineral concretions

Summary Histochemical and cytochemical analyses have been made on the mineral concretions within the midgut cells of Tomocerus minor. The classical histochemical methods are not specific and precise enough and have been supplemented with cytochemical techniques on ultrathin sections. The most interesting of these was the K-pyroantimonate technique combined with glutaraldehyde-osmium fixation. This technique shows the distribution of cations such as Ca⁺⁺, K⁺, Mg⁺⁺ and Na⁺ on the concentric layers of the concretions. Chloride ions can be detected by means of the silver lactate technique. The action of calcium chelators such as E.D.T.A. shows an important distribution of calcium ions in the concretions. The spectra obtained by electron probe microanalysis from areas of fresh, dried and carbon coated midguts as well as from carbon coated semithin or ultrathin sections reveal the presence of Ca, K, Mg, S, Cl and P principally. Other elements such as aluminium, silicon and manganese have also been detected. Iron is not always present. The chemical and X-ray analytical investigations indicate that the midgut concretions are mainly built up of calcium, potassium, magnesium and sodium phosphates, perhaps associated with chlorides and carbonates. An organic matrix formed by polysaccharides seems to join the different mineral layers. These concretions may be formed within the vesicles of rough endoplasmic reticulum. The midgut cells are highly differentiated and very active in transport. Extensive basal infoldings and apical microvilli as well as lateral membranes are a site of small cationic deposits. The possible pathway of ion transport in the cell and the physiological significance of the concretions are discussed. The principal function of these concretions seems to be the maintenance of the mineral balance and to trap foreign and excess ions.

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Résumé L'analyse chimique des sphérocristaux de l'intestin moyen de Tomocerus minor a été réalisée. Les méthodes histochimiques courantes manquant souvent de spécificité et de sensibilité ont été complétées avec des méthodes cytochimiques sur coupes ultrafines. La plus intéressante a été la technique du pyroantimonate de K montrant la distribution des cations Ca⁺⁺, K⁺, Mg⁺⁺, Na⁺ sur les couches concentriques des sphérocristaux. La technique au lactate d'argent permet de déceler les ions Cl^-. L'action d'agents chélateurs du Ca tels l'E.D.TA. montre une importante distribution du calcium dans les sphérocristaux. L'analyse spectrographique d'étalements de mésentérons séches, carbonés et de coupes semi-fines ou ultrafines carbonées montre la présence de Ca, K, Mg, S, Cl, P, Na. D'autres éléments tels l'Al et le Si ont pu être détectés. Le Fe n'est pas toujours présent. Les sphérocristaux semblent formés essentiellement de phosphates de calcium, de potassium, de magnésium, de sodium associés peut-être à des chlorures ou des carbonates. Une matrice organique constituée essentiellement par des polysaccharides semble lier les différentes couches minérales. Ces sphérocristaux prennent naissance à l'intérieur des vésicules de l'ergastoplasme. Les cellules de l'intestin moyen sont très différenciées et sont le siège de nombreux transports actifs. Les replis basaux de la membrane plasmique, les microvillosités apicales, de même que les membranes latérales sont le siège de dépôts de cations. Le transport des ions dans les cellules ainsi que le rôle physiologique des sphérocristaux sont discutés. Le maintien de la balance hydrique ainsi que le piégeage d'ions étrangers ou en surplus semblent être la principale fonction des sphérocristaux.

     

Dicyrtotnina and Ptenothrix (Insecta:Collembola) of the Solomon Islands

This paper deals primarily with the collembolan species of the subfamily Dicyrtominae collected during the Royal Society Expedition to the Solomon Islands of 1965 by P. N. Lawrence, supplemented by material from these islands collected by Dr and Mrs P. J. M. Greenslade, all of which is in the British Museum (Natural History). Additional supporting material described here has been derived from the Noona Dan Expedition of the Zoological Museum, Copenhagen and from the Bernice P. Bishop Museum, Hawaii. I should like to express my thanks to the individuals and Museum sconcerned.     

Ecology of Cryptopygus sverdrupi (Insecta: Collembola) from Dronning Maud Land,Antarctica

Summary Observations on the ecology of Cryptopygus sverdrupi Lawrence (Collembola, Isotomidae) were made with specimens from the Mühlig-Hofmannfjella, Dronning Maud Land, Antarctica. At an elevation of 1600 m a.s.l. the species was numerous in association with the green alga Prasiola on gravel fields and in crevices of large boulders. The distribution of size-classes in field samples suggested that the population comprised several overlapping generations. Growth and development is probably very slow due to long winters and daily periods of subzero temperatures in their microhabitat during the summer. Specimens collected in mid-January had a mean supercooling point of-24.6°C with small individual variations. The lack of high supercooling points in the summer suggests that the springtails feed on a nuleatorfree diet. The ability to supercool was increased during prolonged starvation and acclimation at 0,-4 and-8°C. Glycerol and other potential low molecular weight cryoprotective substances were demonstrated in specimens acclimated at-4 and-8°C. The species possessed a relatively high tolerance to desiccation.Publication No. 81 of the Norwegian Antarctic Research Expeditions (1984/85)     

Ultrastructure of the male accessory glands ofAllacma fusca(Insecta, Collembola)

The accessory glands ofAllacma fusca(L.) (Insecta, Collembola, Sminthuridae) consist of a series of secretory units that are arranged in parallel and open into the ejaculatory duct. Each unit is composed of microvillate cells stacked around a common cavity. Basal cells are involved in ion-control of fluids from the hemocoel to the cavity. The intermediate and apical cells, which have a laminar appearance and contain many microtubules, are involved in the structural integrity of the unit. Supporting cells ensheath the most apical cells. Large openings in the cuticle allow the gland secretion to flow into the ejaculatory duct lumen. These openings are protected by a porous cuticle different from that lining the epithelium of the ejaculatory duct. Conspicuous muscle fibers run along the lateroventral side of the ejaculatory duct beneath the insertion of the accessory glands. The fine structure of the accessory glands indicates that they are type I ectodermic glands as defined by Noirot & Quennedey (1974). Their function could be to control the fluidity of the material for spermatophore formation and to ensure the proper physiological conditions for spermatozoa stored in the ejaculatory duct lumen.     

Molecular evidence for a fourth species within the Isotoma viridis group (Insecta, Collembola)

For identification of single species within the Isotoma viridis group, we present polymerase chain reaction–restriction fragment length polymorphism (PCR–RFLP) as a fast and efficient DNA-based molecular method. We used five PCR primers amplifying the cytochrome oxidase II (COII) region (760 bp) of the mitochondrial DNA. The sequences clearly separated four species ( I. viridis , I. riparia , I. anglicana and I. caerulea ) out of samples from Norway, Sweden, Germany and Switzerland. Examination of genetic variation and phylogenetic relationship did not support the separation of two colour pattern forms of I. viridis into distinct species. For RFLP, several restriction enzymes were tested for their ability to produce not only species-specific restriction fragment patterns but to discriminate more than one species per enzyme used with as few cleavage sites as possible. Such a design should render a clear fragment pattern when performing a double digest. These demands appear to be fulfilled best by the combination of the restriction enzymes Mfe I, Nci I and one of Aci I, Bst EII, Nde I, or Sfc I. From the enzymes tested in a previous study, Ase I proved to be reliable, whereas Mbo I can no longer be recommended.     

A peculiar cave species of Tomocerus (Collembola,Tomoceridae, Tomocerinae) from Vietnam,with a discussion of the postantennal organ and prelabral chaetae in Tomocerinae

The first Tomocerus species with a postantennal organ (PAO) in the adult stage is described from Vietnam. Tomocerus postantennalis sp. n. differs from the other PAO-possessing tomocerid, Tomolonus reductus Mills, 1948, mainly in the morphology of PAO, the number of ocelli, the number of chaetae in trochantero-femoral organ and several features of the furca. The new species is placed in Tomocerus because of the presence of a toothlet on the outer basal mucronal tooth and the absence of the diagnostic character states of Plutomurus Yosii, 1956 and Aphaenomurus Yosii, 1956. Besides the presence of PAO, the new species is peculiar in having six prelabral chaetae, instead of four as in other Tomocerus species. The new species is similar to Tomocerus folsomi Denis, 1929 and Tomocerus ocreatus Denis, 1948 in the type of dental spines but different from them in the body colour, the relative length of antennae to body, the number of unguis inner teeth and the number of mucronal intermediate teeth.     

High diversity and community permeability: the riparian Collembola (Insecta) of a Pyrenean massif

This work deals with the structure of Collembolan communities in riparian habitats. Sixty samples collected along running waters in a Pyrenean massif have been analysed. Diversity was much higher than for any comparable habitat studied in the literature. The proportion of rare species was particularly high, and explained a large part of the observed diversity. The contribution of specialized hydrophilous species to overall diversity was low: they were 5 times less numerous than non-hydrophilous species for a similar global abundance. Correspondence analysis showed that the riparian community was loosely structured, without any strong determining factor. Epigeomorphic species of hydrophilous Collembola were, however, clustered in a well-defined group. The forest type, of major importance for soil fauna diversity in the area, was a poor predictor of diversity in the riparian habitats. Conversely, richness was significantly related to distance from water, in spite of the samples having been all collected from permanently water-saturated substrates. The importance of non-hydrophilous species for the diversity of riparian habitats along running water is interpreted as reflecting the ecological permeability of the hydrophilous community, resulting from the spatial organisation of the habitat in narrow strips, and its frequent disturbance by flooding. Finally, the riparian habitat may act as a refuge for a significant proportion of the soil species affected by reafforestation, presently the most severe disturbance of Pyrenean ecosystems.     

Chromosome elimination and sex determination in springtails (Insecta, Collembola).

A post-zygotic mechanism of sex determination is described in the two symphypleonans Dicyrtomina ornata (Nicolet) and Ptenothrix italica Dallai. The process consists of the loss of two sex chromosomes from the male embryo. At the end of the first meiotic division of spermatogenesis, a second chromosome elimination occurs, allowing half the secondary spermatocytes, later transformed into spermatids, to receive a complete haploid set of chromosomes. The secondary spermatocytes, which receive an incomplete set of chromosomes, degenerate. Males of the two collembolan species, therefore, produce a reduced number (50%) of spermatozoa. Females of D. ornata have 2n = 12 and males 2n = 10 chromosomes; females of P. italica have 2n = 14 and males 2n = 12 chromosomes. In both species, oogenesis proceeds normally and chromosomes pair and form chiasmata in meiotic prophase. The adaptive significance of this post-zygotic mechanism of sex determination is discussed. The mechanism seems to be a characteristic feature of the suborder Symphypleona. The neanurid Arthropleona Anurida maritima (Guérin), which was studied for comparative analysis, has 2n = 8 chromosomes and normal spermatogenesis producing haploid nuclei with four chromosomes. J. Exp. Zool. (Mol. Dev. Evol.) 285:215-225, 1999.     

A novel method of self-righting in the springtail Sminthurus viridis (Insecta: Collembola)

Photographic evidence shows that the ventral tube can be used as a tentacle-like mechanical aid for self-righting from an inverted position. It is proposed that the extraordinary length of the vesicles of the ventral tube in globular springtails has been evolved in order to combat postural instabilities resulting from spherical symmetry of body form.     

Population structure, gene flow and evolutionary relationships in four species of the genera Tomocerus and Pogonognathellus (Collembola, Tomoceridae)

Genetic diversity and gene flow at 10 allozyme loci were investigated in 13 natural populations of four species of the collembolan genera Tomocerus and Pogonognathellus. Levels of observed heterozygosity were found to vary from 0.033 in P. longicornis to 0.120 in T. vulgaris. Average Nei's genetic distances (D) ranged from D = 0.222 between populations of P. ftavescens to D = 0.647 between populations of T. vulgaris. Genetic distances between species were always 1.000 and the highest value (D = 4.321) was between P. longicomis and one population of T. vulgaris. Values of F_sr were very high in all species (from 0.567 to 0.696) and levels of gene flow (Nm) derived from F_ST and the private allele method were low (Nm <1). Gene flow was significandy higher only in two subsets of populations of T. vulgaris. The Plio-Pleistocenic geological rearrangements and the effect of stochastic events, such as genetic drift, are invoked to explain the different levels of genetic divergence between and within species. Distance- and character-based approaches were used to reconstruct evolutionary relationships between and within species. While the monophyly of all species was confirmed, the results did not univocally support the monophyly of the two genera, leaving the question of their generic or subgeneric status unresolved.     

Degeneration of the midgut epithelium in Allacma fusca L. (Insecta, Collembola, Symphypleona): apoptosis and necrosis

Apoptotic and necrotic changes in the midgut epithelium cells of Allacma fusca (Collembola, Symphypleona) are described at the ultrastructural level. The morphological sign indicating the beginning of the apoptotic process in these cells is their shrinkage and the transformation of their mitochondria. The nucleus assumes a lobular shape and finally undergoes fragmentation. The intercellular junctions between an apoptotic cell and adjacent epithelial cells gradually disappear. Apoptotic cells are discharged into the midgut lumen just beneath the peritrophic membrane, where they are initially distributed singly but ultimately form a single layer. No phagocytosis was observed, so no apoptotic bodies are formed. Only young midgut epithelium shows apoptosis; as cells age, necrosis accompanies apoptosis, and necrosis finally completely replaces apoptosis.     

Ultrastructural changes in the midgut epithelium in Podura aquatica L. (Insecta, Collembola, Arthropleona) during regeneration

Data considering the degeneration and regeneration of the midgut epithelium in the primitive wingless insects, such as Collembola, are rather poor. Also information, which treats the regenerative cells as the primordial cells, is poorly known. The midgut epithelium of Podura aquatica L. (Insecta, Collembola, Arthropleona) is formed by the epithelial and regenerative cells. The epithelial cells show distinct regionalisation in the organelles distribution. The ultrastructure of the basal, perinuclear and apical regions of the epithelial cells is described. As in insects without Malpighian tubules, structures which resemble urospherites occur in the cytoplasm of the epithelial cells. After degeneration of the entire midgut epithelium, a new epithelium is formed from regenerative cells. During the process of regeneration, the degenerated epithelium gradually is separated from the basal lamina by the newly formed one. Finally, the detached epithelium is moved into the midgut lumen. Regenerative cells play a role of primordial cells during epithelial regeneration.