Embryogenesis of the dipluran Lepidocampa weberi Oudemans (Hexapoda,Diplura, Campodeidae): External morphology

The external features of the embryo of the dipluran, Lepidocampa weberi Oudemans are described. The long germ band is formed, and blastokinesis is a simple flexion of the germ band. The primary dorsal organ is formed between the cephalic and abdominal ends by concentration of serosal cells. The mouth fold is formed by ventral extension of the intercalary, mandibular, and maxillary terga, through which entognathy is completed. The posteroventral region of the mouth fold develops into the admentum. Rotation of the labial anlagen is involved in labial formation, and the glossa, paraglossa, and labial palp acquire a tandem arrangement. The postmentum is formed by fusion of the labial subcoxae and is appendicular in origin. The styli and exertile vesicles are derived from the distal parts of bifurcated appendicular anlagen of the second to seventh abdominal segments. The columnar appendage of the first abdominal segment is serially homologous with the exertile vesicles of the following segments. The abdomen is composed of ten segments, and the cercus is the appendage of the tenth, last abdominal segment. Embryogenesis of Lepidocampa weberi resembles that of the rhabduran Campodea staphylinus (Uzel, 1898) as well as that of the dicelluratan Japyx major (Silvestri, '33). It may be emphasized that the rhabduran and dicelluratan diplurans share important features such as entognathy formation and abdominal organization, and the resemblance between them seems to be close enough to postulate their close affinity. Some embryogenetic features, which Diplura and Collembola share, are recognized as plesiomorphic and the manner of entognathy formation may significantly differ. J. Morphol. 237:101–115, 1998. © 1998 Wiley-Liss, Inc.     

Microbial Community Structure in Midgut and Hindgut of the Humus-Feeding Larva of Pachnoda ephippiata (Coleoptera: Scarabaeidae)

The guts of soil-feeding macroinvertebrates contain a complex microbial community that is involved in the transformation of ingested soil organic matter. In a companion paper (T. Lemke, U. Stingl, M. Egert, M. W. Friedrich, and A. Brune, Appl. Environ. Microbiol. 69:6650-6658, 2003), we show that the gut of our model organism, the humivorous larva of the cetoniid beetle Pachnoda ephippiata, is characterized by strong midgut alkalinity, high concentrations of microbial fermentation products, and the presence of a diverse, yet unstudied microbial community. Here, we report on the community structure of bacteria and archaea in the midgut, hindgut, and food soil of P. ephippiata larvae, determined with cultivation-independent techniques. Clone libraries and terminal restriction fragment length polymorphism analysis of 16S rRNA genes revealed that the intestines of P. ephippiata larvae contain a complex gut microbiota that differs markedly between midgut and hindgut and that is clearly distinct from the microbiota in the food soil. The bacterial community is dominated by phylogenetic groups with a fermentative metabolism (Lactobacillales, Clostridiales, Bacillales, and Cytophaga-Flavobacterium-Bacteroides [CFB] phylum), which is corroborated by high lactate and acetate concentrations in the midgut and hindgut and by the large numbers of lactogenic and acetogenic bacteria in both gut compartments reported in the companion paper. Based on 16S rRNA gene frequencies, Actinobacteria dominate the alkaline midgut, while the hindgut is dominated by members of the CFB phylum. The archaeal community, however, is less diverse. 16S rRNA genes affiliated with mesophilic Crenarchaeota, probably stemming from the ingested soil, were most frequent in the midgut, whereas Methanobacteriaceae-related 16S rRNA genes were most frequent in the hindgut. These findings agree with the reported restriction of methanogenesis to the hindgut of Pachnoda larvae.     

Embryogenesis of the dipluran Lepidocampa weberi Oudemans (hexapoda: diplura, campodeidae): formation of dorsal organ and related phenomena

The developmental changes of embryonic membranes of a dipluran Lepidocampa weberi, with special reference to dorsal organ formation, are described in detail by light, scanning, and transmission electron microscopies. Newly differentiated germ band and serosa secrete the blastodermic cuticle at the entire egg surface beneath the chorion. Soon after, the serosal cells start to move dorsad. All the serosal cells finally concentrate at the dorsal side of the egg and form the dorsal organ. During their concentration, the serosal cells attenuate their cytoplasm to form filaments. The extensive area from which the serosa has receded is occupied by a second embryonic membrane, the amnion, which originates from the embryonic margin. The embryo and newly emerged amnion then secrete three fine cuticular layers, "cuticular lamellae I, II, and III," above which the filaments of the (developing) dorsal organ are situated. With the progression of definitive dorsal closure, the amnion reduces its extension, the dorsal organ is incorporated into the body cavity of the embryo, and the amnion and dorsal organ finally degenerate.The dorsal organ of diplurans is formed by the concentration of whole serosal cells, while that of collembolans is formed by the direct differentiation of a part of serosal cells. However, the dorsal organs of diplurans and collembolans closely resemble each other in major aspects, including that of ultrastructural features, and there is no doubt regarding their homology. The amnion, which has been regarded as being a characteristic of Ectognatha, also develops in the Diplura. This might suggest a closer affinity between the Diplura and Ectognatha than previously believed.     

Midgut epithelium formation in Thermobia domestica (Packard) (Insecta, Zygentoma)

The origin of midgut epithelium may begin either from yolk cells (energids), tips of stomo- and proctodaeum (ectoderm), inner layer (endoderm) or from both kinds of the above mentioned cells. The origin of the midgut epithelium in wingless insects (Apterygota) has still not been determined. In Thermobia domestica the formation of midgut is much delayed, and it completes in the post-embryonic stage, while the stomo- and the proctodaeum are well-developed in the embryonic period. The energids, which remain inside the yolk, start to migrate to its periphery, where they arrange singly close to cell membrane. The yolk mass with the energids at the 14th day of embryogenesis are referred to as the primary midgut. During the first instar larval stage more and more energids migrate to the yolk periphery and the cell membrane starts to form numerous foldings surrounding the groups of energids, which in turn lead to formation of isolated regenerative cell groups. Eventually the cell membrane invaginations reach the center of the yolk mass. Large cells of the primary epithelium, surrounding the newly formed midgut lumen are formed. The cells of the primary epithelium are filled with yolk and are equipped with microvilli pointing to the midgut lumen. As the yolk is being digested, the process of the primary epithelium cells degeneration begins. The cells are getting shorter and start to degenerate. The definitive midgut epithelium is formed from proliferating regenerative cells. It consists of regularly spaced regenerative cell groups as well as the epithelial cells. The ultrastructure of both these cell groups has been described.     

Oogenesis in Campodea sp. (Insecta,Diplura): Chorion formation and the ultrastructure of follicle cells

During advanced vitellogenesis the follicle cells of Campodea sp. are well developed and contain numerous electron-dense secretory vacuoles. In the postvitellogenic phase the contents of these vacuoles are released from follicle cells and give rise to a thin chorion on the oocyte surface. Concurrently, segregation of yolk spheres takes place in the ooplasm: small spheres migrate to the oocyte periphery and come to lie in the so-called peripheral zone of cytoplasm, whereas large spheres remain in the cell centre.     

Ultrastructure du rein labial céphalique deCampodea chardardi Condé (Diplura,Insecta)

Résumé L'ultrastructure du rein labial céphalique deCampodea chardardi Condé a été étudiée. Cette néphridie comprend trois parties: le saccule terminal, le labyrinthe et le canal excréteur.Les cellules du saccule sont des podocytes typiques, contenant de nombreuses vacuoles de pinocytose et des inclusions diverses. La lumière est envahie par des micro-organismes bacilliformes.Le labyrinthe possède des cellules à indentations basales profondes avec de nombreuses mitochondries, et des microvilli distaux.Le canal excréteur débouchant sur la face ventrale du labium est caractérisé par la présence d'une intima cuticulaire.Le rein labial des Diploures a été comparé avec des organes segmentaires néphridiens d'autres Arthropodes, et avec le néphron des Vertébrés.

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The fine structure of the labial cephalic kidney ofCampodea chardardi Condé (diplura, insecta)

Summary The ultrastructure of the cephalic labial kidney ofCampodea chardardi Condé has been studied. Each nephridium is subdivided into three segments: end-sac, coiled tubule and efferent-duct.The cells of the sacculus are typical podocytes, and contain numerous pinocytotic vesicles and various vacuoles. The lumen contains micro-organisms.The cells of the coiled tubule bear basal infoldings with numerous mitochondria and distal microvilli.The efferent duct terminates close to the ventral face of labium, and possesses characteristic cuticular intima.The labial kidney of Diplura is compared with published data on the nephridial organs of other Arthropods and Vertebrate nephron.

Nous tenons à remercier Monsieur le Professeur Noirot pour ses encouragements et conseils.     

Fine structure of the ladybird spermatozoa (Insecta,Coleoptera, Coccinellidae)

The sperm structure of several ladybird species belonging to different subfamilies of Coccinellidae was studied. Three main sperm types were clearly recognized, and were characterized by differences in acrosomal length, the presence of a dense coat around the acrosome, the length of the basal body, the amount of the centriole adjunct material, and the diameter of the mitochondrial derivatives. However, the whole group shares a pattern of the posterior sperm region uncommon for insects, in which the axoneme and other flagellar components are running parallel with the nucleus. As a general conclusion, this study has revealed an inconsistency between the sperm structure and the systematics of the group, indicating that the generic concepts within the group do not reflect a natural classification, a statement also shared by molecular studies.     

Ultrastructure des glandes dermiques à petits pores des Diploures Campodéidés (Insecta,Entognatha, Diplura)

Summary The integument ofCampodea- species, both endogenous and subterranean, shows dermal glands in males and females. Each of them consists of 3 cells: one secretory, one intermediary and one reservoir cell. The secretory product passes through a sieve located at the lower part of the reservoir.     

Cross-Epithelial Hydrogen Transfer from the Midgut Compartment Drives Methanogenesis in the Hindgut of Cockroaches

In the intestinal tracts of animals, methanogenesis from CO₂ and other C₁ compounds strictly depends on the supply of electron donors by fermenting bacteria, but sources and sinks of reducing equivalents may be spatially separated. Microsensor measurements in the intestinal tract of the omnivorous cockroach Blaberus sp. showed that molecular hydrogen strongly accumulated in the midgut (H₂ partial pressures of 3 to 26 kPa), whereas it was not detectable (<0.1 kPa) in the posterior hindgut. Moreover, living cockroaches emitted large quantities of CH₄ [105 ± 49 nmol (g of cockroach)⁻¹ h⁻¹] but only traces of H₂. In vitro incubation of isolated gut compartments, however, revealed that the midguts produced considerable amounts of H₂, whereas hindguts emitted only CH₄ [106 ± 58 and 71 ± 50 nmol (g of cockroach)⁻¹ h⁻¹, respectively]. When ligated midgut and hindgut segments were incubated in the same vials, methane emission increased by 28% over that of isolated hindguts, whereas only traces of H₂ accumulated in the headspace. Radial hydrogen profiles obtained under air enriched with H₂ (20 kPa) identified the hindgut as an efficient sink for externally supplied H₂. A cross-epithelial transfer of hydrogen from the midgut to the hindgut compartment was clearly evidenced by the steep H₂ concentration gradients which developed when ligated fragments of midgut and hindgut were placed on top of each other—a configuration that simulates the situation in vivo. These findings emphasize that it is essential to analyze the compartmentalization of the gut and the spatial organization of its microbiota in order to understand the functional interactions among different microbial populations during digestion.     

First Alaskan records and a significant northern range extension for two species of Diplura (Diplura,Campodeidae)

Species in the class Diplura are recorded from Alaska for the first time. Two species, Tricampa rileyi Silvestri from Dall and Prince of Wales Islands in the Alexander Archipelago of Southeast Alaska and Metriocampa allocerca Conde & Geeraert from near Quartz Lake, southeast of Fairbanks, both in the family Campodeidae, are documented based on recently collected specimens deposited in the University of Alaska Museum Insect Collection. A brief review of the history of the documentation of the Alaskan soil microarthropod fauna is provided, as well as discussion of possible glacial refugia.     

Structure and function of the arolium of Mantophasmatodea (Insecta)

All species of the insect order Mantophasmatodea characteristically keep the 5th tarsomere and pretarsus (arolium plus two claws) turned upwards and off the substrate. The unusually large arolium was studied in two species of Mantophasmatodea using bright field light microscopy, reflection microscopy, fluorescence microscopy, TEM, SEM, and Cryo‐SEM. It contains an epithelial gland, numerous tracheoles, and nerves. The gland consists of enlarged epithelial cells with large nuclei, mitochondria, RER, golgi complexes, microtubules, and numerous secretion vesicles. Evidence for exocytosis of the vesicles into the gland reservoir between the epithelial gland and the thick cuticle could be observed. Cryo‐SEM revealed that the ventral side of the arolium and distal part of its dorsal side are covered with a liquid film. Fluid footprints of arolia of individuals walking on a glass plate also indicate the presence of secretory fluid on the arolium surface. Behavioral experiments using animals with ablated arolia showed that representatives of Mantophasmatodea do not need their arolia to detect and respond to vibratory communication signals nor to catch small to medium‐sized prey. Individuals with ablated arolia were not able to move upside down on a smooth glass plate. We conclude that Mantophasmatodea use their arolia for attachment when additional adhesion force is required (e.g. windy conditions, handling large prey, mating). They can bring their arolia in contact with the surface in a very fast reflex (18.0 ± 9.9 ms). The secretory fluid found on the surface is produced by the gland and transported to the outside, presumably through small pore channels, to enhance adhesion to the substrate. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

Fine structure of the Nassonow's gland in the neotenic endoparasitic of female Xenos vesparum (Rossi) (Strepsiptera, Insecta)

Nassonow's gland consists of a number of cells with ducts that open on to the ventral surface of the brood canal in the cephalothoracic region of a neotenic female strepsipteran. The structural organization of the gland is reminiscent of the class 3 of the epidermal gland cells as defined by Noirot and Quennedey [Ann. Rev. Entomol. 19 (1974) 61], which consists of secretory and duct forming cells. The ultrastructure of the Nassonow's gland is described in female Xenos vesparum (Rossi) parasitic in the social wasp Polistes dominulus Christ. The large secretory cells are clustered in groups of three to four, rich in smooth endoplasmic reticulum and produce a secretion made up of lipids. In young females, just before mating, the ultrastructure of the cells and their inclusions indicate that they are active. In old-mated females the Nassonow's gland degenerates. Microvilli line an extracellular cavity and there are pores present in the irregularly thick cuticle of the efferent duct. The small duct forming cells, intermingle with epidermal cells, overlap secretory cells and produce a long efferent duct, the cuticle of which becomes thick close to its opening in the brood canal. Nassonow's gland could be the source of a sex pheromone, which might be capable of attracting the free-living male to a permanently endoparasitic female.     

Oogenesis in Campodea sp. (Diplura)

Summary The egg chamber of Campodea consists of a group of nurse cells and an oocyte, and is surrounded by a simple, markedly flattened follicular epithelium. Three types of yolk occur in the oocytes: type I appears within elements of the rough endoplasmic reticulum; type II is produced by specific complexes of endoplasmic reticulum and dictyosomes; type III is incorporated by micropinocytosis. Histochemical tests show that mature yolk spheres contain proteins and polysaccharides. The main function of the nurse cells is to synthesize RNA, but they also produce small amounts of type I yolk. Phylogenetic conclusions are drawn from this and other studies of oogenesis in apterygote insects.The author is grateful to Dr. F. Kaczmarski of the Medical Academy, Kraków, for the use of EM facilities in his laboratory     

Fine structure of the pyloric region and Malpighian papillae of protura (Insecta Apterygota)

The pyloric region of Eosentomon and Acerentomon (Insecta, Protura) is described. In both species the posterior cells of the midgut carry short microvilli. Beneath the epithelial cells there is a muscular pyloric sphincter for closing the intestinal lumen. Behind the sphincter is a wide pyloric chamber lined by cells with very long microvilli which point anteriorly toward the midgut. These cells regulate the passage of the intestinal contents into the hindgut. Secretions from the Malpighian papillae are emitted into the gut at this level. In Eosentomon three regions (R₁, R₂ and R₃) are visible in the Malpighian papillae, whereas in Acerentomon region R₁ is lacking. The R₁ region contains secretory cells with elaborate glycoprotein-containing granules. The R₂ region is composed of cells somewhat resembling the secretory cells of Malpighian tubules of insects. Presumably R₁ and R₂ cells emit secretions into the central cavity of each papilla. Cells of R₃ form a duct for the secretion. It is suggested that the R₂ region represents a basic excretory region, common to Protura, whereas the R₁ region, in Eosentomon, may be a specialized area performing supplementary excretory functions.     

The compound eye of Parnara guttata (Insecta,Lepidoptera, Hesperiidae): Fine structure of the ommatidium

Summary The fine structure of an ommatidium of a skipper butterfly, Parnara guttata, has been studied using the electron microscope. Each ommatidium has nine retinula cells, which were classified into three groups: two distal, six medial and one basal retinula cells. The rhabdomeres of the distal retinula cells are localized in the distal part of the rhabdom, while those of the six medial retinula cells appear throughout most of the rhabdom. The rhabdomere of the basal retinula cell occupies only the basal part of the rhabdom. The rhabdomeres of four medial cells are constructed of parallel microvilli, while fan-like microvilli form the rhabdomeres of other two medial retinula cells. The distal and basal retinula cells have rhabdomeres consisting of both parallel and fan-like microvilli. This is the first time the construction of the rhabdomeres of the distal and basal retinula cells has been described in such fine detail for a skipper butterfly. Nine retinula cell axons of each ommatidium extend to the first neuropile of the optic lobe, the lamina ganglionaris. No difference was found in the number of retinula cells of an ommatidium or the shape of the rhabdom between the dorsal and ventral regions of the compound eye.     

Fine structure of a mandibular stretch receptor in Thrips physapus L. (Thysanoptera,Insecta)

Summary Only one non-ciliary proprioceptor is developed on the mandible of Thrips. It consists of two bipolar, multiterminal sense cells, the dendrites of which form a strand extended between the tentorium and the back mandibular margin. The dendritic terminals are embedded in an electron-dense, homogeneous matrix, which obviously represents the stimulus transmitting structure. The strand is stretched, if the mandible is moved forwards and upwards. In Thrips, the system of proprioceptors monitoring the mandibular movement is reduced extensively compared with other pterygote insects. This is linked up with the far-reaching reduction of the mobility of the mandible itself.This study is dedicated to Mrs. Signe Ulmer, in memory of her winning nature, her invaluable assistance, and her disciplineSupported by a grant from the Deutsche Forschungsgemeinschaft (Ho 826/2-2)     

ACTIVE TRANSPORT BY THE CECROPIA MIDGUT : II. Fine Structure of the Midgut Epithelium

A morphological basis for transcellular potassium transport in the midgut of the mature fifth instar larvae of Hyalophora cecropia has been established through studies with the light and electron microscopes. The single-layered epithelium consists of two distinct cell types, the columnar cell and the goblet cell. No regenerative cells are present. Both columnar and goblet cells rest on a well developed basement lamina. The basal portion of the columnar cell is incompletely divided into compartments by deep infoldings of the plasma membrane, whereas the apical end consists of numerous cytoplasmic projections, each of which is covered with a fine fuzzy or filamentous material. The cytoplasm of this cell contains large amounts of rough endoplasmic reticulum, microtubules, and mitochondria. In the basal region of the cell the mitochondria are oriented parallel to the long axes of the folded plasma-lemma, but in the intermediate and apical portions they are randomly scattered within the cytoplasmic matrix. Compared to the columnar cell, the goblet cell has relatively little endoplasmic reticulum. On the other hand, the plications of the plasma membrane of the goblet cell greatly exceed those of the columnar cell. One can distinguish at least four characteristic types of folding: (a) basal podocytelike extensions, (b) lateral evaginations, (c) apical microvilli, and (d) specialized cytoplasmic projections which line the goblet chamber. Apically, the projections are large and branch to form villus-like units, whereas in the major portion of the cavity each projection appears to contain an elongate mitochondrion. Junctional complexes of similar kind and position appear between neighboring columnar cells and between adjacent columnar and goblet cells as follows: a zonula adherens is found near the luminal surface and is followed by one or more zonulae occludentes. The morphological data obtained in this study and the physiological information on ion transport through the midgut epithelium have encouraged us to suggest that the goblet cell may be the principal unit of active potassium transport from the hemolymph to the lumen of the midgut. We have postulated that ion accumulation by mitochondria in close association with plicated plasma membranes may play a role in the active movement of potassium across the midgut.     

Structure of the coxa and homeosis of legs in Nematocera (Insecta: Diptera)

Construction of the middle and hind coxae was investigated in 95 species of 30 nematoceran families. As a rule, the middle coxa contains a separate coxite, the mediocoxite, articulated to the sternal process. In most families, this coxite is movably articulated to the eucoxite and to the distocoxite area; the coxa is radially split twice. Some groups are characterized by a single split. The coxa in flies is restricted in its rotation owing to a partial junction either between the meron and the pleurite or between the eucoxite and the meropleurite. Hence the coxa is fastened to the thorax not only by two pivots (to the pleural ridge and the sternal process), but at the junction named above. Rotation is impossible without deformations; the role of hinges between coxites is to absorb deformations. This adaptive principle is confirmed by physical modelling. Middle coxae of limoniid tribes Eriopterini and Molophilini are compact, constructed by the template of hind coxae. On the contrary, hind coxae in all families of Mycetophiloidea and in Psychodidae s.l. are constructed like middle ones, with the separate mediocoxite, centrally suspended at the sternal process. These cases are considered as homeotic mutations, substituting one structure with a no less efficient one.